NEW YORK
BOTANICAL

- FLORA
MALESIANA

mn band

APR 27 1961

THY MELAEACEAE (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-Mal. spp.),
homomorphic, rarely heteromorphic, regular, tubular, campanulate or in-
fundibuliform, tube very short in Gonystyloideae, or with almost free sepals in
extra-Mal. spp., mostly caducous, some circumsciss in the lower part, or persistent
(sometimes enveloping the ripe fruit in extra-Mal. spp.), sometimes slit lengthwise
in fruit, 4-5(—6)-lobed, the lobes imbricate (rarely valvate in some extra-Mal. 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-~>, 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-Mal. spp.), dehiscing length-
wise, usually free, sometimes the lower 14-14 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-Mal. 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-Mal. 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. of Aquilarioideae; albuminous or exalbuminous. Embryo
straight; cotyledons plano-convex; radicle short, superior.

(1)

2 FLORA MALESIANA [ser. I, vol. 61

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.

Aquilaria, 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 of 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 Aguilaria 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 (fide RENDLE, Classif. Fl. Pl. 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. 10e). 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, /.c. 92-93). Though the structure is most
peculiar, its effect (for longer distances) must not be overrated as winds are scarce in the tropical rain-
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 of 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. Docrers 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.

W ood-anatomy. DEN BERGER, Determinatietabel houtsoorten van Malesié, Veenman, Wageningen
(1949) 20 (Aquilaria); DEscH, Mal. For. Rec. 15? (1954) 607; LEANDRI, Ann. Sc. Nat. Bot. X, 12 (1930)
125 (hand lens); METCALFE & CHALK, Anat. Dic. 2 (1950) 1169 & 1178; MoLL & JANssonius, Mikr.
Holzes 5 (1934) 413.—By JANssonius /.c. Gonystylus is referred to the Thymelaeaceae, mainly because of
the characteristics shown by the pit pairs; METCALFE & CHALK /.c., 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 (/.c. 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 (/.c. 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 different ways. GiLG (in E. & P. Pfi. Fam.I11,
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.111,
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 of Pimelea
(cf. 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 is found in Wikstroemia, Gnidia, and Daphne; Edgeworthia has n = 18. In Wikstroemia indica
FAGERLIND found also an apomictic triploid 2 n = 27 (Hereditas 26, 1940, 1-50).

Taxonomy. Subdivisions—According to DoMKE (Bibl. Bot. 111, 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—Microsemmatidae, 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 So/msia has the typical
parallel nervation, venation, and leaf texture as in Gonystylus, and the nervation and texture of the leaves
of Microsemma 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 Arry 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—S(-8)-celled. Fruit a capsule. Seeds without
chalazal fold, usually with aril. Endosperm 0. (Gonystylus, Amyxa, and Aétoxylon.)

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 Mal. spp.) at most 10, diplostemonous or haplostemonous;
filaments (in Mal. spp.) partly or entirely adnate to the tube. Disk 0, or ring-shaped. Ovary (in Mal.
spp.) 2-celled. Fruit a capsule. Seeds usually with a conspicuous chalazal fold, and a thin funicle, without
aril. Endosperm 0, or present. (Aguilaria and Gyrinops.)

3. Subfam. Thymelaeoideae. Leaves not pellucid-punctate. Floral tube funnel-shaped or cylindric.

4 FLORA MALESIANA [ser. I, vol se"

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, Wikstroemia, 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 A. cumingiana; the same phenomenon is
observed in Gyrinops where they are free, but in G. moluccana 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 f. 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 of 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 HALLER f. 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 STAPF
(Bot. Mag. 156, 1933, sub t. 9313, 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 Aguilaria 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. STAPF, l.c.). 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 1 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. 111, 1934, 1-3, 16) and
HEInic (Am. J. bot. 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. Dict. 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 bancanus 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.
2. Ligneous, perennial. Inflorescences without involucral bracts, or (in Phaleria and Daphne composita)
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 (1—)2-seeded. (Petaloid
aAppencaressnone., Ol Obscure and rim-lke)) sy) an ee eee en te ee eae aleria
5. Ovary always |-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
Senleeaves: alternate, Disk ring-like or cup-shaped) *.)'3) = 9 2)) 2 ee, = Gs Daphne
3. Stamens the same number as the calyx lobes.
9. Shrubs or trees. Leaves ovate-oblong to lanceolate, 114-24 by (14-)1-3 cm. Ovary densely hairy,
2-celled; style terminal. Fruit a loculicidal ci sears protruding either from the top or from the

split side of the floral tube . . . 7. Gyrinops

9. Dwarf-shrub. Leaves linear, 3-5 by ay x mm. Ovary hairy at ‘the upper half or r only at the top,
l-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. Aétoxylon
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,
De OO ek et) tee AY de kat AEF eo ARTES TUSSLE eye a cee en ONY SOV IOS

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 gat acim te ones. Liana, very rarely erect shrubs or
smiallfirees* =" . 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 24 mm), with 7-9 more or less parallel, longitudinal nerves, sessile, linear.

Subalpmevdwart-shrub, ee) Ss es Pee ery Tee et .) 2) 3 2... “SS Drapetes

2. Annual herbs . . Is PRL AS ce es Gh BPI tat i Tt OO Ee
1. Leaves pellucid-dotted.

8. Leaves opposite or subopposite. See vol. 4, p. 365. . . . . . . +. +. 411. Aétoxylon

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; Domkg, Bibl.
Bot. 111 (1934) 118, map 2; Quis. J. Arn. Arb. 27 (1946) 402.—Agallochum
RUMPH. ex LAMK, Encycl. 1 (1783) 48, nom. gen. rejic—Ophispermum Lovur. FI.
Coch. 1 (1790) 281.—Gyrinopsis DecNE, Ann. Sc. Nat. Bot. II, 19 (1843) 41;
Quis. J. Arn. Arb. 27 (1946) 404.—Decaisnella O.K. Rev. Gen. Pl. 2 (1891)
584.—Aquilariella VAN TieGH. Ann. Sc. Nat. Bot. VII, 17 (1893) 216; Bull. Soc.
Bot. Fr. 40 (1893) 77.—Aquilaria sect. Agallochum HALLIER f. Med. Rijksherb.
n. 44 (1922) 15.—Aquilaria sect. Gyrinopsis HALLIER f. /.c. 16.—Aquilaria sect.
Amphinoman HALLER f. /.c. 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 slightly 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, umbelliform
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 slightly below them, sometimes behind them,
sessile or filamentous, equal in length or sepalous ones longer than the others;
filaments filiform, sometimes slightly swollen at the upper end; anthers linear-
oblong, dorsifixed, free (but in A. cumingiana the lower 1-4 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-Mal. 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)’. i

Fig. 1. Aquilaria beccariana VAN TiEGH. a. Habit, « 24, 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. f. d. Opened flower, x 3.—A. hirta RIDL. e. Opened flower, x 3, f. dehisced fruit, nat.

size.—A. microcarpa BAILL. g. Dehisced fruit, nat. size—A. malaccensis LAMK. h. Dehisced fruit, nat.

size.—A. cumingiana (DECNE) RIDL. i. 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, f 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
laterally adnate to the septum, with an obscure or a distinct funicle; in the latter
case the seeds dangle out of the fruit on the end of the thin funicle in open capsules;

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.%65

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 214 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, 34-114 by 1-114 cm. Seed with a caruncle-like, glabrous appendage c.

2 mm long. Episepalous stamens usually shorter or as long as the petaloid appendages.

2. A. microcarpa

2. Calyx lobes always erect. Ovary slightly Nea a style distinct, filiform and almost as long as
the ovary . . 3. A. brachyantha

1. Flowers short- tubular to cylindric, (Ge Se 15 mm long, the lobes usually 1/5— 4), the length of the tube.

Stamens sessile or subsessile, filaments rarely up to !4 as long as the anthers, in A. urdanetensis the

episepalous ones as long as the anthers.

4. Calyx lobes c. 14 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, 4-4 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 3-514 cm, the nerves scarcely distinguishable from the
intermediate veins . . ~» Te Apanlanra

8. Fruits globose, contracted at the base into a short, slender stipe. Floral tube usually flat in fruit
and lobes erect. Leaves 414-15 by 1-414 cm, nerves 7-12 pairs, distinct from the intermediate
VeInS=., = '. : . . 8. A. parvifolia

4. Calyx lobes 1/,—/; the length ‘of the tube, c ais 2 Io in a beccariana. Seeds with an elongated or tail-
like appendage (except in A. enone

9. Fruits oblanceolate, 2-314 by 1-134 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, 14-14 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 mm long
by a short, thin stipe-like constriction. Petaloid appendages ae almost as long as the
stamens, shortly puberulous . . . . 11. A. beccariana

9. Fruits subglobose, globose, slightly obovoid o or ellipsoid, 134, by 1, cm, contracted at the base,
sessile or with an obscure stipe. Seeds broadly ovoid, planoconvex, glabrous, c. 1 by 34 cm, witha
small caruncle-like appendage. Petaloid appendages short, usually united in a ring. Lower 14-14 of
the anthers usually adnate to the floral tube . . - . . . . +. +. 12. A. cumingiana

Dec. 1960]

THYMELAEACEAE (Ding Hou) nie a 9

1. Aquilaria malaccensis LAMK, Encycl. | (1783)
49, t. 356; DC. Prod. 2 (1825) 59; MEIsN. in DC.
Prod. 14 (1857) 602, excl. citat. of BENTH.; Mia.
Fl. Ind. Bat. 1, 1 (1858) 883; Kurz, Nat. Tijd.
N.I. 27 (1864) 171; For. Fl. Burm. 2 (1877) 236;
RIDL. Trans. Linn. Soc. Lond. II, 3 Bot. (1893)
341; GiLtGc, Bot. Jahrb. 18 (1894) 506, f. 8, B;
ibid. 28 (1900) 145; Borer. Handl. 3 (1900) 112;
Ripe str. Br. Re Ass Soc: 7: 35 (1901)) 73;
GAMBLE, J. As. Soc. Beng. 75, ii (1912) 264;
Koorp. Exk. FI. Java 2 (1912) 656 (erron. record);
Merk. Philip. J. Sc. 10 (1915) Bot. 44; Int. Rumph.
(1917) 381; BRown, Minor Prod. Philip. Forests |
(1920) 403; Merr. En. Born. (1921) 417; HALL. f.
Med. Rijksherb. n. 44 (1922) 16; Merr. En.
Philip. 3 (1923) 130; Rip. Fl. Mal. Pen. 3 (1924)
147; Burk. & HENDeERS. Gard. Bull. S.S. 3 (1925)
417; HeYneE, Nutt. Pl. ed. 2 (1927) 1149; HENDERs.
Gard. Bull. S.S. 4 (1928) 314; METCALFE, Kew
Bull. (1933) 5; CorNER, Ways. Trees (1940)
632; Quis. J. Arn. Arb. 27 (1946) 403; Merr. J.
Arn. Arb. 31 (1950) 270.—Agallochum secundarium
coinamense & A. malaicense RUMPH. Herb. Amb.
2 (1741) 34-35, t. 10.—A. ovata Cav. Diss. (1789)
377, t. 224.—A. secundaria DC. Prod. 2 (1825) 59;
Mia. FI. Ind. Bat. 1, 1 (1858) 883; BorerL. Handl.
3 (1900) 112.—Agallochum malaccense O.K. Rev.
Gen. Pl. 2 (1891) 583.—Aquilariella malaccensis
VAN TIEGH. Ann. Sc. Nat. Bot. VII, 17 (1893) 216;
Bull. Soc. Bot. Fr. 40 (1893) 77.—Fig. th.
Tree up to 40 m by 60 cm. Bark smooth,
whitish; branchlets slender, pale brown, pubes-
cent, glabrescent. Leaves chartaceous, sub-
coriaceous, glabrous, sometimes pubescent be-
neath, glabrescent, shining on both surfaces,
elliptic-oblong to oblong-lanceolate, 7144-12 by
24%4-5'44 cm; base acute, attenuate, or obtuse;
apex acuminate, acumen up to 2 cm; nerves 12-16
pairs, rather irregular, often branched, curving
upward, elevated beneath, plane or obscure above,
veins distinct beneath, invisible above; petiole
4-6 mm. /nflorescences terminal, axillary or supra-
axillary, sometimes on internodes, usually branch-
ed with 2 or 3 umbels and each with about 10
flowers, more rarely a simple umbel; peduncle or
common peduncle 5-15 mm; pedicels slender,
3-6 mm. Flowers green or dirty-yellow, cam-
panulate, 5-6 mm long, scattered puberulous
outside. Floral tube nearly glabrous within,
distinctly 10-ribbed. Calyx lobes ovate-oblong,
2-3 mm long, densely puberulous within, almost
as long as the tube, reflexed. Petaloid appendages
oblong or slightly ovate-oblong, c. 1 mm long,
slightly incurved, densely pilose. Stamens 11%4-2
mm long, the episepalous ones longer than the
others; anthers linear, obtuse, as long as or shorter
than the filaments. Ovary ovoid, 1-114 mm long,
densely pubescent; style obscure; stigma capitate.
Fruits obovoid or obovoid-oblong, rounded at the
apex, cuneate to the base, 3-4 by 214 cm, usually
compressed, pubescent outside, glabrescent; peri-
carp woody, the suture face c. 6 mm wide. Seed
proper ovoid, including the beak 10 by 6 mm,
densely covered with red hairs, the beak c. 4 mm
long. the appendage twisted and as long as the

seed, separated from it by a short, thin, stipe-like
constriction.

Distr. India (Bengal and Assam), Burma
(Tenasserim), and Malaysia: Sumatra (Simalur,
Sibolangit, Palembang, and Banka), Malay
Peninsula (common), N. & E.-Borneo, and the
Philippines (Luzon).

Ecol. Primary forests at low and medium
altitudes up to 270 m.

Uses. According to HEyYNE, /.c. and RIDLEY
(1901, /.c.) the tree yields a celebrated incense
wood which is obtained from the center of an old
or dying tree. It is said to be caused by a disease
which gains entry through the old decaying
branches. Its greatest use has always been for
fumigating and it is highly valued in the Orient
for ceremonial purposes. It also furnishes a
beautiful, silvery bast used for making ropes and
clothes. The bast is highly prized for its strength
and durability.

Vern. Calambac, ching karas, gaharu, galoop,
garu, karas, kayu gaharu, kékaras, képang, laroo,
méngkaras, tabak, taras gharu, téngkaras, M,
sigsigi, Borneo; Sumatra: alim, Batak, halim,
Lamp., karéh, Minangk.; Malayan eaglewood tree,
E.

Note. The vegetative parts of this species are
similar to those of A. microcarpa in the herbarium,
and I cannot find any good character to separate
them.

2. Aquilaria microcarpa BAILL. Adansonia 11
(1875) 304; Gmc, Bot. Jahrb. 28 (1900) 145;
Merr. En. Born. (1921) 417; Domxe, Bibl. Bot.
111 (1934) t. 4 f. 43f; Quis. J. Arn. Arb. 27 (1946)
403.—Aquilariella microcarpa VAN TiEGH. Ann.
Sc. Nat. Bot. VII, 17 (1893) 216; Bull. Soc. Bot.
Fr. 40 (1893) 77.—Aquilariella borneensis VAN
TreGH. Ann. Sc. Nat. Bot. VII, 17 (1893) 217;
Bull. Soc. Bot. Fr. 40 (1893) 77.—A. borneensis
Gite in E. & P. Pfl. Fam. III, 6a (1894) 224;
BoerL. Handl. 3 (1900) 112; Merr. En. Born.
(1921) 417.—Fig. 1g.

Tree up to 40 m by 80 cm. Bark grey, super-
ficially fissured; branchlets light brown, puberu-
lous, glabrescent. Leaves subcoriaceous, shining
and glabrous above, rather dull, glabrous or
sometimes scattered hairy beneath, elliptic-oblong
to obovate-oblong or oblanceolate, 414-10 by
1144-414 cm; base cuneate to attenuate; apex acute
to acuminate, the acumen up to 1 cm; nerves
12-19 pairs, sometimes branched, slightly curved
and ascending to the thickened margin, pro-
minent beneath, visible above; veins and veinlets
rather irregular, subparallel, distinct beneath,
obscure above; petiole 3—5 mm, pubescent.
Inflorescences axillary or supra-axillary, terminal,
or on short lateral branchlets, usually branched,
rarely simple, peduncle short or up to 1 cm,
6—-11-flowered. Flowers white, light-yellow or
yellow, 5 mm long; pedicels c. 5 mm, puberulous.
Floral tube puberulous outside, sparsely puberu-
lous inside. Calyx lobes ovate or oblong, obtuse,
densely puberulous on both surfaces. Petaloid
appendages almost as long as the stamens or

10 FLORA MALESIANA

[ser. I, vol) 16

sometimes slightly longer, ovate or oblong, densely
hairy. Stamens 1-114 mm, alternately long and
short; anther c. 14 mm, usually shorter than the
filament, rarely as long or longer. Pistil ovoid,
114-2 mm long. Ovary densely pubescent; style
obscure or rarely very short; stigma capitate.
Fruits subcordate, slightly compressed, 8—12(—16)
by 10—-12(-15) mm, 1-(2-)seeded; persistent floral
tube sometimes splitting on one side. Seed ovoid,
6 by 4 mm, densely brownish pubescent; caruncle-
like appendage 2 mm long.

Distr. Malaysia: Malay Peninsula (Singapore),

Sumatra (Sidjungdjung, Palembang, and
Lampongs), Billiton, Banka, and throughout
Borneo.

Ecol. Lowland forests up to 200 m.

Vern. Sumatra: réngkaras, M, hepang, Banka;
Borneo: éngkaras, Dayak, karas or sigi-sigi, Bugis,
kumbil, garu, tulang, M.

3. Aquilaria brachyantha (MerRR.) HALL. f. Med.
Rijksherb. n. 44 (1922) 16; Quis. J. Arn. Arb. 27
(1946) 403.—Gyrinopsis brachyantha MERR. Philip.
J. Sc. 7 (1912) Bot. 313; E_m. Leafi. Philip. Bot. 5
(1913) 1629; Merr. En. Philip. 3 (1923) 130;
DomkgE, Bibl. Bot. 111 (1934) t. 2, f. 8; t. 5, f. 43e.
—Fig. ld.

Small tree or shrub up to 2 m. Branchlets
glabrous, yellowish brown to brownish when dry,
the tips usually pubescent. Leaves chartaceous to
subcoriaceous, shining on both surfaces, oblong,
elliptic-oblong, or lanceolate, 8-16 by 2-414 cm;
base acute or obtuse; apex acuminate; nerves and
veins numerous, homogeneous, slightly elevated
beneath, obscure or invisible above; petiole c.
5 mm. Flowers greenish, small, axillary, 1 to
several in a fascicle on a very short peduncle;
pedicels 1-3 mm, pubescent. Floral tube cam-
panulate, 3-4 mm long, pubescent or puberulous
on both surfaces, usually glabrescent outside.
Calyx lobes 5, slightly oblong or ovate-oblong, as
long as the tube or sometimes slightly longer.
Petaloid appendages linear or oblong, c. | mm long,
densely pubescent. Stamens 1-14, mm long,
filamentous, the episepalous ones slightly longer
than the others. Ovary ovoid, c. 144 mm long,
slightly pubescent; style distinct, filiform, c. 1 mm;
stigma capitate. Fruits narrowly obovoid, com-
pressed, 114-114 by 34-1 cm. Seed including the
caruncle-like appendage c. 1 cm long, pubescent,
persistent floral tube splitting on one side.

Distr. Malaysia: Philippines (Luzon: Cagayan
Prov.), twice collected.

Ecol. In primary forests at low altitudes.

4. Aquilaria urdanetensis (ELMER) HALL. f. Med.
Rijksherb. 1. 44 (1922) 16.—Gyrinopsis urdanetense
Emer, Leafi. Philip. Bot. 5 (1913) 1630; MerrR.
En. Philip. 3 (1923) 131; Quis. J. Arn. Arb. 27
(1946) 405.

Slender shrub, up to 7 m. Bark dull grey and
smooth. Young branchlets whitish pubescent,
glabrescent. Leaves chartaceous, shining on both
surfaces, young leaves pubescent beneath es-
pecially on the nerves and veins, glabrescent,

elliptic-oblong to broadly lanceolate, 4-9 by
114-3 ¥, cm; base cuneate to attenuate; margins
slightly thickened and shining on both surfaces;
apex acuminate, the acumen up to | cm, pointed
or obtuse at the tip; nerves and veins undistin-
guishable, numerous, divaricate, subparallel, some
of them branched, distinct beneath, obsolete
above; petiole 3-4 mm. Inflorescences axillary,
sessile or very short peduncled, usually with a few,
very small, caducous bracts, 2- or 3-flowered;
pedicels c. 314 mm, sparsely pubescent. Flowers
short-tubular, 5-6 mm long, yellowish. Floral
tube 314-4 mm long, sparsely pubescent outside,
pubescent towards the base and at the mouth
inside. Calyx lobes 5(-6), ovate and obtuse,
sparsely pubescent outside, densely puberulous
inside, 114-2 mm long. Petaloid appendages
obscure, rim-like. Stamens free from the tube
slightly below the petaloid appendages, 24-114
mm, episepalous ones with a filament shorter than
the anther or as long as it, the others sessile.
Pistil c. 2 mm long. Ovary ellipsoid, densely
puberulous, narrowed into a short style c. 1 mm;
stigma nipple-like. Fruits globose or slightly
obovate, glabrous when mature, c. 8 mm diam.,
with a slender stipe 3—6 mm;; persistent floral tube
splitting on one side. Seed ovoid, plano-convex,
black, c. 1 cm long, with a short caruncle-like
appendage.

Distr. Malaysia: Philippines (Mindanao: Mt
Urdaneta), twice collected.

Ecol. In the mossy forest on exposed ridges,
c. 1700 m (cf. MerRR. l/.c.).

Vern. Majfgod, makolan, Mbo.

Note. Known only from the authentic collec-
tions, ELMER 14195 (lectotype) and 13742 (para-

type).

5. Aquilaria citrinaecarpa (ELMER) HALL. f. Med.
Rijksherb. n. 44 (1922) 18.—Gyrinopsis citrinae-
carpa ELMER, Leafl. Philip. Bot. 5 (1913) 1631;
Merk. En. Philip. 3 (1923) 130; Quis. J. Arn. Arb.
27 (1946) 405.

Small tree up to 8 m. Young branches densely
olivaceous tomentose, glabrescent. Leaves sub-
coriaceous, dull, olivaceous, and densely pubes-
cent beneath, shining, reddish-brown, and glabrous
above, elliptic-oblong, or slightly obovate-oblong,
6-10(—12) by 214-4(-514) cm; base cuneate; apex
acute; margins slightly recurved; nerves 15-20
pairs, on the lower surface obscure, rarely distinct,
slightly ascending towards the margin; veins and
veinlets obscure or visible beneath, obscure above;
petiole c. 3 mm. Inflorescences terminal and
axillary, sessile or with a very short peduncle,
densely hairy, usually with a few small bracts,
3-6-flowered; pedicels 2-4 mm, _ pubescent.
Flowers 8-10 mm long, greenish. Floral tube
cylindric, densely pubescent outside and towards
the base inside. Calyx lobes oblong or ovate,
3-314 mm long, densely puberulous on both sur-
faces. Petaloid appendages oblong or ovate, about
14 the length of the anthers, densely villose.
Stamens sessile, 114 mm long. Ovary slightly
obovate-oblong, densely puberulous, c. 3 mm

Dec. 1960]

THYMELAEACEAE (Ding Hou) is. eet

long, style none; stigma pyramidal, black.
Capsules 174 by 1% cm, bright yellow or citrine,
obtuse to subtruncate at the apex, attenuate to the
base, sometimes slightly compressed, persistent
floral tube splitting on one side. Seeds deltoid, 8
by 7 mm, plano-convex, acute to the apex, almost
truncate at the base, with a very short caruncle-
like appendage at the base.

Distr. Malaysia: Philippines (Mindanao), once
collected.

Ecol. On moist, compact soil of forested ridges,
c. 1300 m.

Vern. Agododan, Mbo.

6. Aquilaria apiculata Merr. Philip. J. Sc. 20
(1922) 411; En. Philip. 3 (1923) 130; Quis. J. Arn.
Arb. 27 (1946) 403.

Shrub or small tree up to 3 m. Branchlets
reddish brown, pubescent, glabrescent. Leaves
papery, glabrous above, sparsely or scattered
pubescent beneath, elliptic-lanceolate, 8-14 by
214-4 cm; base cuneate to attenuate; apex acu-
minate, the acumen up to 2 cm, tip obtuse; nerves
8-16 pairs, curved and ascending to the margin,
prominent beneath, visible above; veins distinct,
sometimes visible beneath, obscure above; petiole
3-5 mm. Inflorescences axillary, sessile or short-
peduncled, usually with a few small bracts,
3-6-flowered; pedicels 6-7 mm, _ puberulous.
Flowers 5—6 mm long. Floral tube short-tubular,
4-5 mm long, puberulous outside and inside,
glabrescent. Calyx lobes ovate to oblong, 114-2
mm long, densely puberulous on both surfaces.
Petaloid appendages semi-orbicular to ovate,
14-4 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 14-1 cm long stipe, protruding
from the floral tube, 124 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.

Distr. Malaysia: Philippines (Mindanao:
Bukidnon Prov.).

Ecol. In dry and mossy forests, 1100-1800 m.

7. Aquilaria filaria (OKEN) Merr. J. Arn. Arb. 31
(1950) 283, excl. syn. Gyrinopsis brachyantha
MeERR.—Cortex filarius RumMpH. Herb. Amb. 7
(1755) 13.—Pittosporum ferrugineum var. B
filarium DC. Prod. 1 (1824) 347, excl. RUMPH. t. 7
cit.; DON, Gen. Hist. 1 (1831) 374.—Pittosporum
filarium OKEN, Allg. Naturgesch. 32 (1841) 299.—
A. tomentosa GILG, Bot. Jahrb. 28 (1900) 145.—
Gyrinopsis brachyantha (non MERR. 1912) MERR.
Int. Rumph. (1917) 380, quoad specim.—Gyrinopsis
acuminata MERR. Philip. J. Sc. 17 (1920) 294;
En. Philip. 3 (1923) 130.—A. acuminata Quis. J.
Arn. Arb. 27 (1946) 403.

Shrub or tree up to 17 m by 50 cm. Young
branchlets light-brown, pubescent and glabres-

cent. Leaves subcoriaceous, glabrous or scattered
hairy rarely pubescent beneath, oblong, elliptic-
oblong to lanceolate, rarely oblanceolate-oblong,
10-20 by 3-514 cm; base obtuse to cuneate; apex
shortly acuminate; nerves and veins usually homo-
geneous, slightly elevated beneath, obscure above;
petiole 3-5 mm. Inflorescences axillary and extra-
axillary, umbelliform or condensed paniculiform,
rarely cauliflorous, very short-peduncled, (1—)3-7
(—cv)-flowered; pedicels 2-5 mm, _ pubescent.
Flowers yellowish-green or white, infundibular,
5-614 mm long. Floral tube sparsely pubescent
outside, glabrescent. Calyx lobes oblong to slightly
ovate, c. 2 mm long, densely puberulous on the
upper part and the margins outside, and the whole
surface inside. Petaloid appendages oblong or
deltoid, c. 1 mm long, densely villous. Stamens c.
1 mm long, the episepalous ones with short, fleshy
filaments, the others sessile or subsessile. Ovary
obovoid, c. 34% mm long, densely villous; style
very short or obscure; stigma capitate. Fruits
ellipsoid to obovoid or subglobose, slightly com-
pressed, rugose, 1 44-1 % by 1 4 cm, sparsely hairy,
glabrescent, narrowed to the base, sometimes on a
very short stipe, yellow. Seeds deltoid, including
the appendage c. 44 by 34 cm, plano-convex, black,
with a very short caruncle-like appendage.

Fig. 2. Localities of Aquilaria filaria (OKEN)
MERR.

Distr. Malaysia: Philippines (Dinagat I. and
Bucas Grande I.), Moluccas (Morotai, Ceram, and
Ambon), and New Guinea (Sorong, Babo, and
Kapor). Fig. 2.

Ecol. In lowland forests, once collected in open
swamp forest (Sorong: PLEYTE 393), up to 130 m.

Vern. Agé, Sorong, bdékuin, Morotai, Jason,
Ceram, kasjik, Tehid lang., malowassi, Uliassers.

Notes. In the description of Cortex filarius
RumpPH. (Herb. Amb. 7, 1755, 13) RUMPHIUs re-
corded the bark with strong bast and the leaves
with more or less parallel veins which agree with
the characters of Aquilaria. He described the fruit
as 2-celled; one of the cells being empty and filled
with pulp, the other having two seeds. However,
in Aquilaria the ovary is 2-celled and each has
only one ovule. Based on the description, the

12 FLORA MALESIANA

[ser. 1, vo

common name ‘Malowassi’ and the usage, HEYNE
(Nutt. Pl. 1927, p. 1151) identified it as a Gyrinopsis
sp. (= Aquilaria).

MERRILL (Int. Rumph. 1917, 380) in inter-
preting RUMPHIUS’ Cortex filarius, with the re-
presenting specimen (ROBINSON’s PI. Rumph. Amb.
n. 274), referred it to Gyrinopsis brachyantha
Merr. (= Aguilaria brachyantha). The leaves of
these two species are very similar to each other.
See also BAKKER (FI. Mal. I, 5, 1957, 359-360).

J. SMITH initiated the error to combine the plate
of Cortex foetidus RumpH. (t. 7) with the de-
scription of Cortex filarius RuUMPH. referring them
both to Pittosporum ferrugineum (in Rees, Cycl.
27 art. Pittosporum, 1814), although he remarked
already that the ‘“‘thready bark”’ ascribed to it by
RuMpPHIus does not occur in Pittosporum. This
error was continued by DE CANDOLLE, /.c.

In reviewing OKEN’s work in 1950, MERRILL
(J. Arn. Arb. 31, 1950, 283) pointed out that the
description of Pittosporum filarium OKEN was
wholly taken from Rumpuius’ Cortex filarius and
concluded that Gyrinopsis brachyantha MEeERR.
should be added to the synonymy of Aquilaria
filaria (OKEN) MERR.

On examining ROBINSON’s specimen indicated
above and another specimen collected by Trys-
MANN (s.n., Bo) from Soja, Ambon, however, it
appears that A. filaria is distinct from the Phi-
lippine A. brachyantha. A. filaria is characterized
by calyx lobes about half the length of the tube,
stamens sessile or very short-filamentous, ovary
densely villous, and the style obscure, while A.
brachyantha is characterized by calyx lobes about
as long as the tube, stamens distinctly filamentous
and the filaments as long as or longer than the
anthers, ovary sparsely hairy, and the style distinct
and as long as the ovary.

8. Aquilaria parvifolia (Quis.) nov. comb.—
Gyrinopsis parvifolia Quis. J. Arn. Arb. 27 (1946)
405.

Shrub c. 1 m tall, branches light brown or
reddish-brown. Leaves subcoriaceous, slightly
pubescent beneath, glabrous above, elliptic-
oblong, ovate-oblong, or lanceolate, 414-15 by
1-414 cm; base acute to cuneate; apex narrowly
acute to acuminate; nerves 7-12 pairs, slightly
curving upward, distinctly elevated beneath, slightly
elevated above; veins distinct beneath, invisible
above; petiole c. 5 mm, sparsely pubescent, gla-
brescent. Infructescences axillary, terminal, some-
times extra-axillary, umbelliform, short-peduncled,
sometimes almost sessile, each with 1 to 4 fruits;
pedicel c. 3 mm, puberulous. Persistent flower
short-tubular, 5—6 mm long. Floral tube sparsely
puberulous on both surfaces. Calyx lobes ovate or
ovate-oblong, 114-2 mm long, densely puberulous
inside and on the margins and tips outside. Petaloid
appendages orbicular or deltoid, c. 14-4 the length
of the stamens, villous, the hairs slightly longer
than the appendage. Stamens sessile, c. 1 mm long.
Fruits slightly obovoid or globose, 1—11/4 by 1-114
cm, yellowish, rugose when dry, constricted at the
base into c. 2 mm long stipe. Seeds broadly ovoid,

8-9 by 614-7 mm, dark-brown, smooth, shining.
with an obscure caruncle-like appendage at the
base.

Distr. Malaysia: Philippines (Luzon: B.S.
26876, 41562, 76441—type (A), WENZEL 1201).

Ecol. On forested slopes, 1000 m.

Note. Very closely related to A. filaria, but
easily separated from the latter by the persistent
floral tube with erect calyx lobes and the smaller
leaves with distinct lateral nerves.

9. Aquilaria hirta RIDL. J. Str. Br.R. As. Soc. n. 35
(1901) 78; GAMBLE, J. As. Soc. Beng. 75, ii (1912)
265; Rip. Fl. Mal. Pen. 3 (1924) 148; DomKE,
Bibl. Bot. 111 (1934) t. 5 f. 43g.—A. moszkowskii
GiLG, Notizbl. Berl._Dahl. 5 (1908) 84; Quis. J.
Arn. Arb. 27 (1946) 403.—Fig. le-f.

Tree up to 14 m with whitish and rather smooth
bark. Young branchlets light brown, covered with
silky hairs, glabrescent. Leaves subcoriaceous,
dull and pubescent beneath especially on the
midrib, nerves and veins, sometimes glabrescent,
shining on the upper surface, elliptic-oblong,
ovate-oblong, 614-14 by 214-514 cm; base
cuneate to obtuse or rounded; apex acuminate,
the acumen up to 114 cm, mucronate, pointed at
the tip; nerves 16-30 pairs, irregular, sometimes
branched, elevated beneath, visible to obsolete
above, slightly curved upward and towards the
margin; veins distinct or visible beneath, obscure
or not visible above; petiole S-7 mm, thickened,
curved, pubescent. Jnflorescences sessile or up to
10 mm peduncled, pubescent, 5—14-flowered;
bracts small. Flowers white (fide Kep. 71521) or
light yellow (fide BUNNEMEIJER 7575), up to 2 cm
pedicelled, pubescent. Floral tube cylindric 6-8
mm long, densely pubescent outside and towards
the base inside, ribbed and sparsely villose within
at the upper part. Calyx lobes ovate and obtuse,
2-3 mm long, densely pubescent outside and
densely puberulous inside. Petaloid appendages
inserted slightly behind the stamens, ovate or
semi-orbicular, densely villous, almost as long as
the stamens, sometimes even slightly longer.
Stamens sessile, c. 1 mm long, oblong, connective
dark-brown. Pistil clavate, 5 mm long; ovary
densely puberulous; style absent; stigma capitate.
Fruits protruding from the floral tube, oblan-
ceolate, abruptly acute at the apex, attenuate to
the base, including the stipe 314-5 by 1 cm, densely
golden puberulous; pericarp coriaceous. Seeds
ovoid, 10 by 6 mm, puberulous, glabrescent,
shortly beaked at the apex, cuneate at the base,
black and shining, with a long glabrous appendage
c. 10 mm long.

Distr. Malaysia: Malay Peninsula (Trengganu,
Pahang, Johore, and Singapore; lectotype:
MuRTON 2, SING; paratypes: RIDLEY 3837, SING
and RipLey 11020, K, SinGc), E. Sumatra (Sena-
maninik), Riouw, and Lingga.

Ecol. Hill slopes, from the lowland up to
300 m.

Vern. Chamdan, changang,
sahare, M; Sumatra: karas.

Note. A. moszkowskii GiLG was described on a

kayu _ chandan,

Dec. 1960]

THYMELAEACEAE (Ding Hou) 13

sterile specimen collected by MoszkowskI (12, B)
at Senamaninik, eastern Sumatra. I have not seen
the type, but the locality and GILG’s detailed des-
cription agree very well with the present species,
especially the silky hairs occurring on the under-
side of the leaf which is peculiar to this species.

10. Aquilaria rostrata RipL. Fl. Mal. Pen. 3 (1924)
148.

Tree. Branchlets pubescent, glabrescent. Leaves
subcoriaceous, glabrous, rather shining on both
surfaces, lanceolate, rarely ovate-oblong, 614-10
by 214-414 cm; base obtuse, cuneate to attenuate;
apex acuminate, the acumen up to 114 cm; nerves
16-many pairs, simple or rarely branched, spread-
ing or slightly curved and ascending, elevated
beneath and visible above; veins visible beneath
and obscure above; petiole 314-7 mm. Pedicels c.
3 mm, brownish hairy. Floral tube cylindric, 6 mm
long, splitting on one side, glabrous outside,
sparsely puberulous inside. Calyx lobes slightly
oblong, c. 114 mm long, puberulous on both sur-
faces. Petaloid appendages unknown. Stamens
sessile. Fruits (young) obovate-oblong or oblan-
ceolate, including the stipe 3 by #4-114 cm,
brownish hairy outside, long-narrowed towards
the base, apex beaked. Seeds slightly ellipsoid-
oblong, 10 by 4 mm (excl. the appendage),
brownish, puberulous, acuminate, base attenuate
and elongate into a slender appendage, glabrous.

Distr. Malaysia: Malay Peninsula (Pahang,
Wray’s Camp, Gunong Tahan, RIDLEY 16264,
type, K, SING).

Note. As mentioned by RIDLEY the specimens
are poor. No material has been collected since the
type. I have seen two sheets of the type number
and one other sterile unnumbered sheet. Only
young fruits are available, with the persistent floral
tube. Unfortunately, the petaloid appendages and
stamens of them were eaten by insects except the
basal parts of two sessile stamens in one flower.
From the available material, it is impossible to
verify the number and shape of the petaloid
appendages and the number of stamens.

This species, as pointed out by RIDLEY, is
characterized by the long-beaked fruits. In ad-
dition, the floral tube is longer than the lobes,
and the stamens are sessile.

11. Aquilaria beccariana VAN TIEGH. Ann. Sc. Nat.
Bot. VII, 17 (1893) 217; Bull. Soc. Bot. Fr. 40
(1893) 77; Gite, Bot. Jahrb. 28 (1900) 145; BoeRL.
Handl. 3 (1900) 112; Becc. Nelle Foreste (1902)
592; MerR. En. Born. (1921) 416.—A. grandifolia
DomKE, Notizbl. Berl.—Dahl. 11 (1932) 348.—
A. cumingiana var. parviflora Airy SHAW, Kew
Bull. (1940) 261.—Gyrinopsis grandifolia Quis. J.
Arn. Arb. 27 (1946) 406.—Fig. 1a-c.

Tree up to 20 m tall and 36 cm diam. with grey
and smooth bark. Young branchlets pubescent.
Leaves papery to subcoriaceous, glabrous on both
surfaces, sometimes scattered pubescent beneath,
oblong, oblong-lanceolate, or elliptic-oblong,
rarely elliptic, (7—-)11-27 by (3-)6-814 cm; base
cuneate to attenuate; apex acute to acuminate;

nerves (10—)15—25 pairs, curving and ascending
towards the margin, elevated and prominent
beneath, distinct above; veins loosely reticulate;
petiole 5-7 mm. Inflorescences axillary or extra-
axillary, branched and up to 1!4 cm peduncled,
short-paniculiform, pubescent; pedicels 3-7 mm,
pubescent. Flowers 7-12 mm long, yellowish,
greenish or yellowish-white. Floral tube cylindric,
10-costate, sparsely hairy outside. Calyx lobes
slightly ovate, puberulous inside, 2-3 mm long,
densely puberulous on both surfaces, sometimes
glabrescent on the outside. Petaloid appendages
oblong, c. 1 mm long, densely short-hairy. Stamens
usually sessile, rarely with very short filaments,
almost as long as the petaloid appendages. Disk
ring-like to cupular, densely puberulous. Pistil c.
5 mm long, with a distinct stipe c. 2 mm long, the
stipe accrescent and elongated. Ovary ellipsoid,
attenuate to the base, gradually narrowed at the
apex; stigma capitate. Fruit protruding from the
top of the floral tube, ellipsoid or obovoid, 2-314
by 134 cm, slightly puberulous and glabrescent,
narrowed to the base into an elongate stipe up to
114 cm, acuminate to the apex, usually slightly
contracted in the middle; floral tube entire, very
rarely splitting on one side (KADIR A 3601). Seeds
black, ovoid, 10 by 5 mm, sparsely puberulous,
acuminate to the apex, with an elongate tail c.
5mm long, attached at the center of the appendage,
the appendage slender, c. 1 cm long, densely
reddish-brown pubescent.

Distr. Malaysia: Sumatra (Palembang), Malay
Peninsula (Johore), and common in Borneo.

Ecol. Primary forests, rarely in swampy forest
(Johore: S.F. 29008, K), from the lowland up to
825 m.

Vern. Meérkaras puti, Sum., gaharu, gumbil,
njabak, M, tanduk = garu, Born.

Notes. This species is characterized by the
cylindric floral tube, the oblong and puberulous
petaloid appendages which are almost as long as the
sessile or subsessile stamens, and the stiped pistil
with a short, puberulous, ring-like disk at its base.

The type specimen of the present species was
collected by BEccARI (PB 2339, Fr) from Sarawak.
It has rather small leaves (814-1314 by 14-4 cm)
and young flowers. The type of A. grandifolia
(GRASHOFF 693, Bo) collected in the swamp forest,
Palembang, S. Sumatra, has larger leaves (17—27
by 6-814 cm) and young flowers. Many specimens
collected in the Malay Peninsula (e.g. S.F. 29008,
29195, 29381, 29470) and Borneo (e.g. bb 34916,
ENDERT 3319, 4035, C.F. 34453, PURSEGLOVE
P 4752, RUTTEN 68, PATRICK PING San A 1726,
and Woop San 15218) have flowers and fruits in
different stages of development and their leaves
show a variable size. From this additional material
we can clearly infer that only one species is
represented.

Aquilaria cumingiana var. parviflora was based
on HAVILAND 3092 (type) and several other
collections from western Borneo. All the specimens
cited in the original description agree with the
present species and are quite different from 4.
cumingiana.

14 A FLORA MALESIANA [ser. I, vol. 6!

a
iia Wp —h
W/ Y yh i

Ey, é OU if Yi jf y, Wns
Vu: py, (MDM ML MU Mp ee

Fig. 3. Phaleria capitata JACK. a. Habit, 24, b. opened flower, x 4/3, c. ovary with disk, x 7, d. stamens,
x 7, e. fruit, nat. size, f. seed, x 2, g. longitudinal section of fruit, one seed removed to show meshes of
endocarp, xX 2.—P. elegans L. M. PERRY, h. Opened flower, x 4/3, i. punctate bract, x 24.—P. macro-
carpa (SCHEFF.) BOERL. j-k. Opened flowers, dimorphous, x £/;, /. longitudinal section of fruit, one seed
removed, x 24.—P. octandra (L.) BAILL. m. Opened flower, x 4/3, 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-43, h Brass 24484, i Brass 24483, 7 ATAsrip 139, kK—-/ C.H.B. VIII-G—93, m-—o
WALSH 36).

Dec. 1960]

THYMELAEACEAE (Ding Hou) 15

12. Aquilaria cumingiana (DECNE) RIDL. J. Str. Br.
R. As. Soc. n. 35 (1901) 80; HALL. f. Med. Rijks-
herb. n. 44 (1922) 17.—Gyrinopsis cumingiana
Decne, Ann. Sc. Nat. Bot. II, 19 (1843) 41, t. 1
f. B, 13-21; MEeEIsn. in DC. Prod. 14 (1857) 603;
Mia. FI. Ind. Bat. 1, 1 (1858) 883; F.-VILL. Nov.
App. (1880) 183; VipaL, Phan. Cuming. (1885)
140; Rev. Pl. Vasc. Filip. (1886) 230; Merr. Bull.
Bur. For. Philip. 1 (1903) 41; ELM. Leafl. Philip.
Bot. 5 (1913) 1629; En. Philip. 3 (1923) 131;
Ho.LtHuis & LAM, Blumea 5 (1942) 216; Quis. J.
Arn. Arb. 27 (1946) 405; Med. PI. Philip. (1951)
636.—Decaisnella cumingiana O.K. Rev. Gen. PI.
2 (1891) 584.—Gyrinopsis cumingiana var. pubes-
cens Eo. Leafli. Philip. Bot. 5 (1913) 1629; MerrR.
En. Philip. 3 (1923) 131.—Gyrinopsis decemcostata
HALL. f. Med. Rijksherb. n. 44 (1922) 17; DomKE,
Bibl. Bot. 111 (1934) t. 2 f. 9; t. 4 f. 36p.—Gyrinop-
sis pubifolia Quis. J. Arn. Arb. 27 (1946) 406.—
Fig. 1i.

Shrub or small tree up to 5 m. Bark ashy grey,
mottled and smooth. Young branchlets densely
pubescent, glabrescent. Leaves chartaceous to
subcoriaceous, glabrous on both surfaces or some-
times pubescent on the lower surface; oblong-
lanceolate, elliptic-oblong, or ovate-oblong, rarely
obovate-oblong, 14-18 by 5!4-814 cm; base
cuneate, rarely rounded; apex acute to acuminate,
acumen up to 114 cm; nerves 12-18 pairs, slightly
curved and ascending to the margin, elevated and
distinct beneath, slightly elevated above; veins and
veinlets numerous, irregularly forked; petiole 4-6
mm. Inflorescences simple or sometimes branched,
few- to many-flowered; peduncle short, c. 5 mm
long, rarely subsessile; pedicels 3 mm, setose.
Flowers whitish, 13-16 mm long. Floral tube
cylindric, puberulous outside, glabrescent, densely
or sparsely pubescent inside, the retrorse hairs
appressed, sometimes distinctly 10-costate inside,
usually with irregular, sulphureous, wart-like
excretions. Calyx lobes ovate or oblong, obtuse,
2-3 mm long, densely puberulous on both surfaces,
sometimes glabrescent. Petaloid appendages short,
usually united in a ring, rarely free or united at
the base, about half as long as the stamens, den-
sely hairy, the hairs longer than or as long as the
appendages. Stamens sessile, 1-2 mm long, free
from the tube, slightly below the appendages or at
the same level with them, the lower 14-14 of the
anther usually adnate to the tube. Pistilc. 7 mmlong,

short-stiped, densely hairy. Ovary obovate, at-
tenuate to the base; style continuous with the
ovary, obscure or distinct; stigma capitate. Fruits
globose, slightly obovoid, or ellipsoid, rugose,
protruding laterally from the split floral tube,
134 by 114 cm. Seeds broad-ovoid, plano-convex,
1 by 34 cm with a short caruncle-like appendage.

Distr. Malaysia: S. Borneo (Sampit region),
Philippines (common), and Moluccas (Morotai
and Halmaheira).

Ecol. In primary forests at low and medium
altitudes.

Uses. According to QUISUMBING, /.c., the bark
and roots are used to stop bleeding from wounds.
The bark, wood, and fruit are used as a substitute
for quinine.

Vern. Alahan, maga-an, palisan, Tag., bago,
Mbo., binukat, Ak. Bis., butlo, Neg., dalakit, S.L.
Bis., magwalem, Sub., pamaluian, Bag.; giba
kolano, Halmaheira.

Note. The type specimen of Gyrinopsis pubifolia
Quis. is B.S. 75314 (A). According to QUISUMBING
(/.c.) it was collected at Mt Abucay, Catanduanes,
at c. 1600 m, September 11, 1928 (the field data
on the label of this specimen are simply indicated
as ““Catanduanes, M. RAMos & G. EDANo, July-
Sept. 1928”); it has rather young flowers with the
lower part of the anthers united with the floral
tube. There is another specimen (B.S. 75516,
SING) which is similar to the above one and bears
the same field data; it has both young and mature
flowers and has been distributed as Gyrinopsis
cumingiana DECNE (= A. cumingiana) with which
I agree. In comparing these two specimens, it
appears that Gyrinopsis pubifolia is conspecific
with A. cumingiana.

Excluded

Aquilaria? bancana Mia. Sum. (1861) 355 is ac-
cording to Airy SHAW (FI. Mal. I, 4, 1953, 361)
= Gonystylus bancanus (MiQ.) Kurz (Thyme-
laeac.).

Aquilaria? macrophylla Mig. Sum. (1861) 356
is according to Airy SHAW (FI. Mal. I, 4, 1953,
354) = Gonystylus macrophylla (MiqQ.) Airy SHAW
(Thymelaeac.).

Aquilaria pentandra BLANCO, FI. Filip. ed. 1
(1837) is according to BAKKER (FI. Mal. I, 5,
1957, 355) = Pittosporum pentandrum (BLCOo)
MerR. (Pittosporac.).

2. PHALERIA

JACK, Mal. Misc. 2 (1822) 59; reimpr. HooK. Comp. Bot. Mag. 1 (1835) 156; DoMKE,
Bibl. Bot. 111 (1934) 123, t.4 f.36h, map 6; Merr. J. Arn. Arb. 33 (1952) 239.—
Drimyspermum REINW. Syll. Pl. Ratisb. 2 (1825) 15; RcusB. Nom. 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) 231.—Dais (non LINNE)

auct.—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 cylindric, 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-Mal. 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 l-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), SE. 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 lowland up to 1400 m.

Note. The generic name Drimyspermum 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

1. 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 314 by 2 cm; pericarp thin, less than 14 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, 214-314 by 1-2 cm. Floral tube wide, 12-15 mm diam. at the
throat =s) . I. P. elegans

3. Involucral bracts usually 4 or By smaller, ¥,— 1/, by if if cm. . Floral tube narrow, 2-6 mm diam.
at the throat.

4. Inflorescences 8—10(—15)-flowered, very rarely many-flowered. Flowers 114-2(-214) cm long;
calyx lobes !4(—'4) the length of the tube. Ovary glabrous. Leaves elliptic-oblong, elliptic-
lanceolate, or obovate, (4-)13-26 by (114-)3-8 cm; nerves 9-11 pairs . . 2. P. octandra

4. Inflorescences 20-many-flowered. Flowers 3-414 cm long; calyx lobes %4-—% the length of the tube.
Ovary usually hairy at the top, rarely glabrescent. Leaves oblong-lanceolate, oblanceolate or rarely
ovate-oblong, 1114-33 by 314-14 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 in P. sogerensis).

5. Stamens and style always included. Stamens sessile or short-filamentous. Style usually not longer
than the tube . . BAGS SE nee yee) ea eee

5. Stamens and style exserted. Stamens long filamentous.

6. Flowers 214-414 cm long. Calyx lobes 4-4 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, 1-114 cm in diam.; endocarp perforated (fibrous strands interlaced, leaving
distinct meshes). . . . -. 5S. Peicapais

7. Inflorescences 20-many- -flowered, rarely cauliflorous. Ovary usually hairy or puberulous at the
top. Fruits ellipsoid, 1!4—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 114-2 cm long. Calyx lobes usually 14(—14) 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 aes cm. Stamens c. 5—6 mm exserted. Calyx
lobes c. 2-214 times as long as wide. . . SLOVO Le BE SP Ssogerensis

Dec. 1960]

_ THYMELAEACEAE (Ding Hou) f RZ

1. 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, 3-514 by 3-414 cm; pericarp

thick, 1-114 cm.

1. Phaleria elegans L. M. Perry, J. Arn. Arb. 39
(1958) 422, f. c & d.—Fig. 3h-i.

Shrub or small tree, 2-3 m, sparsely branched.
Branchlets reddish-brown, terete, glabrous, hollow.
Leaves chartaceous to subcoriaceous, glabrous,
oblanceolate, rarely elliptic-lanceolate, 18-30 by
514-9 cm; base attenuate; apex acute to short-
acuminate; margins recurved; nerves 8—12 pairs,
elevated on both surfaces, spreading, curved to-
wards the margin; veins slightly elevated on both
surfaces, rather widely reticulate; petiole stout,
about as thick as the branchlet, red when fresh
(fide Brass), reddish-brown to black when dry.
Inflorescences terminal and/or in the axils of the
terminal node, subsessile to short-peduncled
(peduncle 5-9 mm in fruit), with small, decussate
bracts towards the base. Involucral bracts cream-
coloured (fleshy in bud), 8 or more, arranged in
whorls, those of the inner whorl longer, oblong or
ovate, 214-314 by 1-2 cm, obtuse or acute,
slightly puberulous towards the upper part inside,
especially near the margins, glabrescent, some-
times pellucid-dotted, 6—20-flowered. Flowers in-
fundibular, 334-414 cm long. Floral tube 144-114
cm diam. at the throat, densely pubescent outside
and towards the base inside. Calyx lobes sparsely
pubescent outside and densely pubescent inside,
+ oblong, c. 9 by 5 cm, obtuse. Petaloid ap-
pendages rim-like. Stamens included, filamentous,
6-10 mm, free from the tube at the throat or
slightly below it; anthers oblong, about 144 mm
long. Disk cup-shaped, crenulate, c. 2 mm long.
Pistil included. Ovary glabrous, ellipsoid, c. 3 mm
long, gradually narrowed towards the apex; style
filiform, c. 3 cm; stigma globose. Fruit ellipsoid to
fusiform, 234-314 by 114-134 cm, slightly com-
pressed, acuminate on both ends. Seeds broadly
ovate or obovate, or semiglobose, plano-convex,
10 by 7-10 mm, sharply pointed at the apex.

Distr. Malaysia: New Guinea (Goodenough
ayy.

Ecol. Undergrowth of an oak forest and oc-
casional in gullies in the forest, 1600-1750 m.

Note. This species is characterized by 8 or
more, large involucral bracts which are caducous
after anthesis, a wide floral tube which is densely
pubescent outside, and the included stamens and
style.

2. Phaleria octandra (L.) BAiLL. Adans. 11 (1875)
321; Merr. Philip. J. Sc. 19 (1921) 367.—Dais
octandra LINNE, Mant. PI. 1 (1767) 69; Burm. f.
Fl. Ind. (1768) 104, t. 32, f.2.— Dais dubiosa
(non BL.) Decne, Herb. Timor. (1834) 41;
SPANOGHE, Linnaea 15 (1841) 335.—Drimysper-
mum laurifolium Decne, Ann. Sc. Nat. Bot. II,
19 (1843) 39, t. 1, f. 1-12, as Drymispermum;
BLEEKER, Nat. Geneesk. Arch. N.I. 2 (1845) 75;
ZOLL. Syst. Verz. 2 (1854) 117; Mia. Fl. Ind. Bat.

8. P. macrocarpa

1, 1 (1858) 885; BLUME & DE VrRigeseE, Ann. Hort.
Bot. (Fl. Jard.) 2 (1859) 33, with pl.—Drimysper-
mum burmanni DECNE, Ann. Sc. Nat. Bot. II, 19
(1843) 40, as Drymispermum; BLEEKER, Nat.
Geneesk. Arch. N.I. 2 (1845) 75; MEISN. in DC.
Prod. 14 (1857) 605.—Drimyspermum blumei
(non DECNE) Hassk. Nat. Tijd. N.I. 10 (1856) 885.
—Drimyspermum ambiguum MEIsN. in DC. Prod.
14 (1857) 605, as Drymispermum.—Drimysper-
mum longifolium Mia. Fl. Ind. Bat. 1, 1 (1858)
885, as Drymispermum.—P. laurifolia Hook. f.
Bot. Mag. (1869) t. 5787; VAL. Ic. Bog. 4 (1913)
211, t. 368, f. 1-3.—P. ambigua Hook. f. Bot. Mag.
(1896) t. 7471.—P. longifolia Boeri. Handl. 3
(1900) 111.—P. laurifolia var. javanica VAL. Ic.
Bog. 4 (1913) 212, t. 368, f. 4-9; in K. & V. Bijdr.
13 (1914) 46; Hav. f. Med. Rijksherb. n. 44
(1922) 23.—P. octandra var. laurifolia WARB. ex
VON MALM in Fedde, Rep. 34 (1934) 282.—P.
parvifolia BAcK. Blumea 5 (1945) 494.—Fig.
3m-o.

Shrub up to 5 m by 5 cm. Leaves chartaceous to
subcoriaceous, elliptic-oblong, narrowly elliptic
or obovate, (4-)13-26 by (114-)3-8 cm; base
attenuate, rarely short-acute; apex acuminate;
nerves 9-11 pairs, slightly ascending and curved;
veins rather widely reticulate, slightly elevated
below, plane and visible above; petiole 6 mm.
Inflorescences usually terminal and/or in the axils
of the terminal node, sometimes in the axils along
the branchlets (BEUMEE 2405, Bo), 8—10(—15—cy)-—
flowered; peduncles very short to up to 1!4 cm,
usually with decussate, small, lanceolate scales at
the base gradually increasing in size apically. In-
volucral bracts 4, rarely 5, ovate or obovate,
8-12 by 4-11 mm, puberulous on the upper part
of both surfaces, persistent, rarely caducous after
anthesis. Flowers (1-)114-214 cm long. Floral
tube cylindric, slightly swollen at the base, usually
pubescent on both surfaces, sometimes glabres-
cent. Petaloid appendages sometimes rim-like.
Calyx lobes 4 or 5, oblong or slightly elliptic or
obovate, 4-7 by 2-3 mm, densely puberulous out-
side and at the upper part inside. Stamens and
pistil long-exserted (up to 8 mm) in anthesis.
Disk cup-shaped, membranous. Pistil 2—2!4 cm
long. Ovary glabrous, ellipsoid, narrowed into the
filiform style; stigma capitate. Fruits ellipsoid,
ovoid, sometimes slightly compressed, 11-16 by
9-15 mm, acute or attenuate and pointed towards
both ends, usually 2-celled, 2— or 1-seeded, once
found 3-seeded (VALETON 123, Bo).

Distr. Australia: North Queensland (MICHAEL
1250, Bo), and Malaysia: throughout Java,
Madura, Bawean, and Lesser Sunda Islands (Bali
to Timor and Tanimbar), Moluccas (Halmaheira),
and South New Guinea (Daru I.). Fig. 4.

Ecol. In beach-forest, common on sandy soil
of teak forest in E. Java, rarely in primary and

18 FLORA MALESIANA

[ser. I, ‘vol:

secondary mixed forests, from the lowland up to
600 m, in Jamdena up to 800 m, in Timor up to
1000 m.

—

Fig. 4. Localities of Phaleria octandra (L.) BAILL.
(+) and P. perrottetiana (DECNE) F.—VILL. (@).

Vern. Kopinan, mritja sunda, pantjal pamor, J,
kaju pateng, Kangean, mandalika, manpulang,
Bawean; Lesser Sunda Islands: daun wémpé, nu
impi, Jamdena, koffifui, Timor, lolong, Sumba;
pépigéow, Halmaheira, Sawai lang.

Note. Dais octandra was first published by
LINNAEUS in 1767 (Mant. PI. 1, p. 69). There is a
specimen in the Linnean Herbarium which bears
LINNAEuUs’s handwriting and agrees with the
original description with the exception that some
of the flowers are sparsely pubescent outside.

In 1768, one year later, Dais octandra appeared
in BuRMAN’s FI. Ind. p. 104, t. 32 f. 2. There are
two specimens of BURMAN’s in the Herb. Delessert
at Geneva; one of them bears in BURMAN’S
handwriting “‘Dais octandra’’ and the other has a
label ‘“‘Java Kleinhoff”. These two specimens are
similar to each other and may belong to one
collection. They are also similar to the specimen
in the Linnean Herbarium mentioned above but
the flowers appear more pubescent on the outside
of the floral tube. I assume BURMAN has sent his
drawing with one of his specimens — which might
have been the one which he had used for the des-
cription — to LINNAEUS, as LINNAEUS cited “‘Burm.
Ind. t. 33, f. 2”? in the description.

In 1876 BAILLON (Adansonia 11, p. 321) rightly
transferred Dais octandra to Phaleria as P.
octandra. He mentioned only BURMAN’s publica-
tion and overlooked that of LINNAEus. According
to priority we should accept LINNAEUS as the
author of Dais octandra and the specimen in the
Linnean Herbarium as the holotype.

Because of some minor discrepancies between
BuURMAN’s (/.c.) description and drawing, and both
of them also not exactly agreeing with the spec-
imens preserved in BURMAN’s herbarium, DECAIs-
NE (Ann. Sc. Nat. Bot. II, 19, 1843, 40) based a
new species, Drimyspermum burmanni, on the
specimens in the BURMAN Herbarium. However,
the discrepancies have no value for defining
species: the floral tubes are usually pubescent
outside but rarely glabrous and, as pointed out by

VALETON (in K. & V. Bijdr. 13, 1914, 40), one can
occasionally find glabrous flowers among the
hairy ones (cf. Koorpers 30200 f, 30145 8, 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 different 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-
and mesocarps form a rather thick layer, with a
smooth, compact, shining inner surface, more
loose than the peripheral parts but leaving no
meshes among them.

In 1893 H. HALLIER (320, Bo) once collected it
along the border of the Palace Garden (Herten-
kamp) adjoining the Botanic Gardens at Bogor;
it must have escaped or been derived from
cultivation (cf. VALETON in K. &. V. Bijdr. 13,
1914, 47). One specimen was collected by
HOOGERWERF (42, Bo) in the beach forest of
Udjungkulon, W. Java, which is apparently the
western limit of this species.

P. octandra is closely related to P. perrottetiana.
Some specimens collected at Sumbawa (ELBERT
3903, 3949, 3994, 4099, 4127) have 15—cv-flowered
inflorescences. One specimen collected in the
Tanimber Is. (S. Moluccas) (BUWALDA 4369), has
large leaves (20 by 9 cm) which are similar to
those of P. perrottetiana both in size and shape.

3. Phaleria perrottetiana (DECNE) F.—VILL. Noy.
App. (1880) 183; Merr. Sp. Blanc. (1918) 378;
Brown, Minor Prod. Philip. For. 1 (1920) 403;
Merr. En. Philip. 3 (1923) 131; Philip. J. Sc. 29
(1926) 404; Merr. & PeERRY, J. Arn. Arb. 22
(1941)265.—Drimyspermum perrottetianum DECNE,
Ann. Sc. Nat. Bot. II, 19 (1843) 40, as Drymisper-
mum; BLEEKER, Nat. Geneesk. Arch. N.I. 2 (1845)
75; MEISN. in DC. Prod. 14 (1857) 605; Mia. FI.
Ind. Bat. 1, 1 (1857) 886.—Dais laurifolia (non
Jacq.) BLANCO, FI. Filip. (1837) 375, ed. 2 (1845)
263, ed. 3, 2 (1878) 125.—Drimyspermum urens
(non REINW.) SCHEFF. Ann. Jard. Bot. Btzg 1
(1876) 46.—Drimyspermum coccineum (non Dais
coccinia GAUDICH.) BeEcc. in d’Albertis, New
Guinea 2 (1880) 398, as Drymispermum, quoad
specim.—P. blumei (non BENTH.) HEMSL. Bot.
Chall. 3 (1885) 244.—P. splendida VAL. Ic. Bog. 4
(1913) 219, t. 370 A-B.

Shrub, sometimes a tree, up to 8 m. Branches
and branchlets glabrous and dark-brown. Leaves
chartaceous, oblong-lanceolate, oblanceolate, or
ovate-oblong, 1114-33 by 314-14 cm; base
cuneate, rounded; apex acuminate; margins
slightly recurved; nerves (8—)13—22 pairs, distinct

Dec. 1960] THYMELAEACEAE (Ding Hou) ae 19

Fig. 5. A tree of Phaleria sp. in bridal attire.

Kebun Raya Indonesia, Bogor.

and elevated on both surfaces, curving and as-
cending towards the margin, veins and veinlets
slightly reticulate; petiole 114 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 3144 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-)3-414 cm
long. Floral tube pubescent outside, villose at
the lower half or lower 7 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,
114-3 by 1144-134 cm. Seed ellipsoid, plano-
convex, 10 by 8 mm.

Distr. Louisiade Archipelago (Sudest 1.),
Admiralty Is. and Malaysia: New Guinea (through-
out), Moluccas (Kai Is. and Ceram), Philippines
(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
Proy., 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 134 cm)
with rather thick pericarps (c. 2 mm). Their
enormous size might have been caused by the
attack of insects, as there are always hole(s) on
the pericarps and excrements of insects inside the
fruits.

4. Phaleria nisidai KANEHIRA, FI. 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, elliptic-oblong,
rarely lanceolate, 10-18 by 214-614 cm; base
obtuse, acute or cuneate; apex acuminate; nerves
6-10 pairs, curving and ascending towards the
margin, elevated beneath, plane and distinct
above, veins reticulate, usually rather dense,
elevated beneath, distinct or obscure above.

20 FLORA MALESIANA

[ser. I, vol. 6!

Inflorescences terminal and/or in the axils of the
terminal node, umbelliform, 10—12-flowered; pe-
duncles c. 2-4 mm with a pair of opposite, green,
and obovate-oblong bracts (4 by 2 mm) at the
upper part; involucral bracts 4, green, glabrous,
caducous after anthesis, oblong or elliptic-oblong,
8-10 by 4-6 mm. Flowers 2-314 cm long. Floral
tube cylindric, slightly dilated towards the top,
glabrous outside, puberulous or pubescent inside.
Calyx lobes 5(-4), oblong, ovate or orbicular,
2-414 by 2-3 mm, puberulous on the margins
and top outside and the whole surface inside.
Stamens included; filaments 14-3 mm; anthers c.
1 mm long. Disk cup-shaped, crenate, mem-
branous. Pistil usually shorter than the tube,
rarely exserted. Ovary ovoid, c. 244 mm long,
hairy at the apex; style filiform; stigma oblong or
slightly globose. Fruits globose or slightly obo-
void, 114-2 by 114-2 cm, slightly compressed,
constricted at the base into a 3 mm long stipe.
Seeds broadly ellipsoid, 5144 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
flowers with included stamens and pistils.

5. Phaleria capitata Jack, Mal. Misc. (1822) 59;
reimpr. in Hook. f. Comp. Bot. Mag. 1 (1835) 156;
K. & V. Bijdr. 13 (1914) 41; S. Moore, J. Bot. 63
(1925) Suppl. 89; HEYNE, Nutt. Pl. (1927) 1152.—
Dais dubiosa Bu. Cat. (1823) 69; Bijdr. (1826) 651;
HaAssk. Cat. Hort. Bog. (1844) 94; Fiver, Pl. Bot.
Tuin Weltevr. (1855) 50.—Drimyspermum urens
REINW. Syll. Pl. Ratisb. 1 (1825) 15; DeEcNe, Ann.
Sc. Nat. Bot. II, 19 (1843) 39; BLEEKER, Nat.
Geneesk. Arch. N.I. 2 (1845) 74; MEIsN. in DC.
Prod. 14 (1857) 604; HoL_tHuis & LAM, Blumea 5
(1942) 216.—Drimyspermum blumei DECNE, Ann.
Sc. Nat. Bot. II, 19 (1843) 39, as Drymispermum;
BLEEKER, Nat. Geneesk. Arch. N.I. 2 (1845) 74;
ZOLL. Syst. Verz. 2 (1854) 117; MeIsn. in DC.
Prod. 14 (1857) 604; Mio. FI. Ind. Bat. 1, 1 (1857)
885.—P. dubiosa ZoLL. Nat. Geneesk. Arch. N.I.
1 (1844) 616.—Drimyspermum laurifolium (non
DEcNE) HAssk. Nat. Tijd. N.I. 10 (1856) 155.—
Drimyspermum phaleria MEIsN. in DC. Prod. 14
(1857) 604, as Drymispermum; Mia. Fl. Ind. Bat.
1, 1 (1858) 884.—Drimyspermum cauliflorum
THw. En. Ceyl. Pl. (1860) 251, as Drymispermum.
—P. cumingii F.-VILL. Nov. App. (1880) 183;
VIDAL, Phan Cuming. Philip. (1885) 140; Rev.
Pl. Vasc. Filip. (1886) 230; BRown, Minor Prod.
Philip. For. 1 (1920) 403; Merr. Bull. Bur. For.
Philip. 1 (1903) 43; En. Philip. 3 (1923) 131;
KANEHIRA, Bot. Mag. Tokyo 45 (1931) 331; FI.

Micron. (1933) 248.—P. urens KoorpD. Minah.
(1898) 577; Suppl. 2 (1922) t. 98; Suppl. 3 (1922)
48.—P. cauliflora BEDpD. For. Man. Bot. (= FI.
Sylv. vol. 3) (1873) 180, t. 25, f. 5; Trim. Fl. Ceyl. 3
(1895) 459; Hook. f. Fl. Brit. Ind. 5 (1886) 199.—
Fig. 3a-g, 6.

Fig. 6. Phaleria capitata JACK. Botanic Gardens,
Singapore, Febr. 1952 (Photogr.
M. R. HENDERSON).

Shrub or small tree, up to 9 m by 16 cm.
Branchlets reddish-brown. Leaves chartaceous,
glabrous, in dry state reddish-brown above, pale
brown beneath; elliptic-oblong, (11—)1514—21(—26)
by (3!4-)514-7(-10) cm; base acute to attenuate,
rarely rounded; apex narrow acute to acuminate,
acumen 114-214 cm; margins sometimes recurved
in dry state; nerves 8-10 pairs, elevated beneath,
slightly elevated above; veins loosely reticulate,
distinct beneath, obscure above; petiole 5 mm.
Inflorescences usually terminal and/or in the leaf
axils of the terminal node, solitary, sometimes
cauliflorous, subsessile or on very short (c. 3 mm)
peduncles, with decussate, small bracts at the
base. Involucral bracts 4, oblong, ovate or
obovate, 6 by 3 mm, usually caducous after
anthesis sometimes persistent, usually 8-flowered.
Flowers 24-44 cm long. Floral tube cylindric,
gradually enlarged towards the top, glabrous on
both surfaces. Calyx lobes oblong or elliptic, 6-7
by 2-314 mm, puberulous inside and towards the
upper part and margins outside, sometimes
glabrous outside. Stamens and style usually ex-
serted sometimes up to 5 mm. Pistil sometimes
shorter than the tube or about as long as it
(BAKHUIZEN VAN DEN BRINK 5063, BAKHUIZEN
VAN DEN BRINK f. 688, and San A 3140). Disk

Dec. 1960] i

eee eee &

cup-shaped. Ovary ellipsoid, glabrous, apex
narrowed into the filiform style; stigma capitate,
144 by 1 mm. Fruits subglobose, 1-144 cm in
diam., sometimes short-acute at the apex, usually
2-celled, 2-seeded; endocarp inside with distinct
meshes.

Fig. 7. Localities of Phaleria capitata JACK.

Distr. Ceylon, Carolines (Palau, fide KANEHIRA),
and Malaysia: Malaya (cult. and natur.), Sumatra,
Java, Borneo, Philippines, Celebes, Moluccas
(Buru), and New Guinea (Waigeo I.). Fig. 7.

Ecol. In primary and secondary forests, from
the lowland up to 1200 m.

Wises-(G@j- HIBYNE, Ic); BuRK. Dict: 2, 1935,
1703). Tough fibres of the bark have been used for
cordage and tying material. The fruits are sweet
and edible. The seeds are used for scurfy eruptions
in children.

Vern. Sumatra: suwa lansat, Simalur, rimbo
suloh, Lampong; Java: godong-laweh, kakapassan,
ki-angkrieng, ki tangkieh, S, kojoian, lawé, lawéan,
lawé-lawé, ulati, J; Borneo: djarum djarum;
Celebes: suka, Bug., sunsuan, susuan, Minah.;
Moluccas: /a’awan’a, Sangi & Talaud Is.

Notes. The mesocarp of the fruit is fleshy and
rather soft. The endocarp consists of interwoven
fibres which form a characteristic network;
through the meshes one can see the testa of the
exposed seed (cf. VALETON, Ic. Bog. 4, 1914, 217).
Fig. 3g.

There are two specimens collected by Corner
(S.F. 28481 and 32776, SING) at Johore, Malay
Peninsula, which might have escaped or been
cerived from cultivation.

6. Phaleria coccinea (GAuUDICH.) F. v. M. Descr.
Not. 2 (1885) 9, quoad basionym; K. Scu. &
Laut. Fl. Schutzgeb. (1900) 459.—Dais coccinea
GAUDICH. Voy. Uranie (1826) 443, t. 44.—Pseudais
coccinea DECNE, Ann. Sc. Nat. Bot. II, 19
(1843) 41; BLEEKER, Nat. Geneesk. Arch. N.I.
2 (1845) 77; MEIsN. in DC. Prod. 14 (1857) 603;
Mia. FI. Ind. Bat. 1, 1 (1858) 883.—Drimysper-
mum revolutum T. & B. Nat. Tijd. N.I. 27 (1864)
30.—Drimyspermum cumingii MEISN. in DC. Prod.
14 (1857) 605, as Drymispermum.—P. vriesii
BAILL. Adansonia 11 (1875) 329.—P. zippelii
BAILL. /.c.—Drimyspermum coccineum BECcc. in

D’Albertis, New Guinea 2 (1880) 398, quoad
basionym, as Drymispermum.—P. revoluta BOERL.
Handl. 3 (1900) 111; VAL. Ic. Bog. 4 (1913) 215,
t. 369.—P. amboinensis MeErRR. Philip. J. Sc. 11
(1916) Bot. 294.—P. platyphylla MerR. Philip. J.
Se. 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, 1514-26 by 6-1114
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. S-7 mm
long. Inflorescences terminal or/and axillary at the
terminal node, rarely cauliflorous, (15—)20-many-
flowered; peduncle up to 2'44 cm. Bracts small,
lanceolate, ovate to obovate, 3-10 mm long;
involucral bracts 4, elliptic- or obovate-oblong,
obtuse, 4-12 by 2-5 mm, caducous after anthesis,
rarely persistent. Flowers 2'!4-3'4 cm long. Floral

tube glabrous outside, glabrous or sparsely
puberulous to pubescent inside. Calyx lobes
obovate-oblong or oblong, 5-7 by 2-3 mm,

reflexed, usually puberulous on both surfaces.
Stamens and style up to 8-10 mm exserted. Disk
cup-shaped. Ovary villous or puberulous at the
top, 2-celled, very rarely 1-celled. Fruits ellipsoid,
114-2 by 1 cm, blunt at both ends, sometimes
apiculate at the apex. Seeds ovoid, 644 by 5 mm.

Distr. New Britain and Malaysia: Philippines
(Panay and Mindanao), Moluccas (Sula Is.,
Ambon, Ceram, and Key Is.), New Guinea
(Sorong, Waren, Hollandia, Central and South
Division, Nabire, Sepik region).

Ecol. In rain-forests from the lowland up to
300 m.

7. Phaleria sogerensis S. Moore, J. Bot. 61 (1923)
Suppl. 43.

Shrub up to 2 m. Branchlets reddish- to dark-
brown, glabrous. Leaves papery, rarely sub-
coriaceous, glabrous on both surfaces, sometimes
brownish or brownish-green when dry, elliptic-
oblong, oblong-lanceolate to lanceolate; 7-16 by
214-7 cm; cuneate to attenuate towards the base;
acuminate at the apex, sometimes with an acumen
c. 2 cm long; nerves 6-10 pairs, spreading towards
the margin and then curved upwards, prominent
and elevated beneath, slightly elevated above;
veins reticulate, distinct beneath, obscure above;
petiole c. 5 mm. Jnflorescences terminal, and/or
in the leaf axils of the terminal one or two nodes,
6-10-flowered; peduncle none to 12 mm; in-
volucral bracts 4, slightly obovate, c. 5 by 3 mm,
usually caducous, sometimes persistent. Flowers
114-2(-214) cm long. Floral tube glabrous on both
surfaces, rarely puberulous inside. Calyx lobes
slightly oblong, 5-8 mm long, densely puberulous
inside and on the margins outside. Stamens and

22 FLORA MALESIANA

Iser. I; voliab*

Fig. 8. Phaleria macrocarpa (SCHEFF.) BOERL.
Kebun Raya Indonesia, Bogor
(C.H.B. VIII-G—93).

pistil slightly exserted. Disk cup-shaped, crenate,
c. ¥% mm long. Ovary usually pubescent or
puberulous at the apex, rarely glabrescent or
glabrous. Fruits ellipsoid, acute or acuminate
towards both ends, 114 by 1 cm, rarely the basal
part of the endocarp inside with small meshes.
Seeds ellipsoid, plano-convex, 8 by 5 mm with a
caruncle-like appendage, c. 114 mm long.

Distr. Malaysia: New Guinea (Sogere, Kanosia,
Koitaki, Boridi, and Hollandia).

Ecol. In forests at low and medium altitudes,
sometimes up to 1400 m.

8. Phaleria macrocarpa (SCHEFF.) BOERL. Handl. 3
(1900) 111; L. M. Perry, J. Arn. Arb. 39 (1958)
420, fig. a-b.—Drimyspermum macrocarpum
ScuHerrF. Ann. Bot. Gard. Btzg 1 (1876) 46, as
Drymispermum,—P. octandra (non (L.) BAILL.)

Fig. 9. Phaleria macrocarpa (SCHEFF.) BOERL.

with its beautiful, glossy, bright-red fruits.
Cultivated in Kebun Raya Indonesia, Bogor.

K. Sco. & Hoxie. Fl. Kais. Wilh. Land (1889)
93; WaRB. Bot. Jahrb. 13 (1891) 337.—P. papuana
Wars. ex K. Sco. & Laut. Fl. Schutzgeb. (1900)
460; Gitc, Nova Guinea 8 (1910) 411.—
P. sp. Gitc, l.c.—P. calantha GiLG, l.c.—P.
wichmannii VAL. Ic. Bog. 4 (1913) 222, t. 371.—
Fig. 3j-], 8-9.

Shrub or small tree, up to 18 m by 15 cm.
Young branches hollow. Leaves chartaceous to
subcoriaceous, glabrous, ovate-oblong, elliptic-
oblong, lanceolate, oblong-lanceolate, 10-25 by
3-10 cm; base cuneate or rounded; apex shortly
acute to acuminate, acumen up to 2 cm; nerves
6-11 pairs, spreading towards the margins, some-
times their distal end curving upwards and united
loop-like, elevated and distinct on both surfaces;
veins loosely reticulate, sometimes subperpendi-
cular to the midrib, slightly elevated on both
surfaces. Inflorescences terminal and in the axils

Dec. 1960] i

| _ THYMELAEACEAE (Ding Hou) i 23

along the branchlets, sometimes cauliflorous, 1 to
5, rarely more peduncles in each axil; peduncles
0 or up to 244 cm (cf. bb 25746), each 2—5(—8)-
flowered; involucral bracts 4, small, obovate,
oblong, caducous, 7 by 2 mm. Flowers 114-4 cm
long. Floral tube glabrous on both surfaces, some-
times puberulous and hairy on the inside towards
the base. Calyx lobes oblong, 4 by 2 mm, reflexed,
densely puberulous inside and on the margins
outside. Stamens sessile or up to 6 mm exserted.
Ovary glabrous; style shorter than or as long as
the tube or c. 5-10 mm exserted. Fruits sub-
globose to broadly ellipsoid or rounded, some-
times slightly obovate and stipe-like narrowed
towards the base, 3-5!4 by 3-414 cm, exocarp
woody when dry. Seed subglobose or slightly
ovate, c. 1144 by 144 cm.

Distr. Malaysia: common in western New
Guinea.

Ecol. Primary and secondary forests, from the
lowland up to 550 m, once at 1260 m.

Uses. It is cultivated in Sabron near Hollandia
on clay soil at 120 m. The bark is used by the
Papuans for making bags (BW 5468).

Vern. Dalom, Sentani, kotteh, Djair, matoniek,
Andjai.

Note. Phaleria macrocarpa was first described
as Drimyspermum macrocarpum by SCHEFFER
(l.c.) based on fruiting specimens collected by
TEYSMANN (7786, L, Bo) near Doré, western New
Guinea. These young fruits are ellipsoid or
slightly obovoid (15 by 12 mm) and narrowed
stipe-like towards the base. This shape is not un-

common and can also be observed in for example
bb 25746, BW 5468 from Hollandia, and NGF
7298 from Morobe Distr. The vegetative and
morphological characters of the type agree with
those of P. calantha and P. wichmannii.

The type of P. wichmannii was collected by
ATASRIP (139, Bo, L; cult. in Hort. Bog. under
n. VIII. G. 75) in northern Dutch New Guinea;
its flowers have a long, exserted style and almost
sessile stamens.

The type of P. calantha was collected by
VERSTEEG (1939, Bo, L. K; cult. in Hort. Bog.
under nv. VIII. G. 93) at Merauke, southern Dutch
New Guinea; its flowers have long filamentous
stamens and a style shorter than or as long as the
floral tube. The fruits of these two ‘species’
collected from the cultivated plants mentioned
above are very similar, large and fleshy. I assume
that the flowers are heteromorphous and these
two ‘species’ represent two forms of one species.
Moreover, two other forms have been found,
viz one with both stamens and style exserted as
represented for example by KLoss s.n. (13/1 &
16/1, 1912 [1913], BM) and HooGLanp 4520 (Bo,
G, L), and another one with short-filamentous
stamens and a short style, represented by NGF
9564 (L).

Excluded
Phaleria axillaris E-mer, Leafl. Philip. Bot. 8
(1915) 2840 is according to MERRILL (En. Philip. 3,
1923, 535) = Tricalysia tinagaoensis ELMER (Ru-
biac.).

3. ENKLEIA

GriFF. Calc. J. Nat. Hist. 4 (1844) 234, in note; VAN TigGH. Bull. Soc. Bot. Fr.
40 (1893) 69; Domke, Bibl. Bot. 111 (1934) 121, t.4 f.36n & map 3.—Linostoma
subg. Linostoma Kurz, J. As. Soc. Beng. 39, ii (1870) 83; reimpr. Flora 53 (1870)
372.—Linostoma WALL. ex ENDL.: BENTH. & Hook. f. Gen. Pl. 3 (1883) 197,
p.p.—Macgregorianthus MerRR. 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-
Mal. sp.). Flowers 5-merous, articulated at the base of the pedicel. Floral tube
cylindric, 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 —, linear 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. 61

aa hy ; ‘
\ ee 1
. A
SORE
Roy f
i

UN i ie

Fig. 10. Enkleia malaccensis Grirr. a. Habit, x 2, b. part of stem with hook-like branchlets, x 24,

c. opened flower, x 6, d. stamens, x 13, e. infructescence with two leafy bracts, x 24, f. fruit, x 2,

g. cross-section showing structure of pericarp, x 4.—E. paniculata (MerR.) HALL. f.. 4. Opened flower,

x 6,7. stamens, x 13 (a KEITH 9234, b MAINGAY 1308/2, c-d HEYNE s.n., €@ ELMER 20834, f KOSTERMANS
7034, h-i ZipPELIUS 1488).

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. Seed the

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. malaccensis

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

1. Enkleia malaccensis GriFF. Calc. J. Nat. Hist. 4
(1844) 235, in note; Gi_tG in E. & P. Pfl. Fam. III,
6a (1894) 231; GamBLeE, J. As. Soc. Beng. 75, ii
(1912) 262, excl. syn.; HALL. f. Med. Rijksherb.
n. 44 (1922) 24; Ripi. Fl. Mal. Pen. 3 (1924) 147;
FIscHER, Kew Bull. (1932) 182; Burk. Dict. 1
(1935) 925; LEANDRI, Rev. Int. Bot. Appl. &
Agr. Trop. 29 (1949) 505; Proc. 8th Pac. Sc.
Congr. Manila 4 (1957) 585.—Lasiosiphon scan-
dens ENDL. Gen. Pl. Suppl. 4, 2 (1847) 67, nom.
illegit.; MEISN. in DC. Prod. 14 (1857) 598; Mia.
Fl. Ind. Bat. 1, 1 (1858) 881.—E. malayana GRIFF.
Not. As. 4 (1854) 363.—Linostoma scandens
Kurz, J. As. Soc. Beng. 39, ii (1870) 83; reimpr.
Flora 53 (1870) 371; Hoox. f. Fl. Brit. Ind. 5
(1886) 198, excl. syn.; BOERL. Handl. 3 (1900) 111;
HEYNE, Nutt. Pl. (1927) 1152.—E. riouwensis
Hai. f. Med. Rijksherb. n. 44 (1922) 25.—
E. coriacea HALL. f. /.c.—Fig. 10a-—g.

Climber up to 30 m by 10 cm. Branchlets some-
times transformed into hook-like organs, reddish
brown. Branchlets, inflorescences, and young
leaves always ferrugineous-pubescent. Leaves sub-
coriaceous to coriaceous, upper surface dull, oli-
vaceous when dry, undersurface usually brownish,
or reddish brown, pubescent on both surfaces,
sometimes glabrescent; usually broad-elliptic and
bluntish, 514-14 by 3-7 cm, obtuse or rounded at
both ends, rarely short-acute; nerves 12-20 pairs,
distinct and plane beneath, visible and impressed
above, veins distinct beneath, obscure above;
petiole 6-12 mm, pubescent. Inflorescences ter-
minal, up to 30 cm long, flowers (4-)6—8(—14) on
each ramification; leafy bracts chartaceous,
oblong, 4-5 by 1-2 cm. Flowers c. 8 mm long,
yellowish or whitish, short-pedicelled. Calyx lobes
c. 2mm long, 3 of them larger and ovate, 2 smaller
and lanceolate. Petaloid appendages 10, linear, ec.
3, mm. Stamens c. 8 mm, sessile or shortly
filamentous. Pistil included, c. 3 mm long; ovary
c. 2 mm; style short; stigma slightly capitate.
Fruits ovoid, 1% by % cm.

Distr. Andaman Is., Burma (Tenasserim and
Prome, fide Kurz), Indo-China (Laos and
Cambodia), and Malaysia: Sumatra (Palembang),
Riouw, Banka, Malay Peninsula (Singapore and
Malacca), and Borneo.

Ecol. In lowland forests.

2. E. paniculata

Uses. Said to give an inferior scented ‘gaharu’
wood ( fide HEYNE, /.c.). The bast fibers can be used
for tying purpose.

Vern. Akar kareh hitam, akar panas, akar
puchong kapur, garu buaja, kapang akar, M,
témentak akar, Banka, térap akar, Sum.; Borneo:
aka dian, Kaya, akar garu, Dusun, tuba-tuba,
Bajau.

2. Enkleia paniculata (MerRR.) HALL. f. Med.
Rijksherb. n. 44 (1922) 26.—Macgregorianthus
paniculatus Mere. Philip. J. Sc. 7 (1912) Bot. 312;
GitG in E. & P. Pfil. Fam. Nachtr. 4 (1915) 212;
Merr. En. Philip. 3 (1923) 132.—E. zippeliana
HALL. f. /.c.—Fig. 10h-i.

Climbing shrub. Branchlets puberulous, glabres-
cent. Leaves chartaceous to subcoriaceous, when
dry the upper surface light-brown, glabrous and
shining, the lower surface somewhat paler, dull,
sparsely puberulous, glabrescent; ovate-(rarely
elliptic-)oblong, 514-11 by 3-5 cm; base acute to
obtuse; apex acute to + acuminate; nerves 11-15
pairs, slightly ascending towards the cartilaginous
margin and united with it, elevated beneath,
slightly elevated or plane above; veins slightly
elevated beneath, plane or slightly impressed
above; petiole 5-8 mm, densely puberulous.
Inflorescences in the upper axils, up to 28 cm long,
densely puberulous; leafy bracts oblong, 2!—6 by
34-2 cm. Pedicels 3-5 mm. Flowers pale-green, c.
1 cm long. Floral tube cylindric and distinctly
costate inside, c. 8 mm long, densely puberulous
outside, glabrous inside, twisted after anthesis.
Calyx lobes oblong, c. 2 mm long, densely pu-
berulous outside and on the margins or sometimes
the whole surface inside. Petaloid appendages
oblong, membranous, emarginate or slightly erose
at the top, 14-1 mm long. Stamens of the upper
series 114-2 mm long with filaments as long as or
longer than the anthers, those of the lower series
c. 1 mm long, sessile or shortly filamentous;
anthers c. 14 mm long, obtuse or slightly apiculate.
Ovary ovoid or ovoid-oblong, c. 244 mm long,
densely pubescent; style filiform, c. 144 mm;
stigma obovoid, papillose. Fruit ovoid, 14% by
Jem:

Distr. Malaysia:
western New Guinea.

Philippines (Luzon) and

26 FLORA MALESIANA

[ser. 1; vol=e

Ecol. In hill-side forests (Philip.) or lowland
rain-forest, 50 m.

Note. According to HALLIER f. (/.c.) E. zippe-
liana is similar to E. paniculata, but would differ
in leaves shortly and sparsely puberulous beneath,
with conspicuous and slightly prominent network
of veins on both surfaces, branchlets, petioles and
panicles minutely rusty (not gray-)tomentose, and

geographic distribution. However, from the two
sheets of the type of E. paniculata (B.S. 12360, Bo,
L) as compared with the type of E. zippeliana
(ZIPPELIUS 148/a, L), it appears that these dif-
ferences are only quantitative. There are only
young flowers on the Philippine specimens and the
floral characters are similar to those in the New
Guinea specimen in the same stage.

4. LINOSTOMA

WALL. [Cat. (1831) no 4203, nomen] ex ENDL. Gen. Pl. (1837) 331; Suppl. 4,
2 (1847) 67; BENTH. & Hook. f Gen. Pl. 3 (1883) 197, p.p.; DoMKE, Bibl. Bot.
111 (1934) 120, map 3.—Nectandra (non BERG. 1767) Roxs. [Hort. Beng. (1814)

NM

Y Z
K——

Fig. 11. Linostoma longiflorum HALL. f.. a. Habit, x 24.—L. pauciflorum GrirF. b. Habit, x 24, 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, f. disk at base of ovary, * 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 Rotts.—Lino-
stoma sect. Eulinostoma MEISN. in Mart. FI. Bras. 5, 1 (1855) 71.—Psilaea Maia.
Sum. (1861) 355.—Linostoma subg. 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.
Petaloid appendages 10, long club-shaped or filiform, 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 f. (Med. Rijksherb. n. 44,
1922, 27) and the two species in our region both belong to the sect. Psilaea (MiqQ.) HALL. f. /.c. 28.

KEY TO THE SBRECIES

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 2-214 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 3-314 cm long .

1. Linostoma pauciflorum GrirF. Calc. J. Nat.
Hist. 4 (1844) 234, in note; GAMBLE, J. As. Soc.
Beng. 75, ii (1912) 261; MeIsN. in DC. Prod. 14
(1857) 600; Mia. FI. Ind. Bat. 1, 1 (1858) 882;
Kurz, J. As. Soc. Beng. 39, ii (1870) 83; For. FI.
Burm. 2 (1877) 334; Hook. f. Fl. Br. Ind. 5 (1886)
198; BoerL. Handl. 3 (1900) 107, 111; RipL. J.
Str. Br. R. As. Soc. n. 59 (1911) 164; Fl. Mal. Pen.
3 (1924) 146.—Psilaea dalbergioides Mig. Sum.
(1861) 355.—L. leucodipterum Hau. f. Med.
Rijksherb. n. 44 (1922) 28; Airy SHAw, Kew Bull.
(1940) 262.—Fig. 11b-g.

A climber up to 24 m, rarely a shrub or small
tree up to 7 m (cf. H. M. BurKILL & SHAH 240).
Branches long, slender, black when dry. Leaves
chartaceous, glabrous, rather glaucous beneath,
elliptic, 114-4(-6) by 1—2(-314) cm; apex obtuse
and mucronate or shortly acute; base acute,
cuneate, obtuse, or rounded; petiole c. 2 mm.
Inflorescences terminal or rarely axillary, (1—)2—4-
flowered, nodding, umbelliform, provided with
2 opposite or subopposite leafy bracts at the lower
half of the peduncle, besides sometimes a small
linear bract (5-7 mm long) at the top of the
peduncle; leafy bracts usually smaller than but
sometimes as large as the ordinary leaves, ovate or
oblong-ovate, 2 14-4 by 114-2 cm, whitish when dry,

2. L. longiflorum

translucent and with less lateral nerves than the
leaves. Peduncles 5-10 mm; pedicels c. 7 mm.
Flowers green to greenish-white. Floral tube
slender, slightly narrowed towards both ends,
12-15 mm long, usually glabrous on both
surfaces. Calyx lobes linear. Petaloid appendages
club-shaped or filiform, c. 5 mm long. Stamens
7-10 mm, usually exserted. Ovary including the
stipe 4-6 mm long; style 15-18 mm, terminal or
slightly sublateral; stigma capitate. Fruits ellip-
soid, c. 1144 cm long, narrowed to both ends.

Distr. S. Siam (Singora, Dulit), Burma
(Tenasserim and E of Tounghoa in the Martaban
Hills), and Malaysia: Sumatra (Simalur), Malay
Peninsula (Perlis, Kedah, Dindings, Penang, and
Singapore), and Borneo (W. Borneo).

Ecol. In primary and secondary forests, from
the lowland up to c. 1300 m.

Vern. Bébora, kakat bétul, kakrat butu or butol,
pérakat bétul, tuba bara, M.

Use. The Burmese use it medicinally (cf. BURK.
Dict. 2, 1935, 1352; field note on Curtis 3197).

2. Linostoma longiflorum HALL. f. Med. Rijks-
herb. n. 44 (1922) 29.—Fig. 11a.

A slender climbing shrub. Leaves chartaceous,
glabrous, dull on both surfaces, ovate, 3-314 by

28 FLORA MALESIANA

[ser. I, vol

114-214 cm; apex obtuse and mucronate; base
rounded, sometimes shortly acute. Flowers
solitary or sometimes 2, axillary, or terminal on
the short branchlets in the inferior leaf-axils of the
branch; peduncle c. 5 mm, provided with 2 small
bracts; pedicels c. 7 mm. Flowers green. Floral
tube slightly ellipsoid, 2-214 cm long, glabrous
outside, sparsely pubescent inside. Calyx lobes

oblanceolate, c. 10 mm long. Petaloid appendages
10, club-shaped, c. 6 mm long. Stamens c. 10 mm,
usually exserted. Ovary c. 7 mm long, surrounded
at the base by a very short disk; style long filiform,
314 cm; stigma capitate. Fruit unknown.
Distr. Malaysia: Borneo (Sarawak).
Ecol. Primary peat-swamp forest,
altitude (fide J. A. R. ANDERSON 9047).

at low

5. WIKSTROEMIA

ENDLICHER, Prod. Fl. Norfolk. (1833) 47, as Wickstroemia, nec SCHRADER 1821
(Theac.), nec SPRENGEL 1821 (Comp.), nom. gen. cons.; Gen. Pl. (1837) 332,
Suppl. 4 (1847) 68; Domke, Bibl. Bot. 111 (1934) 124, t.4 f.36r & s, map 6, excl.
syn. Stellera L. n. 2.—Capura LINNE, Mant. Pl. 2 (1771) 149, nom. gen. rejic.—
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 slightly 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 cylindric 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 (1857) 543.—Subg. Euwikstroemia DomKE, Bibl. Bot. 111 (1934) tab. facing p. 58 (Type species:
W. australis ENDL.).

Nomencl. In LInNAEuSs’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 Ste/lera L. and its delimitation has caused much
controversy and confusion. As far as I 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. MeyYER (Bull. Ac. Imp. Sc. St. Pétersb. 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),
STAPF (in Curtis’s Bot. Mag. 1924, t. 9028), and HircHcock & GREEN (Proposals by British Botanists
1929, 150).

DomKeE merged Stellera L. with Wikstroemia, in transferring its type species, S. chamaejasme L.,
to Wikstroemia, retaining the taxon as a separate subgenus Chamaejasme [AMMAN] Domke (cf. Notizbl.
Berl.—Dahl. 11, 1932, 362; Bibl. Bot. 111, 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 if Ste/lera 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

Fig. 12. Wikstroemia brachyantha MerR. a. Habit, x 24, b. opened flower, x 4/,.—W. androsaemifolia

DECNE. c. Habit, x 24, d. opened flower, x 3.—W. tenuiramis Mia. e. Habit, x 24, f. opened flower,

3, g. pistil with scale-like disk at base, x 7 (a—b CLEMENS 32439, c CoeRT 41, d RANT s.n., e—f For.
Dep. N. Borneo 4173).

REHDER (/.c.) and Popepim (Fl. U.S.S.R. 15, 1949, 502) have, however, retained Wikstroemia and
Stellera as two 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. BRowNn, Min. Prod. Philip. For. 1, 1920, 403).

30 FLORA MALESIANA

[ser. I, vol "G

Notes. FAGERLIND (Hereditas 26, 1940, 38 & 48) found an agamogenic clone of W. indica which is

an intraspecific triploid (2n = 27).

The tropical African Englerodaphne GiLG, reduced by DomkeE (Bibl. Bot. 111, 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 (3-814 by 14-214 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 214-5 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, (134-1514 by 34-5 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 | 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 314-5 cm; base usually obtuse or

cuneate, rarely subcordate. Flowers 1144-2 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, 14-4 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 panies 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. Inflorescences umbelliform;
flowered.

6. W. venosa

rachis not elongating, very short or less than 14 cm long, few-

8. Flowers (15—)18—-22 mm long. Stamens usually sessile; anthers 114-2 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= 1h mm long. Leaf base

acute, attenuate or obtuse.

9. Flowers 9-12 mm sti puberulous outside. Leaves acute at both ends, 134-514(-8) by 4-244

(-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-)6-1514 by (214-)3-5 cm

9. W. ridleyi

3. Leaves usually obovate- or elliptic-oblong, oblanceolate, elliptic, or rarely ovate, 114-414(-7) by
14-2(-314) 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).

1. Wikstroemia lanceolata MerRR. Publ. Govt Lab.
Philip. 29 (1905) 31; Philip. J. Sc. 1 (1906) Suppl.
101; 5 (1910) Bot. 366; Brown, Min. Prod.
Philip. For. 1 (1920) 404; Merr. En. Philip. 3
(1923) 133.—W. angustissima MERR. Philip. J. Sc.
7 (1912) Bot. 92; En. Philip. 3 (1923) 132.

An undershrub up to 4 m. Young branchlets
densely appressed-pubescent and _ glabrescent.
Leaves membranous to papery, glabrous, rarely
sparsely pubescent on the midrib beneath,
lanceolate to narrowly lanceolate, 3-814 by
14-214 cm; base obtuse; apex acuminate; nerves
10-16 pairs, sometimes branched and irregular,
slightly elevated beneath, visible or obscure above,
obliquely spreading to the margins and united into

10. W. indica

an intramarginal vein; veins obscure or distinct,
spreading and loosely reticulate; petiole short,
1-214 mm, appressed-hirtellous. Inflorescences
umbelliform to shortly spicate, terminal, very
rarely axillary; peduncles very short, sometimes
up to 114 cm, (1—)3—5(—20)-flowered; pedicels c.
1-114 mm, appressed-hirtellous. Flowers green or
yellowish-green, 6-15 mm long, puberulous,
glabrescent outside. Calyx lobes ovate or oblong,
obtuse, c. 1 mm long. Stamens sessile or short-
filamentous, c. 34-114 mm long. Disk 2 free scales,
linear or slightly oblong. Ovary ovoid, slightly
hairy at the apex; style obscure; stigma sub-
globose. Fruits short-ovoid, c. 8 by 5 mm, usually
glabrous sometimes sparsely hairy at the top,

Dec. 1960]

THY MELAEACEAE (Ding Hou) Je ¥ => Aa

pericarp fleshy. Seeds 6 mm long.

Distr. Malaysia: Philippines (Palawan, Min-
doro, and Luzon).

Ecol. Common on forested slopes at low and
medium altitudes up to 1300 m.

Vern. Philippines: Aaranpinig, Neg., maragawa,
salagip, salagé, Tag., suka, tuka, I\k.

2. Wikstroemia brachyantha Me_rr. Philip. J. Sc.
13 (1918) Bot. 313; En. Philip. 3 (1923) 132.—
W. crassifolia MERR. ex DOMKE, Bibl. Bot. 111
(1934) tab. facing p. 58.—Fig. 12a—b.

Shrub or small tree up to 3144 m by 244 cm.
Branchlets puberulous, light brown. Branches
reddish-brown and usually transversally fissured.
Leaves subcoriaceous to coriaceous, rarely
chartaceous, in dry condition both surfaces
olivaceous-brown to brownish, glabrous and
shining, elliptic-oblong to ovate-oblong, rarely
lanceolate, 5-15 by 214-5 cm; base obtuse to
cuneate; apex acuminate; nerves 12-15 pairs,
elevated and prominent beneath, slightly elevated
or plane above, obliquely spreading towards the
margin and united with the intramarginal vein;
veins anastomosing, almost as prominent as the
nerves; petiole 2-3 mm, glabrous. Inflorescences
terminal or/and in the leaf axils at the terminal
node, distinctly peduncled; peduncles stout,
brownish-pubescent, sometimes angular or flatten-
ed and gradually thickening towards the top,
usually bent slightly downward, sometimes with 1
or 2 bracts below the 6 to 8 apical flowers. Flowers
yellowish, yellowish-green or green, subsessile.
Floral tube 10-12 mm long, sparsely puberulous
outside, glabrous inside. Calyx lobes 4, ovate-
oblong, 114-3 mm long, obtuse. Two series of
stamens close to each other, on very short fila-
ments; anthers oblong, obtuse, 1144 mm long.
Disk 2 free, oblong, 2-lobed scales. Ovary ellipsoid
or slightly obovoid-oblong, 3 mm long, hairy at
the top; style distinct, filiform, c. 1 mm; stigma
globose, papillose. Fruits red, broadly ellipsoid,
12 by 9 mm.

Distr. Malaysia: Borneo (Kinabalu) and
Philippines (Luzon and Catanduanes).

Ecol. In the Philippines in primary forests at
low and medium altitudes, in Borneo (Mt Kina-
balu) in damp and mossy forests at 1400-2800 m.

Note. This species is characterized by its
usually thick leaves with prominent venation es-
pecially on the lower surface, a marginal vein on
each side, and the stout peduncle.

3. Wikstroemia tenuiramis Mig. Sum. (1861) 141
& 354; BoorsMa, Bull. Dép. Agric. I.N. 1. 7 (1907)
19; HeyNeE, Nutt. Pl. (1927) 1152; Burk. Dict.
(1935) 2258.—W. acuminata MerR. J. Str. Br. R.
As. Soc. n. 76 (1917) 99; En. Born. (1921) 417;
Un. Cal. Publ. Bot. 12 (1929) 218.—W. clementis
Me_rr. J. Str. Br. R. As. Soc. n. 76 (1917) 99; En.
Born. (1921) 417; Herne, Pfi. Clemens Kinabalu
(1953) 69.—Fig. 12e-g.

Shrub or small tree, up to 10 m. Branchlets
light-brown to dark-brown, sparsely pubescent,
glabrescent. Branches smooth, reddish-brown,

glabrous. Leaves membranous to papery, glabrous,
in the dry state the upper surface subolivaceous or
light-brown, rather shining, lower surface dirty-
white or light-green, rather dull, sometimes light-
brown on both surfaces, ovate-oblong, elliptic-
oblong, broadly-elliptic, or lanceolate, rarely
ovate, 6-12 by 114-414 cm; base cuneate, acute or
obtuse; apex acuminate, the acumen up to c.
1 cm; nerves 7—12 pairs, rather irregular, slightly
elevated, rarely indistinct on both surfaces,
obliquely ascending close towards the margin and
then curved upward; veins loosely anastomosing,
reticulations usually obscure on both surfaces;
petiole c. 4 mm. Jnflorescences usually axillary
and occurring in several leaf axils along the
branches or branchlets, sometimes also terminal
or on the top of a reduced or very short branchlet
with bract-like reduced leaves, 1—5-flowered;
peduncle very short to 114 cm, appressed-pu-
berulous. Flowers 10-13 mm long, yellowish, or
cream (fide CLEMENS 20980), subsessile. Floral
tube scattered-puberulous outside, glabrescent.
Calyx lobes ovate-oblong, 2-3 mm long. Stamens
with c. 14 mm space between the two whorls,
those of the upper series sessile or sometimes some
of them shortly filamentous, those of the lower
series always shortly filamentous; anthers linear,
1-114 mm long, acute or slightly apiculate. Ovary
oblong or slightly obovoid-oblong, c. 2 mm long,
glabrous or a few hairs at the top; style very short
or sessile; stigma capitate, papillose. Fruits yellow,
green or orange, ovoid, c. 8 by 5 mm.

Distr. Malaysia: Sumatra (Menggala), Banka,
Borneo (N. Borneo, Brunei, Sarawak, and S.
Borneo: Sampit).

Ecol. In forests, swampy land, and hills, from
the lowland up to 1600 m.

Uses. According to BoorsMa (/.c.) it provides a
scented wood which is used only occasionally.
The wood is harder than that of Aquilaria and
scentless, but when burned it gives forth a frag-
rance similar to that of Aloe-wood. In Banka the
bark is used for making ropes.

Vern. IJnjat, Brunei, kaju lingau, Menggala,
ménameéng, téméntak tindat, Banka.

Note. This species can easily be distinguished
from related ones by the axillary inflorescences
which occur in several leaf axils along the branch-
lets or branches, the more or less discoloured
leaves, and the usually indistinct venation.

Mrs CLEMENS once noted this species to be a
vine (n. 31292) but I believe this to be due to er-
roneous information by her native collectors.

4. Wikstroemia ovata C. A. Mey. [Bull. Ac. Imp.
Sc. St. Pétersb. Cl. Ph.—M. 1 (1843) 357; reimpr.
Ann. Sc. Nat. Bot. II, 20 (1843) 50, nomen] ex
MEISN. in DC. Prod. 14 (1857) 544; Mia. FI. Ind.
Bat. 1, 1 (1858) 880; F.—ViLL. Nov. App. (1880)
182; VIDAL, Phan. Cuming. (1885) 140; Rev. Pl.
Vasc. Filip. (1886) 230, excl. syn. Daphne aquilaria
BLANCO; Merr. Philip. J. Sc. 1 (1906) Suppl. 101;
Sp. Blanc. (1918) 279; Brown, Min. Prod. Philip.
For. 1 (1920) 404; Merr. En. Philip. 3 (1923) 133,
excl. citation of VIDAL, Synopsis; Quis. Med. Pl.

32 FLORA MALESIANA

[ser. I, vol. 6!

Philip. (1951) 637.—Daphne indica (non LINNE)
BLANCO, FI. Filip. (1837) 309, ed. 2 (1845) 215, ed. 3,
2 (1878) 38.—Daphne foetida (non LINNE) BLANCO,
lc. 308, as phaetida, Il. cc. 217, 37.

Shrub up to 5 m by 7-8 cm. Young branchlets
appressed-hirtellous, glabrescent or glabrous.
Leaves membranous or papery, glabrous, rarely
sparsely pubescent on the midrib beneath; ovate to
ovate-oblong, 4-14 by 314-5 cm; base usually
obtuse or cuneate, rarely subcordate; apex acumi-
nate; nerves 8-12 pairs, curved and ascending,
slightly elevated below, distinct above; veins
reticulate, distinct beneath, visible or obscure
above, petiole 3 mm, sparsely appressed-hirtellous.
Inflorescences terminal, short-spicate or umbelli-
form, peduncled, sparsely puberulous, sometimes
with 1 or 2 caducous bracts, 7—20-flowered.
Flowers 14%4—2 cm long, greenish, yellowish, at the
upper end of the peduncle, short-pedicelled.
Floral tube cylindric, sparsely puberulous outside.
Calyx lobes oblong, obtuse, 2-4 mm long. Stamens
sessile or on short filaments; anthers 1-114 mm,
slightly apiculate. Disk 2 free, oblong, scales.
Ovary ellipsoid, 2-3 mm long, hairy at the apex;
style distinct, filiform, 34-114 mm; stigma capitate.
Fruits subglobose to slightly ellipsoid, 8-10 by

Malaysia: Borneo (North Borneo,
Sebattik I., Sampit, and Pulu Lampei) and the
Philippines (Palawan, Mindoro, Luzon, Negros,
and Mindanao). Fig. 13.

Ecol. In thickets, primary and secondary
forests at low and medium altitudes up to 800 m.

Fig. 13. Localities of Wikstroemia ovata C. A. MEY.
ex MEISN.

Uses. This plant has been used by the Filipinos
as a purgative. The leaves are a strong purgative
when chewed and swallowed and one bowel
movement is produced for every taken. The fresh
bark or branches of this plant are tied round
about the neck of a patient to relieve bronchial
catarrh (cf. GarciA, Philip. J. Sc. 51, 1933,
485-494; Quis. /.c.).

Vern. Philippines: arandoén, Ik, dapnit, suka,
Bon., salagé, Tag.; Borneo: gélam hutan, Brunei,
pait-pait, Bajau.

5. Wikstroemia polyantha Me_rr. Philip. J. Sc. 10
(1915) Bot. 332; En. Philip. 3 (1923) 133.—W.
candolleana (non MEISN.) RIDL. Trans. Linn. Soc.
Bot. II, 3 (1893) 341, as candollei, corr. p. 456;
J. Str. Br. R. As. Soc. n. 35 (1901) 180; Fl. Mal.
Pen. 3 (1924) 145; BurK. & HEND. Gard. Bull.
S.S. 3 (1925) 417; Heyne, Nutt. Pl. (1927) 1152;
Burk. Dict. (1935) 2258; SyminctT. J. Mal. Br.
R. As. Soc. 14 (1936) 358.—W. junghuhnii (non
Mia.) K. & V. Bijdr. 13 (1914) 58, sphalm. jung-
huhniana.—W. ridleyi (non GAMBLE) GrBBs, J.
Linn. Soc. Bot. 42 (1914) 132; Merr. En. Born.
(1921) 417.—W. calva BAck. Blumea 5 (1945) 494.

Shrub or small tree up to 7 m by 714 cm.
Branchlets reddish to dark-brown, sparsely pu-
berulous, glabrescent. Leaves membranous, char-
taceous, rarely subcoriaceous, glabrous on both
surfaces, rarely scattered hairy beneath especially
on the midrib, in dry condition light-brown to
dark-brown above, paler beneath, ovate-oblong,
elliptic-oblong, or lanceolate, 6—9(—-12) by 114-314
(-414) cm; base acute to cuneate, sometimes ob-
tuse or rounded; apex acuminate, rarely acute;
margins sometimes slightly recurved; nerves 8-15
pairs, irregular, often branched, elevated beneath,
distinct or plane above, obliquely ascending tow-
ards the margin; veins obscure on both surfaces,
sometimes as distinct as the nerves; petiole 2—4
mm, sparsely pubescent. Inflorescences terminal
or/and in the axils at the terminal node, spicate,
gradually elongating, up to 4 cm, rarely to 6 cm,
erect or slightly curved, very rarely nodding
(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, 14-1 mm, puberulous, articulated at
the base. Floral tube scattered puberulous outside,
glabrous inside. Calyx lobes oblong or ovate-
oblong, 1!4-3'4 mm long. Anthers linear, c.
1 mm long, the two series close to each other;
filaments c. 14 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, 114-214 mm long, glabrous or
sparsely hairy at the top; style distinct, as long as
or slightly longer than the stigma; globose or
slightly oblong, c. % mm long, papillose. Fruits
ovoid, red, c. 844 by 5144 mm.

Distr. Malaysia: Malay Peninsula (Pahang,
Kedah, Perak, Johore, Kelantan, Selangor, and
Gunong Korbu), Java (western part), North
Borneo, and Philippines (Luzon).

Ecol. In forests, from the lowland up to 2200 m.

Use. The wood yields incense.

Vern. Chandan pélaudok, M.

Notes. BACKER (/.c.) has pointed out that the
name “Wikstroemia junghuhniana”’ (non MiQ.)
given by KOoRDERS & VALETON (/.c.) was an error
because they intended to identify their specimens
(KoorpDERS 26824 6 and J. J. SmitH 292, Bo) as
W. junghuhnii Mie. (= W. androsaemifolia
Decne). The sheets cited by KOORDERS & VALETON
under “W. junghuhniana”’ as mentioned above
differ from W. junghuhnii Mig. and belong to a

Wee; 1960]

different species; therefore, BACKER provided
them with a new name ‘‘Wikstroemia calva’ to
indicate the ‘completely glabrous ovary tip’.
However, whether the ovary is glabrous or hairy
at the apex is not a constant character as some
specimens (e.g. VAN STEENIS 4187 & 12930) have
both kinds of ovaries. Specimens collected at high
altitude have thicker and reddish-brown leaves.

The species is characterized by the long-spicate
inflorescences which are gradually elongating
during flowering time. It is closely related to W.
nutans CHAMP. from Kwangtung, China, differing
from it by the larger leaves, distinct style, and the
approximate insertion of the two whorls of
stamens.

The inflorescences are erect or slightly curved
with the exception of one specimen (S.F. 20726)
collected in Pahang at c. 2100 m, which has both
erect and nodding ones.

6. Wikstroemia venosa Merr. & PERRY, J. Arn.
Arb. 22 (1941) 266.

Shrub, c. 1 m. Young branchlets densely yellow-
ish-brown puberulous; older ones reddish- or
dark-brown, puberulous, glabrescent. Leaves
chartaceous or subcoriaceous, lower surface of the
young leaves densely pubescent on the midrib and
scattered pubescent on the lamina, glabrescent;
in dry condition light brown or brown above and
shining, glaucous beneath; ovate-oblong to
lanceolate, 2-7 by 1—3 cm; base obtuse or cuneate;
apex acute to acuminate; nerves 7—10 pairs, rather
irregular, obliquely spreading towards the margin
and then slightly curved ascending, elevated
beneath, plane or slightly depressed above; petiole
c. 2 mm long, puberulous. /nflorescences terminal,
nodding, sessile or up to 1 cm peduncled; spicate,
rachis elongated, 14-2 cm, puberulous. Flowers
crowded, c. 8 mm, green; pedicel c. 1 mm, ar-
ticulated at the base, puberulous. Floral tube
scattered-puberulous outside and glabrous inside,
sometimes glabrous outside. Calyx lobes oblong
or ovate-oblong, 2—2!4 mm long. Stamens shortly
filamentous; anthers linear, c. 1 mm long, obtuse,
the space between the two series c. 34 mm. Disk
2 free linear scales, bilobed at the top, sometimes
disk cup-shaped and erose at the top, 14-1 mm
long. Ovary slightly obovate-oblong, c. 144 mm
long, sparsely strigose at the top; style short, c.
1/, mm; stigma capitate. Fruits ellipsoid, 6-8 mm
long, slightly narrowed at both ends. Seed similar
in shape to the fruit.

Distr. Malaysia: New Guinea (Balim Valley,
Manokwari, Humboldt Bay, Hollandia, and Rona
(Central Div.)).

Ecol. Deforested slopes, grassland, occasionally
on grassy banks of streams, lowland up to 1900 m.

7. Wikstroemia meyeniana WARB. in Perk. Fragm.
Fl. Philip. (1905) 171; Merr. Philip. J. Sc. 1 (1904)
Suppl. 101; BRown, Min. Prod. Philip. For. 1
(1920) 404, t. 23; Merr. En. Philip. 3 (1923) 133.
—Daphne cannabina (non Lour.) SCHAUER, Nov.
Act. Caes. Leop.—Car. 19 (1843) Suppl. 1, 411.—
W. longifolia LECoMTE, Not. Syst. 3 (1914) 128;

THYMELAEACEAE (Ding ow 33

Fl. Gén. I.-C. 5 (1915) 167; LEANDRI, Proc. 8th
Pac. Sc. Congr. 4 Bot. (1957) 582, incl. var.—
W. fenicis MeRR. Philip. J. Sc. 13 (1918) Bot. 312;
En. Philip. 3 (1923) 132.

Shrub up to 3 m. Young branchlets sparsely
pubescent, glabrescent, sometimes glabrous. Leay-
es papery, glabrous, olivaceous, shining, lanceolate
rarely elliptic-lanceolate, (5—)914-13 by (114-)
314-414 cm; base obtuse occasionally shallow-
cordate or attenuate; apex acuminate; nerves
9-14 pairs, slightly curved and ascending, elevated
beneath, visible or obscure above, veins reticulate,
slightly elevated beneath, obscure above; petiole
3-4 mm. /nflorescences terminal and axillary, um-
belliform, sometimes occurring on short, reduced
branchlets associated with reduced leaves (2-3 cm
long) and resembling a leafy panicle, (2—)5—6
(-10)-flowered; peduncle up to 3 cm, densely ap-
pressed-hirtellous. Flowers greenish-yellow, (15—)
18-22 mm long, densely puberulous outside.
Floral tube cylindric. Calyx lobes narrow-
oblong, 2-3'4 mm long. Stamens usually sessile
rarely some of them on short filaments, 114-2 mm
long. Disk 2 free, oblong scales. Ovary slightly
obovoid, 2-214 mm long, hairy at the apex; style
distinct, 14-1 mm long; stigma oblong, 14—!4 mm
long. Fruit ovoid, c. 8 by 6 mm.

Distr. Indo-China and Malaysia:
(Luzon to Mindanao).

Ecol. In primary humid forests at low and
medium altitudes, up to c. 400 m.

Vern. Philippines: sagu, Tag., salago, Bik., Tag.

Philippines

8. Wikstroemia androsaemifolia DECNE, Ann. Sc.
Nat. Bot. II, 20 (1843) 50; in Jacq. Voy. Bot.
(1844) 146; BLEEKER, Nat. Geneesk. Arch. N.I. 2
(1845) 74; MEIsN. in DC. Prod. 14 (1857) 546;
Mia. FI. Ind. Bat. 1, 1 (1858) 879; Back. Bekn. FI.
Java (em. ed.) 4A (1942) fam. 77, p. 5.—W.
spanoghii Decne, Ann. Sc. Nat. Bot. I, 20 (1843)
50; in Jacq. Voy. Bot. (1844) 146; BLEEKER, Nat.
Geneesk. Arch. N.I. 2 (1845) 74; MEIsN. in DC.
Prod. 14 (1857) 545; Mia. FI. Ind. Bat. 1, 1 (1858)
879.—Eriosolena viridiflora ZOLL. & Mor. Nat.
Geneesk. Arch. N.I. 1 (1844) 615, excl. syn. Hassk.
Cat. 117; ZoLi. Syst. Verz. 2 (1854) 116.—W.
candolleana MEIsN. in DC. Prod. 14 (1857) 544;
K. & V. Bijdr. 13 (1914) 56; Mia. FI. Ind. Bat. 1,
1 (1858) 878; Hai. f. Med. Rijksherb. n. 12
(1912) 26; K. & V. Bijdr. 13 (1914) 56; HEYNE,
Nutt. Pl. (1927) 1152; Burk. Dict. 2 (1935) 2258.
—W. junghuhnii Mia. FI. Ind. Bat. 1, 1 (1858) 879;
HALL. f. Med. Rijksherb. n. 44 (1922) 30.—
Fig. 12c—d.

Shrub up to 2!4 m by 4 cm. Young branchlets
slightly flattened at the nodes, densely appressed-
pubescent, glabrescent. Branches terete, reddish-
brown, glabrous; axillary buds densely covered
with golden-coloured hairs. Leaves papery,
glabrous, rarely sparsely hairy on the lower
surface and especially on the nerves and veins of
young leaves, in dry state light-greenish, light-
brown or greenish-brown to brownish and shining
on the upper surface; pale-greenish, light-yellowish
-green or light-brown and dull on the under-

34 FLORA MALESIANA

[ser. I; vol ae

surface; elliptic, elliptic- or ovate-oblong, rarely
broadly elliptic, 134-S14(-8) by 34-21%4(-4) cm;
base acute; apex acute to narrow-acute, very rarely
obtuse; nerves 8-11 pairs, elevated below and
slightly depressed above, obliquely spreading
towards the margin and then curved upward;
veins almost as distinct as the nerves, loosely
reticulate beneath, obscure above; petiole c. 2mm.
Inflorescences umbelliform or spicate, 5—10-flower-
ed, terminal and in the axils of the terminal node,
so usually 3 inflorescences at the top of the
branchlet, of which usually the middle one (some-
times also the lateral ones) is provided with a
pair of bracts or reduced leaves; peduncle obscure
to 314 cm, erect or slightly curved; pedicels c.
1 mm, articulated at the base. Flowers light-green
or yellowish-green. Floral tube slightly pubescent
outside, 9-12 mm long. Calyx lobes oblong, or
slightly ovate, fleshy, 2-314 mm long, obtuse.
Disk 2, rarely 3 free, linear scales. Stamens
filamentous; anther c. 1 mm long, obtuse, the
space between the anthers of upper and lower
series 1-114 mm. Ovary ellipsoid or slightly
obovoid, 114-214 mm long, pilose at the top,
sometimes glabrescent; style obscure to 1 mm
long; stigma globose, c. 14 mm in diam. Fruits red,
oblong, rounded.

Distr. Malaysia: Central and East Java,
Madura, Kangean Arch., Lesser Sunda Islands
(Flores and Timor), Borneo (North Borneo,
Koetai and Balikpapan), Celebes (Bonthain,
Manado, and G. Pangararan), and W. New
Guinea.

Ecol. In lowland forests from near the beach
up to 1800 m, in Celebes at 2200-2400 m.

Note. There are two authentic sheets of W.
androsaemifolia in the Nat. Hist. Mus., Paris; one
is labelled as ‘Java Leschenault’ and the other as
‘Daphne n. 341’. Apparently these specimens may
belong to one collection, as they are very similar.
The leaves are papery and rather discoloured,
brownish above, light-brown beneath, and not
larger than 6 by 214 cm. The flowers are about
10 mm long, sessile, and fascicled or crowded on
a short peduncle (c. 10 mm). The ovary is sparsely
hairy or glabrous at the top.

The type of W. spanoghii was collected by
SPANOGHE (s.n., L) in Timor. Its leaves are rather
membranous, pale-greenish beneath, light-green-
ish above, and the size is up to 8 by 214 cm. It has
longer, spicate inflorescences and the longest
peduncle is c. 2!4 cm. The ovary is densely hairy
at the top, and the style is obscure.

When one compares the type specimens of these
two species they do not seem to be conspecific.
However, after examining a large range of spec-
imens, there are too many intermediate forms in
which the differential characters break down.
Consequently I have interpreted them as forms of
one variable species.

The type specimen of W. junghuhnii Mia.
collected by JUNGHUHN (s.n., L) on Mt Ungaran
(Central Java) and some specimens (e.g. Koor-
DERS 43148 6, 43875 6, 43876 B, and VAN STEENIS
17974) collected on mountains between 1100-—

1800 m have leaves with distinct, densely reticulat-
ed venation, and flowers with distinct peduncles
and short styles.

HANDEL-MAzZZETTI described a different species
from Yunnan, China, as W. androsaemifolia in
1923. If this proves to be a good species it must be
renamed.

9. Wikstroemia ridleyi GAMBLE, Kew Bull. (1912)
200; J. As. Soc. Beng. 75, ii (1912) 260; Rpt. FI.
Mal. Pen. 3 (1924) 146, f. 147; Burk. Dict. (1935)
2258:

Shrub up to 2 m. Branchlets reddish-brown,
sparsely puberulous and glabrescent. Leaves mem-
branous to chartaceous, usually olive-brown when
dry, glabrous on both surfaces, rarely sparsely
puberulous on the midrib beneath, elliptic-oblong,
lanceolate, ovate-oblong or ovate, 4-15!14 by
214-414 cm; base attenuate, acute or obtuse; apex
acuminate; nerves 7-12 pairs, slightly curved
towards the margin and then upward, slightly
elevated on both surfaces; veins obscure on both
surfaces; petiole 2-3 mm, sparsely puberulous
when young. Flowers c. 15 mm long, yellow or
greenish-yellow, 6-14, umbelliform on a terminal,
very short, slightly puberulous peduncle; pedicels
c. 1 mm, articulated towards the base, puberulous.
Floral tube sparsely puberulous outside, glabres-
cent, glabrous inside. Calyx lobes ovate-oblong,
obtuse, 314-4 mm long. Stamens shortly filamen-
tous, free from the tube at the upper half, the two
series c. 1 mm apart; anthers linear, obtuse or
slightly apiculate, c. 144 mm long. Disk 2 free
linear, c. 1 mm long, 2-lobed scales. Ovary
ellipsoid or slightly obovoid, 114-2 mm long,
hairy at the apex; style very short or obscure;
stigma globose and papillose. Fruits red, ellipsoid
or ovoid, 8 by 5 mm. Seeds ovoid, the same shape
as the fruit.

Distr. Lower Siam (Telok Udang) and
Malaysia: Malay Peninsula, chiefly on the east
coast (Kelantan, Trengganu, Pahang, Burau Bay).

Ecol. Sandy open coastal country.

Uses. The species contains a purgative sub-
stance and the leaves are eaten as an aperient. The
bark is used as entering into a compound potion
against small-pox; it is pounded and converted
into a poultice for applying to boils, or merely tied
round the neck to stop vomiting. It is also used as
a fish-poison. The fruits are poisonous (cf.
BURKILL).

Vern. Dépu, dépu pélandok, M; dalu pélandok
is a misprint (cf. BURKILL, /.c.).

Note. According to Rm.ey this plant was
brought from Pekan (not: Penang!) to Singapore
in 1890 and “ran wild for some time in Tanglin,
Singapore”.

Three specimens have been cited in the original
description, collected by RipLey at Pahang
(Kwala Brawas: RIDLEY 1583, lectotype and
Pekan: RIpLey s.n., Aug. 1889, paratype) and
Tringganu (Pulo Katan: RIDLEY s.n., Aug. 22,
1899, paratype).

10. Wikstroemia indica (L.) C. A. Mey. Bull. Ac.
Sc. St. Pétersb. 1 (1843) 357; reimpr. Ann. Sc. Nat.

Dec. 1960]

THYMELAEACEAE (Ding Hou) pate WP mi ee

Bot. II, 20 (1843) 50; MEISN. in DC. Prod. 14
(1857) 543; Mig. FI. Ind. Bat. 1, 1 (1858) 880;
BENTH. FI. Austr. 6 (1873) 37; F.-VILL. Nov. App.
(1880) 182; Via, Rev. Pl. Vasc. Filip. (1886)
229; Fors. & HeEmMsL. J. Linn. Soc. Bot. 26 (1894)
398; BoeRL. Handl. 3 (1900) 111; BAILEY, Queens].
Fl. pt 4 (1901) 1369; BoLp. Zakfl. (1916) 171;
Me_rr. Sp. Blanc. (1918) 279; Brown, Min. Prod.
Philip. For. 1 (1920) 404; Merr. En. Philip. 3
(1923) 132; ReHper, J. Arn. Arb. 15 (1934) 103;
MeErR. Comm. Lour. (1935) 278; BAcK. Bekn. FI.
Java (em. ed.) 4A (1942) fam. 77, p. 4; HOLTH. &
Lam, Blumea 5 (1942) 216.—Daphne indica LINNE,
Sp. Pl. (1753) 375.—Daphne aquilaria BLANCO, FI.
Filip. (1837) 310; ed. 2 (1845) 216; ed. 3, 2 (1878)
39.—W. viridiflora MEISN. Denkschr. K. Bayer.
Bot. Ges. Regensb. 3 (1841) 286; DECNg, in Jacq.
Voy. Bot. 4 (1844) 145; MEeEIsn. in DC. Prod. 14
(1857) 546; Mia. FI. Ind. Bat. 1, 1 (1858) 879;
VIDAL, Rey. Pl. Vasc. Filip. (1886) 229; MerrR.
Philip. J. Sc. 3 (1908) Bot. 422; BAck. Ann. Jard.
Bot. Btzg Suppl. 3 (1909) 419; K. & V. Bijdr. 13
(1914) 54; Rip. Fl. Mal. Pen. 3 (1924) 145;
GUILLAUMIN, J. Arn. Arb. 13 (1932) 88; BurRK.
Dict. (1935) 2259; LEANDRI, Proc. 8th Pac. Sc.
Congr. Manila 4 (1957) 582.—W. ovata (non C. A.
Mey.) VIDAL, Synopsis (1883) 229.—W. indica var.
viridiflora Hook. f. Fl. Br. Ind. 5 (1886) 195.—
W. linearifolia ELM. Leafl. Philip. Bot. 2 (1910) 680;
Merr. En. Philip. 3 (1923) 133.—W. pulgarensis
Eto. Leafl. 5 (1913) 1844; Merr. En. Philip. 3
(1923) 133.—W. pachyphylla MeErR. Philip. J. Sc.
12 (1917) Bot. 297; En. Philip. 3 (1923) 133.—
W. subcoriacea MERR. J. Str. Br. R. As. Soc. n. 76
(1917) 100; En. Born. (1921) 417; Herne, Pfil.
Clemens Kinabalu (1953) 69.—Daphne sp. STEEN.
Bull. Jard. Bot. Btzg III, 13 (1933) 254.

Shrub up to 3 m. Branchlets black-brown,
scattered puberulous, glabrescent, sometimes
transversally fissured. Internodes usually very
short or even obscure. Leaves chartaceous to sub-
coriaceous, in dry condition usually brown to
reddish-brown, sometimes glaucescent, sparsely
puberulous beneath, glabrescent, or glabrous,
shining above and rather dull beneath, obovate-
or elliptic-oblong, oblanceolate, elliptic, rarely
ovate, 114-414(-7) by 14-2(-314) cm; base cuneate
to attenuate; apex rounded, obtuse, sometimes
slightly emarginate, or acute; margins usually
cartilaginous; nerves 5—12 pairs, irregular, and
often branched, obliquely ascending towards the
margin, rarely the basal | or 2 nerves on each side
ascending along the margin towards near the top,
usually distinct beneath obscure above, sometimes
obscure on both surfaces; veins obscure or in-
visible on both surfaces. Petiole c. 2 mm. Jn-

florescence terminal, sometimes 1-2 additional
ones in the axils of the terminal node, few-
flowered, subsessile, sometimes on a very short
peduncle; pedicels 114-2 mm, articulated at the
base. Flowers green, 10-12 mm long, sparsely
puberulous outside, glabrous inside. Calyx lobes
2-3 mm long, broadly ovate or oblong, obtuse.
Stamens very shortly filamentous, rarely sessile;
anthers linear, c. 1 mm long, sometimes those of
the lower series slightly shorter, obtuse rarely
apiculate at the apex, the two series c. 1 mm apart.
Disk 2 free, linear, c. 74 mm long scales with
narrowed or obliquely truncate top, sometimes
lobed or crenate at the apex. Ovary slightly
obovoid or elliptic, c. 114 mm long, sparsely hairy
or glabrous at the top; style very short or obscure;
stigma globose, c. ¥4 mm diam. Fruits broadly
ellipsoid, c. 6 by 4 mm, red.

Distr. India, SE. Asia, through Malaysia to
Australia (N. Australia, Queensland, and N.S.
Wales) and Melanesia (as far E as Fiji), in
Malaysia not found in the seasonal parts: absent
from the Lesser Sunda Islands, in Java only found
in the vicinity of Bogor and once found at 1100 m
near Sindanglaja as an escape from the Botanic
Gardens, now locally thoroughly naturalized and
slowly spreading along roadsides and in other
anthropogenous terrain (BACKER, 1909 /.c.).

Ecol. In thickets and secondary growths, ob-
viously very soil-tolerant and occurring in various
biotopes, for example on sandy soil near the
beach, on limestone of a ridge top, on granite
peaks, along river-banks, and on open hill-sides,
from the lowland up to 1300 m, a few above 2200
m even up to 2700 m (in Celebes and New Guinea).

Vern. Borneo: lajak, M; Philippines: inyam,
P. Bis., arandén, baleo, I\k., palupo, titpuho, Iv.,
salagé, Tag., Bis., Bik., talo, Bik.; Celebes:
pérapata or posi-posi, Manado.

Note. W. indica is a widely distributed species
and is very variable in its vegetative parts. As
pointed out by BENTHAM in a note under W. indica
(Fl. Austr. 6, 1873, 37), “‘it is, however, not always
easy to determine the limits to be assigned to it’’.
FAGERLIND (/.c.) has found apomixis in this spe-
cies, which may give an explanation of its great
vegetative variability and difficulties involving in
specific demarcation. With a large number of
specimens of this species available, no sharply
defined infraspecific taxa or forms can be dis-
tinguished.

Excluded

W. amplifolia (SCHLTR) DomMKE (Bibl. Bot. 111,
1934, 60) of New Caledonia (isotype: SCHLECHTER
14749, L) has erroneously been recorded for New
Guinea by DoMkKE (i.c.).

6. DAPHNE

LINNE, Gen. Pl. ed. 5 (1754) 167; Sp. Pl. (1753) 356; MEISN. in DC. Prod. 14
(1857) 530; GILG, in E. & P. Pfl. Fam. 3, 6a (1894) 237; Domkg, Bibl. Bot. 111
(1934) 130, t.4 f.i & map 8.—Scopolia Linnf£ f. Suppl. (1781) 409, non Jaca. 1764,
nec al.—Eriosolena Bi. Bijdr. (1826) 651; VAN TiBGH. Bull. Soc. Bot. Fr. 40
(1893) 67; Domxg, Bibl. Bot. 111 (1934) 70-83, 130, t.4 f.36 C & map 10.—

36 FLORA MALESIANA [ser. I, vol

\

J
oh
- |

Af
2 iy
fo 7 |

Fig. 14. Daphne luzonica C. B. Ros. a. Habit, 24, b. opened flower, x 4.—D. composita (L. f.) GILG.
c. Habit, x 24, d. opened flower, 4, e. fruit, x 2, f. seed, x 2, g. longitudinal section of fruit, x 2
(a—b BS 40335, c-d RAHMAT sI BOEEA 11238, e-—g SF 51832).

Daphne sect. Eriosolena MrIsn. 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. /nflorescences

Dec. 1960] gee _THYMELAEACEAE (Ding Hou) a 24)

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-Mal. 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-
filiform, terminal, sometimes slightly lateral (in extra-Mal. 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. luzonica belongs to a section Daphnanthoides GiLG (l.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 JAcaQ. nec al.) LINNE f. 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
(/.c.) reduced Eriosolena BL. to a section of Daphne and in 1857 (/.c.) he reduced also Scopolia L. f. to
Daphne. The reduction of Eriosolena has been adopted e.g. by BAILLON (Hist. Pl. 6, 1877, 131), BENTHAM
& HOOKER (Gen. Pl. 3, 1880, 190), Gi_G (in E. & P. Pfl. Fam. 3, 6a, 1894, 238), Bo—ERLAGE (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. n. 44,
1922, 30), DoMKE (Bibl. Bot. 111, 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

1. Flowers in distinctly peduncled (2!4-614 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 . 2 fo 6s) le DEZcomposita
1. Flowers in sessile or short peduncled (0- 3 mm) heads without involucral bracts. Floral tube glabrous
oquisides Disk ‘obscure, ring-likesOvary glabrous). 93) 5) 4 2) eee De lozonies

1. Daphne composita (L. f.) Gitc in E. & P. Pfl. Tjib. 2 (1923) 201; BurK. & Hottr. Gard. Bull.
Fam. 3, 6a (1894) 238; Koorp. 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; Hocnr.
258; K. & V. Bijdr. 13 (1914) 49; Koorp. FI. Candollea 2 (1925) 443, incl. var. montana HOCHR.

38 FLORA MALESIANA

[ser. Ij’ vokeG*

and var. montana f. macrophylla HocurR.; BuRK.
Dict. (1935) 765; CoRNER, Ways. Trees (1940) 633,
f. 240; Back. Bekn. Fl. Java (em. ed.) 4A (1942)
fam. 77, p. 5; HEYNE, Nutt. Pl. (1927) 1152.—
Scopolia composita L.f. Suppl. (1781) 409.—
D. javanica THUNB. Mus. Nat. Acad. Upsal. App.
11 (1806) 4, nomen; FI. Jav. (1825) 13.—D. pendula
So. Pl. Ic. ined. 2 (1790) 34, t. 34, nom. illegit.;
Wikstr. Kongl. Vet. Acad. Handl. (1818) 296;
Meisn. Denkschr. K. Bayer. Bot. Ges. Regensb. 3
(1841) 285; in DC. Prod. 14 (1857) 540, incl. B
montana MEISN. and y concolor MEIsn.; Mia. FI.
Ind. Bat. 1, 1 (1858) 877; Kurz, For. Fl. Burm.
2 (1877) 333; Hook. f. Fl. Br. Ind. 5 (1886) 194;
Boer. Handl. 3 (1900) 111; RipL. Fl. Mal. Pen.
3 (1924) 144.—Eriosolena montana BL. Bijdr.
(1826) 651; HaAssx. Cat. Hort. Bog. (1844) 92,
incl. « macrophylla Hassk. and f minor HAssk.;
ZoLi. Nat. Geneesk. Arch. N.I. 1 (1844) 616;
Syst. Verz. 2 (1854) 116.—D. montana MEISN.
Denkschr. K. Bayer. Bot. Ges. Regensb. 3 (1841)
284.—Eriosolena composita VAN TIEGH. Ann. Sc.
Nat. Bot. VII, 17 (1893) 196; Bull. Soc. Bot. Fr.
40 (1893) 68; Merr. Contr. Arn. Arb. 8 (1934)
111.—Eriosolena pendula BL. ex LrEcomTE, Not.
Syst. 3 (1914) 101.—Fig. 14c-g.

Shrub or small tree up to 10 m by 16 cm.
Leaves chartaceous to subcoriaceous, usually
brownish above and glaucous beneath when dry,
elliptic-oblong to lanceolate, (314—)7—14(—-20) by
(114-)2-5 cm; base attenuate; apex acuminate;
nerves 9-14 pairs, distinct and elevated beneath,
visible or obscure above, sometimes distinct on
both surfaces; petiole 3-5 mm. Inflorescences
axillary, solitary or very rarely 2 inflorescences in
an axil (cf. KING’s coll. 6940); involucral bracts 2,
ovate-oblong to oblong, 1-114 cm long, minutely
pubescent outside; peduncle 214-6 % cm, usually
nodding, with several small linear bracts at the
base, (4-)7—12-flowered. Flowers light-yellowish
or white, fragrant, 10-15 mm long, sessile, densely
covered with appressed, golden-yellowish or whit-
ish hairs outside. Calyx lobes convolute, 2 longer
and 2 shorter, lanceolate or ovate-oblong, rarely
oblong, 2-4 by 1 mm. Stamens sessile or with short
filaments; anthers linear, 1-114 mm long. Ovary
ellipsoid, 114-2 mm, densely hairy; style c. 114
mm; stigma globose. Fruits ellipsoid or ovoid,
10-15 by 5 mm, black (BACKER s.n.) or red
(BACKER 14479).

Distr. India, Burma (Southern Shan States,
Tenasserim and Tounghoo), Indo-China (Annam),
China (Yunnan), and Malaysia: Sumatra, Malay
Peninsula, Borneo, and West Java.

Ecol. In rain-forests (900—1000—)1200—2000 m.

Use. The bark is used as binding material.

Vern. Kakapasan (also used for Phaleria),
kémanden, S, ki-salam, J; Sum.: kulei manis rimbo.

Note. Daphne javanica THUNB. (1806, /.c.) is a
nomen nudum. There is a specimen under that
name bearing THUNBERG’s handwriting in his her-
barium at Uppsala, kindly sent on loan from there.

This specimen might have come from Java as the
epithet indicated (cf. also Wixstr. Kongl. Vet.
Acad. Handl. 1818, 297). It is distinctly D. com-
posita. On the back of the sheet at the upper left
corner is written in THUNBERG’s handwriting
“e Ceilona. Thunberg’’, for this reason the species
has also been listed in THUNB. FI. Ceilan. (1825) 5.
However, the genus does not occur in Ceylon and
the record of this specimen from Ceylon is ap-
parently an error.

2. Daphne luzonica C.B. Ros. Bull. Torr. Bot. Cl.
35 (1908) 72, 75; MerR. Philip. J. Sc. 5 (1910) Bot.
366; En. Philip. 3 (1923) 132; STEEN. Bull. Jard.
Bot. Btzg 13 (1934) 254.—Fig. 14a-b.

Slender shrub up to 1144 m. Branchlets light
brown to reddish-brown, glabrous. Leaves charta-
ceous to subcoriaceous, glabrous on both surfaces,
narrow elliptic-oblong, 8-9 by 214-214 cm; base
cuneate to attenuate; apex acuminate; nerves 6-8
pairs, distinct beneath, obscure above, curved as-
cending; petiole almost absent, up to 3 mm.
Inflorescences sessile or up to 3 mm peduncled,
few- to many-flowered; pedicels 1 or 2 mm. Buds
very acute. Flowers c. 10 mm long, articulated at
the base. Calyx lobes ovate, 244-3 mm by 2 mm,
very acute. Stamens subsessile, or with a very short
filament; anthers 114-2 mm long. Disk ring-like.
Ovary ellipsoid-oblong, 3 by 1 mm, glabrous;
style obscure; stigma globose. Fruit (young) ovoid,
7 by 4 mm.

Distr. Malaysia:
Benguet Prov.).

Ecol. In the mossy forest on the higher moun-
tains, 2000-2500 m.

Note. The leaves are similar to those of D.
odora THUNB., from which it differs by obviously
smaller flowers, absence of bracts below the umbel,
and absence of the typical yellow tomentum on
peduncle and pedicels. From D. kiusiana Mia. it
differs in the glabrous flowers, from the Formosan
D. arisanensis HAYATA by the absence of floral
bracts, the very acute calyx lobes, shorter pedicels,
longer anthers, and larger calyx tube.

Though MERRILL (1908, /.c.) suggested that
there is a closely allied form in Yunnan, I have not
succeeded in identifying it with a Chinese species.

Excluded

Daphne decandra BL. Bijdr. (1825) 650 is according
to SLEUMER (FI. Mal. I, 5, 1954, 91) = Casearia
velutina BL. (Flacourt.) (cf. also Miq. Fl. Ind. Bat.
1, 1, 1858, 709).

Eriosolena affinis ZOLL. Syst. Verz. 2 (1854) 116.
I have seen two sheets, one from Paris, with
ZOLLINGER’s Own handwriting and the type num-
ber 3209, collected in Lombok, and one at Leyden.
MIQUEL (FI. Ind. Bat. 1, 1, 1858, 878) already had
suggested it to represent a Rubiaceae and I have to
thank Dr BAKHUIZEN VAN DEN Brink for the final
reduction to Antirrhoea hexasperma (ROXxB.) MERR.
En. Philip. 3 (1923) 540 (Rubiaceae).

Philippines (N. Luzon:

Dec. 1960] THYMELAEACEAE (Ding Hou) —‘siséiSDYd

7. GYRINOPS

GAERTN. Fruct. 2 (1791) 276, t.140 f.6; DomKE, Bibl. Bot. 111 (1934) 119, map 2;
Quis. J. Arn. Arb. 27 (1946) 404.—Lachnolepis Mig. Ann. Mus. Bot. Lugd.-Bat.
1 (1863) 132.—Brachythalamus GILG, Bot. Jahrb. 28 (1900) 146.—Aquilaria sect.
Brachythalamus HALL. f. Med. Riksherb. n. 44 (1922) 19.—Aquilaria sect.
Gyrinops HALL. f. l.c.—Aquilaria sect. Lachnolepis HALL. f. l.c.—Fig. 15.
Trees or shrubs. Leaves spirally arranged, usually with distinctly parallel veins
joining the several intramarginal veins; margin thickened. /nflorescences 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 glabroiis. 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, linear, 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 Trichadenia (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. 2'4 times as long as wide. Flowers in an umbel.
3. Nerves prominent and spaced beneath, 16-20 pairs. Infructescences 12—14-flowered. Pedicels c.
2mm. Petaloid appendages united in a ring. Fruits ovoid-oblong, c. 2214 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. 17/4 mm
long. Valves #/,—4/; mm thick at the suture. . . 3. G. ledermannii
Floral tube cupular, 2-5 mm long. Fruits emerging from the top ‘of the intact floral tube.
a Leaves usually narrow-lanceolate, 114-10 by '/;-1 cm. Petaloid appendages oblong, as long as
the stamens . . . . 4. G. salicifolia
4. Leaves elliptic- -oblong or ovate- oblong, very rarely lanceolate, ae 15 by 114-5 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 (MiQq.) Batt. Adansonia Aguilaria moluccana HA. f. Med. Rijksherb.
11 (1875) 326; Gite in E. & P. Pfil. Fam. 3, 6a on. 44 (1922) 19.—Fig. 15a-d.

(1894) 225; BoerL. Handl. 3 (1900) 111; Quis. J. Shrub. Leaves chartaceous, glabrous, oblong-
Arn. Arb. 27 (1946) 404.—Lachnolepis moluccana lanceolate, (8—)18—24 by (114-)2-3 cm; base ob-
Mia. Ann, Mus. Bot. Lugd.—Bat. 1 (1863) 132.— __ tuse; apex acuminate; nerves 23-32 pairs, slightly

40 FLORA MALESIANA [ser: I, volagae

Fig. 15. Gyrinops moluccana (MiQ.) BAILL. a. Habit, x 24, b. opened flower, one anther removed, x 3,
c. attachment of stamen, < 7, c’. stamen, x 7, d. fruit bulging out of floral tube, x 2.—G. versteegii
(GitG) DoMKE. e. Opened flower, x 5.—G. podocarpus (GILG) Domke. f. Opened flower, * 5, g. frontal
and lateral view of fruit protruding from floral tube, x 2.—G. decipiens DiNG Hou. h. Habit, x %,
i. dehisced fruit bulging out of floral tube, x 2, j. opened floral tube of fig. i, x 3.—G. salicifolia RIDL.
k. Habit, x 24.—G. caudata (G1LG) Domxe. /. Habit, x 24, m. opened flower with characteristic long
pedicel, x 5, n. young fruit protruding from floral tube, * 2 (a—d DE VRiESE & TEYSMANN S.n., @ VER-
STEEG 1381, f-g PLEYTE 567, h—j KJELLBERG 889, K KANEHIRA & HATUSIMA 12443, /-m BEccCARI PP 911).

Dec. 1960]

_THYMELAEACEAE (Ding Hou) fi 41

curved and ascending, at c. 60° to the midrib,
distinct or visible beneath, indistinct above; veins
+ parallel. Inflorescences axillary, sometimes on
the branches, simple, rarely branched, 3—S-flower-
ed; peduncle almost none up to c. 10 mm, some-
times 2 or 3 in an axil; pedicels c. 4 mm. Flowers
long-tubular, c. 15 mm. Calyx lobes oblong, c.
3 mm long, erect, the apex slightly incurved.
Petaloid appendages usually united behind the
stamens, with hairs almost as long as themselves.
Stamens inserted slightly below the appendages,
sometimes the lower part of the anthers adnate
to the tube. Pistil c. 4 mm long. Ovary ovoid or
ellipsoid, gradually narrowed into a distinct style,
pilose; stigma ovoid. Fruits emerging through the
lateral slit of the floral tube, ovoid, shortly stalked,
sparsely pilose, ovoid, 1 '4 by 1 cm, I- or 2-seeded.
Seeds ovoid, with a short and thick appendage
at the base.

Distr. Malaysia: Moluccas (Buru: Kajeli and
Halmaheira), thrice collected. Also cultivated in
Hort. Bog. (from seed of Buru).

Note. One specimen collected from a cultivated
plant in Hort. Bog. has smaller leaves (8-14!4 by

114-214 cm).

Fig. 16. Distribution of Gyrinops GAERTN.

2. Gyrinops decipiens, nov. sp.

Differt ab G. moluccana foliis elliptico- vel sub-
obovato-oblonga, pedunculis brevibus_ validis
apiceque incrassatis, floribus umbellatis, antheris
liberis, pericarpio late suturato. Typus G. KJELL-
BERG 889, Bo, L.—Fig. 15h-j.

Small tree, c. 4m. Leaves chartaceous, glabrous,
rarely sparsely hairy beneath, shining on both
surfaces when dry, elliptic- or slightly obovate-
oblong, 14-17 by 5-7 cm; base narrowly cuneate;
apex shortly acuminate; nerves 16-20 pairs,
slightly curved or obliquely spreading towards the
margin, elevated beneath, visible sometimes ob-
scure above. /nfructescences terminal and axillary,
umbelliform, 12—14-flowered; peduncle very short
to 214 cm, thick, accrescent, knob-like thickened
at the top; pedicels c. 2mm. Flowers long-tubular,
c. 15 mm long. Floral tube almost glabrous inside.
Calyx lobes oblong, 3-4 mm long. Petaloid ap-
pendages united behind the stamens with hairs as
long as themselves. Stamens inserted slightly below
the appendages. Fruits ovoid-oblong, c. 214 by
1144 cm, acuminate to the apex, suture surface c.
3 mm wide. Seeds unknown.

Distr. Malaysia: Central Celebes (Wavatoli,
Palarahi).

Ecol. In rain-forest, at 100 m.

Note. This species is closely related to G.
moluccana by the long tubular flowers with
petaloid appendages united into a ring and fruits
emerging through the lateral slit, but differs from
it by the characters shown in the key.

The leaves are similar to those of Aquilaria
beccariana.

3. Gyrinops ledermannii DomMKE, Notizbl. Berl.—
Dahl. 11 (1932) 349.

Shrub. Leaves subcoriaceous, glabrous except
sparse hairs at the lower parts on both surfaces
and the midrib beneath, oblong- or ovate-lanceo-
late, sometimes obovate-lanceolate, 614-12 by
2!14-5 cm; base rather acute or shortly narrowed
towards the petiole; apex acute or acuminate;
nerves spreading, obsolete, close to each other,
among which c. 25-35 stronger pairs, curved and
ascending towards the apex. IJnfructescences
pseudo-lateral or terminal, subsessile, 2—3-flower-
ed; pedicels thin, 3-5 mm. Floral tube cylindric,
indistinctly ribbed, 13 mm long, |'4 mm in diam.
Calyx lobes ovate, 114-2 by c. 14 mm, the outer
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. 3/; by 4 mm, obtuse, connected at the
base, villose. Stamens sessile, oblong, 1-114 by
1/, mm. Fruits + pyriform, c. 134 by 14 cm (in-
cluding stipe 3 mm and acute and cuspidate apex
4 mm), short pilose, compressed, irregularly,
transversely -+ rugose. Seeds 2 or | by abortion,
c. 9 mm long (including an appendage c. 3 mm
long), woolly.

Distr. Malaysia: New Guinea (Sepik R.,
Station Mt Pfingst: LEDERMANN 7401).

Ecol. Slope in dense, virgin forest, at the foot
of the mountain, 0-200 m altitude.

Note. The type specimen of this species is not
available and no additional material has been
collected in that area. The description above is
extracted from the original one. From DOMKE’s
detailed description, this is a distinct species
characterized by the long floral tubes, distinct
petaloid appendages and the leaves with many
obscure nerves. It is closely allied to G. moluccana
and G. decipiens by the flower with a tubular
floral tube and the fruit emerging from the lateral
slit of it, but differs from both of them by the
characters given in the key.

4. Gyrinops salicifolia Ripit. Trans. Linn. Soc.
Bot. II, 9 (1916) 145.—Gyrinopsis salicifolia Quis.
J. Arn. Arb. 27 (1946) 407.—Fig. 15k.

Slender shrub, c. 1 m. Branchlets light brown,
pubescent. Leaves sparsely pubescent on the mid-
rib and sometimes on the nerves and veins
beneath, lanceolate to linear-lanceolate, 114-10
by '/,-1 cm; base cuneate; apex acuminate and
pointed; nerves and veins similar and equally
strong, slightly visible beneath, obscure above;
petiole c. 4 mm. Inflorescences terminal, sessile,
3—-5-flowered; pedicel c. 2 mm. Flowers pale

42 FLORA MALESIANA

[ser. ne vol. 6!

yellow, c. 3 mm long. Floral tube c. 24% mm long,
cupular, pilose outside. Calyx lobes oblong, c.
1 mm long, puberulous. Petaloid appendages ob-
long, c. 14 mm long, shortly hairy. Stamens sessi-
le, as long as the petaloid appendages. Disk
obscure, ring-like. Pistil c. 2 mm long, densely
short-hairy. Ovary obovate, 1 mm long; style
filiform, c. 1 mm; stigma obscure. Fruit unknown.

Distr. Malaysia: western New Guinea (Utakwa
and Nabire).

Ecol. In fringing rain-forest, 300 m.

5. Gyrinops caudata (GILG) DoMKE, Notizbl. Berl.—
Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27 (1946)
404.—Brachythalamus caudatus GILG, Bot. Jahrb.
28 (1900) 147; in E. & P. Pfl. Fam. Nachtr. 3
(1903) 238.—Fig. 15]-n.

Shrub or tree up to 17 m by 36 cm, fide BW
6738. Branchlets greyish, whitish pubescent and
glabrescent. Leaves chartaceous, glabrous, dull
beneath and shining above, elliptic-oblong, ovate-
oblong, rarely lanceolate, 6-13 by 114-4 cm; base
cuneate; apex up to 114 cm, acuminate; nerves
and veins scarcely distinguishable, numerous,
parallel, visible beneath, obscure above; petiole c.
3 mm. Jnflorescences axillary or terminal, 3—10-
flowered, sessile and peduncled, peduncle up to
8 mm. Flowers c. 5 mm pedicelled. Floral tube
cupular, 2 mm long. Calyx lobes oblong, 1 mm
long. Petaloid appendages transverse-oblong, c.
14 mm long. Stamens subsessile, slightly longer
than the appendages. Ovary ovoid, densely pilose;
style very short; stigma capitate.

Distr. Malaysia: New Guinea (Sidai and. Mt
Arfak).

Ecol. Primary forest, 5-20 m (fide BW 6738).

Vern. Niwawur, Amberbaken language.

Note. This species is easily recognized by the
pedicel usually c. 2 times as long as the floral tube.

6. Gyrinops versteegii (GILG) DomkeE, Notizbl.
Berl._Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27
(1946) 404.—G. walla (non GAERTN.) Koorb.
Minah. (1898) 577; Boer. Handl. 3 (1900) 111.—
Brachythalamus versteegii G1LG, Nova Guinea 8
(1910) 410.—Aquilaria versteegii HALL. f. Med.
Rijksherb. n. 44 (1922) 19.—G. sp. HALL. f. /.c. 20.
—Fig. 15e.

Shrub up to 6 m, or tree up to 21 m by 65 cm
(fide bb 21394, Bo). Leaves chartaceous to sub-
coriaceous, pubescent, especially on the nerves
and veins beneath, glabrescent or glabrous, dull
and yellowish-brown beneath, shining and reddish-
brown above, elliptic-oblong, ovate-oblong, or
obovate-oblong, 5-14 by 114-5 cm; base cuneate;
apex up to 2 cm narrow-acuminate; nerves and
veins similar, numerous, slightly oblique and par-
allel; petiole short, 3-5 mm. Inflorescences sessile,
usually terminal, consisting of 6 to 8 flowers,
rarely axillary, or on the branchlets; pedicels

1-3 mm. Flowers white, yellowish, light greenish,
or yellowish-green. Floral tube cupular, c. 344 mm
long. Calyx lobes oblong, c. 1 mm long. Petaloid
appendages deltoid, about half as long as the
anther, densely hairy. Stamens sessile, c. ¥, mm.
Disk scale-like. Pistil c. 244 mm long, densely
puberulous, except the stigma. Ovary ovoid, c.
1 mm long, narrowed towards the apex; style ab-
sent; stigma ovoid. Fruits yellow, slightly obovoid
or ellipsoid, 244 by 1 cm, shortly acuminate to the
apex, attenuate to the base. Seeds ovoid, plano-
convex, 9 by 6 mm, with a caruncle-like appendage
at the base, c. 2 mm thick.

Distr. Malaysia: Lesser Sunda Islands (Lom-
bok, Sumbawa, Flores, and Sumba), NE. Celebes
(Minahassa), and West New Guinea (Alkmaar
Bivouac and Somula).

Ecol. In forests, scattered, from the lowland up
to 900 m.

Vern. Kétémunan, Lombok, ruhu wama, Sumba,
seke, Flores.

Note. Closely related to G. podocarpus; more
material is needed to verify whether it deserves
specific distinction.

7. Gyrinops podocarpus (GILG) DoMKE, Notizbl.
Berl.—Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27
(1946) 404.—Brachythalamus podocarpus GILG,
Bot. Jahrb. 28 (1900) 146; in E. & P. Pfi. Fam.
Nachtr. 3 (1908) 238.—Aquilaria podocarpus HALL.
f. Med. Rijksherb. n. 44 (1922) 19; Domke, Bibl.
Bot. 111 (1934) t. 2 f. 10.—G. ledermannii (non
DomMKE) Merr. & PERRY, J. Arn. Arb. 22 (1941)
264.—Fig. 15f-g.

Slender shrub, 14-2 m. Leaves chartaceous,
pubescent beneath especially on the nerves and
veins, glabrescent or sometimes glabrous, glabrous
above, elliptic-oblong, narrow-oblong, slightly
obovate-oblong, 10-15 by 3—5 cm, base cuneate,
apex up to 2!4 cm acuminate; nerves 25—40 pairs,
distinct beneath, obscure above; veins and veinlets
visible beneath, indistinct above, sometimes nerves
and veins similar. Inflorescences terminal or
axillary, sessile or with a short peduncle up to
6 mm, 2—6-flowered; pedicels 2-3 mm, pubescent.
Flowers white, 4-5 mm long. Floral tube cupular.
Calyx lobes ovate, 1-114 mm long. Petaloid ap-
pendages deltoid or slightly oblong, 4-4 mm
long, densely whitish hairy. Stamens sessile. Disk
shortly cup-shaped, crenate. Pistil shortly stipitate,
densely hairy except the stigma, c. 444 mm long.
Ovary oblanceolate; style distinct, c. 14% mm;
stigma capitate. Fruit (young) green, pyriform,
15 by 6 mm, densely puberulous, acute to the apex
and crowned by the persistent, curved style, stipe-
like, cuneate towards the base.

Distr. Malaysia: West New Guinea (Ramoi,
Sorong, Monep, and Idenburg R.).

Ecol. In primary forests, from the lowland up
to 750 m.

Vern. Kokkoree, Asmat language.

Mec 960) sy SC THYMELABACEAE (Digg Hou) = a a

8. DRAPETES

BANKS ex LAMK, J. Hist. Nat. Paris 1 (1792) 188, t.10 f.la—d; PERSOON, Syn. |
(1805) 148; GAERTN. Fruct. 3 (1807) 199, t. 215; ENDL. Gen. Pl. (1837) 33; Suppl.
4 (1847) 61; Bentn. FI. Austr. 6 (1873) 35; BENTH. & Hook. f. Gen. Pl. 3 (1880)
196; DomkKE, Bibl. Bot. 111 (1934) 138, map 17.—Kelleria ENDL. Gen. Pl. Suppl.
4 (1847) 61; Domke, Bibl. Bot. 111 (1934) 137, map 16.—Daphnobryon MEIsN.
in DC. Prod. 14 (1857) 566.—Drapetes sect. Daphnobryon BoeRL. Handl. 3
(1900) 106.—Fig. 17.

Dwarf shrubs with creeping, radiant, or -+ 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
mouth of the tube, consisting of 1-2 episepalous scales (or 0). Stamens 4, free from
the tube at the mouth. alternate 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-celled, pilose or hairy in the upper half or at the apex; style
linear, lateral, usually longer than 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 (/.c.) with only one species, D. muscosus, 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 (/.c.) based a new genus Kelleria on D. dieffenbachii Hook. from New Zealand,
which should differ 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 f. (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 (/.c.) proposed a new genus Daphnobryon for Drapetes ericoides HooK. f. from Borneo
and D. tasmanicus Hook. f. 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 different 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 (/.c.) 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 (l.c. 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 they

44 FLORA MALESIANA [ser. 1; voleape

are distinct or confluent into a single entire or notched one, is not constant, as has been pointed out by
BENTHAM (/.c. 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 flower! Because of the
great resemblance in habit and the number of stamens being the same as that of the perianth lobes,
BENTHAM (/.c. 36) and BENTHAM & Hooker f. (Gen. Pl. 3, 1880, 196) reduced Kelleria and Daphnobryon
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. f. Ic. Pl. (1852) t. 895;
Stapr, Trans. Linn. Soc. II, 4 Bot. (1894) 221;
Boer. Handl. 3 (1900) 111; Gisss, J. Linn. Soc.
Bot. 42 (1914) 132; Merr. En. Born. (1921) 417. —
Daphnobryon ericoides MEIsN. in DC. Prod. 14
(1857) 566; Mia. Fl. Ind. Bat. 1, 1 (1858) 881.—
Kelleria papuana Domke, Bot. Jahrb. 62 (1929)

484; STEEN. Bull. Jard. Bot. Btzg 13 (1934) 254.—
Kelleria ericoides DomMKE, Bot. Jahrb. 62 (1929)
485; STEEN. Bull. Jard. Bot. Btzg 13 (1934) 254.—
Kelleria patula MERR. & PERRY, J. Arn. Arb. 22
(1941) 267.—Fig. 17.

Stem reddish- or dark-brown, up to 50 cm;
young branchlets villous, glabrescent. Leaves 3—5

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

by 24 mm. Inflorescences (1-)4—9-flowered; pedi-
cels short, c. 14 mm. Flowers white, cylindric,
slightly expanded towards the apex, 3-5 mm long,
12-costate. Calyx lobes ovate, obtuse, 1-144 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 mm long, 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.

Ecol. Alpine plant, on dry grasslands, shallow

Fig. 17. Drapetes ericoides Hook. f.. a. Habit,

nat. size, b. flower, x 6, c. opened flower, pistil
removed, x 6, d. pistil, x 6, e. stamens, x 13,
f. stem showing leaf-scars, x 6, g. leaf, x 6

soil over rocks on sheltered grasslands, plentiful
on sandy banks of streams, and in cracks of
granite, usually occurring from 3000 to over

(RosBBINS 315). 4000 m.

9. PIMELEA

BANKS & SOLAND. ex GAERTN. Fruct. 1 (1788) 186, nom. gen. cons.; MEISN. in
DC. Prod. 14 (1857) 496; Bent. FI. Austr. 6 (1873) 1; Domxe, Bibl. Bot. 111
(1934) 138, map 18, non Banksia Forst. 1776.—Thecanthes WiksTrR. 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 24, b. inflorescence, 2, c. longitudinal section of
inflorescence, x 2, d. flower, x 4, e. opened flower, x 4, f. stamen, 13, g. seed, x 13.—P. concreta

F. v. M. A. Inflorescence, x 2, i. longitudinal section of inflorescence, x 2, j. flower, x 4, k. pistil, x 4
(a WOMERSLEY & VAN ROYEN 5853, b—g WOMERSLEY 8767, h—k VAN STEENIS 3304).

Herbs, shrubs or undershrubs (extra-Mal. spp.). Leaves opposite or decussate,
sometimes subopposite, or alternate. /nflorescences terminal or axillary (extra-
Mal. spp.), capitate, rarely spicate or solitary (extra-Mal. spp.); involucral bracts
partially united, or free (extra-Mal. spp.), 4, or in extra-Mal. spp. 4-6(—8) or more,
sometimes wanting. Flowers bisexual, sometimes unisexual by abortion or
dioecious (extra-Mal. spp.), at least in Mal. spp. developing centripetally. Floral
tube tubular, often silky villous outside, in Mal. 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, filiform or

46 FLORA MALESIANA

[ser. I, voltae

club-shaped lobes or wanting (extra-Mal. 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.,

to Malaysia.
Ecol.
to 1000 m.

two of which extend

In Malaysia in grassland and savannahs in the seasonal areas, mostly at low altitude, ascending

Note. The two Malaysian species belong to subg. Thecanthes (WIKSTR.) MEISN. ex GILG (in E. & P. Pfl.

7°

Fam. 3, 6a, 1894, 243); in the text “‘Untergen.
DALLA ToRRE & HARMS p. 340).

was erroneously printed as “‘Unterfam.

”. cf. also DE

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 ‘invelueral bracts. Calyx lobes

usually erect. Stamens included; filaments almost as long as the anthers.

1. Pimelea concreta F. v. M. Fragm. 5 (1865) 73;
BENTH. FI. Austr. 6 (1873) 6.—P. brevituba Fawc.
in Forbes, Wand. (1885) 516; Gitc in E. & P.
Pfil. Fam. 3, 6a (1894) 243, f. 84, D, E.—P. sp.
DAMMERMAN, Nat. Tijd. N.I. 86 (1926) 45, f. 1.—
Fig. 19h-k, 20.

Fig. 20. Pimelea concreta F. v. M. Sumba
(Photogr. DE Voocp).

2. P. cornucopiae

Annual, simple or branched, up to 50 cm,
glabrous throughout. Leaves membranous, nar-
rowly oblong or oblong-lanceolate, or obovate-
oblong, 134-314 cm by 4-8 mm; base acute to
obtuse; apex acuminate and minutely apiculate;
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 mm long,
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 414(-7) cm; pedicels flat, usually dilated
at the base, very short to 114 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, 2-214 mm long. Disk scale-like,
small, c. 14 mm long. Stamens 214-314 mm. Ovary
ovoid, 1-114 mm long; style exserted; stigma sub-
capitate. Fruits ovoid, 2-4 by 1 mm, short-stalked.

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. Ros. Philip. J. Sc. 6 (1911)
Bot. 345; Merr. En. Philip. 3 (1923) 134.—
Fig. 19a-g.

Annual up to 50 cm tall. Leaves membranous,

Dec. 1960]

ay

lanceolate, narrowly oblong, rarely obovate-
lanceolate, (114-)2-314 cm by 114-6(-714) mm;
apex acuminate; base obtuse; nerves obscure,
3-6 pairs; petiole very short, c. | mm. J/nflores-
cences terminal, 15—40-flowered; peduncles vari-
able in length, very short, or up to 514 cm; pedi-
cels 1-5 mm, flat but not dilated at the base,
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 214—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 mm long. Disk
small and obscure. Stamens 1-114 mm, slightly
shorter than the calyx lobes. Ovary ovoid, 1144 mm
long; style slightly protruding beyond the floral
tube; stigma small, globose. Fruits ellipsoid, 3 by
114 mn, slightly stiped.

Distr. Australia (Queensland), D’Entrecas-
teaux Is. (Fergusson I.), New Britain, Louisiade

Archipelago (Misima I. and Sudest I.), and
Malaysia: Philippines (extreme N of Luzon,
Cagayan Prov. near Sanchez Mira, once collected,
B.S. 7410, US) and New Guinea (Western and
Central Divisions). Fig. 21.

Ecol. Common on savannah ridges and grass-
lands, from the lowland rarely up to 570 m.

Fig. 21. Localities of Pimelea concreta F. v. M.
(+) and P. cornucopiae VAHL. (®) in Malaysia;
both species also occur in Australia.

10. AMYXA
Cf. Atry SHAW, FI. Mal. I, 4 (1953) 363, f.5.—Fig. 22.

1. Amyxa pluricornis (RADLK.) DoMKE, Bibl. Bot.
111 (1934) 116; Atry SHAw, FI. Mal. I, 4 (1953)
363.—Fig. 22.

SHAw questioned whether the fruit of this small
Bornean tree is dehiscent. Judging from fruiting
material collected in Sept. 1958 by M. JAcoss
(n. 5376) near Belaga, in the Third Division of
Sarawak, this seems indeed to be the case, though
the spontaneous dehiscence of the almost mature
fruit happened during the process of drying the
specimens. But the dehiscence of the valves ap-
pears so regular along distinct sutures that there
seems to be hardly any doubt that dehiscence will

also take place in nature. The seed colours are
marked: fruit green, softish, seed dark glossy
brown, funicle partly very fleshy, aril-like, white.
How these colours will be at complete maturity is
unknown; they may change at the last moment.
Tropical fruits are often devoured by animals
before maturity. When mature the attractive seed
with its contrasting colours will dangie out of the
dehisced capsule hanging on the thin basal part
of the funicle from the apex of the valve; it is
likely to be dispersed by animals.

Distr. Now also twice collected by KOsTER-
MANS in West Kutai, East Borneo.

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 (JAcosBs 5376).

48 FLORA MALESIANA

Excluded and Doubtful

Gnidia oppositifolia (non L.) BLANCO, FI. Filip. (1837) 299, and Gnidia? philippinica MEIsN. in DC.

Prod. 14 (1857) 592 are according to ExeLt (FI. Mal. I, 4, 1954, 555) = Terminalia polyantha Pr.
(Combret.).

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

H. glomerata JUNGH. /.c. 316.—Japan.

H. umbellata JUNGH. /.c. 316.—Japan.

No type specimen of either of these species has been as yet located. Dr VAN STEENIS and Dr HATUSIMA
(in litt.) could not clarify them from the description. They might not be natives of Japan (cf. BENTH. &
Hook. f. Gen. Pl. 3, 1880, 188-189). Mr Airy SHAW (inlitt.) assumes them to be Lauraceous.

Passerina javanica THUNB. FI. Jav. 2 (1825) 19. The type specimen could not be found in THUNBERG’s
herbarium at Uppsala (cf. BACKER c.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 Euscaphis 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, efc.

LINDLEY (1835) kept it as a separate family in the Hypericineae together with Cunoniaceae.
HALLER f. has repeatedly stressed their close affinity to the Cunoniaceae (Uber Juliania, etc.
1908, 74, 116, 182; Arch. Néerl. Sc. Ex. Nat. III 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 differences 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 differentiates 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 + 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
fs t0, 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. Gén. 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 (Dict. Suppl. 10, 1959, 213)

50 FLORA MALESIANA [ser. I, volLog

Fig. 1. Details of Malaysian species of Turpinia.—T. borneensis (MERR. & PERRY) VAN DER LINDEN.
a. Habit, x 14, b. open nerves and venation, x 1/4, c. flower, x 5,d. ditto, insection, < 5, e. fruit with
cross-section, nat. size, f. stamen, * 10.—T. montana (BL.) Kurz. g. Leaf showing its closed nervation,
x \y,h-i. stamens, * 10.—T. pomifera (RoxB.) DC. j-k. Fruit and its section, nat. size.—T. sphaerocarpa
HAssk. /—-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,
f CLEMENS 30070, b, e CLEMENS 28840, i KERR 2527, j-k LAM 2855, 0 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
these pairs must have been the tropics from where their ancestors have been branched off with a
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.

Vegetatively Turpinia shows a marked, structural resemblance with Sambucus, both possessing
decussate, simply-pinnate, frequently herbaceous, toothed leaves, with stipules and gland-like
stipels (though stipules are not always well represented in the latter genus and some Turpinias
have simple leaves). Properly the sympetaly and inferior ovary of Sambucus 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. TURPINIA

VENT. Choix (1803) 31, t.31, nom. gen. consery. prop.; MERRILL & PERRY, J. Arn.
Arb. 22 (1941) 543; J. Krause!, in E. & P. Pfl. Fam. ed. 2, 20b (1942) 306;
BAKH., V. D. LINDEN & VAN 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. Pl. 5 (1874) 342, 343,
excl. Euscaphis; MAZzA, Dicc. Bot. Nom. Vulg. Cub. & Puerto-Rig. (1889) 15; cf.
also O.K. Rev. Gen. Pl. 1 (1891) 148.—Dalrympelea Roxs. [Hort. Beng. 1814, 17,
nomen, ‘Dalrympelia’ | P|. Corom. 3 (1820) 76, t.279.—Ochranthe LINDL. Bot. Reg. 8
(1836) t.1819.—Hasskarlia MEISN. Pl. Vasc. Gen. 2 (1843) 348; cf. WALP. Ann. 1
(1849) 753.—Maurocenia § Triceros O.K. Rev. Gen. 1 (1891) 149.—Kaernbachia
SCHLTR, Bot. Jahrb. 52 (1914) 151, nom. illeg., non O. KUNTZE, Rey. Gen. | (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. 1.

Evergreen 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, ear/y caducous
(except in 7. 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 | 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) Krause, /.c., has cited all subsequent different spellings of Dalrympelea, 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: Levoleige

sepalous, equal; filaments linear, gradually widened to the base, (in Mal. spp.)
glabrous, inserted close to the disk, caducous; anthers rounded or ovate, with
spreading cell-bases, dorsifixed, dehiscing lengthwise, introrse, sometimes dis-
tinctly apiculate. Disk annular, glabrous and crenate, fleshy. Ovary superior,
(2-)3(-4)-celled, the three cells and styles closely appressed but not connate, the
combined stigmas 3-lobed. Ovules 1-o in each cell, anatropous, attached on the
dissepiment very close to the axis, in 2 vertical rows. Fruit up to 21/4 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-co 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 Staphylea.

Some species have a wide distribution, for example T. montana (BL.) KURZ, from Java and Sumatra
northward to Indo-China, Hongkong, and Yunnan, and 7. pomifera (RoxsB.) DC. in SE. Asia and
Malaysia (rare) where it has often been confused with 7. sphaerocarpa 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 Turpinias 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 other 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. LiertTINcK. LORZING repeatedly noted the flowers of 7. sphaerocarpa to be sweet-scented.

Morph. Stipules. MERRILL & PERRY mention in their key (/.c. 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 7. montana has always 2, T. pentandra 4(-S), and T. brachy-
petala 8(—7). In T. sphaerocarpa 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 7. 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. 7. stipulacea 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 7. simplicifolia.

(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 smail 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 7. montana; two leaf galls caused by gall midges and
a witches’ broom caused by a gall mite on J. 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 different 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 Turpinia has been tried out as a pioneer
for reafforestation on devastated mountain slopes in Central Java (Pangentjongan on Mt Galunggung
and Klédung on Mt Sindoro) which were promising, according to KOoRDERS’s report.

Wood Anat. DEN BERGER, Meded. Proefstat. Boschw. 13 (1926) 95, Determinatietabel houtsoorten
van Malesié, Veenman, Wageningen (1949) 30; Descu, Mal. For. Rec. 15? (1954) 524 (hand lens);
Hemmscu, Lilloa 8 (1942) 163; METCALFE & CHALK, Anat. Dic. 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
/.c. recognized the large differences between the two families; HEIMSCH /.c. p. 182, 189 (erroneously cited
by METCALFE & CHALK /.c. p. 446) suggested affinity to Ce/astrales but critical studies are necessary;
DescuH /.c. 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 insufficient basis: the types of T. parviflora, T. simplicifolia, T. unifoliata, and of T. laxiflora were
described without fruit. Admittedly I 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 javanica 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: T-yokas:

KEY TO THE SPECIES

1. Leaves all and always simple, lanceolate-oblong, 10-15 by 314-514 cm, on both sides with prominent
venation. Sepals and petals c. 244-3 mm long. Filaments up to 2 mm. Ovules 2 per cell.
1. T. simplicifolia
1. Leaves 1-cv-jugate, the upper ones under the inflorescence exceptionally 1-foliolate.

2. Stipules large, ovate-triangular, c. 1% cm long, persistent, stout (fig. 1p). Leaflets (7—)9, rather thin,
elliptic, c. 9 by 5 cm. Venation on both sides fine-reticulate (tending to be tessellate) and prominent
(in the herb.). Petals c. 214-3 mm long. Stamens c. 214 mm long; anthers ?/, 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. 1-114 cm diam.;
pericarp c. 14-1 mm 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. 114-214 mm long; anthers c. 14 mm). (Mature fruits not crowned by
3+ distinct horns).
5. Pericarp (mostly much) thinner than 1 mm (fig. le). Stamens 114-2 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 !/;-'’4 mm diam.. Nerves
patent and + straight, connected by regular loops of equal strength and prominence (fig. 1o).
Leaflets mostly thin and herbaceous (mostly pale green) in the herb., rather narrow-elliptic
oblong, c. 5—10(—16) by 1!4—5(—7) cm. Inflorescences delicate . . . 4. T. montana

6. Stipules c. 444-9 mm long, glabrous, entire. Ovules 4-8 per cell. Fruit c. 8 15 mm diam. Testa
14-1 mm 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.
Anthers distinctly apiculate, c. 14 mm long. Ovules 4 in each cell. Seeds large, c. 9 mm averagely;
testa 4/,-1 mm diam.. Leaflets mostly sped widest below the middle, often towards the
base: 2 . . 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 134-2 mm. Anthers not apiculate, c. 44-14 mm
long. Ovules 6-7-8 in each cell. Seeds c. 5 mm, testa 14 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. 11). 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. 14-1 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 34 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 4-814 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. 14-34 mm diam..Ovules 4(—5) per cell. Testa
34-1 mm diam. . . . . . §. 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 S—7, prominent below, spreading, ascending from the base, not dis-
tinctly looped, chartaceous. Anthers 34 or 1-114 mm, not (or slightly) apiculate. Ovules 5 or
6(-8) per cell. Testa 0.3-0.4 mm diam. 52 eo . . . 9. T. ovalifolia

8. Venation in the herbarium not distinctly prominent at both sides.
10. Pericarp thick, at least 5 mm diam. ne 1j). 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.

. Ovules 4(—5) per cell. Fruit with 3 distinct horns, at least when young. Leaflets 214-414
(-614) cm wide. . . 11. T. pentandra

. Ovules 8(—7) per cell. Fruit without ‘style horns. Leaflets GB iyi asaee 8 cm wide.
12. T. brachypetala

Dec. 1960]

STAPHYLEACEAE (van der Linden)

»D

1. Turpinia simplicifolia Merr. Philip. J. Sc. 27
(1925) 34.—T. unifoliata MERR. & CHUN, Sunyat-
senia 2 (1934) 37.

Tree up to 4 m. Leaves simple, lanceolate-
oblong to oblong-elliptic, glabrous, acuminate,
decurrent at the base, slightly dentate, 10-15 by
314-6 cm; petiole up to 4 cm long, glabrous.
Stipules small, glabrous, 2-3 mm long. Panicles
axillary, up to 18 cm long. Sepals elliptic, rounded,
scarcely ciliate, c. 244 mm long. Petals oblong-
elliptic, ciliate, 2!4,—3 mm long. Stamens c. 214 mm
long; filaments 2 mm. Ovary 3-celled, each cell
with 2 ovules. Fruit globular, c. 8 cm diam.,
without style-horns; pericarp very thin, 0.1—0.2
mm thick.

Distr: Hainan; in
(Luzon), thrice collected.

Ecol. In forests.

Notes. The type (LoHER 12992) bears an in-
florescence and slightly immature foliage. In
comparing the characters it seems not specifically
different from that of T. unifoliata MERR. & CHUN
(S. P. Ko 52249, NY) from Hainan, which has
more mature leaves but an inflorescence which is
in bud; the sizes of the floral parts (stamens) are
obviously for this reason somewhat smaller than
those of 7. simplicifolia.

Another (fruiting) specimen from Hainan (F. C.
How 73403, SING) was preliminarily referred by
MERRILL to the same species, but from its venation
and leaf-shape I conclude this to represent T.
indochinensis MERR. This seems to be extremely
close to 7. formosana NAKAI which has only 4
ovules per cell, T. indochinensis having 8. It might
probably be better to reduce the latter as a variety
to T. formosana.

MASAMUNE (FI. Kainant. 1943, 178) reduced
T. unifoliata to T. formosana NAKAI, J. Arn. Arb.
5 (1924) 80. In my opinion this is a different
species by possessing 4 ovules per cell, larger
flowers (petals 4 mm), lesser nerves (7—9 pairs),
and hairy stamens. I have not seen the type
(WILSON 10130), but TANAKA & SHIMADA 13554
and 11180, which agree with the description of
T. formosana.

The leaves are astonishingly resembling those
of certain specimens of 7. borneensis, but this
species has 5 leaflets, petals less than 2 mm long,
and 4 ovules per cell.

Its closest alliance is, as MERRILL & CHUN
already remarked, not with other 1-foliolate
species (such as 7. arguta), but with T. nepalensis
from which it differs by a more regular nervation,
finer inflorescence, 2 ovules per cell, and simple
leaves.

Malaysia: Philippines

2. Turpinia stipulacea v. Dp. LINDEN, nov. sp.—
Fig. 1p.

A speciebus omnibus differt stipulis maximis
persistentibus lignescentibus. Folia 4—5-jugata,
foliolis 6-10 * 244-5 cm. Inflorescentiae robustae,
usque ad 25 cm longae. Sepala 2-2\4 «x 114-2
mm. Petala 2,-3 mm longa. Stamina c. 2% mm
longa, antheris suborbicularibus subapiculatis.
Ovula 4 in utroque loculo. T: SF 27516 Carr, Sing.

Tree c. 18 m high. Leaves 4-5 jugate; leaflets
ovate, glabrous, acuminate, rounded at the base,
margin denticulate, 6-10 by 2!4-5!4 cm; petiole
up to 8 cm; petiolules of the lateral leaflets 3-7 mm.
Stipules large, 17 by 10 mm, persistent, woody,
glabrous. Panicles axillary up to 25 cm long;
pedicels 1-2 mm long. Sepals ovate-oblong, green,
suffused red down to the middle of the segments,
2-244 by 114-244 mm. Petals obovate-oblong,
creamy-yellow, 214-3 mm long. Stamens c. 21/4
mm long; filaments c. 2 mm; anthers round,
slightly apiculate, */; mm long. Ovary 3-celled,
with 4 ovules per cell. Fruit unknown.

Distr. Malaysia: North Borneo (Mt Kinabalu,
near Tibabah R.), once collected, 2100 m; f/. June.

Notes. A remarkable species deviating from all
others by its obviously persistent, very large
stipules. The large number of leaflets and the 4
ovules per cell remind of the Papuan T. pentandra,
but this species has another venation and much
larger anthers. It is possibly more allied to T.
borneensis with which it shares the prominent
venation and 4 ovules, but this species has nar-
rower, more pointed leaf-shape, smaller flowers,
and finer, shorter inflorescences, besides lacking
the unique stipules.

3. Turpinia grandis v. Dp. LINDEN, nov. sp.

Arbor 10-18 m alta. Internodia ultima, petioli et
infructescentiae indumento denso brevi muniti.
Folia 1-3 foliolata, foliolis magnis grosse-serratis,
15—23 * 10-16 cm, venulis utrinque prominentibus.
Infructescentiae 30 cm longae. Fructus globosus
ecorniculatus, 1-1\% cm diam., pericarpio c. 4-1
mm crasso. Semina flavescenti-brunnea, 10-18 in
fructo singulo. T.: ENDERT 4669, L.

Tree 10-18 m. Upper internodes, stipules,
petioles, and infructescences with a very short
(puberulous) but continuous indument. Leaves
1-3-foliolate; leaflets ovate to ovate-oblong,
acuminate, rounded at the base, sharply dentate,
15-23 by 10-16 cm; venation on both sides dense
and prominent; petiole up to 7 cm long; petiolules
of the lateral leaflets 14-114 cm. Stipules small, c.
5 mm long. Flowers unknown. Infructescences
axillary, sub-terminal or terminal, up to 30 cm
long. Fruit globular, without horns, c. 1-114 cm
diam.; pericarp 14-1 mm thick. Seeds pale to pale-
brown, rather large, 10-18 in each fruit.

Distr. Malaysia: Central E. Borneo (W. Kutai:
Kiau; Mt Kemul; Long Petah), thrice collected.

Ecol. On forested river-banks and in deep
ravines, 450-1000 m.

Note. Superficially somewhat resembling 7.
laxiflora but with 1-3 leaflets. Besides, T. /axiflora
has a loose, only slightly prominent venation,
smaller fruits, and sepia-coloured seeds. It is
probably closest allied to T. borneensis, but it
differs from that species by its indument, leaf-size,
and number of ovules.

4. Turpinia montana (BL.) Kurz, J. As. Soc. Beng.
44, ii (1875) 182, x genuina; Koorp. Exk. FI. Java
2 (1912) 528; Atlas Baumarten 1 (1913) t. 92;
FI. Tjib. 2 (1923) 149; Hocur. Candollea 2 (1925)

56 FLORA MALESIANA

[ser. I, vol36*

412, incl. f. arborescens et f. scandens; MERR.
Contr. Arn. Arb. 8 (1934) 93; BACKER, Brittonia 3
(1938) 81; Merr. J. Arn. Arb. 19 (1938) 42; MerRR.
& Perry, J. Arn. Arb. 22 (1941) 552; Back. Bekn.
Fl. Java (em. ed.) 7 (1948) fam. 152, p. 1.—
Zanthoxylum (Xanthoxylum) montanum BL. Bijdr.
(1825) 248; Mia. Fl. Ind. Bat. 1, 2 (1859) 670, ef.
RADLK. Sitz. Ber. K. Bay. Ak. Wiss. 16 (1886)
305-306 (1887).—Zanthoxylum serrulatum BL.
Bijdr. (1825) 249; Mie. Lc.—Triceros cochin-
chinensis (non Lour.) Moritzi, Syst. Verz. (1846)
15.—Maurocenia zollingeri O.K. Rev. Gen. Pl. 1
(1891) 147, 150, cf. BACKER, Brittonia 3 (1938) 81.
—T. parva K. & V. Bijdr. Booms. 9 (1903) 249;
Back. Schoolfl. (1911) 272; Crais, Fl. Siam. En.
1 (1926) 338.—Evodia parviflora CRAIB, Kew Bull.
(1915) 425.—? T. gracilis NAKAI, J. Arn. Arb. 5
(1924) 79.—T. parviflora Craip, Fl. Siam. En. 1
(1931) 339.—Fig. 1g-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 114-714 cm; nerves straight, very
regularly connected by a looped intramarginal
vein; petiolules of the lateral leaflets 14-3 cm.
Stipules c. 3 mm, puberulous, with a short but
sharply bifid apex. Panic/es axillary, open, up to
c. 18 cm long. Sepals ovate, glabrous, ciliate,
1-144 by 114-2 mm. Petals obovate, glabrous,
ciliate, 1-114 by 1144-2 mm. Stamens c. 14% mm
long; filaments 1-114 mm; anthers 14-14 mm, not
or only very slightly apiculate. Ovary (2—)3(-4)-
celled, each cell with 2 ovules. Fruit globular,
sometimes with 3 radial lines from the top, 8-10
mm diam.; pericarp thin, to 4% mm thick; more
than one seed developed; testa 0.2—0.3 mm thick.

Distr. Deccan Peninsula (Pulney Hills), Burma,
Siam, Indo-China, China (Yunnan, Kwantung,
Hainan), Hong Kong, in Malaysia: Sumatra to
Central and W. Java (most frequent in W. Java).

Ecol. In primary montane rain-forest, 750-2300
m. Fl. Oct.—Jan., fr. March—Aug.

Vern. Puhun putag, ki bantjét leutik, S.

Notes. By its thin, very regularly looped nerves
and delicate inflorescence easily recognized. On
the mainland a few specimens have been found
with somewhat thicker, shorter leaves and more
contracted inflorescences (in drier climate?);
venation and ovules are however exactly matching
those of T. montana. This form has been described
as a distinct species T. parviflora CRAIB from Siam
(KERR 2527, K) and is also found in Indo-China
(PIERRE 907, L). The size of the flower parts is
slightly smaller than that in T. montana, but this is
due to the fact that both specimens cited above
are in bud. There is no question that this species
is ever scandent. The number of leaflets may
occasionally be up to 11 (GARRETT 792, Thailand).

A collection from the Pulney Hills (A. SAULIERE
115, Bo, K, L) certainly represents 7. montana;
it has the same venation, glabrous stamens,
constantly 2 ovules per cell, and 3-4 mm long bifid
stipules.

There is a distinct affinity with 7. nepalensis

which, however, has never only 2 ovules per cell.

T. montana var. borneensis MERR. & PERRY
I have found to represent a distinct species under
that epithet.

Whether the Chinese 7. glaberrima MERR.
(Linen. Sc. J. 7, 1931, 312: thid. 14, 1935, 272:
Arn. Arb. 22, 1941, 552) differs from T. montana
is not certain as several numbers MERRILL &
PERRY mentioned to belong to it (TsIANG 2715,
How 71654, 73218, CHUN & Tso 43918) represent
in my opinion 7. montana.

Fig. 2. Demarcation of the Indo-Malaysian part

of the distributional area of Turpinia, showing

also the number of species in each island or island

group; above the hyphen the number of endemic
species, below it other species.

5. Turpinia borneensis (MERR. & PERRY) V. D.
LINDEN, nov. comb.—T. montana var. borneensis
Merr. & Perry, J. Arn. Arb. 22 (1941) 553. T:
CLEMENS 29391 bis.—Fig. 1a-f.

Tree up to 15 m. Leaves 3-—5(-7)-foliolate;
leaflets ovate-oblong to almost lanceolate, glab-
rous, distinctly acuminate, rounded at the base
(in the Philippine specimens sometimes cuneately
decurrent), dentate, 7-17 by 214-6 cm; primary
nerves and the reticulating veins distinctly pro-
minent on both sides; petiolules of the lateral
leaflets c. 14-2 cm long. Stipules c. 5mm, glabrous.
Panicles axillary, open, up to c. 20 cm. Sepals
ovate, ciliate, 2 by 114 mm. Petals obovate, ciliate,
134 by 114 mm. Stamens 134 mm long; filaments c.
114 mm; anthers 14 mm long, apiculate. Ovary
(2-)3-celled, each cell with 4 ovules. Fruit globular,
sometimes with 3 radial lines from the top, 8-15
mm diam.; pericarp thin, 0.1-0.2(—0.5) mm diam.
Seeds large brown; testa 34-1 mm thick.

Distr. Malaysia: Borneo and the Philippines.

Ecol. In rain-forests, up to 1600 m.

Note. MERRILL & PERRY already remarked that
their 7. montana var. borneensis might deserve
specific rank. Its venation, prominent on both
sides, ovate-oblong leaf, its curved nerves, and
4 ovules per cell (in T. montana always 2) remove it
distinctly from 7. montana. The latter is an ex-

Dec: 1960]

STAPHYLEACEAE (van der Linden) ‘ere 57

clusively montane species distributed from SE.
Asia along the Sumatran track to Java.

6. Turpinia laxiflora Ripv. J. Str. Br. R. As. Soc.
n. 82 (1920) 179; Fl. Mal. Pen. 1 (1922) 512;
Merr. & Perry, J. Arn. Arb. 22 (1941) 553.—
T. pomifera var. sphaerocarpa (non HaAssk.)
KiNG; Kina, J. As. Soc. Beng. 65, ii (1896) 453,
pro parte.

Tree, up to 24(-35) m by 50 cm. Leaves 3-7-
foliolate; leaflets ovate-elliptic, glabrous, dentate,
acuminate, rounded at the base, 8-24 by 314-12
cm; petiolules of the lateral leaflets 4-20 mm,
glabrous. Stipules 5-9 mm long, glabrous. Panicles
axillary, lax, long, up to 45 cm, the ultimate
branches minutely puberulous. Sepals ovate,
sparsely ciliate, 114-2 by 1-2 mm. Petals oblong,
distinctly ciliate, 214-214 by 1-144 mm. Stamens
134-2 mm long; filaments 114-134 mm; anthers
1%4,-l4 mm long, roundish, not apiculate. Ovary
3-celled, each cell with (6—)7—8(-9) ovules. Fruit
globular, in dry state mostly wrinkled, 14-1 cm
diam.; pericarp (very) thin, 0.2(-0.9) mm diam.
Seeds several in each fruit; testa c. 4 mm thick.

Distr. Malaysia: N. Sumatra (also Simalur I.)
and Malay Peninsula (Perak: Larut).

Ecol. In rain-forests, at low altitudes, up to
150 m, once at 900 m. 7. /axiflora has been noted
by RIDLEY (n. 6214, coll. a. 1894) to change its
foliage with new flush appearing simultaneously
with young inflorescences. Young leaflets, when
they first appear, are narrow elliptic-lanceolate!
As no later similar data have been reported it
seems premature to conclude that it is deciduous.

Vern. Arilan-buluh, arilan pajé étem, arilan
sitobudlung, (awa)arilan uding, awa mdatdadn nanas,
lahulung, matan-nanas pajo, Simalur, kaju long-
gakan, k. songgak, Sumatra, k. rébung, Pasemah
(Palembang).

Note. Closely allied to 7. sphaerocarpa but
obviously distinct by a different pericarp, larger
inflorescences, and generally wider leaflets which
are brown in dry state.

7. Turpinia sphaerocarpa HaAssx. Flora 25, ii
(1842) Beibl. 1, p. 42; Mig. Fl. Ind. Bat. 1, 2
(1859) 593; Ripv. J. Mal. Br. R. As. Soc. 1 (1923)
58; BAKER f. J. Bot. 62 (1924) Suppl. 24; MeRR. &
Perry, J. Arn. Arb. 22 (1941) 548.—Dalrymplea
javanica Hassk. Pl. Jav. Rar. (1848) 439.—T.
pomifera [non (Roxs.) DC. 1825]: Mia. Sum.
(1861) 201, 513; K. & V. Bijdr. Booms. 9 (1903)
245; Back. Schoolfl. Java (1911) 272; Koorp.
Exk. Fl. Java 2 (1912) 528; Atlas Baumart. 1
(1913) t. 93; Fl. Tjib. 2 (1923) 149; Hocur.
Candollea 2 (1925) 412; BAck. Bekn. FI. Java
(em. ed.) 7 (1948) fam. 152, p. 1; Herne, Pfi.
Kinabalu (1953) 57.—Maurocenia sphaerocarpa
O.K. Rev. Gen. Pl. 1 (1891) 147, 149.—T. pomifera
var. sphaerocarpa KING, J. As. Soc. Beng. 65, ii
(1896) 453, pro nomen, partim.—T. latifolia WALL.
[Cat. 4939] ex Rip. J. Str. Br. R. As. Soc. n. 82
(1920) 1783 FIs) Mal: “Penw 1) (1922) 512 —72
sambucifolia ELMER, Leafl. 9 (1934) 3217.—T.
nepalensis (non [WALL. Cat. 4277] ex W. & A.

1834): Merr. Contr. Arn. Arb. 8 (1934) 94.—
Fig. 11-n.

Large shrub or tree, up to 20 m high, 55 cm
diam. Leaves 3—S—7(—11)-foliolate, brownish-red
when young; leaflets ovate, oblong or elliptic,
glabrous, more or less acuminate, base obtuse or
obtusely rounded or decurrent, more or less
(sometimes not) dentate, 3-18 by 2-10 cm; petio-
lules of the lateral leaflets 3-15 mm, glabrous or
puberulous. Stipules large, 8-9 mm long, glabrous.
Panicles axillary, open, up to 30 cm long. Sepals
ovate, ciliate, 14-2 by 114-214 mm. Petals oblong
to elliptic, thin, more or less ciliate, 214 by 114 mm.
Stamens 214-3 mm long; filaments 2-214 mm;
anthers 14 by 14 mm, generally more or less
apiculate. Ovary (2—)3(-4)-celled, each cell with
5—6(—8) ovules. Fruit globular, mostly with three
grooves from the top, c. 1-114 cm diam.; pericarp
1-3 mm thick. Seeds brown, mostly several per
fruit; testa 14-1 mm thick.

Distr. Malaysia: Sumatra, Malay Peninsula,
Borneo, Java, Lesser Sunda Islands (Bali, Flores),
Celebes, Philippines, Moluccas (Ceram, Ambon).

Ecol. Frequent in rain-forests, on various soil
types, 50-2000 m.

Vern. Ki bangkong, ki bantjét, ki keujeup, ki
pongpasang, ki tjéhai, tjawané soré, S; godong
bantjét, J; langkiang étém, rébung, tutuh sirawi,
Simalur, aé, kua, Endeh.

Notes. This species has often been confused
with T. pomifera but the differences in size of the
flowers and thickness of the pericarp make it
desirable to distinguish these taxa as two species;
their areas overlap. No attempt has been made to
unravel all confusions in the references.

In the Malay Peninsula and Sarawak T.
sphaerocarpa is mostly represented by a slightly
distinguishable form described as T. /atifolia but
I cannot find any fitting characters to delimit it
against 7. sphaerocarpa.

A specimen from Ceram (EyMA 2150) shows a
remarkable resemblance to the Papuan species but
it has small flowers (244 mm) and constantly 6
ovules per cell. In fruit these species can hardly be
distinguished.

In Sumatra some specimens have been found
which are shortly but rather stiffly hairy on both
leaves and inflorescences; they represent a minor
variety, var. pubescens, nov. var. No great impor-
tance can be attached to this variety, although it is
occupying a geographically coherent area in
Central Sumatra. A specimen from Mt Kinabalu
(CLEMENS 34462) and one from Celebes (Palu:
bb 28283) deviate in having on/y the inflorescence
puberulous, besides having stamens 114-134 mm,
anthers 34 mm.

Of MERRILL’s record of 7. nepalensis in N.
Sumatra I saw only one cited number (BANGHAM
981) which I refer to T. sphaerocarpa. This is a
fruiting specimen; the leaf is typically discoloured
as in T. sphaerocarpa. The ripe fruits are distinctly
too large for T. nepalensis and may well fit the
size and structure of 7. sphaerocarpa, but they
have a distinctly 3-horned tip which is unusual
for the latter species. The horns are not widely

e-

58 FLORA MALESIANA

[ser. I, vol.ogt

spaced as in 7. cochinchinensis which species has
also no discoloured leaves.

8. Turpinia nitida MeRR. & PERRY, J. Arn. Arb. 22
(1941) 549.

A recumbent treelet, 444-6 m high. Leaves
1—5-foliolate; leaflets coriaceous, oblong to elliptic,
glabrous, denticulate, acuminate, rounded at the
base, upper surface shining, 11-25 by 514-1014
cm; nerves prominent, arcuately ascending, then
parallel with the margin; petiole up to 6 cm long,
glabrous; petiolules of the lateral leaflets 12—20
mm. Stipules small, glabrous, c. 5 mm long.
Panicles axillary, subterminal (or terminal), up to
22 cm long. Pedicels 2-214 mm. Sepals oblong-
elliptic, ciliate, 2-4 by 114-214 mm. Petals oblong,
ciliate, 4 by 2 mm. Stamens 214-314 mm long;
filaments c. 3 mm; anthers c. 1 by 1 mm, dis-
tinctly apiculate. Ovary (2—)3-celled, each cell with
4(-5) ovules. Fruit globular, with 3 distinct, some-
times closely appressed horns; pericarp 14-34 mm
thick. Seeds large, two or more in each fruit;
testa 34-1 mm thick.

Distr. Malaysia: North Borneo (Mt Kinabalu).

Ecol. Fairly frequent on wet forest ridges,
900-1800 m.

Note. In size of the leaflets resembling T.
grandis (also occurring on Mt Kinabalu), but
differing in the shape of the fruit, the venation of
the leaflets, and the absence of an indument.

9. Turpinia ovalifolia ELMER, Leafi. Philip. Bot. 2
(1908) 490; Merr. & Perry, J. Arn. Arb. 22
(1941) 544.—T. trifoliata RipLey, J. Str. Br. R.
As. Soc. n. 82 (1920) 178; Fl. Mal. Pen. 1 (1922)
511.—T. lucida Naxal, J. Arn. Arb. 5 (1924) 80.—
? T. pachyphylla Merr. Philip. J. Sc. 27 (1925) 33,
ex descr.—Fig. 1o.

Tree c. 8 m or sometimes higher. Leaves 3—5
(—7)-foliolate; leaflets rounded to elliptic, glabrous,
acuminate, sometimes with an abrupt acute point,
rounded at the base, margin crenate with fine,
whitish, callous points, (2!14—)4-20 by (1-)4-10
cm; petiolules of the lateral leaflets 4-15 mm,
glabrous. Stipules small, 2!4—3 mm long, glabrous.
Panicles mostly short, c. 10-15 cm long (in fruit
to 30 cm), rather dense. Sepals ovate, ciliate,
2-414 by 114-3 mm. Petals obovate-oblong, thin,
ciliate, 314 by 2 mm. Stamens 2!4 mm long;
filaments 2-214 mm; anthers 34 or 1-114 mm long,
not (or slightly) apiculate. Ovary (2—)3-celled,
each cell with 5 or 6(—8) ovules. Fruit with 3 short
horns on top, 4-12 mm diam.; pericarp thick
1-4 mm; testa 0.3-0.4 mm thick.

Distr. Malaysia: Malay Peninsula (Nyalas,
Selangor), Philippines (Luzon, Palawan).

Notes. I cannot well separate T. ovalifolia and
T. trifoliata; the type of the former is extremely
poor; the flowers ELmer described are not present
in any of the isotypes I have had on loan for study.
Though there is a slight difference in the size of the
anthers I cannot discriminate the Malayan spec-
imens from those of the Philippines. The species
can be expected to occur in Borneo.

T. robusta CraieB from Siam is habitually very

similar to Malay Peninsula specimens of “TJ.
trifoliata’”’ but its flowers are smaller and there are
only 4 ovules per cell; it is unfortunately only
described after flowering material which in all its
characters is extremely close to T. cochinchinensis
(Lour.) MerRR., all species having 4 ovules per
cell. The latter species has much larger fruits
(1-2 cm diam.) with spaced horns; T. nepalensis
has small fruits c. 4 cm diam., also provided with
small remains of the style bases.

10. Turpinia pomifera (Roxs.) DC. Prod. 2 (1825)
3; WALL. Cat. 4267; Kurz, J. As. Soc. Beng. 44,
ii (1875) 182, excl. syn.; HAYATA, Ic. Pl. Form. 1
(1911) 160; ?KiNG, J. As. Soc. Beng. 65, ii (1896)
453, pro nomen, excl. var. sphaerocarpa; KANSILAL
& Das, Flora of Assam 1, 2 (1937) 309; MErRR. &
Perry, J. Arn. Arb. 22 (1941) 546; HoL_tHuis &
Lam, Blumea 5 (1942) 205; GAGN. FI. Gén. I.-C.
Suppl. 1 (1950) 993.—Dalrympelea pomifera
Roxs. [Hort. Beng. (1814) 17] Pl. Corom. 3
(1819) 76, t. 279 (Dalrympelia).—Maurocenia
pomifera O.K. Rey. Gen. Pl. 1 (1891) 147, 149.—
Turpinia sp. Merr. For. Bur. Bull. (Philip.) 1
(1903) 34.—Fig. 1j-k.

Tree, c. 10-20 m. Leaves 3—5—7(—9)-foliolate;
leaflets elliptic-oblong, glabrous, distinctly acu-
minate, decurrent (sometimes a little rounded) at
the base, dentate, 12—25 by 6-10 cm; petiolules of
the lateral leaflets 3-10 mm. Stipules triangular,
4-5 mm. Panicles terminal, subterminal or axillary,
10-30 cm long, sometimes slightly puberulous.
Sepals ovate, unequal, fleshy, scarcely ciliate,
2-314 by 114-2 mm. Petals oblong, thin, scarcely
ciliate, 3-314 by 1-114 mm. Stamens 3 mm long;
filaments c. 244 mm; anthers 4/;-1 mm long,
mostly subapiculate. Ovary 3-celled, each cell with
(5—)6 ovules. Fruit globular, mostly with 3 grooves
from the top, up to 25 cm (perhaps to 37 mm);
pericarp very thick, to c. 5 mm diam. Seeds small
and brown, glossy.

Distr. Continental Asia from the East
Himalaya eastward; in Malaysia rare: Sumatra,
Java, Central Celebes (Nuha Distr.: KJELLBERG
2303) and Minahassa, Talaud Is, and Philippines
(Mindanao: AHERN 354; Luzon: Camarines Sur:
BS 76375).

Ecol. In forests, 0-2100 m; f7. March, fr. Sept.—
Oct.

Vern. Ki bangkong, ki rénggang, S, lampasia,
Minahasa.

Notes. 7. pomifera which is a common species
on the Asiatic continent is rare in Malaysia. It is
characterized by a very thick pericarp closely
enveloping the seeds, and besides it differs from
T. sphaerocarpa, with which it has frequently been
confused in Malaysia, by larger flowers and much
larger anthers. In sterile state it is impossible to
tell them apart.

The only specimen of the Malay Peninsula
which might represent true T. pomifera is KING’s
coll. 4243 (SING), but the material is too inade-
quate to be conclusive.

In the W. Deccan (Ghats and Nilgiris) and
Ceylon the records of 7. pomifera have been

Dec. 1960]

STAPHYLEACEAE (van der Linden) 59

straightened out by GAMBLE (Kew Bull. 1916, 135;
Fl. Madras pt 2, 1918, 241); it appears that two
different species are concerned, viz T. nepalensis
and a new species, 7. malabarica GAMBLE, which
is also the single one occurring in Ceylon, charac-
terized i.a. by hairy filaments.

11. Turpinia pentandra (SCHLTR) v. D. LINDEN,
Nova Guinea n.s. 10 (1959) 212.—Kaernbachia
pentandra SCHLTR, Bot. Jahrb. 52 (1914) 151, f. 5
H-N; ENGLER, in E. & P. Pfil. Fam. ed. 2, 18a
(1930) 241.—T. papuana MERR. & PERRY, J. Arn.
Arb. 22 (1941) 554.—? T. papuana HArMs in E. &
P. Pfi. Fam. ed. 2, 20b (1942) 312, descr. germ.,
sine typ., homon. illeg.

Tree, 15—20(-25) m. Leaves (1—)2-3-jugate
(rarely paripinnate by absence of the terminal
leaflet); leaflets oblong or ovate-oblong, glabrous,
shallowly serrate, distinctly obtusely acuminate,
obtuse to rounded at the base, 6—12(—15) by
24%4,414(-614) cm; petiole 2-414(-7) cm, very
short-hairy to glabrous; petiolules of the lateral
leaflets 3-5(-8) mm. Stipules triangular, glabrous
or sparsely hairy outside, 5—7 by 3-4 mm. Panicles
axillary, open, up to 25 cm long. Pedicels 14-3 mm
long. Sepals broad-obovate, ciliate, 3-314 by
114-2 mm. Petals obovate, rounded or obtuse at
the apex, thin, ciliate, 4-414 by 134-2 mm.
Stamens 314-414 mm long; filaments 214-3 mm;
anthers 34-1 mm long, apiculate. Ovary 3-celled,
each cell with 4(—5) ovules. Fruit globular, mostly
with 3 distinct horns (remains of the styles), c.
144 cm diam.; pericarp c. 2 mm thick. Seeds
mostly 1(—2).

Distr. Malaysia: New Guinea.

Ecol. In primary rain-forests, locally frequent,
1200-2500 m.

Vern. Naun, Waria (Mt Hagen).

Note. Very close to TJ. brachypetala, but

constantly differing in the number of the ovules,
the shape of the fruit (horns), and width of the
leaflets.

12. Turpinia brachypetala (SCHLTR) v. D. LINDEN,
Nova Guinea n.s. 10 (1959) 212.—Kaernbachia
brachypetala SCHLTR, Bot. Jahrb. 52 (1914) 153,
f. 5 A—G; ENGLER, in E. & P. Pfil. Fam. ed. 2,
18a (1930) 241, f. 140 A-G.—T. versteeghii MERR.
& Perry, J. Arn. Arb. 22 (1941) 554.

Tree, 4-26 m; bark brownish grey, flaky.
Leaves (1—)2—3-jugate (rarely paripinnate by ab-
sence of the terminal leaflet) ; leaflets ovate-oblong,
serrate, glabrous, obtuse, more or less acuminate,
obtuse to rounded at the base, (614—-)8—-17 by
414-814 cm; petiole 314-9 cm long, very short-
hairy or glabrous; petiolules of the lateral leaflets
4-9 mm long; young foliage glossy brownish
green. Stipules triangular, glabrous or sparsely
hairy outside, 4-7 by 3-6 mm. Panicles axillary,
open, up to 30 cm long. Pedicels 14-3 mm long.
Sepals ovate, ciliate, white, 314-4 by 2-234 mm.
Petals obovate-oblong, spathulate, thin, sparsely
ciliate, white, 314-414 by 114-134 mm. Stamens
314-414 mm long; filaments 3—4 mm; anthers 34-—
0.9 mm long, distinctly apiculate. Ovary 3-celled,
each cell with (7—)8 ovules. Fruit globular, some-
times with 3 radial lines from the top, 1-1'4 cm
diam.; pericarp c. 2 mm thick. Seeds mostly 3
or more.

Distr. Malaysia: New Guinea.

Ecol. In primary rain-forests, locally frequent,
300-2000 m.

Note. We have at present rather numerous
collections from various places all over New
Guinea from both T. pentandra and T. brachypetala.
They are doubtless closely allied, but there appear
to be no intermediates. If only fruiting material is
available T. brachypetala can hardly be dis-
tinguished from T. sphaerocarpa Hassk.

rs eavar is.
“pata: aye
ual

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 Stixis pellucid-
dotted. Stipules thorny, or minute, or wanting. /nflorescences 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-Mal. 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 c, 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 cy, 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 ~, rarely | (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 tropical and subtropical regions.
The largest genera are Capparis (over 250 spp.), Cleome (over 150 spp.), Maerua (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 xerophilous, 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 Stixis.
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. voli age

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, 111-116). 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 gynandra, 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 Mal. 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 C/eome 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 C. chelidonii), 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 (/.c. 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 different 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 effective 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 different 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. FI. Pl. 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 Pittosporales, Tamaricales, Violales, 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. Dic. 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 Malesié, Veenman, Wageningen
(1949) 48 (Crateva) & 51 (Capparis); DescH, Mal. For. Rec. 15! (1941) 70 (hand lens); METCALFE &
CHALK, Anat. Dic. 1 (1950) 91; MoLL & JANsSsonius, 1 (1906) 175.—C.A.R.-G.

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 sei eed after anthesis leaving no scars

on its base. Fruit a berry with leathery pericarp . . A tt ee 2. aN ALIS
2. Stamens at the base connate with the gynophore, leaving scars which 1 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, co-seeded. . . ee ais 3. Cadaba
. Androgynophore | to a few mm long. ‘Stamens ~w. Sepals reflexed. Petals absent. Fruit ellipsoid,
PeSeededaarn mr) | =: ue cease FN se 5 en en aoa LINAS

1. Leaves palmately compound or dissected.
4. Ligneous plants. Leaves compound. Flowers opening in a very early stage. Fruit globular to ellipsoid,

indehiscent, fleshy . . . . I. Crateva

4. Herbaceous plants. Leaves dissected. Flowers mostly in 1 bud anti anthesis. Fruit cylindrical,

CEMISCEMUMGEYA Mes tics sh oe” Sees 95 Sa cc eth cre er cae, 1) De ee Se Oleome
1. CRATEVA

LINNE, Gen. Pl. ed. 5 (1754) 203; Sp. Pl. 1 (1753) 444; HAMILTON, Trans. Linn.
Soc. 15 (1827) 116 (Crataeva); Kurz, J. Bot. 12 (1874) 193; Corner, Gard. 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. 46%

Fig. 1. Crateva nurvala Ham. var. nurvala (Cult. Hort. Bog. IV—F—76; JAcoss, 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) oa | 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 11 pairs of nerves. Rachis up to 10 cm long, with up to 20, rarely 40, fiowers.
2. Leaflets generally sessile or subsessile, occasionally on petiolules up to 5 mm, very rarely 13 mm.
Blade thin-herbaceous, green when dry, (5!4—)8!4—16(—27) cm. Fruit 6—12(—15) cm long, papillate,

greyish when dry .

1. C. religiosa!

2. Petiolules 4—S(—6) mm. Blade herbaceous to subcoriaceous, mostly red-brownish when dry, 514-10

(-14) cm. Fruit 314-4 cm long, smooth, red-brownish when dry

2. C. odora f. 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 .

1. Crateva religiosa Forst. f. Pl. Escul. Ins. Oc.
Austral. (1786) 45, (Crataeva); Fl. Ins. Austral.
Prod. (1786) 35; DC. Prod. 1 (1824) 243, p.p.;
non Hook. f. Fl. Br. Ind. 1 (1872) 172; Laur. Bot.
Jahrb. 52 (1914) 110; Merr. Philip. J. Sc. 11
(1916) 272; WALKER, Imp. Trees Riukiu (1954) 95,
f. 45.—C. membranifolia Mia. Sum. (1861) 387,
158; Ilustr. (1870) 21; Koorp. Minah. (1898) 343;
Laut. Bot. Jahrb. 52 (1914) 111; Merr. En. Born.
(1921) 280; CorNER, Ways. Trees (1940) 181.—
C. macrocarpa Kurz, J. Bot. 12 (1874) 195, t. 148
f. 8-10; Kina, J. As. Soc. Beng. 58, ii (1889) 397;
Ripi. Fl. Mal. Pen. 1 (1922) 125; MeErr. Philip.
J. Sc. 29 (1926) 371; Corner, Gard. Bull. S.S. 10
(1939) 16 (Crat. B).—C. hansemannii K. Scu. Bot.
Jahrb. 9 (1888) 201; K. Sco. & Ho utr. FI. Kais.
Wilh. Land (1889) 50; Wars. Bot. Jahrb. 13
(1891) 318; K. Sco. & Laur. FI. Schutzgeb.
(1901) 335, p.p.; VALETON, Bull. Dép. Agric. Ind.
Néerl. 10 (1907) 15; Laur. 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 (1-)5—15(-30) m. Stipules 14-1 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, (5!4-)
814-16(-27) by (3-)4-10!14 cm, central leaflet
oblong, obovate, the base narrowly decurrent, the
apex shortly (incidentally up to 214 cm) acuminate,
often mucronulate; nerves 7-11 pairs; petiole
(3'4-)614-10 cm, on sterile twigs often longer,
up to 22 cm; petiolules 0O—S(—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 1-114 mm, 3—5 mm
petioled. Sepals ovate, obtuse to acute, 4-7 by
114-3 mm. Petals once recorded orange, 5—20 mm
stalked, blade broadly ovate to elliptic, acute to
obtuse, upper pair 2—3(-4) by 1—2(—2.3) cm, lower

3. C. nurvala var. nurvala

pair 114-2 by 1-114 cm, nerves 4—6 pairs. Stamens
(10—)13—18(—30); filaments 414-1114 cm, pink or
purple towards the top; anthers 214-6 by 114 mm,
sometimes recurved. Gynophore 4-7 cm; ovary
4-6 by 1!4-2'4 mm, subcylindrical, sometimes
ovoid, contracted below the stigma 114 mm in
diam. Pedicel in fruit (4-)5—714(-8) cm, 3—4(—5)
mm thick; torus 7-11 mm wide; gynophore
514-814(-14) cm long, sometimes cylindrical, c.
3-5 mm diam., or gradually thickened and up to
1 cm thick at the top. Fruit subglobular to (ob-)
ovoid, 6—12(—15) by 514-914 cm, wall in the unripe
stage up to 7 mm thick, at maturity probably not
thicker than 1-114 mm, smooth when very young
but soon covered with flat, pale, dry papillae,

Fig. 2. Lateral leaflets of Crateva, a. C. religiosa
Forst. f.,b. C. odora Ham. f. axillaris (PRESL)
JAcoss, c. C. nurvala Ham. var. nurvala, all X 2/;.

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

66 FLORA MALESIANA

fser. I, vol. 6!

sometimes giving the impression of a thin, dull,
yellowish grey crust. Seeds more or less asymme-
trically cordate, 10-19 by 5-17 mm diam. and
4-8 mm thick, dorsally with a keel rather narrow
and sparsely tuberculate to very broad and
densely tuberculate (sometimes rather smooth),
the sides smooth to shallowly grooved.

Distr. India (E. Himalaya), Burma, Lower
Siam, Indo-China, Ryukyus, through Micronesia
(Marianes & Carolines) and Melanesia (Solomons)
to Polynesia (Fiji, Samoa, Society Is., Gambier
Is.); in Malaysia: Sumatra, Riouw Arch., Malay
Peninsula, W. Java (twice collected), Borneo (also
Banguey I.), S. Philippines (Palawan, Sulu Arch.),
Celebes (also Kabaena and Muna Is), Moluccas
(Talaud, Sula, Buru, E. Ceram, Ambon, Kai, Aru,
and Tanimbar), New Guinea (also Normanby I.
and Salawati), New Ireland.

Ecol. The species seems to be frequent in
Borneo, New Guinea, and the Solomon Is where
it is often found in periodically inundated forest
along rivers, rarely in secondary or primary dry-
land forest; one record from the beach in Sarawak,
where it seems to attain smaller size. Mostly below
100 m, but up to 700 m. Fi. fr. in all months.
Incidentally cultivated.

Vern. Képayan (ayér), Malaya, wujesta, S,
tigarun, tjigaron, S. and W. Borneo, kénohan
séguntu, Kutei, makendem alus, malasut, sang-
kiauw, Minahasa, tjandaule, SE. Celebes: Toko-
laki, kamfooiju, Mangoli, bala-lehe, Muna,
papangi-nasu, Talaud, ombo-ombo, SW. New
Guinea, ai-yumba, bam-baimovi, Solomons, pua
veoveo, Tahiti.

Uses. In the Solomon Is the leaves are heated
and applied in case of ear-ache, and the fruits are
used against constipation. In Yap the fruits are
eaten. The raw fruit is used as fish-bait in W.
Borneo.

BURKILL briefly refers (under C. macrocarpa) to
occult power ascribed to Crateva species in India
and Polynesia where it is planted round temples,
to which also ForRsSTER’s epithet refers (Dict. p.
676).

Notes. The species is very variable in floral
parts, in the shape of the fruit, and the size of the
seeds.

The leaves are mostly (in Malaya to a less
degree) so thin in texture that a herbarium spec-
imen with undamaged leaves is extremely rare
and there is no difference in colour between the
upper and the lower surface. Besides, the seeds,
however variable, have always a more or less
developed crest of warts on the dorsal side.

The fruit is once recorded to be compressed. In
West Malaysia the subglobular to ellipsoid shape
seems to prevail, whereas in eastward regions the
fruits are more elongate. In Malaya the seeds also
seem to be smaller than in the eastern specimens.
From the Solomon Is the fruit is reported to be
sausage-shaped 14 by 4cm, with unpleasant odour
when cut; in W. New Guinea VAN ROYEN noted
that “‘the smell of the fruit fills the forest with a
soury scent not unlike durian’. From Yap
VOLKENS mentioned a fruit 18 by 10 cm and added

that a variety exists with fruits as small as a
thumb’s length. Another record about such small
fruits could not be found. The few field notes give
the colour of the young fruit as pale green, once
dark mauve, and of the ripe fruit white. According
to VAN ROYEN in New Guinea the fruits are light
green with yellowish scales, hard pericarp, spongy
endocarp, and possess floating capacity. In
Bornean specimens seeds were measured 1 by
34-1 cm in diam. and 14 cm thick; in one specimen
from Java the seeds were 114 by 1-114 cm in diam.
and 14 cm thick.

About the identity of the Palawan specimens I
am not certain. The material available, ELMER
12650 and CENABRE c.s. FB 27861, is not in a very
good state and seems to be intermediate in charac-
ters between C. religiosa and C. odora. Pro-
visionally, on account of the large seeds, the com-
paratively large leaflets with a base somewhat
decurrent on the petiolule and a short-acuminate
top, I assume them to belong to C. religiosa,
notwithstanding the brownish colour of the leaves
in the ELMER specimen and the almost smooth
surface of the seeds.

Mr N. G. Bisset observed in Tanimbar that
the fruit pulp had a burning taste, and that the
seeds contained a very high amount of alkaloids,
while starch was practically absent.

2. Crateva odora Ham. Trans. Linn. Soc. 15 (1827)
118.

f. axillaris (PRESL) JACOBS, stat. nov.—C. tapia
(non L.) BL. Bijdr. 2 (1825) 54; Mia. FI. Ind. Bat.
1, 2 (1858) 102.—C. axillaris PREsL, Rel. Haenk. 2
(1835) 85; F.-ViLL. Nov. App. (1880) 10.—C.
religiosa (non Forst. f.) BLANCO, FI. Filip. (1837)
399, ed. 2 (1845) 279, ed. 3, 2 (1878) 154, t. 176;
F.-VILL. Nov. App. (1880) 10; VIDAL, Sinopsis
Atlas (1883) 13, t. 6 f. C; Phan. Cuming. (1885) 94;
Rev. Pl. Vasc. Filip. (1886) 48; Merr. Sp. Blanc.
(1918) 158; En. Philip. 2 (1923) 210; Quis. Med.
Pl. Philip. (1951) 341.—C. tumulorum Mia.
Illustr. (1870) 21, t. 11; K. & V. Bijdr. 4 (1896)
269; Back. Schoolfl. (1911) 64; Bekn. Fl. Java
(em. ed.) 4A (1942) fam. 45, p. 6.—Fig. 2b, 3.
Small tree, 3-10(—30?) m. Stipules falcate, small.
Leaflets on slender stalks 4-S(—6) mm long, her-
baceous to subcoriaceous, in herb. dull, red-
brownish; 514-10(-14) by 2!4-S(-—7) cm, elliptic
to oblong, rarely lanceolate; base cuneate, top
abruptly acutely c. 1-114 cm acuminate; central
leaflet mostly the largest, broadest at or somewhat
above, rarely below the middle; lateral leaflets
strongly asymmetrical; nerves generally 5, some-
times up to 7, rarely up to 10 pairs; petiole slender,
(314-)6-8(-10) cm, on top occasionally with a
triangular gland. Jnflorescences on small twigs,
afterwards whether or not growing through,
flower-bearing part c. 3(—-10) cm long, with a few
to about forty flowers partly above the axils of
young leaves or bracteate. Pedicels 4—S(—7) cm.
Sepals ovate, acute, c. 4-6 by 114-214 cm. Petals
suborbicular to broadly ovate-elliptic, (1'14-)
214-3 by 1-2 cm, the top blunt to rounded or

CAPPARIDACEAE (Jacobs) ea 67

Fig. 3. Crateva odora Ham. f. axillaris (PRESL) JAcoBs. Sumbawa, 1934 (DE Voocp 1911).

sometimes notched, base rounded and (rather) ab-
ruptly narrowed into the stalk 6-10 mm. Stamens
21-25, 3-4 cm long, anthers c. 3 by 1 mm, re-
curved. Gynophore (2—)3-4(-6!14) cm, ovary sub-
globular to oblong, c. 2-5 by 2 mm, constricted
below the stigma c. 1144 mm broad. In fruit the
gynophore slightly thicker than the pedicel, c.
2-3 mm thick all over, torus c. 7 mm wide. Fruit
globular, (up to?) 3144-4 cm diam. (in Indo-
Chinese specimens up to 6 cm), pericarp leathery,
c. 1 mm thick, smooth during the whole develop-
ment, at maturity probably red when fresh,
brownred when dry. Seeds rather irregularly
horseshoe to deeply kidney-shaped, about 6 by 2
mm, smooth.

Distr. The species occurs throughout India,
Ceylon, Burma, Indo-China, S. China, Formosa,
and Hainan, the Malaysian form in S. India
(Coimbatore), Ceylon, Malay Peninsula (Malacca:
cultivated), Java (W. and E. part, cultivated),
Madura, Kangean Arch., Lesser Sunda Is.
(Sumbawa), Philippines (N. Palawan, Mindoro,
Luzon, most provinces, Guimaras, Mindanao).

There are far more collections from Luzon than
from any other locality. It looks as if the species is
native in the Philippines. It cannot be verified
whether it is introduced or native in Kangean and
Sumbawa; in Madura, Java, and Malaya there is

little doubt that it has been introduced long ago;
it is sometimes found planted on graves.

Ecol. Prefers dry, shrubby places. Mostly deci-
duous, the flowers then appear just before the
young leaves break out, but the blossoming seems
not to show periodicity. In the Malay Peninsula it
is only rarely found in fruit.

Uses. QUISUMBING, Med. PI. Philip. (1951) 341,
mentioned quite a few minor medical applications.

Vern. Sémpal wadak, J (for the genus, reliable);
kémalo-kémalowan, sékar bulan, Kangean, saling-
bogog, Tag. (Philip.).

Notes. From C. religiosa distinguished vege-
tatively by its leaves being firmer in texture, often
red-brownish in the herbarium, especially the
specimens from Java and the Lesser Sunda Is.,
and giving a more graceful impression, due to the
smaller size, the slender distinct petiolule, and the
acute longer tip of the leaflets. In fruit it is different
by its shorter gynophore and by the surface of the
fruit which is smooth and not papillate; this
character being more reliable than those provided
by the (often immature) seeds.

In one specimen, cultivated in Hort. Bog. under
IV-F-81, the largest leaflet was 17 by 9 cm. Its
adult leaves were only slightly reddish tinged, the
tip being more or less abortive or absent. This
seems to prove once more that species may show

68 FLORA MALESIANA

[ser: I, volieGt

deviations unknown from the wild state merely by
cultivation in a botanic garden.

The closest affinity is with the African C.
adansonii DC. which could even be looked upon
as a subspecies. In C. adansonii the inflorescence
rachis is + 4-5 cm, and finally set with the thick
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-
count of these differences it seems appropriate to
keep them apart.

3. Crateva nurvala Ham. Trans. Linn. Soc. 15
(1827) 121, (Crataeva Niirvala); W. & A. Prod. 1
(1834) 23; Kurz, J. Bot. 12 (1874) 195; GaGn.
Not. Syst. 8 (1939) 213; Fl. Gén. I.-C. Suppl. 1
(1939) 157, t. 14 f. 8.

var. nurvala.—C. nurvala HAm. /.c.; Mia. Illustr.
(1870) 20; K. & V. Bijdr. 4 (1896) 266; Back.
Schoolfl. (1911) 64; HALL. f. in Winkl. Bot. Jahrb.
49 (1913) 369; Merr. En. Born. Pl. (1921) 280;
Back. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45,
p. 6.—WNiirvala RHEEDE, Hort. Malab. 3 (1682)
49, t. 42.—C. religiosa (non Forst. f.) BL. Bijdr. 2
(1825) 54; Ripi. Fl. Mal. Pen. 1 (1922) 125;
RENDLE, J. Bot. 63 Suppl. (1924) 5.—C. magna
[an (Lour.) DC.?] HAssk. Pl. Jav. Rar. (1848) 179;
Mia. FI. Ind. Bat. 1, 2 (1858) 102; Sum. (1862)
158, 387.—C. religiosa non Forst. f. var. nurvala
(Ham.) Hook. f. & THoms. FI. Br. Ind. 1 (1872)
172, p.p.—C. lophosperma Kurz, J. Bot. 12 (1874)
195, t. 147 f. 4-6, an var. propria?; CORNER, Gard.
Bull. S.S. 10 (1939) 16 (Crat. A); Ways. Trees
(1940) 181, f. 48.—Fig. 1, 2c, 4.

Tree, (8—)10—15(—20) m, to 40 cm diam. Branch-
lets slightly zig-zag. Stipules minute, late caducous,
acute with broad base. Leaflets firmly herbaceous
to subcoriaceous, (0O—)3—6(—10) mm stalked, mostly
lanceolate, sometimes oblong, rarely linear,
(414-)9-15(-28) by (11!4-)3-S(-614) cm; base
acute, top acuminate, tip acute; central leaflet
broadest about or below, rarely above the middle,
lateral ones more or less symmetrical; nerves
(7-)10-15(—22) pairs; surface below duller and
paler than above, sometimes on both sides with
minute grey-brown scattered papillae. Petiole
(4-)514-914(-14) cm, vigorous, broadly sulcate,
on top bearing numerous pale to light brown
gland-like appendages up to 1 mm long. Jn-
florescence ultimately c. 10-16 cm long, bearing
20-100 fiowers. Bracts early caducous, 5-9 by
344-1144 mm, acute, their stipules minute, longer
persistent. Pedicels 4-7 cm. Sepals 2-314 by
144-1144 mm, ovate, acute, somewhat narrowed
at the base. Petals 5-12 mm stalked, blade (8-)
15-30 by (5—)15-22 mm, suborbicular or sub-
rhomboid to elliptic, ovate, base rounded, ab-
ruptly narrowed into the stalk, top obtuse.
Stamens 15-25, 3\4-414(-6) cm; filaments purple;
anthers c. 2-3 by 1 mm. Ovary ellipsoid to cylin-
drical, c. 5-6 by 214 mm; stigma dark purple.
Gynophore 344-514 cm, in fruit probably not or

hardly lengthened (rarely up to 10 cm), 3—4(—S)
mm thick, only slightly thicker towards the top;
pedicel mostly thinner, torus 7-8 mm wide. Fruit
unknown in fully mature state, ellipsoid, rarely
ovoid, (up to?) 5-514 by 4-414 cm, pericarp 4-5
mm thick, covered with a thin, dull, yellow-

Fig. 4. Crateva nurvala Ham. var. nurvala. Flower,
front view (Cult. Hort. Bog. IV—F—76; JAcoss.
1955).

greyish crust breaking into minute particles which
seem to peel off sooner or later, leaving the surface
smoothish. Seeds deeply horseshoe-shaped, 6-9
mm long and wide, 2-3 mm thick, dorsally with a
crest of sharp irregular protrusions.

Distr. India (Deccan, Sikkim, Assam), Burma,
S. China (Yunnan, Kweichow?), Hainan, Indo-
China, Siam; in Malaysia: Sumatra (P. Weh,
East and West Coast Res., Palembang), Malay
Peninsula, W. to E. Java, and Borneo. Cultivated
throughout the area, for ornamental or magic
purposes.

Another variety occurs in India (mainly in the
Northeast), Burma, Siam, Indo-China, S. China
(Yunnan, Kwangtung, Fukien), Hainan, and S.
Japan (probably this species).

Ecol. Mostly along streams in shady places,
sometimes behind the sea-shore, at low altitudes
up to 600 m. Seems to be rare everywhere. Flower-
ing and fruiting time irregular.

Vern. Si baluak, Sumatra, badat, dalath), dalur,
Mal. Pen., barun(d)aj, ki howe, S, djaranan, sémpal
wada, J, pingos, sasagah, sébélu, tigarun, Borneo.

Uses. The wood seems to find local application
as timber. In Malaya the root, leaf, or bark are
boiled with oil and applied externally in case of
‘sakit angin’. In Siam the fruit is used as fish-bait
and the young leaves are pickled in salt. See
BuRKILL (Dict. 1, p. 676) for other minor uses.
According to FiLet, cf. HEYNE, Nutt. Pl. p. 682,
the bark contains an acrid, bitter substance and is
pounded in water, used as a skin-irritant against
high fever, efc.

Notes. This species is vegetatively characterized
by its nerf-pattern: the leaves have underneath a
typical aspect by the thin but vigorous and prom-
inent nerves, more numerous than in other
species, while the insertions of the intermediate
veins on the midrib are also far more distinct.
Besides, the lower surface is more pale, dull, and
greyish or glaucous than the upper, the difference

Dec. 1960]

CAPPARIDACEAE (Jacobs) . 69

being far greater than in the other Cratevas.

In none of the herbaria consulted any type
material of C. lophosperma was found. Kurz
named as type specimen ““GusTAvy MANN, from
banks of the Koolsee River, Kamroop, Assam,
Fr. July.”’ From the description it is clear that one
of the two varieties of C. nurvala was meant; pro-
bably it was var. nurvala. CORNER’s interpretation
of KurRz’s species as C. nurvala s. lat. was correct;
his cited material no doubt belongs to var. nurvala,
but perhaps Kurz himself had HAMILTON’s
*C. unilocularis’ at hand, which is more frequent in
Assam than var. nurvala.

Imperfectly known

4. Crateva hygrophila Kurz, J. As. Soc. Beng. 41,
ii (1872) 292; J. Bot. 12 (1874) 196, t. 148 f. 6, 7;
ieeAss soc. Beng. 48, 1 (1874) 33; BI Burm. 1
(1877) 67.

Leaves herbaceous, dull reddish brownish,
largest leaflet 1214 by 3'4 cm, leaflets sessile, not
much asymmetrical, narrowed towards base and

top; nerves ++ 6-7 pairs, as the midrib narrow,
prominent above. /nflorescence rachis practically
none. Flowers unknown. Pedicels in fruit 514-614
cm, gynophore 434-534 cm, both blackish, woody,
+ 2-214 mm in diam. Fruit (immature) cylindrical
umbonate, + 9 by 244 cm, dull brown-purplish
with numerous white lenticel-like specks, c. 11/4
mm thick. Seeds irregularly horseshoe-shaped,
+ 11 by 8 by 3 mm, the outer side with some
coarse sculpture.

Distr. Burma (Pegu); in Malaysia: Penang
(KING’s Coll. 1412).

Note. There are only two collections; both are
inadequate but match very well. They come nearest
to C. religiosa, but I feel reluctant to refer them to
this species. They may represent an abnormal,
deviating form.

Excluded

Crateva marmelos L. Sp. Pl. (1753) 444 =
Aegle marmelos (L.) CorREA in Trans. Linn. Soc.
5 (1800) 223 (Rutaceae).

2. CAPPARIS

TOURN. ex LINNE, Gen. Pl. ed. 5 (1754) 222; Sp. Pl. 1 (1753) 503; LAMk, Encycl.
1 (1785) 604; DC. Prod. 1 (1824) 245; Mia. Ilustr. (1870) 22-36, t. 12-19; GaGn.
in Morot, J. Bot. 21 (1908) 53; PAx & Horrm. in E. & P. Pfl. Fam. ed. 2, 17b
(1936) 172.—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
fairly regularly and interlooping near the margin. Flowers pedicelled, arranged in
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 ~, 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 © ovules; stigma
sessile, small. Berry in Mal. globular to ellipsoid, rarely elongate, with leathery or
corky pericarp, l-celled. Seeds (l1-)-), embedded in pulp, obliquely reniform,
rather large, with circinnate embryo.

70 FLORA MALESIANA (ser, 15 -volaems

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. f., orange in C. floribunda WiGutT, 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. lucida (BANKS ex DC.) BENTH. and C. spinosa var. mariana (JACQ.) K. SCH., are
known to be noctiflorous. C. erycibe HALL. f., 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. 783A, 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. induta (hairs soft and somewhat twisted); 7. C. /anceolaris
(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. Seriales DC. (incl. sect. 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 oplong.,Sepals 5-7 mmlong 9... 4 2a «, + %) .« -es,) 20 are
2. Leaves suborbicular. Sepals longer than 10 m hy sth hae ae 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 244 cm.
5. Pedicels 14-1 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) . ii

8. Gynophore 3-414 cm. Sepals 8-12 mm long. Leaves with a hardened tip.

12. C. callophylla

8. Gynophore 2-314 cm. Sepals 5—6(-10) mm long. Leaf-tip not particularly thickened.

13. C. zippeliana

7. Innovations hairy, aie Leaves obovate, subcoriaceous, subtriplinerved, (6—)10—14 cm

long

2. C. trinervia

4. Sepals shorter ‘than 7 mm. Gynophore shorter ‘than 2Yy cm.
9. Flowers in a panicle. Leaves longer than (8—)10 cm.

10. Gynophore 4-12 mm. Fruit 144 cm diam.
10. Gynophore 2-314 cm. Fruit c. 344 cm in diam.

. 4. C. erycibe
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— eae mm. Twigs vigorous, hairy, zig-zag, mostly with

strong thorns

10. C. sepiaria

. Stamens 12-15. Gynophore c: 15 mm. Twigs slender, glabrescent, straight, with weaker thorns.

11. C. diffusa

3. Inflorescences axillary or supra-axillary, sometimes a/so 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.

14. Sepals longer than 5 mm, ciliate. Fruit smooth. Thorns patent

13. Sepals puberulous outside.

16. C. buwaldae
18. C. micracantha

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
17. Stamens 7-8. Sepals shorter than 5 mm

ba Young leaves glabrous. Fruit elongate.

19. C. zeylanica
20. C. quiniflora
17. C. cucurbitina

Flowers in sessile racemes or in subumbels or panicles.

xin Flowers in short, sessile, axillary racemes.

19. Gynophore at anthesis hairy all over. Rachis of the inflorescence inconspicuous.

19. Gynophore at anthesis glabrous. Rachis of the inflorescence thick,

14. C. pubiflora
1-214 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 3-414 cm. Sepals 8-12 mm long. Leaves with a hardened tip.

12. C. callophylla

21. Gynophore 2-314 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 144 cm. Fruit 44-2 cm diam.

24. Stamens c. 8. Sepals 3-4 mm long. Fruit 144-2 cm diam.

8. C. floribunda

24. Stamens oo. Sepals 4-5 mm long. Fruit 114 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!4 cm. Fruit (in) 2-30 cm ‘diam.
26. Twigs glabrous or with appressed pubescence. Thorns, if present, recurved.
27. Leaves 614—12(—-16) cm long, the ley somewhat recurved. Sepals at least 6 mm long.

Fruit 2-314 cm diam.

27. Leaves 5-6 cm long, flat. Sepals GC 3, mm long. Fruit unknown ‘

7. C. lanceolaris
6. C. longestipitata

26. Twigs generally with short, patent pubescence. Leaves 4—8(—13) cm long, ovate, subcordate.

Fruit c. 144 cm diam. Thorns, if present, straight

1. Capparis scortechinii KING, J. As. Soc. Beng. 58,
ii (1889) 394; Ann. R. Bot. Gard. Calc. 5 (1896)
118, t. 135; Ripe. Fl. Mal. Pen: 1 (1922) 122.

Climbing shrub, 2-10 m. Twigs straight, an-
gular and pubescent, terete and glabrescent when
older; internodes 1-3 cm. Thorns strong, recurved,

9. C. lobbiana

2-4 mm. Leaves (sub)coriaceous, pubescent when
young, soon glabrescent, ovate to obovate, 2-5
times as long as broad, 614-12(-21) by 144-5
(-714) cm; base narrowed, acute; top rounded to
narrowed, more or less acuminate, dark-mucro-
nate; midrib flat above; nerves c. 5-6 pairs;

FLORA MALESIANA [ser.: 1; -volagae

72

Fig. 5. Capparis zeylanica L. a. Flowering twig, the leaves still young, « 24, b. flower, nat. size, c. fruiting
branch with adult leaves, the fruit opened, x %

Direc 1960]

CAPPARIDACEAE (Jacobs) = ; 73

petiole c. 1 cm, brown pubescent, late glabrescent.
Raceme terminal, 214-10 cm long, brown pubes-
cent all over. Bracts 10 by 114-2 mm, sometimes
larger and resembling small leaves. Pedicels
5—10 mm, leaving a prominent scar, in fruit woody.
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 414 mm, densely
pubescent outside. Petals pink, c. 8-9 by 414-6
mm, obovate, notched, with cuneate base, sparsely
pubescent inside, or glabrous. Stamens 35—SO, c.
15 mm. Gynophore c. 5-644 cm (occasionally
abortive), in fruit transversely wrinkled, woody,
thickened up to 16 mm; ovary ovoid, 134 by 34
mm. Fruit globular, c. 10(—12'4) cm diam., peri-
carp woody, 2-214 cm thick, smooth, yellow.
Seeds ~, c. 2!4 by 134-214 by c. 1 cm.

Distr. Malaysia: ? Banka, Malay Peninsula
(Perak, Trengganu, Pahang, Selangor, and
Penang).

Ecol. In rain-forests, up to 1400 m.

Vern. Ménawul, Banka, susoh béruga, Pahang.

Note. An entirely glabrous sterile specimen
collected by TEYSMANN s.n. (Bo) in Banka, pro-
bably belongs here. In KING’s type collection both
flowers with an abortive and a developed gyno-
phore are represented.

basal part; nerves (6—)7—8(—10) pairs, glabrous to
early glabrescent; petiole (7—)10—-14(-18) mm,
hairy as the twig. Flowers (3—)5—10(—-15) in a
terminal raceme. Pedicels rather vigorous, in the
axils of the upper leaves or in the axils of very

r? lw
Iya Cg 2
LY \ Qe
) YA8
oa Se
2) =
SS or, eS
BG ee fe
NON 0% _, 20,4 _)2 e
< 4 = —— 5) ca = a Tamm G ue
Nu! c > 7) =
2 ae : :
os “59° Se
< i2 soon iN —>
f et

Fig. 6. Species density of the genus Capparis in
Malaysia. The number above the hyphen refers
to the number of endemic species, that below the
hyphen to the number of other species in each
province. C. /ongipes MERR. (incompletely known;
from the P.I.) has not been incorporated.

2. Capparis trinervia Hook. f. & Tu. FI. Br. Ind. 1
(1872) 175; Kurz, Fl. Burm. 1 (1874) 64; GaGn.
Fl. Gén. I.-C. 1 (1908) 193; Suppl. 1 (1939) 168. —
C. kunstleri KING, J. As. Soc. Beng. 58, ii (1889)
396; Rip. Fl. Mal. Pen. 1 (1922) 122.
Climber or shrub, up to 4 m. Young parts
ferruginous-tomentose, soon glabrescent. Twigs
almost straight, often angular, internodes c. 3—5
cm; thorns 1—2(-3) mm, patent or slightly re-
curved. Leaves subcoriaceous, elliptic to oblong,
slightly obovate, (6—)10—-14 by (234-)314-514 cm;
base cuneate, often subtriplinerved, top acuminate
with acute tip c.34 cm; midrib above sulcate in the

Fig. 7. Approximate area of distribution of

Capparis trinervia Hook. f. & Tu. (broken line)

and localities of C. quiniflora DC. (dots within
continuous line), the latter also in Fiji.

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 11 mm, outside densely orange-yellow puberu-
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 234-414 cm long,
light red, glabrous, ovary ellipsoid to ovoid, 214
mm long, with umbonate stigma, green, glabrous.
Fruit globular, deep yellow, 314 cm in diam.,
pericarp soft-woody, 4 mm thick. Seeds 15-17 by
14 by 6 mm.

Distr. Indo-China (Tonkin, Laos, Annam),
Burma (Tenasserim); in Malaysia: NE. Sumatra
(Tinggiradja, N of lake Toba), Malay Peninsula
(Perak: G. Booboo). Fig. 7.

Ecol. A creeper reported clinging to trees in
dense jungle, and from dry, almost bare lime-
cinder ledges, at low altitude.

Note. The Malaysian specimens differ slightly
from those of Indo-China where the leaves have
3-5 pairs of nerves and the buds are not
pointed.

3. Capparis brachybotrya HALL. f. in Fedde, Rep.
2 (1906) 59; Laur. Bot. Jahrb. 52 (1914) 112.

Branchlets stout, terete, slightly zig-zag, gla-
brous, internodes 214-5 cm. Thorns slightly re-
curved, minute to 2 mm long. Leaves coriaceous,
elliptic, oblong or sublanceolate, light green when
dry, glabrous, 13-2114 by 5—10!4 cm; base ob-
tusely acutish to rounded; top rounded and 2-10
mm acuminate, with a dark and stiff tip; nerves
(6—)7—9(—10) pairs, depressed above. Petiole stout,

74

FLORA MALESIANA

[ser. I, vol

— aT

(a Lawill)

il
4 \

ry)

STON

Fig. 8. Capparis erycibe HALL. f. a. Habit, x 24, b. flower, x 6, c. fruits, x 24 (after Jacoss 4838).

6

Dec. 1960]

CAPPARIDACEAE (Jacobs) 75

1-114 cm. Racemes axillary, sometimes serially in
twos; rachis stout, (1-)2!4 cm, up to 20-flowered,
pale-puberulous, glabrescent, at the very base
surrounded by conferted bracts. Bracts subulate,
small, early caducous; bracteoles stiff, subulate,
minute, later caducous. Pedicel 1—2!4 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—160, 25—35 mm long,
filaments white. Gynophore 2—3'\4 cm; ovary ovoid,
2-3 by 1 mm, stigma 14 mm, all glabrous.

f. brachybotrya.

Leaves elliptic, 1.7—2.1 times as long as broad,
14-22 by 714-1014 cm, with rounded base. Fruit
ona gynophore | 14-214 cm, ellipsoid with narrow-
ed base and short, abruptly acuminate top, 4-514
by 314 cm, pericarp leathery. Seeds 04, c. 10 by 7
by 5 mm, light brown.

Distr. Malaysia: Moluccas (Kai
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 RoyeEN. 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.

Is), New

f. angustifolia (HALL. f.) JAcoBs, stat. nov.—C.
brachybotrya var. angustifolia HALL. f. in Fedde,
Rep. 2 (1906) 60; Laur. Bot. Jahrb. 52 (1914) 112.
—Fig. 9.

Leaves oblong, 2.4-3.3 times as long as broad,
13-2144 by 5-7 cm, with obtusely acutish to
rounded base. Gynophore 414 cm; fruit (almost
mature?) elongate, 4 by 1!4 cm, pericarp thin.
Seeds o, c. 8 by 7 by 4 mm.

Distr. Malaysia: Celebes (Pangkadjene, Mina-
hasa), New Guinea (Vogelkop).

Ecol. Coastal forest.

Note. In sterile state very similar to C. micra-
cantha, but easily distinguished by the axillary
inflorescence with a thick rachis, C. micracantha
having a supra-axillary row of pedicels or their
scars.

4. Capparis erycibe HALL. f. Bull. Herb. Boiss. 6
(1898) 216; BAck. Schoolfi. (1911) 62; Koorp.
Exk. Fl. Java 2 (1912) 294; Back. Bekn. FI. Java
(em. ed.) 4A (1942) fam. 45, p. 10.—C. paniculata
Riwt. J. Fed. Mal. St. Mus. 10 (1920) 129; RipL.
Fl. Mal. Pen. 1 (1922) 124; Baxer, J. Bot. 63
Suppl. (1924) 5.—Fig. 8.

Climber, about 1!14(-14?) m. Twigs slack,
brownish, brown-puberulous, especially when
young, slightly zig-zag; internodes about 3—5S cm.

Fig. 9. Capparis brachybotrya HALL. f. f. angusti-
folia (HALL. f.) JAcoss, node with leaf and
inflorescence, 24 (KOSTERMANS 2819).

Thorns up to 2 mm long, recurved, mostly wanting.
Leaves herbaceous to subcoriaceous, often reddish
brown when dry, glabrous above, glabrescent
beneath, broadest above, sometimes at the middle,
1.8—2.5(—2.8) times as long as broad, (9!4)—12-16
(-20) by 414-814 cm, base mostly narrowed,
sometimes rounded, acute to obtuse; top rounded
to acuminate, mucronate; midrib above mostly

76 FLORA MALESIANA

[ser. I, vole?

narrowly sulcate; nerves c. 6-8 pairs, glabrous
above, glabrescent beneath; petiole 4-10 mm,
brown-pubescent, late glabrescent. Panicle mostly
brown-puberulous on a c. 5-10 cm long, slender
peduncle terminal on a twig of which the upper
part has mostly lost its leaves, c. 10-20 by 5-15 cm.
Pedicels slender, 4-18 mm. Bracts minute, some-
times wanting. Flowers white to greenish, some-
times tinged reddish. Buds globular, c. 4mm diam.
Sepals 4-6 by 214-3 mm, the outer pair sometimes
sparsely puberulous outside; the inner pair with
broad membranous margin, glabrous. Petals 4
(occasionally single flowers with 5 or 6), 414-6 by
1-4 mm, suborbicular to subspathulate, sometimes
puberulous, especially at the base. Stamens 20-40,
5—6(—8) mm. Ovary ovoid to spindle-shaped, 2 by
1 mm, glabrous; gynophore 2-5 mm, glabrous, in
fruit 4-12 mm long, with the pedicel and torus a
little incrassate. Mature fruit not known, globular,
c. 114 cm diam., pericarp thin, leathery, smooth or
finely papillate. Seeds 1-4, subglobose to angular,
9-10 by 7-9 by 4-5 mm.

Distr. Indo-China (Annam), in Malaysia:
S. Sumatra (Lampongs), Malay Peninsula
(Pahang, Kelantan), Borneo (Sarawak), Java
(scattered).

Ecol. Forests, often on limestone, up to 600 m.
FI. Dec.-June, fr. July-Nov. Apparently a rare plant.

Vern. Endog-dogan, J, lortéloran, Md.

Note. The material of C. paniculata RIDL. is
somewhat different from the Javanese C. erycibe
by its being less hairy, by the longer pedicels, the
glabrous sepals, the more orbicular petals, and the
smaller number of stamens; its fruit is unknown.
The only Bornean specimen, collected by HAvi-
LAND (SAR), and the only Sumatran specimens,
Forses 1696 and 1719A, agree quite well with the
type of C. paniculata.

5. Capparis cantoniensis Lour. FI. Coch. (1790)
331; ed. WILLD. (1793) 404; DC. Prod. 1 (1824)
253; MerR. Comm. Lour. (1935) 173.—C. salac-
censis BL. Bijdr. (1825) 54; Mia. FI. Ind. Bat. 1,
2 (1858) 101; Illustr. (1870) 23, t. 12A (excel. var.
celebica Mig., quae est C. lanceolaris); BACK.
Schoolfl. (1911) 61; Koorp. Exk. Fl. Java 2
(1912) 294; RipL. J. Fed. Mal. St. Mus. 84 (1917)
15; Back. Bekn. FI. Java (em. ed.) 4A (1942) fam.
45, p. 9.—C. pumila CHAMP. in Hook. J. Bot. Kew
Gard. Misc. 3 (1851) 260; Hook. f. & Tu. FI. Br.
Ind. 1 (1872) 177; GaGn. Fl. Gén. I.-C. 1 (1908)
188.—C. hasseltiana Mia. Illustr. (1870) 24, t. 13.
—C. celebica Mia. l.c. 26.

Slack climber, 3-20 m. Twigs almost straight,
angular and puberulous when young, terete and
glabrescent when older; internodes c. 144-4 cm.
Thorns patent to recurved, 2-5 mm long, es-
pecially on flowering branches minute or wanting.
Leaves subcoriaceous, oblong to _ lanceolate,
(2.3-)214-4(-5) times as long as wide, sometimes
ovate, rarely obovate, above glabrous, beneath
sparsely puberulous but soon glabrescent, young
leaves salmon-reddish, (4-)5!4-1014 by (114-)
2-3 1,(-4) cm; base obtuse to acutish, top narrow-
ed, acuminate, tip often blunt, sometimes slender

and acutish, mucronate; midrib above sulcate all
Over; nerves inconspicuous, 6—9(-11) pairs;
petiole 5—6(-10) mm, hairy as the twig. Flowers
sometimes fragrant, in axillary subumbels which
are often arranged in a terminal panicle c. 15—20
cm long, each a few cm peduncled, sparsely hairy.
Pedicels slender, 14-2 cm. Bracts subulate, 1-2
mm long, caducous; bracteoles basal, minute,
sometimes wanting. Buds globular, 4-5 mm diam.
Outer sepals 414-6(-7) mm diam., sometimes
sparsely puberulous at the base outside, inner
sepals elliptic to obovate, 5—7(—8) by 4-6 mm, with
membranous, ciliate margin. Petals white (some-
times greenish or pinkish?), (3!4-)5-614 by
2-3(-4) mm, mostly obovate, pubescent. Stamens
20-45, 15-25(-32) mm long, filaments white.
Gynophore 4-12 mm, ovary approximately ellip-
soid, 114 by 1 mm, both glabrous. Fruit globular
to ellipsoid, 1-114 cm diam., pericarp thin,
leathery, smooth. Seeds one to few, globular and
5-6 mm to elliptic and 10 by 7 by 5 mm.

Distr. India (Sikkim, Khasia, and Assam),
Burma (Dawna Range), S. China, Hainan, Indo-
China, and the Andamans; in Malaysia: Central
Sumatra, Java, Lesser Sunda Islands (Bali,
Lombok), Celebes, Philippines (Mindanao),
Moluccas (Sula Is: Mangoli, Buru). Fig. 10.

Ecol. Forests and forest edges, frequently in the
shade, seems to prefer moist places. Fi. fr. Jan.—
Dec.

In the Himalaya, Sumatra, and Java it is found
between (700—)1200 and 1750(—2000) m. In Hong
Kong and vicinity it occurs near sea-level, and the
same is probably the case in the Philippines and
the Sula Is.

Fig.

10. Distribution of Capparis cantoniensis
Lour. The three areas north of Malaysia form
one whole.

Vern. Endog éndogan, kidjéruk, ségore tjalot
(2), sigar djalak, J, sanik lakik, Md.

Notes. In the two Lombok specimens, collected
by Expert, the leaf-top is rounded to acutish.

A few specimens from Mindanao were originally
identified as C. sepiaria, and indeed show resem-
blance to that species, as the leaves are compara-

Dec. 1960]

CAPPARIDACEAE (Jacobs) d Ud

tively small with a slightly notched top. On ac-
count of the midrib, however, which is narrowly
sulcate, I reckon them to belong here. They are
from altitudes below 300 m.

6. Capparis longestipitata HEINE, Mitt. Bot.
Staatssamml. Miinch. Heft 6 (1953) 210; Pf.
Samml. Clemens (1953) 41.

Young parts with a greyish, very short in-
dument, glabrescent; twigs terete, straight, inter-
nodes c. 1!4 cm; thorns hardly or not developed.
Mature /eaves firmly herbaceous, glabrous, ob-
long to slightly obovate, c. 5-6 by 2'4(-3) cm,
base rounded, top acuminate, tip 4-7 mm long,
acute; midrib above sulcate in the basal half,
otherwise flat; nerves 5—7 pairs, thinly prominent
on both sides, reticulation distinct; petiole c.
1 cm long, hairy as the twig. Subumbels c. 15-
flowered, in the axils of the higher leaves and
some terminal, c. 114-214 cm peduncled. Pedicels
c. 8-15 mm, on small distinct cushions; bracts
subulate, few mm long, very soon caducous. Buds
globular, c. 3 mm diam. Outer pair of sepals c.
3 mm diam., outside densely greyish puberulous,
the inner ones smaller and flattish and hairy only
in the centre. Petals c. 4 by 2 mm, glabrous, white.
Stamens c. 20-30, c. 7 mm long. Gynophore
20-25 mm, glabrous; ovary spindle-shaped, sub-
ovoid, 114 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 m respectively.

Note. The Indo-China specimen, CHEVALIER
38671, differs slightly in having thorns 3—4 mm,
the leaves */,-11/, cm acuminate and brown-pu-
berulous on the midrib underneath, the sepals
5—51/, by 31/,-4 mm, the petals 6 by 2 mm and pu-
berulous inside, + 18 stamens 1'/,—2 cm, and the
gynophore 3 cm long.

I have compared C. Jongestipitata 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. /anceolaris it differs by
the far smaller flowers.

7. Capparis lanceolaris DC. Prod. 1 (1824) 248;
Mia. FI. Ind. Bat. 1, 2 (1858) 101; BAck. Schoolfl.
(1911) 62; Koorp. Exk. Fl. Java 2 (1912) 294;
BACK. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45,
p. 9.—C. subspinosa Roxs. Fl. Ind. ed. CAREY 2
(1832) 568.—C. roxburghii (non DC.) SPAN.
Linnaea 15 (1841) 166.—C. elliptica SPAN. I.c.—
C. platyacantha Turcz. Bull. Soc. Nat. Moscou
27, 2 (1854) 323.—C. callophylla (non BL.) Mia.
Pl. Jungh. (1855) 397; Fl. Ind. Bat. 1, 2 (1858) 101,
p.p.—C. salaccensis var. celebica Mia. Illustr.
(1870) 23, t. 12b.—C. oblongata MERR. Gov. Lab.
Publ. Philip. 7. 35 (1906) 15; Philip. J. Sc. 1 (1906)
Suppl. 58; En. Philip. 2 (1923) 212.—C. copelandii
Ermer, Leafl. Philip. Bot. 2 (1910) 680; Merr. En.
Philip. 2 (1923) 210.—C. torricellensis LAutT. Bot.
Jahrb. 52 (1914) 112.—C. viridis ELmer, Leafl.
Philip. Bot. 8 (1919) 3076.

Scandent shrub, few (up to 20?) m high, rarely
self-supporting and c. 114 m high. Twigs mostly
overhanging, straight, fulvous to red-brownish
puberulous when young, sooner or later (rarely
not) glabrescent but nearly always vestiges of the
pubescence persistent near the leaf axils; inter-
nodes c. 2-5 cm. Thorns mostly present, recurved,
up to 7 mm long. Leaves subcoriaceous, above
glabrous, beneath mostly glabrous, sometimes
fulvous-puberulous, whether or not glabrescent,
(1.5—)2.0-4.0(—4.7) times as long as broad, broad-
est in the middle, (3'14—-)614-12(-16) cm long,
(134-)2-4(-7) cm broad; base acutish to rounded
or subcordate, top acuminate, rarely rounded to
subemarginate, tip mostly acute-mucronate, marg-
in often markedly recurved, especially towards the
top; midrib above sulcate mostly all over, rarely
flat; nerves 6-10 pairs, hardly visible; petiole
(14-)34-1(-114) cm, hairy as the twigs. Subumbels
mostly axillary, sometimes terminal, mostly
simple, sometimes branched; peduncle 2—7 cm,
occasionally leafy and up to 15 cm, more or less
glabrescent. Pedicels (114—)2-214(-3) cm, gla-
brous, sometimes each with a pair of distinct
thorns at the base. Bracts small, caducous, narrow,
hairy. Buds globular, 5—6(-8) mm diam. Flowers
white, yellow-white, pink or red, whether or not

Fig. 11. Distribution of Capparis lanceolaris DC.

fragrant. Sepals c. 6—7(—10) by 5 mm, with mem-
branous margin, glabrous; outer pair herbaceous,
inner pair flatter and thinner, rarely minutely
ciliate. Petals obovate, somewhat oblique, thin,
8-11 by 4-6 mm, puberulous towards the base,
especially inside. Stamens c. 20 (40), 2-3 cm.
Gynophore (2—)3—4(—S) cm, ovary ellipsoid, 1-2 mm
long, both glabrous. Fruits few; pedicel, torus, and
gynophore but little incrassate. Fruit (sub)globu-
lar, 214-31!4 cm, bluish black. Seeds 3 or more, c.
8 by 6 by 5 mm.

Distr. Malaysia: M.-E. Java (also Nusa Ba-
rung), Madura, Lesser Sunda Islands (Timor,
Damar, Jamdena), Celebes (also Salajar and
Buton), Philippines (not known from Palawan),
Moluccas (Ceram, Nusa Laut), New Guinea (N.
part from Manokwari to the Sepik and Papua,
also Schouten I.), New Ireland. Fig. 11.

78 FLORA MALESIANA

[ser. I, vol. 6!

Ecol. Secondary (or primary) forests, thickets,
or hedges, mostly on dry, calcareous, rocky soil,
also in coastal vegetation, up to 700 m, once at
1650 m.

Vern. Gagaan, tjantélan, J, kengkeng, Md,
nonoh mukeh, Timor; Philippines: sulu-sulu, Bag.,
kamit-kabag, Dum.

Notes. In sterile state very similar to C.
floribunda, but this species has commonly some-
what longer petioled, mostly ovate leaves narrow-
ing gradually towards the apex, whereas a sharply
acuminate leaf tip is typical for C. /anceolaris,
especially in the Philippines. Some sterile New
Guinean specimens resemble C. zippeliana (see
there).

In the Philippines some deviating specimens
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 cm long. This latter feature, however,
varies even in different duplicates of the type
collection (R. MeryYER 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
found in the inflorescence and by comparatively
small leaves (314-8 cm).

8. Capparis floribunda WiGur, Ill. Ind. Bot. 1
(1840) 35, t. 14; Hook. f. & Tu. Fl. Br. Ind. 1
(1872) 177; F.-VitLt. Nov. App. (1880) 11.—
Crataeva octandra BLANCO, FI. Filip. (1837) 400,
ed. 2 (1845) 280. ed. 3, 2 (1878) 155, non
Capparis octandra JACQ.—C. luzonensis TURCZ.
Bull. Soc. Imp. Nat. Mosc. 27,2 (1854) 324; MerRR.
Sp. Blanc. (1918) 159; En. Philip. 2 (1923) 212,
incl. var. ampla MERR. I.c.—C. andamanica KiNG,
Ann. R. Bot. Gard. Calc. 5 (1896) 119, t. 137.
—C. oligostema Hay. Ic. Pl. Form. 3 (1913) 22.

f. floribunda.

Young parts glabrous, rarely very soon glabres-
cent.—Shrub or climber (?), few m high. Twigs
straight or slightly zig-zag, terete, smooth, inter-
nodes 2—5 cm; thorns small, recurved, mostly
wanting. Leaves firmly herbaceous, broadest at
the middle or sometimes below, mostly narrowed
in the upper half, mostly 214-3 times as long as
wide, (4~-)6—10(-13) by (114-)214-4(-6) cm; base
mostly rounded and more or less acute, top
variable; midrib above often narrowly sulcate;
nerves 7—9 pairs; margin often slightly recurved;
petiole 14-114(—2) cm. Flowers numerous, white,
fragrant, in small subumbels, c. 1-2 cm stalked,

arranged in a terminal panicle up to 15 by 10 cm,
with some additional smaller axillary panicles.
Bracts linear, 2-6 mm, sooner or later caducous.
Buds globular, 3-4 mm diam. Sepals 2-4 by
114-214 mm, ovate, patent and persistent for
some time after anthesis, outer ones patent, not or
narrowly membranous-margined, inner ones
sometimes broader, with a broad, membranous
margin. Petals very thin, oblong or ovate, 3-5 by
114-2 mm. Stamens 7—9(-12), 6-8 mm. Gynophore
4-6 mm (in Ceylon up to 10 mm); ovary ovoid,
acutish, 1-114 by 14-1 mm, both glabrous. In
fruit the pedicel, torus and gynophore only

slightly thickened. Fruit globular, + 114-2(-2'4)
cm diam., soft, fleshy, pericarp coriaceous, smooth
orange. Seeds 1-3, + 13 by 10 by 6 mm.

Fig. 12. Localities of Capparis floribunda W1iGHT

(e) and of its f. induta Jacoss (+). The locality

near the Malay Peninsula (Koh Ha) is approx-
imate. Also in the Deccan and Formosa.

Distr. Ceylon, Deccan, Indo-China (Annam),
Formosa, Peninsular Siam, Andamans; in Malay-
sia: Java (only Kangean Arch.), Philippines
(Debangan I. near Palawan, Babuyan I., Luzon),
Moluccas (Halmaheira, Tidore, Sula Is, W.
Ceram, Kai Is). Fig. 12.

Ecol. This showy plant seems to prefer dry
country and rocky soil, sometimes in coastal
vegetation, in the lowlands; once at 1600 m. Fi. fr.
Jan.—Dec.

Eimer found “many fruits opened and seeds
eaten by birds’’.

f. induta JAcoss, f. nova.

A f. floribunda differt indumento denso fulvo
puberulo in partibus juvenilibus.

A dense, fulvous-puberulous indument on
twigs, young leaves, and inflorescences. Outer
sepals at the base somewhat hairy and petals
slightly ciliate.

Distr. Indo-China (Annam); in Malaysia:
E. Java (Surabaja Residency, Muning, fl. ix-1927,
DE Voocp 673 (L, type; Bo)).

Vern. Wangon lanang, J. Fig. 12.

Dec. 1960]

CAPPARIDACEAE (Jacobs) : 79

Notes. The species is characterized by the
glabrous appearance, with small or without thorns,
the large number of small flowers with 8 stamens,
subpersistent sepals, and a short gynophore which
is more vigorous than in C. sepiaria; the latter
species has also a lighter coloured fruit.

The only specimen known from the mainland
of Java is the specimen described as f. induta
Jacops. In sterile state it can look like C. lanceo-
laris (see there).

The leaves are variable in width. TURCZANINOW
described C. /uzonensis on the specimen with the
narrowest leaves that I have seen, the leaf index
being 3.7—-4.1. MERRILL’s var. ampla actually
represents the normal leaf-shape; intergrading
specimens are by no means rare.

9. Capparis lobbiana Turcz. Bull. Soc. Nat.
Moscou 27, 2 (1854) 323; RoLFE, J. Bot. 23 (1885)
210; VIDAL, Rev. Pl. Vasc. Filip. (1886) 47.—
C. sepiaria var. acuta VIDAL, 1.c.—C. littoralis
Merk. Philip. J. Sc. 7 (1912) Bot. 270; En. Philip.
2 (1923) 211.—C. loheri Merr. Philip. J. Sc. 7
(1912) Bot. 270; En. Philip. 2 (1923) 211.—C. pala-
wanensis Merr. Philip. J. Sc. 10 (1915) Bot. 304;
En. Philip. 2 (1923) 212.—C. ilocana MEeERR.
Philip. J. Sc. 13 (1918) Bot. 13; En. Philip. 2
(1923) 211.—Fig. 13.

Climber 1!4-3(-4) m high. Twigs slender,
straight or slightly zig-zag, with leaves not seldom
in two rows, densely clothed with up to 14 mm
long, straw-coloured hairs, rarely almost glabrous
or soon glabrescent; internodes 1-—2(—3) cm.
Thorns almost straight, up to 2, rarely 5 mm, thin,
dark, with lighter base, pointing upwards, ex-
ceptionally downwards. Leaves herbaceous, broad-
est below, rarely at the middle, (1.5—)1.8—2.5
(4.3) times as long as wide, glabrous or sparsely
hairy above, hairy beneath, especially on the
nerves, 4-8(—15) by 2—314(—5S) cm; base cuneate,
sometimes rounded, rarely acute, top narrowed,
acute to (rarely long-) acuminate, often with a
nerf-tip; midrib above shallowly sulcate in the
lower part, yellowish, sparsely hairy underneath;
nerves 6-8 pairs; petiole 2-4(-12) mm, hairy as are
the twigs. Subumbels axillary; peduncle 14-6 cm,
thin, bearing sometimes one or more small leaves,
hairy. Pedicels slender, 114-3 cm, mostly with
distant hairs, rarely glabrous. Bracts inconspicu-
ous, narrow. Buds globular to depressed-globular,
5-6 mm diam. Flowers white or pale pink. Sepals
persistent for a short time, outer pair 5-6 mm
diam., herbaceous, hairy (or rarely glabrous),
inner sepals 6-7 mm diam., thinner, ciliate. Petals
very thin, ovate, 2-4 by 2!4-3 mm, ciliate. Stamens
15 to 60, 8-12 mm long, anthers 1-114 by 14-1
mm. Torus obscure. Gynophore 114-3(-434) cm,
glabrous; ovary ovoid, c. 2 by 1 mm. In fruit
the pedicel, torus, and gynophore only slightly
incrassate. Fruit globular, c. 114 cm diam., peri-
carp leathery, thin, smooth, glossy, blackish.
Seeds few, subellipsoid, c. 14 cm long.

Distr. Malaysia: Endemic in the Philippines
(not known from the Sulu Arch.).

Ecol. Prefers dry, rocky conditions, also in

primary forests, bamboo thickets, efc., up to
600 m.

Vern. Manungal-lalaki, Mindoro.

Notes. A variable species which can always be
distinguished from C. Janceolaris by the straight,
though sometimes little developed thorns.

Capparis palawanensis MERR. represents a form
with unusually large leaves, but transitions with
the average leaf-size occur. C. Jittoralis MERR.
represents obviously a glabrous form; in some
specimens of typical /obbiana the leaves are also
soon glabrescent. There is but poor material of
C. ilocana MERR.; the type and another specimen
agree mutually very well and seem to represent
another glabrous paramorph with almost abortive
thorns and subcoriaceous leaves.

10. Capparis sepiaria LINNE, Syst. Nat. ed. 10, 2
(1759) 1071; Sp. Pl. ed. 2, 1 (1762) 720; Burm. f.
Fl. Ind. (1768) 118; DC. Prod. 1 (1824) 247,
incl. B glabrata; Roxs. Fl. Ind. 2 (1832) 568; W.
& A. Prod. 1 (1834) 26; DECNE, Nouv. Ann. Mus.
Hist. Nat. Paris 3 (1834) 427; Herb. Timor.
Descr. (1835) 99; SPAN. Linnaea 15 (1841) 166;
JACQUEM. Voy. Ind. Bot. 4 (1844) 21, t. 24; Gray,
U.S. Expl. Exp. Bot. 1 (1854) 70; Mig. FI. Ind.
Bat. 1, 2 (1858) 101; Sum. (1860) 159; Illustr.
(1870) 27; Hook. f. & Tu. Fl. Br. Ind. 1 (1872) 177,
incl. var. vulgaris; KURZ, Fl. Burm. 1 (1877) 66;
FERN.-VILL. Nov. App. (1880) 11, cum var.;
NAvVEs in Blanco, FI. Filip. ed. 3 (1880) t. 209;
VIDAL, Phan. Cuming. (1881) 94; Sinopsis Atlas
(1883) 13, t. 6 f. A; Hemsv. Bot. Chall. 3 (1884)
120; VIDAL, Rev. Pl. Vasc. Filip. (1886) 47; KING,
J. As. Soc. Beng. 58, ii (1889) 393; Kurz ex
PrAIn, J. As. Soc. Beng. 59, 1i (1890) 277, incl. var.
grandifolia; ibid. 60, 1i (1891) 302; ibid. 62, ii
(1893) 65; BAck. FI. Bat. (1907) 59; Merr. Philip.
J. Sc. 3 (1908) Bot. 77; GaGn. FI. Gén. I.-C. 1
(1908) 191; Back. Schoolfl. (1911) 61; Rip. J. Str.
Br. R. As. Soc. n. 59 (1911) 68; Koorp. Exk. FI.
Java 2 (1912) 294; Merr. Fl. Manila (1912) 215;
Ript. Fl. Mal. Pen. 1 (1922) 122; Merr. En.
Philip. 2 (1923) 212, excl. var. acuta VIDAL in
synon., quae est C. lobbiana; Crats, Fl. Siam. En.
1 (1925) 83; C. T. Wuite, J. Arn. Arb. 10 (1929)
217; GaGn. FI. Gén. I.-C. Suppl. 1 (1939) 165;
BAcK. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45,
p. 9.—C. umbellata R. Br. ex DC. Prod. | (1824)
247; C. T. WuiTeE, Proc. Roy. Soc. Queensl. 34
(1922) 31.—C. emarginata PREsL, Rel. Haenk.
2 (1835) 85, non A. RICH.; FERN.—VILL. Nov. App.
(1880) 11; Merr. En. Philip. 2 (1923) 211.—
—C. retusella THw. En. Pl. Zeyl. (1864) 16
= C. sepiaria var. retusella THw. lI.c. 400; FERN.
—VILL. Nov. App. (1880) 11. —C. subacuta Mia.
Illustr. (1870) 35, p.p., quoad specim. halmah.
—C. trichopetala VAL. Bull. Dép. Agr. Ind.
Néerl. 10 (1907) 14 = C. sepiaria var. trichope-
tala VAL. I.c. 72; Laut. Bot. Jahrb. 52 (1914)
112.—C. affinis MeErRR. Philip. J. Sc. 10 (1915)
Bot. 303; En. Philip. 2 (1923) 210.

Wide, much-branched shrub, few m high, some-
times climbing. Young shoots densely fulvous or
greyish puberulous, sooner or later glabrescent;

80 FLORA MALESIANA [ser. I, vokage

Fig. 13. Capparis lobbiana Turcz. a. Branch with fruits and flowers, x 24, b. bud, * 3 (WILLIAMS 382
and BS 85241).

Dec. 1960]

CAPPARIDACEAE (Jacobs)

81

twigs stout, zig-zag, terete; internodes 114-314
cm. Thorns generally vigorous, recurved, 3—S mm
long. Leaves firmly herbaceous to subcoriaceous,
when dry mostly greyish green (in some Philippine
specimens brownish), elliptic, sometimes obovate,
or ovate, exceptionally linear, hairy as the twigs,
upper surface glabrescent first, often with scattered
minute warts, (114—)1.8—2.3(-4) times as long as
wide, (114-)314-514(-10) by (1-)14-24%4(-4) cm;
top mostly rounded, nearly always notched,
rarely blunt; midrib flattish above, sometimes
slightly sulcate at the base; nerves 4—6(—8) pairs,
not very distinct; petiole 2-4(—7) mm. Subumbels
few- to 20-flowered, at the end of small lateral
twigs, rarely terminal. Bracts small, hairy, early
caducous. Pedicels 34—2(—214) cm, glabrous. Buds
globular, 4(-5) mm diam. Sepals ovate, 4-6 by
3-5 mm, occasionally finely ciliate, outer pair
herbaceous with narrow membranous margin;
inner pair somewhat smaller, very thin, mem-
branous towards the margin. Petals 414-714 by
1144-3 mm, very thin, white, more or less pubes-
cent, especially outside at the base of the upper
pair. Torus inconspicuous. Stamens 30-45, 7-12
mm, filaments white, anthers 114 by 1 mm.
Gynophore (4-)6—-10(-13) mm, glabrous; ovary
ovoid, 114-2 by 1 mm, glabrous. In fruit the torus
and gynophore somewhat incrassate, the pedicel
hardly so. Fruit rather fleshy, (sub-)globular,
1—2-seeded, 1-114 cm diam., pericarp subcoriace-
ous, smooth, whitish-yellowish to almost blackish.
Seeds + 8 by 5 by 4 mm.

Fig. 14. Distribution of Capparis sepiaria L. in
Malaysia.

Distr. Ceylon, India (Deccan, Sind, Punjab)
to SW. Burma (Diamond I.), Andamans, SE.
China (Kwangtung), Indo-China, Siam, and
Hainan to Australia (Arnhem Land, Queensland,
and New South Wales to c. 32° SL), in Malaysia:
Malay Peninsula (Kedah, Kelantan), North
Borneo, Java (in W only twice found at the N.
coast), Madura, Kangean Arch., Lesser Sunda
Islands (Bali, Nusa Penida, Sumbawa, Alor,
Timor, Leti, Babar, Wetar), Celebes (also Salajar
I. and Buton), throughout the Philippines,
Moluccas (Halmaheira, Buru, Ambon, Kai Is),
New Guinea (near Merauke, S. Papua). Fig. 14.

Ecol. Drier places in thickets, hedges, teak-
forests, efc., in the lowlands, often near the seaside,
solitary or in groups, obviously bound to seasonal
climatic conditions. When in fruit, often a good
deal of the leaves is shed.

Vern. Poka(n), Md.

Uses. The plant is said to be have some medi-
cinal use (cf. QUISUMBING, Med. PI. Philip. 340).

Notes. It seems not unlikely that this species
occurs also in Africa under the names C. corym-
bosa LAMK and C. tomentosa LAMK, but this
question deserves further research.

In the eastern part of Malaysia specimens are
found which are intergrading between typical C.
sepiaria and the Australian C. umbellata; they
have straighter stems with less developed or often
wanting thorns, larger, ovate, soon glabrescent
leaves, fewer flowers in a shorter stalked and more
often terminal and (sub)sessile subumbel. In a few
specimens the midrib is sulcate all over.

C. sepiaria and C. cantoniensis are easily dis-
tinguished by the midrib, flattened in the former
and narrowly sulcate in the latter.

Inone sterile specimencollected by GAUDICHAUD
near Manila the leaves are extremely narrow, viz c.
10 times as long as wide (6-7 mm). A similar
phenotypic variation was found in C. quiniflora.

11. Capparis diffusa Ripv. J. Str. Br. R. As. Soc.
n. 59 (1911) 68; FI. Mal. Pen. 1 (1922) 122;
HENDERS. J. Mal. Br. R. As. Soc. 18 (1939) 35.

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 214-4 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. 114 cm,
anthers small, white. Gynophore c. 11% cm,
glabrous; ovary subglobular, acute, c. | mm long.
Fruit unknown.

Distr. Malaysia: N. Sumatra (P. Weh), Malay
Peninsula (Perlis), twice collected.

Ecol. On (limestone) rocks. F/. Dec.

12. Capparis callophylla BL. Bijdr. 2 (1825) 53;
Koorp. Minah. (1898) 342; Back. Schoolfl.
(1911) 62; Bekn. Fl. Java (em. ed.) 4A (1942) fam.
45, p. 10.—C. tylophylla SPRENG. Syst. Nat. 4
(1827) 204; Mia. Choix (1864) t. 2; Fl. Ind. Bat.
1, 2 (1858) 101; Illustr. (1870) 22; Koorp. Exk.
Fl. Java 2 (1912) 294; Merr. En. Philip. 2 (1923)
213.—C. cumingii MERR. & ROLFE, Philip. J. Sc. 3
(1908) Bot. 101.—C. turczaninowii ELmer, Leafi.
Philip. Bot. 5 (1913) 1755.—C. mucronata ELMER,
lc. 1757.—C. robusta HEINE, Mitt. Bot. Staats-

82 FLORA MALESIANA

[ser. I, vol. 6!

samml. Miinch. Heft 6 (1953) 211; Pfil. Samml.
Clemens (1953) 41.

Climbing, glabrous shrub, few m high. Twigs
stout, (reddish-)brown when dry; internodes 2—7
cm. Thorns often wanting, if present recurved, up
to 5 mm long. Leaves coriaceous, often reddish
brown when dry, broadest in the middle, (1!4-)
2-3(-314) times as long as wide, (8—)14!4—20
(—26) by (314-)5-9(-14) cm; margin slightly re-
curved; base more or less acute to rounded, some-
times subcordate, top rounded (or emarginate)
and mostly with a hardened tip or shortly and
bluntly acuminate; midrib above flat to sulcate;
nerves c. 6—7 pairs, subprominent above; petiole
1—2(-214) cm, stout, dark-coloured, rough. In-
florescence a more or less leafy, terminal panicle
of subumbels. Pedicels 2-414 cm. Bracts early
caducous. Buds globular, 8-12 mm diam. Flowers
white, pink or red-brown. Outer sepals coriaceous,
subpersistent, 10-13 by 8 mm, rough; inner sepals
subcoriaceous, 12-14 by 6-10 mm, thinner towards
the margin. Petals oblong to obovate-subspathu-
late, 22-35 by 6-11 mm. Torus 4-5 mm broad.
Stamens c. 50-80, filaments light red, 314-414 cm,
anthers 2-3 by | mm. Gynophore 3-414 cm, ovary
ellipsoid, c. 4 by 2-3 mm, both glabrous. In fruit
the pedicel (especially towards both ends), the
torus, and the gynophore towards the top, con-
siderably incrassate. Fruit globular to ellipsoid,
5-614 cm long, pericarp 2—5 mm thick, smooth,
yellowish orange. Seeds ~%, c. 1 by 4 cm.

Distr. Formosa (a variety), in Malaysia: Sumatra
(Palembang), W. Java, Madura, North Borneo,
Celebes, and throughout the Philippines. Fig. 15.

Except in the Philippines, the species seems to
be rare.

Ecol. Mostly on dry, often calcareous soil, but
also in moister habitats, in secondary forest, etrc.,
from the lowland up to 700, in N. Borneo up to
1700 m. Fi. fr. Jan.—Dec.

Vern. Manunggal, Mindoro, sani, Minahasa.

Fig. 15. Localities of Capparis callophylla BL.;
a variety in Formosa.

13. Capparis zippeliana Mia. Illustr. (1870) 25,
t. 14; ScHerr. Ann. Jard. Bot. Btzg 1 (1876) 5;
K. Sco. & Hour. Fl. Kais. Wilh. Land (1889) 49;
K. Sco. & Laut. Fl. Schutzgeb. (1900) 336;
Laut. Bot. Jahrb. 52 (1914) 114, f. la—d, incl. var.
novohibernica, f. le, et var. novobritannica, f. 1f;
Pax & HoFrM. in E. & P. Pfl. Fam. ed. 2, 17b
(1936) 178, f. 90.—C. dahlii Gita & K. Scu.
Notizbl. Berl.-Dahl. 1 (1896) 208; K. Sco. &
Laut. Fl. Schutzgeb. (1900) 335.—C. carolinensis
KANEH. Bot. Mag. Tokyo 48 (1934) 919, f. 6.

Small climber. Twigs slender, fairly straight,
glabrous, rarely glabrescent; internodes 2-5 cm.
Thorns mostly wanting, if present slightly recurved,
up to 3 mm. Leaves firmly herbaceous, often dull
red-brownish when dry, broadest in the middle,
(1.7-)2.2-2.8(-3.5) times as long as wide, (8-)
13—20(—26) by (4!4-)5—-8(-10) cm, glabrous or
rarely thinly puberulous below; margins sub-
revolute; base rounded or acutish or subcordate,
top (rarely sharply) acuminate, or rarely emar-
ginate; midrib above mostly sulcate in the basal
part, to flat; nerves 5-9 pairs; petiole (34-)
1-114(-134) cm, sulcate above, glabrous, rarely
glabrescent. Subumbels up to 10-flowered, mostly
sparsely puberulous, sometimes axillary but
mostly arranged in a terminal panicle, slender,
2-7(-12) cm peduncled. Pedicels 1—4!4(—6) cm.
Bracts small, subulate, hairy, caducous. Buds
globular, 5—6(—10) mm diam. Sepals subpersistent,
outside sparsely hairy or glabrous, with mem-
branous margin, 5—6(—12) mm diam.; outer pair
subcoriaceous, one often initially enveloping the
bud for 24; inner pair suborbicular, herbaceous.
Petals thin, obovate, 6-8 by 2!4-4 mm, white,
minutely puberulous on both sides, base narrowed,
top rounded (sometimes crisped?). Stamens c.
25-45, (114-)214-3 cm long, white or pale pink;
anthers 114-214 by 14 mm. Gynophore 2-3'4(-4)
cm, often thinly puberulous at the base, soon
glabrescent; ovary ellipsoid, 144 by 1 mm. In
fruit the pedicel, torus, and gynophore are rather
incrassate. Fruit ovoid or ellipsoid, c. 444 by 314
cm, top sometimes umbonate, pericarp thin,
leathery, smooth, red. Seeds oy, c. 12 by 9 by 6mm.

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. Jan-
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, Nouy. Ann. Mus. Hist. Nat. Paris 3 (1834)
246; Herb. Timor. Descr. (1835) 98; DELEss. Ic. Pl.
3 (1837) t. 12; SPAN. Linnaea 15 (1841) 165; Mia.
Fl. Ind. Bat. 1, 2(1858) 100; Illustr. (1870) 27, t. 15,

Dec. 1960]

CAPPARIDACEAE (Jacobs) 83

incl. var. sumatrana et var. moluccana Mia. l.c.
28; KiNG, J. As. Soc. Beng. 58, ii (1889) 394, incl.
var. perakensis ScortT.; HALL. f. in Koord. Minah.
(1898) 343; Riot. J. Fed. Mal. St. Mus. 10 (1920)
129; Back. Bekn. Fl. Java (em. ed.) 4A (1942)
fam. 45, p. 13.—C. nigricans SPAN. Linnaea 15
(1841) 165; Mig. Fl. Ind. Bat. 1, 2 (1858) 100.—
C. cerasifolia A. Gray, U.S. Expl. Exp. Bot. 1

Fig. 16. Capparis pubiflora DC. (Cult. Hort. Bog.
XV-J—B-II-6; JAcoss, 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. lI.c. 322; FERN.—VILL. Nov.
App. (1880) 11; VimpaL, Phan. Cuming. (1885) 94;
Rey. Pl. Vasc. Filip. (1886) 48.—C. myrioneura
HALL. f. in Fedde, Rep. 2 (1906) 60, p.p.—C.
dealbata (non DC.) Back. Schoolfl. (1911) 62.—
C. perakensis (ScorT. ex KING) RipL. FI. Mal.
Pen. 1 (1922) 124.—C. borneensis MeERR. Pl. 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(-4.4) times as long as wide, often
obovate, rarely ovate, (5—)8—16(—27!4) by 214-6
(-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 1—5(—10) in short,
axillary racemes, with a persistent, densely tomen-
tose, subulate bract and 2 smaller basal bracteoles,
or sometimes this inflorescence replaced by a
short young twig with axillary flowers; racemes
and young twigs with a number of minute subulate
bract-like leaves at the base. Pedicels 14—3(—5)
cm, glabrescent. Buds globular to ovoid, c. 5 mm

long. Sepals herbaceous, concave, 414-7 by
214-4 mm, outside puberulous, outer pair acute
or blunt, sometimes slightly cucullate, inner pair
with blunt or rounded top. Petals thin, mostly
obovate, 7—10 by 3-4 mm, outside soft-hairy at
top and margins. Torus conical, disk c. 1 mm.
Stamens 20—30(—50), filaments 15—20(—25) mm,
anthers elliptic, 1 by 14 mm. Gynophore 15—25 mm,
densely tomentose, in fruit glabrescent; ovary
ellipsoid, 244 by 114 mm, hairy as the gynophore,
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. 17. Capparis pubiflora DC., flower. a. Corolla,
front view, stamens and gynophore removed, x §/;,
b. calyx (one sepal removed) and disk, x 2,
c. calyx and corolla from the back, x 214,
d. flower, near-longitudinal section, x 214 (after

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,
roadsides, teak forests, brushwood, jungle, up to
400(-600) m. F/. June-July, fr. March.

Vern. (Daun) poka, saneg-sanegan, Md,
djénggotan, waan-waanan, J, bangol bangol, Bali;
Minahasa: maha-limu, Ratahan, malemo, Ton-
sawang, tutunean woring, Tontemboan, sahamun-
tei; wama pusu, Halmaheira, holiboi, Mbo.

Notes. Some specimens from Sumatra and the
Malay Peninsula, and generally those from
Celebes, the Moluccas, and the Philippines, have
markedly larger, obovate leaves; they cannot be

Fig. 18. Distribution of Capparis pubiflora DC.

distinguished taxonomically from the population
in Java and the Lesser Sunda Islands.

C. MUELLER, /.c., copied A. GRAyY’s name
C. cerasifolia as C. crassifolia. PAX & HOFFMANN
continued this mistake in a remarkable way when
stating: ““Die Ueberleitung zu den Philippinen
bilden die 2 Arten der Sulu-Inseln: 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 «; DC. Prod. 1 (1824) 246; DeELEss.
Ic. Pl. 3 (1837) t. 11; non W. & A. Prod. 1 (1834)
25, quae est C. grandiflora WALL. ex Hook. f. &
TH. — C. acuminata WILLD. Sp. Pl. 2 (1799)
1131; DC. Prod. 1 (1824) 247; Hook. f. & Tu.
Fl. Br. Ind. 1 (1872) 178; Back. Fl. Bat. (1907) 56;
Voorl. (1908) 13; Schoolfl. (1911) 63; Koorp.
Exk. Fl. Java 2 (1912) 293; Back. Bekn. FI. Java
(em. ed.) 4A (1942) fam. 45, p. 7.—C. zeylanica
(non L.) DC. Prod. 1 (1824) 247, quoad specim.
javan.; W. & A. Prod. 1 (1834) 25.—C. foetida
Br. Bydr 2, (1825)0o2; Mio: BEalnd. Bat. 1592
(1858) 99; Gacn. Fl. Gén. I.-C. 1 (1908) 184;
CRAIB, Fl. Siam. En. 1 (1925) 80; GAGn. FI. Gén.
I.-C. Suppl. 1 (1939) 161.—C. dasypetala Turcz.
Bull. Soc. Nat. Moscou 27, 2 (1854) 322.—C.
oxyphylla Miq. Pl. Jungh. 4 (1855) 397; FI. Ind.
Bat. 1, 2 (1858) 100.—C. horrida (non L. f.) Mia.
Illustr. (1870) 34, pro var. « pro parte et synon. C.
foetida et C. oxyphylla.

Shrub, sometimes climbing, 114—214(-314) m;
twigs straight, terete, with minute, stellate,
ferruginous hairs, glabrescent; internodes 2—5 cm.
Thorns patent, straight or slightly curved upwards,
1-3(-4) mm. Leaves on lateral branches mostly
distichous, more or less firmly herbaceous,
elliptic, when young with minute, stellate, fulvous-
ferruginous indument giving it a farinaceous as-
pect, glabrescent, (1.2—)1.7—2.2(—3) times as long
as wide, often ovate, sometimes obovate, 5-914
(-15) by 214-414(-614) cm; base rounded to blunt,
top acuminate with a mostly blunt tip 14-114 cm
long; nerves yellow to light-brown, midrib some-
times sulcate above; nerves c. 5 pairs, veins reti-
culate; petiole c. 14 cm, densely greyish brown
tomentellous. Flowers 2-4, serial. Pedicels (1-)
114-2(-214) cm, thin, densely hairy, glabrescent
except for the somewhat broadened top. Buds
globular, 4-5 mm diam. Sepals elliptic-ovate, 4—5
by 2!4-4 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, 14 mm high,
both glabrous. In fruit neither the pedicel, nor the
gynophore much incrassate. Fruit about globular,
8-12 mm diam.; pericarp minutely rugose when
dry, glossy, black when ripe (once reported red),
glabrous. Seeds 2-6, 6 by 3-4 by 2 mm, smooth,
glossy brown.

Fig. 19. Distribution of Capparis pyrifolia LAMK.

Distr. Siam, Indo-China (Annam, Cochin-
china); in Malaysia: N. Sumatra (Laubalang?),
Java, Kangean Arch., Madura, Lesser Sunda
Islands (Bali, E. Sumbawa). Fig. 19.

Dec. 1960]

CAPPARIDACEAE (Jacobs) © i 85

Ecol. In the lowlands and hills in dry places,
in teak forests, brushwood, hedges, on limestone
hills, fairly common in Java, up to c. 850 m. Fi.
especially Sept., no flowers collected Febr.-May;
fr. July—Febr.

Vern. Kaju tudjuh, S, gagahan, gédangan,
kédéling, lorowan, risini, tjantélan, waan-waanan,
wan-uwanan, J, sanek, sanik-lakek, Md, kaloang-
kaloangan, pokak-pokadn, Kangean.

Uses. According to HEYNE (Nutt. Pl. 1927,
682) the white wood is sold in Djakarta as ‘kaju
tudjuh’ against bile and stomach ache, and an
extract of the rasped wood is taken against
dizziness. Fruit once reported to be sweet and
edible.

Notes. It cannot always easily be observed
that the flowers are serial because the row is
sometimes very short. This could lead to con-
fusion with C. pubiflora, which is characterized,
however, by the hairy gynophore and a fruit over
12 mm diam.

16. Capparis buwaldae JACOBS, sp. nov.

Glabra. Spinae stipulares breves, recurvatae.
Folia + 6 mm petiolata, oblongata, apice acute
acuminata; costa nervique majores supra Ssulcati.
Flores 2-4 in serie supraaxillare dispositi, minores;
alabastra globosa acuta 3 mm diametro; pedicellus
gynophoriumque fructiferum vix incrassatum. Fruc-
tus globosus vel ellipsoideus, umbonatus, 2-4 cm
longus, irregulariter tuberculatus; semina majora.

Typus: HALLIER f. 2573 (Bo, holotype; L, iso-
type).

Climbing shrub or liana, 2-15 m high, glabrous;
twigs terete, slightly zig-zag; internodes 1—8 cm.
Thorns recurved, 1-3 mm long, often wanting.
Leaves firmly herbaceous, oblong, sometimes
ovate or obovate, (2.2—)2.4-4(—5) times as long as
wide, 6—13(—2314) by (114-)2'4-414(-8) cm; base
rounded to acute, top narrowed, acuminate, tip
acute, often mucronulate, 3-20 mm long; midrib
sulcate above; nerves (4—)6—9 pairs, conspicuously
looped-arcuating; petiole 4-7(-10) mm. Flowers
white, as a rule on slender, short, lateral twigs,
serial, 2-4, supra-axillary. Pedicels slender,
1—3(-4) cm long, glabrous. Buds globular, acute, c.
3 mm diam. Sepals ovate, 3-5 by 2-3 mm, with a
slightly thickened and more or less acute top,
outer pair almost glabrous at the margins, inner
pair slightly ciliate. Petals very thin, 4-6 by 2-3
mm, white, outside more pubescent than inside,
often minutely ciliate. Disk bilobed, fleshy, 1 by
144 mm. Stamens 20-30, c. 2 cm long. Gynophore
13-20 mm, ovary ellipsoid to globular, c. 114 by
¥%4-1 mm, smooth, stigma 14 mm long, all gla-
brous. In fruit the pedicel hardly incrassate and
the gynophore slightly so. Fruit orange or red,
globular to ellipsoid, 3-5'4 by 214-314 cm,
shortly umbonate at the top and sometimes also
at the base; pericarp woody to coriaceous, 2-3
mm thick, more or less tuberculate. Seeds ~,
embedded in whitish pulp, 10 by 6-8 by 5-6 mm,
smooth, brown.

Distr. Malaysia: throughout Borneo (also P.
Laut).

Ecol. Forests, jungle, along rivers, from the
lowland up to c. 1600 m. Fi. fr. Jan.—Dec.

Vern. Dunggol manok, Sarawak.

Uses. Two collectors mention the fruit(pulp)
to be edible.

Note. In vegetative characters much resembling
C. micracantha ssp. korthalsiana but distinguished
by the recurved thorns, the narrower leaves with
long and slender tip, and the nerves being more
prominent towards the margin.

17. Capparis cucurbitina KiNG, J. As. Soc. Beng.
58, ii (1889) 395; Ann. R. Bot. Gard. Calc. 5 (1896)
119, t. 136; Rmpv. Fl. Mal. Pen. 1 (1922) 124.

Scandent, 6-10 m high; twigs slightly zig-zag,
(nearly) glabrous; internodes c. 2 cm. Thorns
recurved, 2-3 mm long, sharp. Leaves herbaceous,
obovate, c. 214 times as long as broad, glabrous,
c. 9-1814 by 314-7!4 cm; base generally rather
abruptly rounded and acute, top rounded and
rather abruptly acuminate, tip narrow and acute,
1-114 cm; midrib above sulcate in the basal part;
nerves 6-8 pairs, thinly subprominent above,
reticulations distinct on both sides; petiole
14,34, cm. Flowers in mature state unknown, pale
green, yellow inside, 2-3, serial. Pedicels 2—3 cm.
Buds ovoid, acute, 4 mm long. Sepals ovate-
lanceolate, acuminate. Petals broadly elliptic,
obtuse. Stamens c. 20. In fruit neither the pedicel
nor the torus incrassate, the gynophore 11-17 mm,
slightly thickened, obviously articulated with the
fruit. Fruit irregularly spindle-shaped, (3'4—)
514-74 by 1'4-21%% cm, the base slightly acu-
minate-tapering, the top umbonate; pericarp 1
mm thick, leathery, yellow to glossy orange.
Seeds oy, ovoid, 9 by 7 by 7 mm.

Distr. Malaysia: Malay Peninsula (Perak),
few collections.
Ecol. Dense mixed lowland jungle, up to

200 m.

Notes. The only flowering material which we
have is KinGc’s Coll. 8824 (CAL). Contrary to
KinG I found the sepals not puberulous but
glabrous.

Differs vegetatively from C. micracantha ssp.
korthalsiana in that the leaves of the latter are
much coarser, and thicker, while in C. cucurbitina
the leaf tip is longer and more slender.

18. Capparis micracantha DC. Prod. | (1824) 247.
KEY TO THE SUBSPECIES

1. Ripe fruit (sub)globular. Stamens c. 20—35(—60).
Sepals mostly obtuse ssp. micracantha

1. Ripe fruit elongate, acute. Stamens 60-100.
Sepals acute to acuminate. West Malaysia.
ssp. korthalsiana

ssp. micracantha.—C. micracantha DC. Prod. 1
(1824) 247; BL. Bijdr. 2 (1825) 52; SPRENG. Syst.
Veg. 4, 2 (1827) 204 (micrantha, sphalma, non
A. RicuH.); Mio. FI. Ind. Bat. 1, 2 (1858) 99;
Hook. f. & Tu. Fl. Br. Ind. 1 (1872) 179; Kurz,
J. As. Soc. Beng. 43, ii (1874) 31; RADLK. Sitz.

86 FLORA MALESIANA

[ser. J, yoaGe

Ber. Bayer. Ak. Wiss. 14 (1884) 118, 128; VipAL,
Rev. Pl. Vasc. Filip. (1886) 47; Kina, J. As. Soc.
Beng. 58, ii (1889) 394; Pax in E. & P. Pfl. Fam.
3, 2 (1891) 231; BRANDis, Ind. Trees (1906) 36;
Merk. Philip. J. Sc. 1 (1906) Suppl. 58; Back. FI.
Bat. (1907) 57, incl. var. callosa (BL.) HALL. f.;
GAGN. FI. Gén. I.-C. 1 (1908) 186; BAck. Schoolfl.
(1911) 63; Rip. J. Str. Br. R. As. Soc. n. 59 (1911)
68; HALL. f. Med. Rijksherb. 12 (1912) 18; Koorp.
Exk. Fl. Java 2 (1912) 293; Merr. FI. Manila
(1912) 215; Sp. Blanc. (1918) 160; RipL. Fl. Mal.
Pen. 1 (1922) 123, f. 11; W. H. BRown, Min. Prod.
Philip. For. 2 (1921) 282; Merr. En. Philip. 2
(1923) 212; PARKINS, For. Fl. Andam. (1923) 82;
GEERTS—RONNER, Trop. Natuur 13 (1924) 167,
f. 8; Crars, Fl. Siam. En. 1 (1925) 82; Merr.
Philip. J. Sc. 29 (1926) 371; GaGn. FI. Gén. I.-C.
Suppl. 1 (1939) 162; CoRNER, Ways. Trees (1940)
180, f. 47; BAcK. Bekn. Fl. Java (em. ed.) 4A
(1942) fam. 45, p. 7; Quis. Med. Pl. Philip. (1951)
339.—C. billardierii DC. Prod. 1 (1824) 247;
Mia. FI. Ind. Bat. 1, 2 (1858) 99.—C. callosa BL.
Bijdr. 2 (1825) 53; Mia. Fl. Ind. Bat. 1, 2 (1858)
99; Illustr. (1870) 29, t. 16; NAves in Blanco, FI.
Filip. ed. 3 (1877-83) t. 180; RADLK. Sitz. Ber.
Bayer. Ak. Wiss. 14, 1 (1884) 131; K. & V. Bijdr.
4 (1896) 262; Koorp. Gedenkb. Jungh. (1910)
167.—C. flexuosa BL. Bijdr. 2 (1825) 53; non L.,
nec VELLOZO; HAssk. Pl. Jav. Rar. (1848) 178;
Mia. Analecta 3 (1852) 1; Fl. Ind. Bat. 1, 2 (1858)
98; Illustr. (1870) 30; RADLK. Sitz. Ber. Bayer.
Akad. Wiss. 14, 1 (1884) 101, 129.—C. odorata
BLANco, FI. Filip. (1837) 439, ed. 2 (1845) 305,
ed. 3, 2 (1878) 201; FEeRN.—-VILL. Nov. App.
(1880) 11; Merr. Philip. J. Sc. 3 (1908) Bot. 77.-
C. forsteniana Mia. Illustr. (1870) 32, t. 18, excl.
syn. C. ovalifolia Zipp., quae est C. zeylanica L.;
HALL. f. in Koord. Minah. (1898) 343.—C.
roydsiaefolia Kurz, J. As. Soc. Beng. 39, ii (1870)
62.—C. myrioneura HALL. f. in Fedde, Rep. 2
(1906) 60, p.p., excl. Koorp. 16341, excl. var.
latifolia.—C. venosa MeErR. Philip. J. Sc. 10 (1915)
Bot. 305, ex descr.; En. Philip. 2 (1923) 213.
Stout shrub or small tree, rarely climbing,
2-6(-10) m; trunk greyish, finely fissured and set
with small knobs each surmounted by a thorn;
branchlets terete, mostly zig-zag, when young
sparsely pubescent; internodes c. 214-4 cm.
Thorns 2-4(-7) mm long, patent or directed up-
wards, straight or slightly curved. Leaves (sub-)
coriaceous, glabrous, 1.7—2.9(-4.1) times as long
as wide, mostly broadest about halfway, sometimes
below, or rarely above the middle, 8—18(—24) by
4-8(-121!4) cm; base mostly rounded, sometimes
blunt to subcordate or acute, top broader or
narrower rounded, sometimes slightly emarginate,
or acute, rarely acuminate, dark-tipped; midrib
subprominent above, nerves 5—7(—10) pairs, light
green when dry; petiole 6—-10(-15) mm. Flowers
up to 6, serial. Pedicels 1(—2) cm. Buds ellipsoid,
acute, 5-12 mm long. Sepals firmly herbaceous,
514-13 by 214-514 mm, + boat-shaped, ovate to
oblong, the margins mostly hairy. Petals very thin,
oblong to lanceolate, 10-25 by 3-7 mm, white,
honey-guide yellow, turning dark red or brownish,

or dark violet. Disk bilobed, c. 1 mm. Stamens c.
20—-35(—60); filaments 18-30 mm, white; anthers
144-244 by 4 mm, grey or bluish. Gynophore
15—30(—35) mm, ovary ovoid to ellipsoid, c. 3 by
2 mm, both glabrous, sometimes vestigial. In
fruit the gynophore 4-6 mm diam., the pedicel
thinner. Fruit globular to ellipsoid, with 4 longitu-
dinal sutures, 3-614 by 3-414 cm, yellow, orange,
or red; pericarp smooth, 2 mm thick, when dry
woody-coriaceous, pulp juicy. Seeds c. 6-8 by
414-7 by 3-5 mm, red to shiny black, smooth.

Fig. 20. Distribution of Capparis micracantha DC.

ssp. micracantha (continuous line, add Formosa)

and of ssp. korthalsiana (MiqQ.) JAcops (broken
line). Localities of C. zippeliana Mia. (dots).

Distr. Burma, Siam, Indo-China, S. China,
Formosa (a variety), Hainan, Andamans, in
Malaysia: S. Sumatra, Malay Peninsula (except
in the S. corner), Java (in W only a few coastal
localities, also Prinsen I., Nusa Kembangan),
Madura, Kangean Arch., Lesser Sunda Islands
(Bali, Lombok, E. and W. Sumbawa, W. Flores,
Komodo, Alor, Timor, Wetar), North Borneo
(also Banguey), throughout the Philippines,
Celebes, Moluccas (Halmaheira). Fig. 20.

Ecol. Especially in light shade (monsoon, teak,
or evergreen forest) on dry, often calcareous soil,
also in thickets, savyannahs, hedges, efc., not
seldom coastal, mostly below 500 m, highest
record 1400 m. Fil. fr. Jan.—Dec.

Anat. RAGHAVAN investigated the vascular
supply of the floral parts in a ‘C. flexuosa’ from
Java, cultivated at Kew (the plant was no longer
living in 1959) (J. Linn. Soc. Bot. 52, 1939, 247).

MOLISCH stated that nearly all the parenchyma
cells of the petiole, the leaves, and the stem
contain a strongly refractive crystal of lime. Also
bodies of silicon are sometimes found (Ber. Deut.
Bot. Ges. 34, 1916, 154-160).

Note. C. hbillardierii, C. flexuosa, and C.
roydsiaefolia have been described on specimens
with a vestigial gynophore and ovary. RADL-
KOFER’s sect. Monostichocalyx comprised C.
flexuosa Bu., C. callosa BL., and C. micracantha
DC. which are in my opinion all synonyms.

ssp. korthalsiana (MiQ.) JAcoss, stat. nov.—C.
korthalsiana Mia. Sllustr. (1870) 31, t. 17; MERR.
En. Born. PI. (1921) 280.—C. finlaysoniana WALL.
[Cat. (1832) 6992 B, nomen] ex Hook. f. & TH.

Dec. 1960] 4 CAPPARIDACEAE (Jacobs) = aa 87

Fl. Br. Ind. 1 (1872) 179; Kina, J. As. Soc. Beng.
58, ii (1889) 395; Rip. Fl. Mal. Pen. 1 (1922) 124;
Kew Bull. (1925) 77.

Fruit oblong, c. 6-17 by 2!4-334 cm, tapering
to the top and sometimes to the base. Leaves
acuminate, not mucronate, mostly recurved, never
cordate at the base. Sepals very acute and slightly
cucullate at the top, the buds therefore (sub-)
acuminate. Flowers comparatively large. Stamens
very numerous (up to 100).

Distr. Malaysia: Malay Peninsula (Perak,
Pahang, Johore, Singapore), Central and S.
Sumatra, Borneo. Nowhere common. Fig. 20.

19. Capparis zeylanica LINNE, Sp. Pl. ed. 2 (1762)
720; DC. Prod. 1 (1824) 247, quoad specim. Ceyl.;
Dunn, Kew Bull. (1916) 62; non Hook. f. & Tu.
Fl. Br. Ind. 1 (1872) 174, quae est C. brevispina
DC. — C. horrida L. f. Suppl. (1781) 264; DC.
Prod. 1 (1824) 246; Wicut, Ic. Pl. Ind. Or. 1
(1839) t. 173; Mra. Illustr. (1870) 34, guoad var. B
erythrodasys; Hook. f. & TH. Fl. Br. Ind. 1 (1872)
178; FeRN.—-VILL. Nov. App. (1880) 11; VIDAL,
Rev. Pl. Vasc. Filip. (1886) 48; Merr. Philip. J. Sc.
1 (1906) Suppl. 58; GaGn. FI. Gén. I.-C. 1 (1908)
185; Back. Schoolfl. (1911) 63; Ripv. J. Str. Br. R.
As. Soc. n. 59 (1911) 69; Koorpb. Exk. FI. Java 2
(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; Cram, FI. Siam. En. 1 (1925) 81;
Gaan. FI. Gén. I.-C. Suppl. 1 (1939) 161; BAck.
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 8;
Quis. Med. PI. Philip. (1951) 338.—C. dealbata
DC. Prod. 1 (1824) 246; DECNE, Nouv. Ann. Mus.
Hist. Nat. Paris 3 (1834) 426; Herb. Timor. Descr.
(1835) 98; SPAN. Linnaea 15 (1841) 165; Mia. FI.
Ind. Bat. 1, 2 (1858) 100; Illustr. (1870) 27.—
C. zeylanica (non L.) Roxs. Fl. Ind. ed. CAREY 2
(1832) 567; GAMBLE, FI. Pres. Madras 1 (1915)
18.—C. aurantioides Prest, Relig. Haenk. 2
(1835) 86; Gray, U.S. Expl. Exp. Bot. 1 (1854) 70;
FERN.-VILL. Nov. App. (1880) 11.—C. Jinearis
(non JAcQ.) BLANCO, FI. Filip. (1837) 438, ed. 2
(1845) 305, ed. 3, 2 (1878) 200.—C. nemorosa (non
JACQ.) BLANCO, FI. Filip. (1837) 438.—C. micra-
cantha (non DC.) BLANCO, FI. Filip. ed. 2 (1845)
305, ed. 3, 2 (1878) 200, t. 188.—C. rufescens
Turcz. Bull. Soc. Nat. Moscou 27, 2 (1854)
321.—C. erythrodasys Mia. Pl. Jungh. (1855) 397;
Fl. Ind. Bat. 1, 2 (1858) 99.—C. ovalifolia Zipp.
ex Mig. Illustr. (1870) 33.—C. viminea (non
Hook. f. & TH.) FERN.-VILL. Nov. App. (1880)
11.—C. myrioneura var. latifolia HALL. f. in Fedde,
Rep. 2 (1906) 61.—Fig. 5.

Climbing shrub 2—5(—10) m. Innovations brown-
red to greyish-tomentose; branchlets mostly zig-
zag, glabrescent; internodes 2—7 cm. Thorns re-
curved, 3-6 mm long. Leaves subcoriaceous,
(1.2—)1.7—2.3(—2.9) times as long as broad, ovate
or elliptic, rarely obovate, above soon glabrescent
then glossy, beneath later or not glabrescent, and
dull, 4-10(-18) by 3-S!4(-9) cm; base rounded,
sometimes subcordate, rarely acute, top acute to
rounded, rarely slightly acuminate, generally with

a recurved, stiff, darker mucro up to 3 mm; midrib
subdepressed above; nerves 3-8 pairs; petiole
14-114(-2) cm, glabrescent. Flowers developing
before the leaves on young twigs, conspicuous,
2-6, serial. Pedicels 4—20(—28) mm, hairy. Buds
globular, c. 8 mm diam. Sepals subcoriaceous,
outside more or less densely tomentellous, outer
pair orbicular to elliptic, mostly acute, 6-11 by
5—9 mm, the posterior one (surrounding the disk)
the largest; inner pair elliptic to oblong with more
rounded top, 6-10 by 3-7 mm. Petals very thin,
oblong with rounded top, 9—12(—16) by 314-5 mm,
white, turning pink, largely glabrous; upper pair
with a pinkish to reddish basal-median spot,
hairy at the base. Disk c. 1 mm diam. Stamens
30-—45(—70), 20-30(—-35) mm, white, turning red;
anthers oblong, slightly broader at the base, c.
2 by | mm, bluish grey. Gynophore slightly ex-
ceeding the stamens, up to 4!4(—514) cm, basal
part pale pubescent, otherwise glabrous; ovary
ellipsoid, 1!4-214 by 1-114 mn, stigma 14 mm
long. In fruit the pedicel sometimes still hairy,
gynophore glabrous, up to 514 cm by 3-6 mm, as
thick as the pedicel. Fruit globular to ellipsoid,
up to 5 by 4 cm, pericarp c. 2 mm thick, woody-
coriaceous, smooth, reddish, orange or purple.
Seeds oo, 5-7 by 5-414 mm, brown.

= =<
0 #0

Fig. 21. Distribution of Capparis zeylanica L. in
Malaysia. Add Hainan.

Distr. Ceylon, India (largely E of the line
Bombay-Delhi-Dehra Dun, S of the Himalaya),
Burma, Siam, Indo-China, Hainan, Andaman
Islands, in Malaysia: Java (east of Djokjakarta),
Lesser Sunda Islands (Lombok, Sumbawa, Semau,
Timor), Celebes (also Salajar I.), Philippines (Min-
doro, Luzon, Mindanao, Sulu Arch.). Uncommon
in Indonesia, common in the Philippines. Fig. 21.

Ecol. Forest edges, bushes, savannahs, hedges,
limestone hills, obviously bound to seasonal
climatic conditions, mainly in the lowlands, up
to 700 m. Fi. fr. Jan.—Dec.

Uses. Of less importance; see Quis. /.c.

Vern. Philippines: Manunggal lalaki, Mindoro,
baralduik, Ibn., dauag, halubdgat-baging, Tag.,
laginau, Bis., talaktak, tarabtab, Uk.

88 FLORA MALESIANA [ser. I, vol. 6!

Fig. 22. Capparis quiniflora DC. a. Flowering branch, x 2, b. flower from the back, x 2, c. fruit, x 7,
d-e. narrow leaves, x 24 (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
easily distinguished by the recurved thorns.

20. Capparis quiniflora DC. Prod. 1 (1824) 247;
BENTH. FI. Austr. 1 (1863) 94; F. v. M. Descr.
Not. Pap. Pl. 1+ (1875) 5; BAILeEy, Queensl. Fl. 1
(1899) 57; Laut. Bot. Jahrb. 52 (1914) 112;
Domin, Bibl. Bot. Heft 89 (1925) 685.—C.
trapeziflora SPAN. Linnaea 15 (1841) 165; Mia.
Fl. Ind. Bat. 1, 2 (1858) 99.—C. subcordata SPAN.
Linnaea 15 (1841) 166; Miq. Fl. Ind. Bat. 1, 2
(1858) 99; Illustr. (1870) 34.—C. richii A. Gray,
U.S. Expl. Exp. Bot. 1 (1854) 69; SEEM. FI. Vit.
(1865) 6.—Fig. 22.

Climbing shrub; twigs slightly zig-zag, densely
minutely grey or ferruginous-tomentose, some-
times glabrescent; internodes up to 9 cm. Thorns
recurved, c. 1—2(—3) mm, sharp, sometimes want-
ing, especially on flowering twigs. Leaves sub-
coriaceous to coriaceous, often brownish green
when dry, ovate to (rarely) obovate, 1.6—4.3 times
as long as wide, initially densely minutely ferru-
ginous-tomentose, soon glabrescent and glossy
above, (514-)7-9(-12) by 134-4(-714) cm; base
rounded, subcordate, blunt, or acute, top mostly
attenuate and acuminate, rarely blunt, acumen
up to 14 cm, acutish with a minute, thickened
mucro; midrib flat to shallowly sulcate above;
nerves 6—8(—11) pairs under an angle of c. 45°;
intermediate veins often wanting, reticulations
distinct; margin sometimes crinkled; petiole
5-17 mm, hairy as the twigs. Flowers white, serial,
sometimes developing before the leaves at the
end of young twigs, 2-10 in a row 2-12 mm long,
pedicels 6-17 mm, hairy as the twigs, finally
glabrescent; floral leaves sometimes abortive.
Buds c. 4 mm diam. Sepals herbaceous, acute,
ferruginous-tomentellous outside, outer pair c.
4-5 by 2-3 cm, the inner pair narrower and
flattish. Petals ovate to elliptic, 5-7 by 2-4 mm,
slightly unequal, puberulous on both sides. Disk
up to | mm long. Stamens 7—8(—12), filaments
20-27 mm. Gynophore 2-214 cm but sometimes
abortive and only a few mm, glabrous or glabres-
cent at the base; ovary ovoid, c. 1!4 by | mm,
stigma knob-shaped. Pedicel, torus, and gyno-
phore slightly incrassate. Fruit (sub)globular, c.
2-234 by 2-214, pericarp corky-leathery, c.
114-244 mm thick, with small scattered warts.
Seeds 4, up to 7-8 by 5-6 by 4 mm.

Distr. Pacific (Fiji and New Caledonia), North
Australia to Cape York Peninsula, in Malaysia:
SE. Celebes, Lesser Sunda Islands (Lombok,
Sumbawa, Flores, Semau, Timor), Moluccas
(Kai and Tanimbar Is), New Guinea (NW. part;
Papua; islands in Torres Strait). Fig. 7.

Ecol. Prefers obviously coastal habitats in
drier areas, rambling on shore trees, bound to a
seasonal climate, in the lowland.

Notes. Although in fertile material the leaves
are variable in shape, the venation (see fig. 22a,
d-—e) is constant and very characteristic.

The Australian specimens are much more
uniform in leaf index than those of Malaysia,
where it does not exceed 1.8.

In the material described by A. Gray from Fiji
as C. richii the leaf-shape is remarkably variable,
one specimen having the average leaf-index, the
second having leaves 914-11 by 114-134 cm, the
third having linear leaves c. 4-10 cm by 1-4 mm.
In West Flores a similar, also sterile linear-leaved
specimen was collected (SOEHANDA, exp. DE JONG
272, Bo, K, L); its thorns are strongly recurved,
1-3 mm, the leaves measure 4-10 by 0,1—0,4 cm.
As I observed in several other species of Capparis
such sterile shoots have larger thorns than the
fertile twigs. Might it be that such shoots have
been developed under exceptionally dry weather
conditions?

The only collection from Bali is a duplicate
specimen from Bogor in Paris with a ZOLLINGER
label ‘““Leg. Teysmann, Balie, Buleling’. As no
other sheets were seen from Bali, the location
seems to be erroneous.

21. Capparis larutensis KING, J. As. Soc. Beng. 58,
ii (1889) 393; Ann. R. Bot. Gard. Calc. 5 (1896)
118; t- 134) Ripr, FIFE Mal! Pen: 1 (1922) 122.

Climber, 10-13 m. Twigs straight, brown-
puberulous, more or less glabrescent; internodes
1—2 cm. Thorns vigorous, recurved, 3-4 mm long,
glabrous, with darker, glossy, very sharp top.
Leaves firmly herbaceous to subcoriaceous,
oblong, sometimes slightly obovate, glabrous,
2-314 by 34-134 cm, base acute to subcordate,
top obtuse to acuminate, tip finely retuse; midrib
narrowly sulcate above; nerves 4—5 pairs, hardly
visible; margin somewhat recurved when dry;
petiole 3-6 mm, brown-pubescent. Flowers
axillary. Pedicel 114-2 cm, glabrous. Sepals
fleshy, 7 by 5 mm, ciliate towards the top, the
outer pair ovate, the inner pair suborbicular.
Petals obovate, c. 8 by 5 mm, white then pink,
puberulous inside. Stamens (10?—)30, 24% cm,
anthers 2 mm long. Torus 2 mm broad. Gynophore
314-4 cm; ovary ovoid, c. 144 mm long. Mature
fruit unknown, at least 13 by 10 mm, globose,
umbonate. Seeds several.

Distr. Malaysia: Malay Peninsula (Perak,
Selangor).

Ecol. Dense jungle, clinging to trees, below
200 m. Fi. Sept., Nov.

22. Capparis spinosa LINNE, Sp. Pl. 1 (1753) 503;
HemMsL. Bot. Chall. 3 (1885) 120 (var.).

var. mariana (JAca.) K. ScuH. Bot. Jahrb. 9
(1888) 201; K. ScuH. & Laur. FI. Schutzgeb.
(1901) 335; Laur. Bot. Jahrb. 52 (1915) 111.—
C. cordifolia LAMK, Encycl. 1 (1785) 609; MEeErR.
Philip. J. Sc. 7 (1912) Bot. 235; Fl. Manila (1912)
216; Sp. Blanc. (1918) 159; En. Philip. 2 (1923)
210; pE VooGp, Trop. Natuur 25a (Jub. Uitg.)
(1936) 72, f. 9; Merr. Philip. J. Sc. 9 (1914)
Bot. 84.—C. mariana JAcQ. Hort. Schoenbr. 1
(1797) 57, t. 109; DC. Prod. 1 (1824) 245; DECNE,
Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 426;
Herb. Timor. Descr. (1835) 98; SPAN. Linnaea
15 (1841); BLANco, FI. Filip. ed. 2 (1845) 305;
Mia. Fl. Ind. Bat. 1, 2 (1858) 100; TEysmM. Nat.

90 FLORA MALESIANA

[ser. Ty. volinge

Tijd. N.I. 34 (1874) 359; NaAves in Blanco, FI.
Filip. ed. 3, 2 (1878) 201, t. 179; FEeRN.—VILL.
Nov. App. (1880) 11; SAFFoRD, Contr. U.S. Nat.
Herb. 9 (1905) 212; Burk. Dict. 1 (1935) 443.—
C. sandwichiana DC. Prod. 1 (1824) 245; A. Gray,
U.S. Expl. Exp. Bot. 1 (1854) 69; DEGENER,
Fl. Hawaii. 1 (1937) fam. 142, with plate.—C.
baducca (non L.) BLANCO, FI. Filip. (1837) 438.
—Fig. 23-24.

Tree? or shrub, mostly prostrate. Twigs terete,
mostly zig-zag; internodes 1—3 cm; innovations
whitish-tomentose, glabrescent. Thorns wanting
in Mal.. Leaves suborbicular, rarely elliptic, often
ovate, (1!14-)214-6(-714) cm diam., subcoria-
ceous (fleshy when fresh); base truncate to round-
ed, top rounded, seldom minutely emarginate to
acute; midrib above shallowly depressed in the
basal part, otherwise flat; nerves c. 5-7 pairs,
thin, subprominent on both surfaces; petiole
34-114 cm. Flowers axillary. Pedicel 414-714 cm,
glabrescent. Buds conical when young, later
bulging at the posterior base, sometimes acumi-
nate, finally 2-214 cm diam. Sepals c. 25-28 mm
long, fulvously pubescent towards the base,

glabrescent, ovate, outer pair 8-18 mm wide,

Fig. 23. Capparis spinosa L. var. mariana (JACQ.)
K. Scu. a. Bud, just before anthesis, from the left,
x %, b. left petals, of the upper (adaxial) and the
lower pair, as seen from within, nat. size, c. stamen
as it is in bud, x 14, d. bud opened, the outer
sepals cut medianly, one inner sepal, the left petals
and the stamens removed, nat. size (DEGENER
26425 from Hawaii).

the posterior one the larger and strongly saccate,
sometimes slightly keeled, inner pair equal,
slightly larger than the outer pair, 11-20 mm wide,
with broad-membranous margin. Petals pure
white, sometimes slightly emarginate, upper pair
more or less rhomboid, (3—)3'4-4'4 by 2-4 cm,
with thick fleshy base, lower pair 3-314 by
114-14%4,(-34%4) cm. Torus conical, 4-5 mm broad,
the disk c. 4 by 5 mm. Stamens more than 100,
white, 414-5 cm, anthers 4 by 34 mm, brownish.
Gynophore 6-7 cm, subpubescent towards the
base; ovary oblong, 5-8 by 1-2 mm, glabrous.
In fruit the gynophore incrassate until it is as thick
as the pedicel, and sometimes coiled as a pig’s
tail. Fruit ellipsoid to spindle-shaped, c. 5 by
114 cm, narrowed towards the apex, with 5 nerves.
Seeds oy, subglobular, c. 4 mm through.

Distr. Scattered over the Pacific within the
quadrangle formed by the Sandwich Is, Marianas
(Guam), Solomon Is, and Henderson I. (ENE of
Pitcairn, 24°20’S, 128°20’W), NW. Australia
(once coll.); a different form of the species in
Australia (see notes), in Malaysia: Lesser Sunda
Islands (Semau, Timor, Leti, n.v.), Philippines
(Luzon, cultivated in a few localities; Bohol,
spontaneous along the sea-shore), New Ireland,
and New Britain (7.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, efc. A
noctiflorous species, in the lowland up to ec.
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,
l.c., 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 in appearance 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.

Many paramorphs have been described in C.
spinosa. The var. mariana is distinct by a com-
bination of the following characters: twigs
unarmed, not too densely tomentose and glabres-
cent, leaves comparatively large and orbicular,
lacking an apical spine, flowers very large with one
saccate sepal. Capparis galeata FrEsENn. of Arabia
and E. Africa has similar flowers, the odd sepal

Dec. 1960]

_CAPPARIDACEAE (Jacobs) g ue 91

being even more irregularly shaped, but this plant
has strong stipular thorns and an apical leaf-spine.

The species is taxonomically widely different
from all native Capparis in Malaysia and Austral-
asia and it had and has a distinctly economic
significance in the Mediterranean. From this it
appears probable that the species was introduced
in the Indo-Pacific as a cultigen in post-Columbian
time. The first records from the Pacific-E. Malay-
sian area are:

Marianas, via SONNERAT!
1781,

Sandwich Is, MENZIES, Vancouver Exp. 1792,

Timor, CUNNINGHAM, 1818,

Society Is, BEECHEY’s Voyage, 1826,

Philippines, BLANCO, before 1837.

The dates of this list are significant, because it
appears clearly that it had not been collected
during the earliest Pacific expeditions such as
“Cook’s Voyages’ and the ‘Boudeuse et |’Etoile’,
1768-1770, on which such trained botanists col-
lected as BANKS, SOLANDER, the FORSTERS,
COMMERSON, efc., who did not discriminate be-
tween cultivated, introduced, and native plants,
as MERRILL rightly observed in his account of
Cook’s Voyages (Chron. 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 Acapulco in West Mexico
(from where, however, the species is not known)
and Manila via Guam in the Marianas; the regular
galleon route was initiated, for return voyages,
in 1565, with 1—2(—3) ships yearly in either direc-
tion, to be abandoned in 1815 (MerRILL, /.c.
193). It is remarkable that the species has never
been reported from Mexico.

In this respect it appears significant that the
earliest record from the Pacific, 1774-1784, is
from the Marianas, from where is also the very
early record of its being cultivated and exported.

In the Philippines there is the indication that
it has been imported there from Guam.

RuMPHIUS makes no mention of it, though he
frequently referred to plants from islands outside
the Moluccas proper.

In Timor there is no indication of such import,
but it is not far-fetched to assume its introduction
by the Portuguese. One may question why it has
not further spread in arid places in East Malaysia
and the Lesser Sunda Islands. It may well be that
the capers, this well-known delicacy to Mediter-

between 1774 and

ranean people, did not appeal to the native taste
and were therefore not further dispersed.

In conclusion, there remains hardly any doubt
of its introduction at an early date from presum-
ably seeds from the Mediterranean into the
Marianas and dispersal by man from thence into
E. Malaysia and Polynesia, as suggested by
HEMSLEY, /.c.

It seems likely that introduction took place once
only, because of the variability pattern of the
species: repeated introduction would presumably
have caused a greater variation of the population.

Fig. 24. Capparis spinosa L. var. mariana (JACQ.)
K. Scu. Flower (VAN DER PIJL).

In addition it should be stated that the species
occurs also on the west coast of Australia and in
Queensland, from where it has been described as
C. nummularia DC., a name also taken up by
BENTHAM. It has, probably rightly, been reduced
to C. spinosa by F. v. MUELLER (Syst. Cens.
Austr. Pl. 1882, 5; 2nd ed. 1889, 8) and considered
to represent a variety of C. spinosa by F. M.
BAILEY (var. nummularia (DC.) F. M. BAILEY,
Syn. Queensl. Pl. 1883, 15). It has even been found
in arid places in Central Australia. The Australian
plants look hardly different from those of the
Mediterranean and they are distinctly thorny.
But there is one specimen from NW. Australia
(PRITZEL 284) which matches the Polynesian
material. For this reason it seems likely that the
Australian form is an introduction which took
place independently from that in the Pacific. As a

(1) LAMARCK stated to have received his material from SONNERAT, but the latter never visited the
Marianas (see vol. 1, p. 494). He attached himself, however, to COMMERSON and worked with him for
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

iser -bwee 6"

matter of fact the case of the rather early record
of this European or West Asian plant in Australia
is matched by some other examples. In all these
cases the Australian population is (mostly
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
Floral, 4, 1949, lii). Different imports of a cultigen
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. FI.
Austr. 1 (1863) 96; BAiLey, Queensl. FI. 1 (1899)
60; Brirren, Ill. Austr. Pl. 1 (1900) 6, t. 6;
Domin, Bibl. Bot. Heft 89 (1925) 688; C. T.
Wuirte, J. Arn. Arb. 10 (1929) 217.—Thylachium
lucidum BANKS ex DC. Prod. | (1824) 254.—C.
subacuta Mig. Fl. Ind. Bat. 1, 2 (1858) 101;
Illustr. (1870) 35, t. 19, guoad specim. javan. (vide
sub C. sepiaria); K. & V. Bijdr. 4 (1896) 260;
Back. Schoolfi. (1911) 63; Koorp. Exk. Fl. Java
2 (1912) 293 (acuta, sphalma); BACK. Bekn. FI.
Java (em. ed.) 4A (1942) fam. 45, p. 10.—C.
nobilis (non BENTH.) F. v. M. Descr. Not. Pap. PI.
28 (1886) 41.

Tree or shrub, c. 2-12 m. Twigs terete, straight,
densely fulvous pubescent, sooner or later glabres-
cent; internodes less than 2 cm. Thorns in flowering
specimens nearly always wanting, if present up to
2 mm. Leaves coriaceous, ovate to obovate,
glabrous above, mostly very soon glabrescent
beneath 1.7—2.5(—-3.5) times as long as wide,
(3-)514-7(-9) by (2-)214-3 cm; base acute to
cuneate, top rounded to obtuse, sometimes
mucronate; midrib above subprominent, nerves
5-9 pairs, subprominent on both sides; petiole
14-114 cm, hairy as the twigs. Raceme pubescent,
terminal, with up to a dozen of sweet-scented
flowers, the lower in the axils of small leaves.
Bracts proper, if present, small, acute. Pedicels
114-6 cm. Buds ovoid, c. 114 cm long, often with
umbonate top. Outer pair of sepals coriaceous,
initially connate and completely enveloping the
bud, then splitting more or less regularly, patent
at anthesis, c. 10-15 by 10 mm, acuminate,
glabrous; inner pair rather thin, flat, ovate, c.
11-13 by 5-6 mm, shortly acuminate. Petals
subobovate, 18-30 by 10 mm, pubescent inside
and outside towards the base, white to yellowish.
Torus comparatively broad, short-conical, with a
small posterior disk. Stamens c. 50-70, 244-5 cm,
filaments white, anthers c. 244 mm long. Gyno-
pPhore 2\4-7 cm, mostly thinly woolly in the basal
half, glabrescent; ovary ellipsoid, acuminate, c.
3-4 mm long, glabrous, stigma dark-purple. In
fruit the pedicel, the torus, and the gynophore

somewhat incrassate. Fruit globular, 244-4144 cm
diam., pericarp c. 3 mm thick, soft, leathery,
smooth, dull purplish brown, pulp spongy. Seeds
7-30, sometimes subangular, c. 9 by 6 by 5 mm.

Fig. 25. Localities of Capparis lucida (BANKS ex
DC.) BENTH. From the New Hebrides one un-

confirmed record. In Australia several other
localities to 30° S, and one unspecified locality
“Gulf of Carpentaria”.

Distr. Australia: NW. coast (BENTHAM), and
NE. coast of Queensland to 20° S. lat.; in Malay-
sia a rare plant: E. Java (one locality), Lesser
Sunda Islands (Komodo, Pada, Timor), SE.
Celebes, New Guinea (Papua), Booby I. in Torres
Strait, ?Bismarck Arch. Fig. 25.

Ecol. Chiefly coastal, but also in savannahs.
Fl. fr. Jan.—Dec.

The sweet-scented, white or pale lemon-yellow
flowers are typically nocturnal. KoorDERS noted
that the fruit pulp has a sourish stench, like that
of rotten fish. In a living plant in Kebun Raya
Indonesia (II-Q—17) I found the spongy, yellow
pulp tasting and smelling as insipid mango. They
might be eaten by bats which could add to their
dispersal.

Notes. In the Bogor Gardens another specimen
is cultivated (XI—-B—V-—117); it is a tree, 5 m by
15 cm; it stands in deep shade and never flowers.
The leaves of its long, overhanging twigs tend to
have a distichous position and are subtended by
straight thorns 2-214 mm long.

Seeds sown from n. II-Q-17 produced seedlings
of which the shoot is very similar to those of the
just mentioned sterile tree. And though I have not
observed the transitional stage wherein the
seedling gets unarmed twigs with normal leaves I
consider the unarmed sterile tree as a retarded
juvenile stage through the effect of shade.

The occurrence of a sterile juvenile stage with
thorns seems to be typical for the mainly Austra-
lian sect. Busbeckea: BENTHAM reported it in
C. canescens BANKS ex DC. and I observed it in
several specimens of C. nobilis F. v. M. and C.
mitchellii LINDL.

DE CANDOLLE referred this species to the genus
Thylachium on account of lacking petals; this
error is perhaps due to their being very easily
caducous.

Dec. 1960]

CAPPARIDACEAE (Jacobs) 93

Incompletely known species

Capparis longipes Merr.! Philip. J. Sc. 13 (1918)
Bot. 12; En. Philip. 2 (1923) 211.

Scandent, glabrous shrub, branches slender,
terete, brownish or olivaceous, the ultimate
branchlets c. 1 mm diam. Thorns straight, usually
2-414 mm. Leaves lanceolate, membranaceous to
chartaceous, green or greenish-olivaceous when
dry, somewhat shining, 7-11 by 2—3 cm, narrowed
upwards to the very slender apex, sharply acute-
acuminate, base acute; nerves c. 15 pairs, slender,
distinct on both surfaces, anastomosing, primary
reticulations lax, ultimate ones close, both
distinct; petiole 2-3 mm. Infructescences axillary,
very slender, sparingly branched, up to 20 cm
long, each branch bearing a single fruit, its pedicel
c. 3. cm. Fruit globose, brown when dry, glabrous,
c. 12 mm in diam.

Distr. Malaysia: Philippines (Luzon: Abra),
only known from the type BS 26980 Ramos (US,
K; L, photogr.).

Notes. The identity of this specimen could not
be established with certainty, as the holotype in
Manila has been lost, and the isotypes have no
fruit. From the description by MERRILL (vide supra)
and the completely sterile isotypes it is not pos-
sible to form an adequate idea about the structure
of the inflorescence.

Though Merrit did not mention the bract-like
minute scales towards the base of the shoots

which are typical for C. pubiflora | am inclined to
compare the material with that species, with
which it agrees in the greyish pubescence of the
innovations, the straight thorns, and the venation
pattern. It may represent a deviating form
(teratological or developed in shade?) though I
have as yet not seen any comparable paramorph
in C. pubiflora.

Excluded

Capparis carandas Burm. f. Fl. Ind. (1768) 118,
119. MERRILL reduced this to Carissa carandas L..,
cf. Rumph. Herb. Amb. (1917) 425. Dr BAEHNI
kindly sent me BURMAN’s authentic material in
the Geneva Herbarium on loan. The entry on
p. 118 may refer to one sheet which is most
probably a Carissa; the entry on p. 119 refers to
four other sheets belonging to Carissa carandas L.
Mant. | (1767) 12 (Apocynaceae). According to
the labels the material was derived from cultivated
plants in Java.

Capparis versicolor GrirF. Notul. 4 (1854) 577;
Hook. f. & TH. Fl. Br. Ind. 1 (1872) 175. Recorded
from Tenasserim and from Java by HooKEer, who
suggested its possible identity with C. salaccensis
BL.

I have seen the type in the Calcutta Herbarium
and agree that it is vegetatively similar to C.
cantoniensis; the buds have disappeared but are
assumed to have measured c. 12 mm diam. on
pedicels c. 3 cm.

3. CADABA

Forsk. Fl. Aegypt.-Arab. (1775) 67; PAx & HorrM. 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
solitary, axillary). Sepals 4, in 2 whorls, unequal, caducous, outer pair enveloping
the bud, valvate, inner pair apert. Peta/s 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, |-celled; placentas 2-4; ovules «~, 2-seriate; stigma sessile, indistinct.
Fruit mostly cylindrical, dehiscent with 2 coriaceous valves (or an indehiscent
berry). Seeds oo, 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. 1 (1824) 244;
Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3
(1834) 427; Herb. Timor. Descr. (1835) 99;
DELEss. Ic. Sel. Pl. 3 (1837) 5, t. 9; SPAN. Linnaea
15 (1841) 164; Mia. FI. Ind. Bat. 1, 2 (1858) 97;
BENTH. Fl. Austr. 1 (1863) 92; Mia. Illustr. (1870)

21; Koorp. Exk. FI. Java 2 (1912) 295; Back.
Onkr. Suiker. (1931) 256, Atlas t. 266; PAx &
HorrM. in E. & P. Pfl. Fam. ed. 2, 17b (1936) 186,
f. 806, 85 F—G; Back. Bekn. FI. Java (em. ed.) 4A
(1942) fam. 45, p. 11.—Fig. 26-27.

Shrub, 14-3 m, procumbent to erect, very dense

(1) C. longipes STANDL. Contr. U.S. Nat. Herb. 23 (1922) 303, non MERR. 1918 = C. discolor STANDL.
op. cit. 20 (1919) 182, non DoNN. SMITH, 1897 = C. renominata Jacoss, nom. nov. It is a Mexican species.

94 FLORA MALESIANA

[ser. I, vol. 6!

Fig. 26. Cadaba capparoides DC. a. Habit, x 24, b. flower from the left, nat. size, c. pedicel with andro-
gynophore, gynophore, fruit, x 24, d. seed, x 3 (a—b type specimen, LESCHENAULT s.n., c-d TEYSMANN
5.n.).

glandular-pubescent to almost glabrous. Twigs
terete, straight; internodes 34-2144 cm. Thorns
straight to slightly recurved, 114-4 mm, yellowish,
sometimes small or wanting. Leaves simple,
firmly herbaceous, ovate (to rarely obovate),
(1.6—)2—3(—S.2) times as long as wide, (3!4—)6—10
(-1314) by (134-)214-414(-814) cm; base rounded
to acute, sometimes obtuse, top narrowed, blunt

to acutish, rarely rounded, mostly mucronate;
nerves c. 7—9 pairs; petiole (14—)14-4 cm. Racemes
terminal, corymbiform; rachis 2—3 cm. Pedicels
(114-)2-3(-314) cm. Bracts caducous, 2-514 mm
stalked, blade 6-10 by 14-214 mm, lanceolate to
ovate, acute, subtended by 2 minute thorns. Buds
ovoid, acute. Sepa/s glandular-puberulous outside,
boat-shaped, (8—)10—15(—20) by (3-)6-9(-12) mm,

Dec. 1960] CAPPARIDACEAE (Jacobs) . Le

ovate, acute to acuminate, inner pair somewhat obovate, acute. Petals pure white on a green claw,
smaller, flattish, sometimes up to | mm clawed, all pointing upwards at the adaxial side, (414-)
8—15(—22) mm clawed, blade orbicular to broad-
elliptic, 5-12 by 444-7 mm. Gland green, 514-15
by c. 1 mm; top yellow, flat, transverse, one-sided,
lanceolate, 444-6 mm long. Stamens (5—)6(—7),
androgynophore 1-214 cm; filaments 1-2 cm,
anthers 314-5 by 34 mm, all glabrous. Gynophore
114-214 cm; ovary cylindrical, 414-6 by 34-1 mm,
both sparsely glandular-puberulous; stigma knob-
shaped, small; placentas 2. In fruit the pedicels
hardly stretched or thickened, androgynophore
(1—3—)2-3(-3 14) cm, gynophore (114-)2-3(-3'4)
cm, more or less densely glandular-puberulous.
Fruit cylindrical, dull grey-brownish, (41—)7—11
cm by 5-13 mm, + glabrescent, dehiscing basis-
copically, the filiform placentas persisting as a
replum. Seeds up to c. 20, 3—4 by 214-3 by 114 mm,
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. Cadaba capparoides DC. Open flower. Vern. Bangol bangol, Bali (also in use for other
The gland is in the centre (VAN DER PIL). Capparidaceae), paumahatas, Timor.
4. STIXIS

Lour. FI. Coch. (1790) 295, ed. WILLD. (1793) 361; PIERRE, Bull. Soc. Linn. Paris
1 (1887) 652.—Roydsia Roxs. Pl. Corom. 3 (1819) 87; Hook. f Fl. Br. Ind.
1 (1874) 180, 409.—Covilhamia Kortn. 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 ~-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, ellipsoid, 244-5 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 A/ytostylis Hook. f.
(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. suaveolens (ROXB.) PIERRE and S. philippinensis

96 FLORA MALESIANA [ser. I, volife

we a b @ Cc}
Fig. 28. Stixis ovata (KORTH.) HALL. f. ssp. ovata. a. Habit, x 24, b. flower, one inner sepal removed, x 4,

c. pedicel, torus, gynoecium, x 4, d. cross-section through ovary, x 3, e. fruit, x 24 (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 S. suaveolens and S. obtusifolia (HooK. f. & TH.) PIERRE,
which two species would belong to § Roydsia. 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 S. ovata, 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

Gynophore shorter than 5 mm, glabrous. Sepals c. 4 mm long.

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.

1. Stixis philippinensis (TURCZ.) MERR. Govt Lab.
Publ. (Philip.) n. 35 (1906) 72; Philip. J. Sc. 1
(i906) Suppl 58; En: Philip. 2 (1923) 213;
ERDTMAN, Pollen Morph. (1952) 97.—Roydsia
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. Vasc. Filip.
(1886) 48.—Roydsia floribunda PLANCH. ex HOOK.
f. Fl. Br. Ind. 1 (1874) 409.—S. floribunda (PLANCH.
ex Hook. f.) 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
(1114) cm, smooth except for a few pustules
above near the base of the midrib, top c. 44 cm
acuminate; midrib above broadly sulcate; nerves
9-11 pairs; petiole 2-3 cm. Panicle ~-flowered,
terminal, c. 25 cm long; branches up to 15 cm,
all axes angular and fulyous-puberulous. Bracts
orbicular 14 mm to linear 3 mm. Pedicels 14 cm.
Buds 10-11 mm long, with cylindrical base and
somewhat swollen, acute top. Torus 1144-2 mm
broad. Sepals during anthesis reflexed halfway,
narrow-spathulate, blunt to acute, obscurely
3—S-nerved, the base not narrowed but slightly
thickened. Androgynophore 2—3 mm, sometimes
hairy at the top. Stamens c. 35—40(—48), glabrous,
filaments 10-15(-16) mm. Gynophore 8-11 mm,
densely puberulous; ovary subglobular, glabrous,
c. 2 by 114 mm;; style 114—2(-214) mm, glabrous,
with 3 small but distinct stigmatic lobes. Pedicel in
fruit c. 1 cm, torus 4 mm broad, gynophore 3 mm
thick, still hairy. Fruit (only some unripe and a
few fragments of a ripe one are known) globular
with persistent style, up to 314 cm diam.; pericarp
thin, leathery. Seed c. 2 by 114 cm.

Distr. Philippines (Luzon: Bataan and Laguna
Proy.; Basilan; Mindanao: Surigao, Cotabato,
and Davao Prov.). Seems to be very local. Fig. 29.

Ecol. Forests at low and medium altitudes.
Fl. Jan.—Dec., fr. March—April.

2. Stixis ovata (KorTH.) HALL. f. Beih. Bot.
Centralbl. 39, ii (1921) 35.—Covilhamia ovata
KortH. Ned. Kruidk. Arch. 1 (1848) 307; Mra.

3. S. scortechinii

Fig. 29. Distribution of the Malaysian species of

Stixis and of Cadaba.—1. 8S. _ philippinensis

(Turcz.) MErRR., 2. S. ovata (KORTH.) HALL. f.

(triangles representing localities), a. ssp. fasciculata

(KING) JAcoss, b. ssp. ovata, 3. S. scortechinii

(KING) JAcoBs.—Cadaba capparoides DC. (dots
representing localities).

Fl. Ind. Bat. 1, 2 (1858) 180; Merr. En. Born.
(1921) 381.—S. fasciculata var. borneensis HEINE,
Mitt. Bot. Staatssamml. Miinch. Heft 6 (1953) 212.

ssp. ovata.—Fig. 28.

Branchlets fulvous-tomentose, finally glabres-
cent. Leaves firmly herbaceous, oblong to lan-
ceolate, mostly somewhat obovate, rather narrow,
344-114 cm acuminate, fulvous-pubescent, glabres-
cent, (8—)12—17(—22) by 4-8(—11) cm; midrib above
sulcate; nerves 7—9 pairs, mostly densely set with
minute whitish pustules. Petiole 1—2(—2!4) cm,
hairy as the twigs. Racemes or panicles axillary
(rarely a terminal panicle), 6-17 cm long, axes
angular, fulvous-tomentose. Bracts linear, 2-3
(-4) mm, tomentose. Pedicels thin, c. 4% cm. Buds
obovate, 4-5 mm long. Flowers greyish green.
Torus 1 mm wide. Sepals reflexed, lanceolate, c. 4
by 1 mm, acutish, obscurely 3-nerved. Andro-
gynophore c. % mm long and wide, glabrous.
Stamens 26-34, filaments 214-314 mm, glabrous.

98 FLORA MALESIANA [ser.. I, vole

SRS ~ N
SSS’ SS
=

So

Uy Y
LZ Nee
LG is > ®
Vig ¢
Ze =— = Se ee Z “ff

a Fa /
ett

ty te

eee ee

Fig. 30. Cleome rutidosperma DC. a. Flowering branch, x 2, b. flower, lower sepal and petals removed,

right 3 stamens omitted, x 6, c. flower from the adaxial side, upper sepal, petals, two stamens removed,

lower petals partly, other sepal and stamens omitted, x 6, d. front view of flower, x 2, e. flower, x 4,

f. flower from the abaxial side, lower petal and 4 stamens removed, upper sepal, petals, and stamens
omitted, x 6, g-h. seed with elaiosome, 6 (living material, Bogor).

Dec. 1960]

CAPPARIDACEAE (Jacobs) : ~ 99

Gynophore \—2 mm, glabrous; ovary ellipsoid, c.
114 by 1 mm, densely appressed-pubescent; style
114-2 mm, slender, glabrous, stigma obscurely
3-lobed. Fruit on a woody stalk %4 by %4 cm,
ellipsoid, 3-314 by 244,-2'4 cm; pericarp 34-1 mm
thick, leathery-corky, irregularly covered with
whitish crust-like fragments, glabrous, crowned
by the style base. Seed oblong, c. 234 by 114 cm.

Distr. Malaysia: Borneo. Fig. 29.

Ecol. In forests, up to 1200 m.

Vern. Sansang, Dusun (N. Borneo).

Note. In continental Asia (Burma, Tenasserim,
and Indo-China) another subspecies occurs,
formerly distinguished as a separate species,
which differs from the type subspecies in having
the gynophore hirsute except at the base, flowers
in terminal panicles 12—35 cm long or racemes over
12 cm long. Bract mostly 4 mm.

3. Stixis scortechinii (KING) JACOBS, nov. comb.—
Roydsia scortechinii KiNG, J. As. Soc. Beng. 58,
ii (1889) 397; Ann. R. Bot. Gard. Calc. 5 (1896)
120; t. 139; Rw. Fl. Mal. Pen. 1 (1922) 121.—
Roydsia parviflora (non GRIFF.) KING, J. As. Soc.
Beng. 58, ii (1889) 396.

Shrub, often twining (to the left), (1—)3—4(—8) m.
Branchlets downy puberulous, glabrescent. Leaves
firmly herbaceous to subcoriaceous, elliptic to
oblong, glabrous, c. (1.4—-)2 times as long as broad,
mostly obovate, (8—)10—-16(—20) by 4-7(-9) cm,
base more or less narrowed and acute, top rounded
and abruptly acuminate with triangular, short,
blunt tip; midrib above flat to finely sulcate,

below mostly densely set with pustules; nerves
6—7(—8) pairs, reticulations prominent; petiole
(1-)1'14-2(-2144) cm. Inflorescence a terminal,
slender, leafy, many-flowered panicle 15—20 cm
long, densely downy-puberulous. Bracts linear, c.
3 mm. Pedicels 2—3(—5) mm. Buds obovoid, 214-3
mm long. Flowers fragrant. Torus c. 114 mm wide.
Sepals reflexed, strap-shaped, somewhat obovate,
acute, inside pink, 3—4(-414) by 34-114(-114) mm.
Androgynophore hardly visible, glabrous. Sta-
mens (20—)25-—40; filaments 114-314 mm; anthers
elliptic, c. ¥% mm long. Gynophore 34-114 mm
long; ovary ellipsoid to ovoid, c. 1-114 by Imm,
both glabrous; style 1-114 mm, slender, glabrous,
after anthesis somewhat recurved, stigmas ob-
scure. Fruit on a woody stalk, 5-7 by 4-5 mm,
ellipsoid c. 3-4 by 2-3 cm; pericarp leathery,
thin, mostly smooth but sometimes spotted with
whitish, crustlike fragments. Seeds c. 2!4 by 114
cm, embedded in pulp c. 3 mm thick.

Distr. Malaysia: Northern Sumatra (East and
West Coast Res.), Malay Peninsula (Wellesley,
Perak, Selangor, Negri Sembilan, Penang). Fig. 29.

Ecol. Dry sunny places, young secondary
forest, landslides, jungles, 100-1100 m. Fi. fr.
Jan.—Dec.

Uses. Sore eyes are treated with the juice from
the roots (NE. Sumatra).

Vern. Simar silaun, Toba, andor si bumbun,
Syuimal, 18. (C

Note. KUNSTLER recorded it on his labels once
as a “tree 40—-SO ft’ and once as a “splendid
climber 80-100 ft’’; this needs verification.

5. CLEOME

Linne, Gen. Pl. ed. 5 (1754) 302; Sp. Pl. 2 (1753) 671; R. Br. in Oudney, Denh.
& Clapp. Narr. Trav. Disc. Afr. (1826) App. 220; DC. Prod. 1 (1824) 238;
SCHULTES, Syst. 7! (1829) 23; PAx & HOFFM. in E. & P. Pfl. Fam. ed. 2, 17b (1936)
210.—Pedicellaria 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 & Horr. 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 leaf. Leaves petioled, herbaceous, in the Mal. spp. palmately
dissected into 3-7 leaflets. Flowers pedicelled, in leafy, terminal racemes or
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 «, in Mal. 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-valved, beaked, dehiscing from the base or from the apex,
the 2 placentae forming a persistent replum. Seeds ~, 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. 61

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. chelidonii the plants are covered, mostly densely,
with appressed, stiff, glassy hairs with a more or less bulbous base. In C. aspera and C. rutidosperma
there are sparse, subpatent, epidermal appendages, too large to be regarded as hairs in the strict sense.

Taxon. DE CANDOLLE (1824) distinguished Cleome, Gynandropsis, and Polanisia. R. BROWN (1826)
suggested the reduction of Gynandropsis as a section to Cleome, except for C. gynandra on which he
based a new section Gymnogonia. The SCHULTESes (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 subsect. 1.
Pedicellaria (DC.) SCHULT., with a long fleshy torus and a long gynophore, to which would belong
C. spinosa, and subsect. 2. Siliquaria (FORSK.) SCHULT. with a small torus and a short or wanting gynophore,
to which would belong C. aspera, C. rutidosperma, 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 I_tis 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. viscosa; sect. Corynandra (SCHRAD.) SCHULT.: C. chelidonii; sect. Rutidosperma I tis:
C. aspera, C. rutidosperma.

Dr Itis (Brittonia 10, 1958, 33 and 12, 1960, in 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 Woopson, 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, ILtTis pointed out that it is satisfactory to restrict Polanisia to the original concept of
RAFINESQUE, the type species being the N. American P. graveolens Rar. (= 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. Ixtis, who kindly put his MS at my disposal and gave additional critical
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 1 cm, or none. Stamens free.

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 stiff scaly hairs. Flowers pink : . . . 4 C. chelidonii
5. Stamens less than 30, filaments almost filiform. Fruit hardly ‘arrowed at the base, valves centri-
petal-nerved. Flowers yellow . . eee ree wo ee SC. SLI,

4. Stamens 6. Ovary on a short synophore.
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 long. Fruit 4 mm diam. Seeds with an open cleft.
7. C. rutidosperma
7. Petals white, 4-5(-7) mm long. Fruit 2 mm diam. Seeds orbicular, with a closed cleft.
8. C. aspera

Dec. 1960]

1. Cleome gynandra LINNE, Sp. Pl. 2 (1753) 671;
C. B. Rosins. Philip. J. Sc. 3 (1908) Bot. 182.—
Lagansa rubra RuMPH. Herb. Amb. 5 (1747) 280,
t. 96 f. 2.—C. triphylla LINNE, Sp. Pl. ed. 2 (1763)
938.—C. pentaphylla LINNE, I.c. 938; R. Br. in
Oudney, Denh. & Clapp. Narr. Trav. Disc. Afr.
(1826) 222.—Pedicellaria pentaphylla SCHRANK in
Roem. & Ust. Mag. Bot. 3, pt 7 (1790) 11; MerRrR.
Publ. Govt Lab. Philip. n. 27 (1905) 18.—Gynan-
dropsis pentaphylla DC. Prod. 1 (1824) 238; Mia.
Pl. Jungh. (1854) 397; Fl. Ind. Bat. 1, 2 (1858) 96;
Sum. (1860) 19; ErcHv. Fl. Bras. 13, 1 (1865) 261,
t. 58 f. 3; Hassk. Neu. Schliiss. Rumph. (1866)
263; Mig. Illustr. (1870) 19; FerRN.—VILL. Nov.
App. (1880) 10; Kinc, J. As. Soc. Beng. 58, ii
(1889) 392; Merr. Philip. J. Sc. 1 Suppl. (1906)
58; BAcK. Fl. Bat. (1907) 53; Schoolfl. (1911) 60;
Merr. FI. Manila (1912) 216; Laur. Bot. Jahrb.
52 (1915) 110; Merr. Int. Rumph. (1917) 241;
Sp. Blanc. (1918) 158; Ripr. FI. Mal. Pen. 1
(1922) 120; v. D. PisL, Trop. Natuur 19 (1930) 162.
—Gynandropsis affinis BL. Bijdr. 2 (1825) 51.—
C. affinis SPRENG. Syst. Veg. ed. 16, 4, 2 (1827) 138,
non DC. 1824.—C. blumeana SCHULT. Syst. Veg.
7 (1829) 23.—C. alliacea BLANCO, FI. Filip.
(1837) 522.—C. blumeana D. Dietr. Syn. Pl. 2
(1840) 1065.—C. alliodora BLANCO, FI. Filip. ed. 2
(1845) 363, ed. 3, 2 (1879) 307, t. 233.—Gynandrop-
sis gynandra Brig. Ann. Cons. Jard. Bot. Genéve
17 (1914) 382; Merr. En. Philip. 2 (1923) 209;
DocTERS VAN LEEUWEN, Zoocecidia (1926) 211;
HeEyYne, Nutt. Pl. (1927) 681; OcCHSE & BAKH. Ind.
Groenten (1931) 96, f. 57; BAck. Onkr. Suiker.
(1931) 253, Atlas t. 264; Trop. Natuur 22 (1933)
112; Burk. Dict. 1 (1935) 1119; PAx & HoOFFM.
in FE. & P. Pfil. Fam. ed. 2, 17b (1936) 218; BAcK.
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 4;
Merr. Pl. Life Pac. World (1946) 135, f. 119.

Erect, mostly widely branched, annual herb,
15-80 cm. Stem glandular-pubescent to glabrous.
Normal /eaves with 5 leaflets, lowest and upper
with 3, towards and in the inflorescence diminish-
ing in size; leaflets thinly herbaceous, about twice
as long as wide, obovate, c. 2-7'4 by 1-334 cm,
base cuneate, top rounded and + distinctly
acuminate, ciliate to denticulate; nerves 5-8 pairs;
petiole 2-10 cm, petiolules 1-3 mm, webbing at
the base, densely glandular puberulous. Flowers
in long, corymbose racemes, nocturnal. Pedicels
thin, 144-214 cm, glandular-puberulous. Sepals
214-5 by 14-14% mm, acute, puberulous, ciliate.
Petals with open aestivation, all pointed upwards
towards the adaxial side, elliptic with narrowed
base and rounded top, 114—5 mm stalked, 7—15
mm long in all, 14-4 mm broad. Androgynophore
9-16 mm. Stamens 6; filaments 114-2 cm; anthers
linear, 2-3 mm long. Gynophore 1-2 mm; ovary
cylindric, c. 3 by 14 mm, in some flowers (es-
pecially the apical ones) sessile and reduced. In
fruit pedicels 1-3 cm, androgynophore 13-20 mm,
gynophore 4-10 mm. Fruit cylindrical, tapering
to both ends, 2-11 cm by 3-4(-6) mm; beak 1-4
mm; valves with longitudinal, centripetal veins.
Seeds depressed-globular, c. 114 mm diam., with
a shallow and narrow cleft, black-brown, with

CAPPARIDACEAE (Jacobs)

101

many superficial concentric ribs and numerous
irregular distinct cross-ribs. No elaiosome.

Morph. RAGHAVAN found in this species (in
S. India) that about 50% of the ovaries are
sessile and sterile with only vestiges of ovules
being present (J. Linn. Soc. Bot. 52, 1939, 239).

MAURITZON studied the embryology (Ark. Bot.
26 ie laee 935" 1):

Distr. From Ceylon and the Punjab through-
out SE. and E. Asia as far as Peiping; throughout
Malaysia. Widely introduced in the New World.

Ecol. A weed, in dry rice-fields, along road-
sides, near houses, from the lowlands up to c.
500 m. Fl. fr. Jan.—Dec.

Vern. Maman(g), mamam, M, S, bobowan,
entjéng-entjéng, J, bhubhuwan, Md; Philippines:
apoi-apoian, balabalanoian, Tag.; halaya, hulaya,
P. Bis., tantandok, t.-a-dadakel, \\k.

Uses. Some minor medicinal applications are
mentioned by BurRKILL, /.c. and HEYNE, /.c.
According to Ocuse, /.c., the bitter leaves are
prepared by boiling and salting as a vegetable,
especially in Java.

2. Cleome speciosa RAF. Fl. Ludovic. (1817) 86;
H.B.K. Nov. Gen. & Sp. Pl. 5 (1821) 84, t. 436.
—Gynandropsis speciosa DC. Prod. 1 (1824) 238;
HAssk. Nat. Tijd. N.I. 10 (1856) 118; Hort. Bog.
Descr. (1858) 11; Mia. Fl. Ind. Bat. 1, 2 (1858) 96;
FERN.—VILL. Nov. App. (1880) 10; Merr. Philip.
Govt. Lab. Publ. n. 6 (1904) 24, n. 27 (1905) 18;
BAcK. FI. Bat. (1907) 54; GaGn. Fl. Gén. I.-C. 1
(1908) 174; Back. Ann. Jard. Bot. Buit. Suppl. 3
(1910) 403; Schoolfl. (1911) 60; Brig. Ann. Cons.
Bot. Genéve 17 (1914) 384; ScHRoo, Trop.
Natuur 4 (1915) 108, f. 1-7; Merr. Sp. Blanc.
(1918) 158; En. Philip. 2 (1923) 209; DocTERS VAN
LEEUWEN, Zoocevidia (1926) 211; HEyYNe, Nutt.
Pl. (1927) 681; Burk. Dict. 1 (1935) 1119; BAck.
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 4.
—C. speciosissima DEPPE ex LINDL. Bot. Reg. 14
(1831) t. 1312; Burxk. Dict. (1935) 580.—Gynan-
dropsis pentaphylla (non DC.) BLANCO, FI. Filip.
(1837) 523.—C. gigantea (non L.) BLANCO, ibid.
ed. 2 (1845) 364, ed. 3, 2 (1879) 307, t. 234.
Little-branched herb, c. 1-114 m, erect, glabrous
or glabrescent. Leaflets 5—7, herbaceous, sub-
sessile, slightly webbing at the insertion, narrowed
towards base and top, lanceolate, c. 10—12(—18)
by 214-414 cm; nerves 14—-16(—25) pairs; petiole c.
8—-12(-17) cm. Raceme up to 40 cm long. Flowers
subtended by subsessile leaves upwards gradually
simplified and diminishing in size. Pedicels thin,
c. 214-314 cm. Buds cylindric, c. 2-244 cm by
3 mm; petals imbricate, androgynophore with the
base of the filaments bulging out at the anterior
side shortly before anthesis; flowers opening at
dusk. Sepals patent, subulate, 5 mm, ciliate. Torus
c. 114 mm wide. Petals during anthesis pointing
upwards adaxially, c. (25—)30(—35) by 5-8 mm,
lanceolate, rounded, narrowed into a claw c.
5 mm, glabrous, pink or white (sometimes referred
to as f. alba). Androgynophore 5-7(-9) mm,
slightly incrassate towards base and top, glabrous.
Stamens 6, glabrous, filaments filiform, c. 5-6 cm,

102

FLORA MALESIANA

(ser. I, -volia64

anthers c. 7 mm, linear. Gynophore c. 6 cm,
glabrous, ovary cylindrical, few mm long, gla-
brous. Fruit cylindric, c. 8-9 cm by 3 mm, valves
indistinctly parallel-nerved. Seeds c. 234 mm
diam., 2 mm thick, with a fairly shallow cleft,
light brown, the surface with small, pale, scattered
scales; no elaiosome.

Distr. South America from Mexico to Peru
and Guyana, cultivated as an ornamental in
SE. Asia, up to 1500 m, in Malaysia collected in
Sumatra (East Coast), Java, Borneo (Sarawak,
W. Borneo), Philippines (Luzon), Moluccas (Aru
Is). According to HAssKARL (1856, /.c. 119) intro-
duced in Malaysia some years before 1855.

Ecol. Scuroo, /.c., observed that in Java the
flowers open between 4 and 5 p.m.; small stingless
bees (Apis indica) collect pollen but do not touch
the nectar; pollination would be effected by
nocturnal butterflies.

3. Cleome spinosa JAcq. En. Pl. Carib. (1760) 26;
LINNE, Sp; Pll ed. 2:1(1762), 9393 DE. Prod: 1
(1824) 239; Ercuu. FI. Bras. 13, 1 (1865) 252;
Merr. En. Philip. 2 (1923) 208; BAck. Bekn. FI.
Java (em. ed.) 4A (1942) fam. 45, p. 2.—C.
sandwicensis A. Gray, U.S. Expl. Exp. Bot. 1
(1854) 65.

Herb 1—114 m. Stems vigorous, glandular-
pubescent. Stipular thorns minute (some petioles
without) to 4 mm long, sharp, recurved. Leaves
much reduced towards the inflorescence; leaflets
5-7, herbaceous, lanceolate, slightly webbed at the
base, sparsely glandular-hairy, central leaflet
largest, 6—8(—10) by 134-214 (—3) cm; base cuneate,
decurrent, top attenuate, acute, mucronate;
nerves 10-15 pairs; petiole c. 5-10 cm, sometimes
with scattered prickles as the midrib beneath.
Racemes up to 40 cm. Flowers subtended by sub-
sessile, ovate-oblong, sparsely glandular-hairy
bracts, gradually diminishing in size from 2 to
14 cm. Pedicels 2—3 cm, short glandular-hairy.
Buds cylindrical, c. 234 cm by 4 mm, glabrous;
petals imbricate, androgynophore with the base of
the stamens bulging out at the anterior side
shortly before anthesis. Sepals patent, narrow-
triangular, (4-)6-7 mm, glandular-hairy outside.
Petals glabrous, on a filiform claw (5—)10—-12 mm,
blade oblong, + asymmetrical, 10-17 by 4-6 mm,
with cuneate base and rounded top. Stamens 6;
androgynophore 1—2 mm, glabrous; filaments
filiform, 314-4 cm; anthers linear, 7-8 mm.
Gynophore glabrous, in flower c. 4 cm, in fruit to
514 cm; ovary linear, c. 4 mm, glabrous. Fruits
patent, cylindrical, blunt at both ends, 514-614
cm by 4 mm, valves finely and densely nerved.
Seeds asymmetrical, 2 mm diam., almost smooth,
concentric and cross ribs subprominent; elaiosome
not present.

Distr. Native in tropical America, also oc-
casionally cultivated in the tropical to the tem-
perate zones, in Malaysia: occasionally cultivated,
known from Luzon and from Java, in the latter
island once found run wild.

Note. According to ILTIs (in /itt.) the Malaysian
material would not belong to C. spinosa, but to the

related C. houtteana SCHLECHT. (Linnaea 8, 1851,
669) which would be recognizable by glabrous
buds and ovaries, and pink flowers.

4. Cleome chelidonii LINNE f. Suppl. (1781) 300;
Hook. f. & Tu. Fl. Br. Ind. 1 (1872) 170.—
Polanisia chelidonii DC. Prod. 1 (1824) 242; Mia.
Fl. Ind. Bat. 1, 2 (1858) 97; Back. Schoolfi. (1911)
60; HEYNE, Nutt. Pl. (1927) 682; Back. Onkr.
Suiker. (1931) 254, Atlas t. 264; Bekn. Fl. Java
(em. ed.) 4A (1942) fam. 45, p. 5.—Polanisia
angulata DC. Prod. 1 (1824) 242; Mia. Fl. Ind.
Bat. 1, 2 (1858) 97; Illustr. (1870) 20.—Fig. 31,
32a-b.

Widely branched herb, 15—80cm; tap-root stout,
white. Stems angular, with sparse, appressed, pale,
stiff hairs with a bulbous base. Leaflets firmly
herbaceous, 6—7 below to apically only 3 or 1,
obovate, mostly densely appressed-hairy, central
leaflet largest, up to 4(-6) by 1144(-134) cm; base
cuneate, top rounded and subacuminate to acute;
nerves 4-5 pairs; petioles upwards gradually
decreasing in length from c. 8-10 cm to almost
zero, hairy as the stem, apex and petiolules white.
Raceme corymbiform, flowers subtended by
reduced leaves, actinomorphic. Pedicels (1—)1 14-3
cm, hairy as the stem. Buds ellipsoid, ++ obovate,
acute, 6-10 mm long. Sepals narrowly imbricate,
appressed, elliptic (to obovate), acuminate, 2—4
mm long, sparsely scaly-hairy outside, margin
membranous. Petals 4(—8), mostly obovate with
narrowed base and rounded top, 7—12(-15, in
India —21) by 3-5 mm, glabrous, light red-purple.
Stamens 30—40(—55), somewhat shorter than the
petals, glabrous; filaments with a thickened top;
anthers c. | mm, yellow. Ovary linear, about as
long as the stamens, glabrous. Fruit linear,
parallel-nerved, narrowed at the very base, gla-
brous, c. 1-3 mm beaked. Seeds asymmetrical
nearly 2 mm, cleft open, dull blackish, not ribbed
but warty by scattered scales mainly on the dorsal
side. Elaiosome wanting.

Distr. India, Burma and Siam (locally), in
Malaysia: Central and East Java.

Anat. T. S. RAGHAVAN investigated the
development of the female gametophyte, embryo,
and seed, and the vascular supply of the floral
parts (J. Linn. Soc. Bot. 51, 1937, 43-72; ibid. 52,
1939, 249).

Ecol. Fallow sawahs, sugarcane-fields on
heavy clays, and marls periodically drying out
during the pronounced dry season, in Java below
100 m, locally sometimes so abundant that the
fields are coloured red-purple by the flowers
(BACKER, 1931). Fl. fr. Jan.—Dec.

Kooper defined a weed community characteriz-
ed by Polanisia chelidonii on constantly moist,
fairly to very heavy clay in sugarcane-fields (Rec.
Trav. Bot. Néerl. 24, 1927, 84 seq.).

MirAsut found it characteristic in the vegetation
of freshwater swamps in India; he discussed also
some anatomical details (Proc. Ind. Ac. Sc. 43B,
1956, 233-236).

Note. The type material of Polanisia angulata
DC., hailing from Java, could not be located in

Dec. 1960]

the Paris Herbarium; his description is vague,
but he records a field note by LESCHENAULT
“flowers violet’. This character occurs only in

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

HEWETSON, 1950).

C. chelidonii; therefore, | agree with BACKER in
referring it here.

5. Cleome viscosa LINNE, Sp. Pl. 2 (1753) 672;
Hook. f. & TH. Fl. Br. Ind. 1 (1872) 170; F. v. M.
Descr. Not. Pap. Pl. 14 (1876) 52; RipL. Fl. Mal.
Pen. 1 (1922) 119.—Lagansa alba RumMpH. Herb.
Amba (Gi747)) 200; t. 96 f 3:—C: icosandra
LINNE, Sp. Pl. 2 (1753) 672; in Stickman, Herb.
Amb. (1754) 21; Burk. Dict. 1 (1935) 581.—
Polanisia viscosa DC. Prod. 1 (1824) 242; BL.
Bijdr. (1825) 52; BLANco, FI. Filip. ed. 2 (1845)
364, ed. 3, 2 (1878) 308; Mia. FI. Ind. Bat. 1, 2
(1858) 97; Illustr. (1870) 20; FerRN.—VILL. Nov.
App. (1880) 10; VipaL, Phan. Cuming. (1885) 94;
Back. Fl. Bat. (1907) 52, quoad var. ~; MERR.
Philip. J. Sc. 3 (1908) Bot. 407; Back. Schoolfl.
(1911) 60; Merr. Fl. Manila (1912) 216; Int.
Rumph. (1917) 240; Sp. Blanc. (1918) 158;
HeEyYngE, Nutt. Pl. (1927) 682; Back. Onkr. Suiker.
(1931) 255, Atlas t. 266, quoad f. typica.—Polanisia
icosandra W. & A. Prod. (1834) 22; Merr. En.
Philip. 2 (1923) 209; Back. Bekn. Fl. Java (em.
ed.) 4A (1942) fam. 45, p. 5, guoad f. typica; MERR.
Plant Life Pac. World (1946) 135, f. 121; Quis.
Med. PI. Philip. (1951) 344.—C. acutifolia ELMER,
Leafil. Philip. Bot. 7 (1914) 2574.—C. chelidonii
(non L. f.) BURK. Gard. Bull. S.S. 3 (1924) 280;
Dict. 1 (1935) 580.—Fig. 32c—d.

Annual, mostly widely branched herb up to
1 m, glandular, yellowish hairy, viscid and stink-

CAPPARIDACEAE (Jacobs)

ing. Leaflets tinhly herbaceous, 5—3, diminishing
upwards in size, subsessile, oblong, the central
leaflet c. 1-3(-514) by (14-)'!4-1(-2) cm; base
cuneate, top acute to obtuse; nerves 3-6 pairs;
petiole ('4—-)1-3(-6) cm. Racemes corymbose;
flowers in the axils of reduced leaves, largely
actinomorphic, opening in the morning, closing
in the afternoon, ephemeral. Sepals oblong,
blunt to acute, (2!4—)6—-7(-8) by (14-)1-3 mm.
Petals yellow (once reported white), thin, glabrous,
oblong, (4-)7-12 by (1144-)3—5 mm, base cuneate
or + clawed, top rounded. Stamens (8—)10—20
(—30), glabrous; filaments (3—)5—7 mm towards the
abaxial side, gradually increasing in length by

Fig. 32. Fruit and seed of Cleome. a—b. C. cheli-
donii L. f., c-d. C. viscosa L. (a, c nat. size, b,
a x10):

104

FLORA MALESIANA

[ser: 1; volags

1—2 mm, not or only very slightly swollen at the
top; anthers linear, c. 112-2 mm. Ovary sessile, c.
3-10 mm _ beaked, minutely glandular-hairy.
Fruit erect, (14-)2-3(-4) cm pedicelled, (1 14—)6-8
(-10) cm by (2-)3-414 mm, beak 214-4(-7) mm;
valves with distinct centripetal nerves. Seeds c.
1144 mm diam., 1 mm thick, red-brown, cleft
narrow, with strong cross-ribs and faint con-
centric ribs. No elaiosome.

Distr. Native in the Old World, from tropical
Africa and §S. Arabia to tropical Australia;
common throughout Malaysia, commonly ad-
ventive in the New World.

Ecol. A common, very tolerant, ruderal plant,
on fallow land, along roadsides, on rubbish-
heaps, in fields, efc., often on sandy, sometimes
on calcareous soil, both under seasonally dry and
everwet climatic conditions, sometimes near the
coast or in savannahs, up to 500 m, mostly lower.
Fl. fr. Jan.—Dec.

According to RipLeY in Malaya dispersed
endozoically by water-buffalos (Disp. p. 368).
FosBerG observed in the Marianas that the seeds
were eaten by birds.

Kooper found on the sugarcane-fields in East
Java a weed community, characterized by Polanisia
viscosa, typical for recent, very pervious, volcanic,
light soils valuable for sugar culture. He recorded
wilting experiments with this species under dry
conditions (Rec. Trav. Bot. Néerl. 24, 1927, 56
seq., 218 seq.).

Mr N. G. Bisset found the seeds containing an
appreciable amount of alkaloid.

Uses. There is a record from Java and one
from Hainan stating that the sap of the leaves
with water or milk is applied on the eyes. In Perak
the herb is rubbed on the body against rheuma-
tism. In Central Sumatra the leaves and seeds are
added to tobacco to stress its narcotic properties.
Besides, the plant finds a number of other minor
medical applications; see HEYNE (/.c. p. 682) and
QUISUMBING (i/.c. p.346).

Vern. Daun maman pantai, machmud panta,
maman panta, maman patih, Perak, poko kutepeng,
Malacca, maman hutan, M, nai velai, Tamil
(Mal. Pen.), dek tau chan, Penang, daun gliengang
ajam, Palembang, maman, mamman, S, antjang
antjang, bobowan, éntjéng éntjéng, étjéng, étjéng
témbéking, J, bhubhuan, Kangean, pupuan loke,
tiongbléntjongan, Md, ahuru, Nenusa Is, susawi
utan, Ambon, poompito, N. Celebes; Philippines:
apoi-apoian, balabalanéian, silisilihan, Tag., huldaya,
tulayag, P. Bis., kabdu, Iv., lampotaki, Tagb.,
tandandok, Ik.

Notes. BurRKILL recorded C. chelidonii for the
Malay Peninsula (Gard. Bull. S.S. 3, 1924, 280;
see also Dict. p. 580). The material he cited
belongs, however, to C. viscosa, partly to its f.
deglabrata.

As far as could be established DE CANDOLLE
was the first to combine C. icosandra and C.
viscosa and, under Polanisia, he chose the epithet
viscosa (1824).

f. deglabrata (BAcK.) JACOBS, Stat. nov.—
Polanisia viscosa var. deglabrata BAck. Fl. Bat.
(1907) 53; Domtn, Bibl. Bot. Heft 89 (1925) 683.—
Polanisia icosandra f. deglabrata BAck. Bekn.
Fl. Java (em. ed.) 4A (1942) fam. 45, p. 5.

Differs from f. viscosa in being entirely glabrous;
not with a distinct smell.

Distr. Malaysia: Malay Peninsula, Java.

6. Cleome aculeata LINNE, Syst. Nat. ed. 12, 3
(1768) 232; DC. Prod. 1 (1824) 241; HAssk. Nat.
Tijd. N.I. 10 (1856) 119; Hort. Bog. Descr. (1858)
12; Mia. Fl. Ind. Bat. 1, 2 (1858) 96; EIcut. FI.
Bras. 13, 1 (1865) 259, t. 58 f. 2; BAck. Ann. Jard.
Bot. Buit. Suppl. 3 (1910) 398; Schoolfl. (1911) 59;
Ript. Fl. Mal. Pen. 1 (1922) 120; BAcKk. Bekn.
Fl. Java (em. ed.) 4A (1942) fam. 45, p. 2.—C.
hulletii KinG, J. As. Soc. Beng. 58, ii (1889) 392;
Ann. Bot. Gard. Calc. 5 (1896) 117, t. 133A.
—Fig. 33a.

Erect, widely branched, annual herb, 44-4 m,
rather densely covered with minute, glandular
hairs, neither viscid nor smelling. Stem with
fulvous, recurved, stipular thorns c. 14% mm.
Leaves upwards gradually reduced in size, ulti-
mately simple and subsessile. Leaflets 3, sub-
sessile, thinly herbaceous, oblong, sometimes
obovate, central leaflet c. 214-3(-314) by 14-134
(—2) cm; base acute to cuneate, slightly decurrent,
top acute; nerves 5-8 pairs; petiole 2—3(—414) cm.
Racemes few-flowered, flowers subtended by
simple, subsessile leaves. Pedicels 114-2 cm.
Sepals linear, 114-2144 by 4-4) cm, minutely
glandular-hairy outside. Petals 134-4 mm (incl.
claw 14-1 mm), obovate, pale yellowish. Stamens
6, slightly exceeding the petals, anthers 114-2 mm.
Ovary subsessile, cylindric, 2-3 mm long, gla-
brous. Fruit on a gynophore 14-214 mm, cylindric
but abruptly tapering to both ends, 314-4 cm by
3 mm; valves finely parallel-nerved. Seeds asym-
metrical, with closed cleft, 2-24%, mm diam.;
concentric ribs faint, obtuse, cross-ribs irregular,
scattered, incomplete, distinctly prominent; elaio-
some distinct.

Distr. Native from Mexico to N. Argentina,
introduced in Malaysia: W. Java (Bogor and
vicinity. According to HAssKARL (1856, /.c.)
incidentally introduced with soil from Suriname
into the Hortus Bogoriensis, whence it escaped
before 1889), Malay Peninsula (Singapore), New
Guinea (Papua: Kanosia, one record of 1935).

7. Cleome rutidosperma DC. Prod. 1 (1824) 241;
ILTIs, Brittonia 12 (1960) in the press.—C. ciliata
ScHuM. & THONN. Dansk Vid. Selsk. Afh. 4
(1828) 67; JocHems, Trop. Natuur 17 (1928) 80,
f. 1, 2; Beumée, Trop. Natuur 18 (1929) 99;
Pax & HorrmM. in E. & P. Pfil. Fam. ed. 2, 17b
(1936) 213; BAck. Bekn. FI. Java (em. ed.) 4A
(1942) fam. 45, p. 3; JoHNSON & TAN, Gard.
Bull. Sing. 17 (1959) 325-330, fig. 1-2.—Fig. 30,
33b.

Erect to spreading, widely branched, annual,
odourless herb, 14-1 m; stem, petioles, and the
nerves underneath with sparse prickle-like, softish

Dec. 1960]

CAPPARIDACEAE (Jacobs)

105

appendages up to 2 mm long. Leaflets 3, upwards
gradually reduced and subsessile, thinly herbace-
ous, ovate to subrhomboid or oblong-lanceolate,
ciliate, central leaflet 3-314(-S) by 1144-21'4 cm;
base cuneate, decurrent, top narrowed, acute to
blunt; nerves 6-9 pairs; petiole generally 3—4(—5S)
cm, slightly webbing between the short petiolules.
Racemes with reduced leaves. Pedicels filiform,
2-3 cm, in fruit c. 3 cm, rather densely set with
minute, gland-like hairs. Sepals 344-4 by 4-34
mm, ovate, acute to acuminate, hairy like the

Fig. 33. Seeds of Cleome. a. C. aculeata L.,
b. C. rutidosperma DC., c. C. aspera KOEN. ex DC.,
all x 8.

nerves below. Peta/s violet-blue, all pointing up-
wards at the adaxial side, 9-12 mm (including a
claw 2-3 mm) by 1!4-2!4 mm. Stamens 6, 7-10
mm; anthers linear, 2 mm, strongly curved after
anthesis. Gynophore c. 1144 mm, ovary linear, c.
8-10 mm, slightly S-curved, minutely glandular,
beak c. 1 mm. Fruit on a gynophore 4-8(—10) mm,
cylindrical, tapering at both ends, 5—7!4 cm by
4 mm, beak 1-4 mm, valves glabrous, parallel-
nerved. Seeds dull black, asymmetrical, with a
pale centre, with faint, obtuse concentric ribs and
stronger cross-ribs, cleft open, 134 by 1144 mm;
ribs sometimes with microscopical bristles; base
of the seed with a white elaiosome.

Distr. West tropical Africa (from Guinea to
Angola and the Congo), introduced in the
Caribbean region and occasionally found in
Malaysia: first records 1920: Belawan, Sumatra
East Coast; 1924: Singapore; 1946: Java (Tand-
jung Priok); 1958: wide-spread from Sarawak to
N. Borneo. In 1946 found in Siam (100 km NW.
of Kanburi), in 1948 in Burma (Insein Distr.) and
in Manila.

Ecol. A lowland ruderal in the course of ex-
tending its area with a tendency to become a
common plant. BEUMEE, /.c., observed the plant

growing in trees, where the seeds were obviously
brought by ants attracted by the fatty elaiosome.
About one third of the ovaries is 2-3 mm
long and sterile.

Vern. Séru walai, Tamil (Malaya).

Note. The exact synonymy I owe to Dr ILTIs
(unpublished) who found that the type specimen
of C. rutidosperma must have actually come from
Sierra Leone, Africa, but was erroneously labelled
by DE CANDOLLE “‘Antilles” and described by him
as from ‘“‘Tabago”’.

8. Cleome aspera KOEN. ex DC. Prod. 1 (1824)
241; Hook. f. & Tu. FI. Br. Ind. 1 (1872) 169;
Back. Onkr. Suiker. (1931) 252, Atlas t. 262;
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 3.
—Fig. 33c.

Erect, widely branched, annual, stinking herb,
14-34 m high. Stem, petioles, and nerves under-
neath sparsely set with small prickle-like, softish
appendages. Leaflets 3(—5), upwards reduced and
becoming subsessile, on short petiolules webbing
at the insertion, thinly herbaceous, ovate-lan-
ceolate, ciliate, central leaflet generally 214-3
(-514) by 34-1(-114) cm; base cuneate, top blunt;
nerves 4-9 pairs; petiole generally 114—214(-8)
cm. Racemes with reduced leaves. Pedicels fili-
form, 14-114 cm, 114-2 cm in fruit, rather
sparsely set with minute glandular hairs. Sepals
2-3 by 14-14 mm, acuminate to obtuse, glandular-
hairy. Petals white, 4-5(—7) (including claw 1-2
mm) by ¥%4-1'44 mm, oblong-lanceolate, top
rounded. Stamens 6(—7), 4-6 mm; anthers c.
1 mm, linear, curved after anthesis. Gynophore c.
1 mm; ovary 3-4 mm long, linear. Fruit on a
gynophore 3—-6(-10) mm, 3-6 cm by 2 mm,
cylindrical, tapering to both ends, the beak 2-8
mm; valves parallel-nerved, glabrous. Seeds sub-
orbicular, c. 144 by 134 mm, dark-brown, with
small paler centre, with very narrow-elevated,
strongly prominent cross-ribs, and distinct obtuse
concentric ribs, cleft closed; no elaiosome.

Distr. Native in India and Ceylon, in Malaysia:
Central and East Java, Madura, and Bali.

The oldest Malaysian collection is by Hors-
FIELD from Java between 1802 and 1819; the
second one is from Pekalongan, in 1912.

Ecol. Obviously bound to a distinctly seasonal
climate.

Vern. Entjéng éntjéng-kébo, J (in use for all
Cleomoideae).

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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 ©, 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 ~©; 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 Campanula, counting
several hundreds of species. In Malaysia the family is sparsely represented although with one endemic
genus, Phyllocharis, in New Guinea, and a subendemic one, Pentaphragma.

Two genera show a remarkable trans-Pacific distribution, viz Laurentia sect. Isotoma with the majority
of species Australian and one species in tropical America, and the Pratia affinity of Lobelia which is a
distinctly South Pacific-Subantarctic group.

Codonopsis on the other hand is typically East Asian, mainly Chinese, and Peracarpa is Sino-Japanese.

Wahlenbergia is worldwide distributed. Sphenoclea 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 confusa
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 Sphenoclea 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 Campanulacea 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 fall
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 ail 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. 61

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 angulata RipLey (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
Phasianinae in East Malaysia and Australia; in South America this group is replaced by the subfam.
Odontophorinae 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. Dic. 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 Pentaphragma, 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 non-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 & WyYLIE’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 evenn = 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. Caffeic 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 «, ~!-disubstituted- N-methylpiperidine base. The alkaloids of all other species of 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 in Centropogon 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 raising the genera

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 I agree with WIMMER and HUTCHINSON (Fam. FI. PI. ed.
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 BrH.: Peracarpa, Wahlenbergia, Codonopsis.

2. Tribus Sphenocleae SCHONL.: Sphenoclea.

3. Tribus Pentaphragmeae SCHONL.: Pentaphragma.

4. Tribus Lobelieae Btu.: Lobelia, Phyllocharis, Laurentia.

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.
lat. 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, Bulbophyllum, 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.

1 The remarkable development of a large number of species of Lobelia cg. genera with arboreous
habit on the high African volcanoes and the Hawaiian Islands point in the same direction.

110 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 angulata, L. nicotianaefolia, Codonopsis 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 rotundifolia. 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.
Pfi. Fam. 4, 5, 1894, 43) there is a great variability in the numbers of the floral parts; see also under
Wahlenbergia marginata.

Uses. HEYNE /.c. 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
Campanulaceae can in the tropics only successfully be grown in the montane zone.

Notes. Mr MoE iono is responsible for revising Lobelia, Codonopsis, and Peracarpa, Mr Tuyn for
Laurentia, 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 Mal. spp.) a berry ; Renny Os 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

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

6. Corolla white, tube 714-11 cm long, narrow-cylindric, not dorsally slit, almost actinomorphic.

Stamens inserted high in the tube (in Mal. 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 .~. . « 9. Phyllockinis
7. Flowers arranged in various ways, not epiphyllous . . . . .. . . . 6. Lobelia

1. PERACARPA

Hook. f. & Tu. J. Linn. Soc. Bot. 2 (1858) 26; Freer, Bot. Jahrb. 12 (1890) 620,
t.: VI -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 0; 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. f. Jahrb. 12 (1890) 620, t. 7B; CRais, Fl. Siam. En.
& Tu. J. Proc. Linn. Soc. Bot. 2 (1858) 26; 2 (1936) 308; YAmAmoTo, Obs. Fl. Form. 13
CLARKE, FI. Br. Ind. 3 (1881) 437; Freer, Bot. (1936) 149; Hara, J. Jap. Bot. 21 (1947) 20;

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

111

MAKINO, Ill. Fl. Japan (1954) 82, incl. var. cir-
caeoides.—Campanula carnosa WALL. in Roxb. FI.
Ind. 2 (1824) 102; Cat. (1828) n. 1282; DC. Prod.
7 (1839) 474.— Campanula circaeoides FR. SCHMIDT
ex Mig. Ann. Mus. Lugd. Bat. 3 (1867) 195,
204; Fors. & HeEmsL. J. Linn. Soc. Bot. 26 (1889)
9.—P. circaeoides Freer, Bot. Jahrb. 12 (1890)
621; Feporov, FI. U.R.S.S. 24 (1957) 380.—P.
luzonica ROLFE, Kew Bull. (1906) 201; Merr.
& Merritt, Philip. J. Sc. 5 (1910) Bot. 392; MerRR.
En. Philip. 3 (1923) 586.

A weak, branching succulent herb up to 16 cm
long. Leaves broad ovate to ovate, 4-30 by 4-20
mm, cordate or truncate, entire or crenate, acute
or obtuse, both sides puberulous, the underside
sometimes glabrous; petiole as long as to a little
shorter than the leaves, 4-25 mm, glabrous.
Flowers 5-12 mm. Pedicels 6-30 mm. Sepals 5,
free, subulate, sometimes triangular, 14—1(—11,)

campanulate, varying from blue to white; lobes
5, for up to 14 connate, nearly equal, elongate
tc elliptic 234-10 by 1-2 mm, entire, acute or
obtuse, glabrous. Stamens: anthers c. 1 mm,
glabrous; filaments linear, broadened to the base,
2-4 mm, finely haired, sometimes glabrous. Ovary
obconical to campanulate, 144-3 by 1-2 mm,
glabrous; style simple, glabrous, rarely hairy,
with narrow stigmas. Fruit ovoid to obovoid,
3-5 by 3-5 mm, with a very thin wall, pendent
on the pedicels. Seeds fusiform-ellipsoid, 1-11,
mm, brown and smooth.

Distr. SE. Asia: N. India (Himalaya), Siam,
S. China (Yunnan, Kweichow), Japan, and
Formosa; in Malaysia: Philippines: Luzon
(Benguet and Lepanto: Mt Osdung; Mt Pulog)
and Panay.

Ecol. In moist mountain (oak) forest, at the
base of trees, 1450-3000 m. Fi. fr. Febr.—June.

by 14 mm, entire, glabrous. Corolla 3-12 mm,

2. WAHLENBERGIA

SCHRAD. [Pl. Sem. Hort. Ac. Gott. 1814, p. 3, nom. nud.] ex Rotu, Nov. Pl. Sp.
(1821) 399, nom. gen. cons., cf. STEEN. Taxon 9 (1960) 125; ?D. Don, Prod. Fl. Nep.
(1825) 156; ScHRAD. Blumenbachia, etc. (1827) 123; Reus. Ic. Pl. Rar. cent. 5°
(1827) 47, t. 480; ScHRAD. Comm. Gott. 6 (1828) 123; DC. Mon. Camp. (1830)
129; BREHMER, Bot. Jahrb. 53 (1915) 9; LOTHIAN, Proc. Linn. Soc. N.S.W. 71 (1947)
201.—Lightfootia Herir. Sert. Angl. (1789) 4, t. 4, non Sw. 1788 (Flac.), nec
SCHREB. 1789 (Rub.); R. S. ADAMSON, J. S. Afr. Bot. 21 (1955) 155.—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. Néerl. Ind. 1 (1844) 484,
nomen; cf. STEEN. Bull. Jard. Bot. Btzg HI, 17 (1948) 463.—Campanopsis O.K.
Rev. Gen. Pl. 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 Mal.) 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 Mal.) superior, lobes 3-6 (in Mal. 3-5), acute, blunt or slightly
acuminate, triangular to linear, persistent. Corolla 5- or 4-partite, or a 3—6- (in
Mal. 3—S-) 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-Mal.
ssp. up to 5), spreading late in anthesis. Ovary inferior (in Mal.) to superior, 2—3-
celled (in extra-Mal. spp. up to 5-celled). Capsule loculicid, opening by as many
apical valves as there are cells in the ovary. Seeds ~, ellipsoid or triangular, shining,
up to 1 mm.

2 FLORA MALESIANA [ser. I, vol. 64

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 forms 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, concealing 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 difficult 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.

Taxon. 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, /.c.) 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 (/.c. p. 12) showed that no
demarcation can be drawn. He maintained both names because of his ‘personal aversion’ to use the
nomenclaturally older name Lightfootia ’ HERIT. 1789 for the Wahlenbergias which he assumed to be
older in the phylogeny, excusing this curious conclusion by stating that in the Campanulaceae genera
are often distinguished by ‘“‘sekundare”’ characters. Dr R. S. ADAmsoN, /.c., 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: ‘‘I 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 scientifically
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’’. F

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 changes 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 the natural subgeneric taxa in various ranks. A similar procedure has been followed within other
natural, but very large genera, as in Astragalus, Bulbophyllum, Carex, Dendrobium, Ficus, Eucalyptus,
Acacia, etc. | 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, /.c. 56) that it would
deviate from Lightfootia by a “‘stigma en téte’’ (capitatum, simplex, l.c. 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 ’ HERIT. is the oldest name, it is a later homonym of Lightfootia Sw.
(see for publication dates Fl. Mal. I, 4, 1954, cxevii, 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
fe Ovary glabrous. Seeds trigonous. Filaments narrow triangular . . . 2. W. hookeri

. 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)

2%4-12 mm (in var. grandiflora 15-25 mm) F 4. W. marginata

1. Wahlenbergia erecta (ROTH ex R. & S.) Tuyn,
nov.comb.—Dentella erecta RoTH ex R. & S.
Syst. Veg. 5 (1819) 25, ex descr.; Nov. Pl. Sp.
(1821) 140; CHam. & SCHLECHTEND. Linnaea 4
(1829) 151; DC. Prod. 4 (1830) 419.—Dentella
perotifolia WILLD. ex R. & S. Syst. Veg. 5 (1819)
25, descr. in syn., nom. illeg.; DC. Prod. 4 (1830)
419.—W. perotifolia W. & A. Prod. (1834) 405,
nom. illeg.; DC. Prod. 7 (1839) 434; Wicut,
Ic. Pl. Ind. 3 (1844-45) 4, t. 842.—Cephalostigma
schimperi Hocust. ex RicH. Tent. Fl. Abyss. 2
(1851) 2; Crarke, Fl. Br. Ind. 3 (1881) 428;
Trim. Fl. Ceyl. 3 (1895) 58.—Cephalostigma
erectum (ROTH) VATKE, Linnaea 38 (1874) 699,
quoad specim. pro parte.—Fig. 1h-i.

Annual, erect herb. Stems 5-10 (—35) cm,
flexuose, branched, blunt-angular, the angles pale
elevated lines, patently -- hispid-hairy. Leaves
5-20 by 114-6 mm, spirally arranged (sometimes
almost distichous), sessile, oblong, sometimes
elliptic, acute, mostly obtuse at the base, sparsely
hairy (hairs especially along the midrib) or almost
glabrous, margin thickened, undulate shallowly
and remotely dentate. Flowers axillary and ter-
minal. Pedicels 5-10 mm, filiform, hairy. Calyx
lobes 5, 114-3 mm, linear to narrow-triangular,
acute, outside sparsely patently hairy. Corolla
5-parted, segments linear acute about as long as

the calyx lobes. Stamens 5, anthers 0.5 by 0.2
mm; filaments 0.8 mm, membranous, broad at
the base, 3-lobed, the side lobes small, the middle
lobe bearing the anther. Ovary 2—2'4 mm, bell-
shaped to obconical, patently hispid-hairy (hairs
up to 14 mm), top conical, apparently lengthening
during anthesis to about 114 mm; style 114 mm,
narrow-cylindric, thickened below the apex, with
3 short stigmatic lobes. Fruit 2-3 mm, obconic
to bell-shaped, 5-nerved, valves 3, before opening
forming a glabrous cone 1—1.7 mm high. Seeds ~,
compressed-ellipsoid, shining, brown, 14 mm.

Distr. East Africa, ?Ceylon, India (the Deccan
and Khasya), in Malaysia: North Sumatra (Karo
and Toba-Batak Highlands). According to
TRIMEN, /.c., probably not in Ceylon.

Ecol. Open, heath-like land and waste places,
750-1400 m.

Vern. Dukut tawar, Karo-Batak.

Note. This species has sometimes been con-
fused with W. hirsuta (EDGEW.) TUYN, nov.comb.*
(Cephalostigma hirsuta EpGew. Trans. Linn. Soc.
20, 1846, 81); the latter has also a hispid ovary
but trigonous seeds.

1 The name Wahlenbergia hirsuta STEUD.
Nomencl. ed. 2, 2, p. 782, is a nomen nudum and
has no nomenclatural status.

114 FLORA MALESIANA [ser. I, vol. 61

2. Wahlenbergia hookeri (CLARKE) TUYN, nov.comb.
—Cephalostigma hookeri CLARKE, Fl. Br. Ind. 3
(1881) 429; DAaNGuy, FI. Gén. I.—C. 3 (1930) 690;
Cras, Fl. Siam. En. 2 (1936) 307.—Cepha-
lostigma paniculatum (non 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—114 mm, bell-shaped to ob-
conic, glabrous; top conic, up to 1 mm high;
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 PiyL 144, BO, L, K), only once collected,
June 1929.

Ecol. In reed stands (?Themeda), probably c.
1000—1200 m alt.

Note. Differs from Wahlenbergia candollei
TuyNn, nom.nov. [Cephalostigma paniculatum DC.
Mon. Camp. (1830) 117, non Wahlenbergia pani-
culata (THuUNB.) DC. /.c. 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 MerrR. & 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,
2-11 by 14-3 mm. Flowers solitary. Pedicels 2—7
cm. Calyx lobes 5, + spreading, linear-lanceolate,
acute to acuminate, usually entire, always longer
than the ovary, 214-4 mm long. Corolla tube
about twice as long as the calyx lobes, sometimes
longer, 614-8 mm; lobes 5, ovate, acute, 5—10 mm.
Filaments linear-triangular, 214-314 mm. Ovary
obconical, 3-celled, 1.2-3 mm; style slightly

Fig. 1. Wahlenbergia hookeri (CLARKE) TUYN.
a. Habit, x 24, b. flower, stamens removed, x 7,
c. fruit, x 7, d. seed, x 13, e. flower, stamens
removed, x 7, f. fruit, x 7, g. seed, x 13.—
Wahlenbergia erecta (ROTH ex R. & S.) Tuyn.
h-i. Flowers, stamens removed, x 7, j. 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

Fig. 2. Wahlenbergia confusa MERR. & PERRY.
a. Habit, x 24, b. longitudinal section of flower,
<x 2, c. filament, x 6 (BRAsSs 9399).

shorter than the corolla tube, 5'4-7 mm; stigma
3-lobed, (1—-)1144-3 mm long. Fruit slightly wider
than long, 3-valved, 2-3 mm diam., crowned by
the recurved calyx lobes.

Distr. Malaysia: West New Guinea (vicinity
of Mt Wilhelmina).

Ecol. Wet, grassy, open places along streams,
once noted on sandstone, 3400-4000 m.

Note. MERRILL compared his new species with
W. eurycarpa DomIN (Bibl. Bot. Heft 89, 1929,
638) but already remarked that the fruit in the latter
would appear too long. DomIN himself compared
his species with W. sieberi and gave the differences
with W. multicaulis which species I have reduced
to W. marginata. As DomMIN neither mentions the
shape of the filaments nor the relative length of
the corolla tube, and as his material is not avail-
able to me, I refrain from giving a definite
opinion.

4. Wahlenbergia marginata (THUNB.) DC. Mon.
Camp. (1830) 143; Prod. 7 (1839) 433; Stes. FI.
Jap. 2 (1845) 179; Koorpb. Exk. Fl. Java 3 (1912)
300; NANNFELDT, Act. Hort. Gothob. 5 (1929)

32; ALSTON, FI: Ceyl.. Suppl: 6 (1931). 176;
DOCTERS VAN LEEUWEN, Verh. Kon. Ak. Wet.
A’dam sect. 2, 31 (1933) 267; Hocur. Candollea
5 (1934) 290, cum var.; STEEN. Bull. Jard. Bot.
Btzg III, 13 (1934) 179; Cras, 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; Hatcu, Trans.
Proc. R. Soc. New Zeal. 79 (1952) 368, cum var.;
YUNCKER, Bull. Bern. P. Bish. Mus. 220 (1959)
261.—Campanula marginata THUNB. FI. Jap.
(1784) 89.—Campanula gracilis Forsr. Prod.
(1786) 84; R.Br. Prod. (1810) 561, incl. var.—
Campanula quadrifida R.Br. Prod. (1810) 561.—
Campanula dehiscens Roxs. [Hort. Beng. (1814)
85] ex WALL. As. Res. 12 (1816) 571, cum ic.;
Fl. Ind. 2 (1824) 96.—Campanula agrestis WALL.
in Roxb. FI. Ind. 2 (1824) 97; Cat. (1829) 1292.—
Campanula lavandulaefolia REINW. ex BL. Bijdr.
(1825) 726.—W. gracilis [SCHRAD. Blumen-
bachia (1827) 38, in obs.] DC. Mon. Camp.
(1830) 142, incl. var.; Prod. 7 (1839) 433; JUNGH.
Nat. Geneesk. Arch. Neérl. Ind. 2 (1845) 48, cum
var. hirsuta JUNGH.; HooK. f. Fl. Tasm. 1 (1860)
239; Bru. Fl. Austr. 4 (1869) 137; CLARKE, FI.
Br. Ind. 3 (1882) 429; Wars. Bot. Jahrb. 18
(1893) 212; Hems-. J. Linn. Soc. 30 (1894) 183;
BAILEY, Queens]. Fl. 3 (1900) 922; CHEESEMAN,
New Zeal. Fl. (1906) 402; DEN BERGER, Trop.
Natuur 6 (1917) 104 f. 4; STEARN, Dict. Gard.
R.H.S. 4 (1951) 2257.—W. quadrifida (R. BR.)
DC. Mon. Camp. (1830) 144; Prod. 7 (1839) 433;
N. E. Brown, Gard. Chron. 54 (1913) 316;
Domi, Bibl. Bot. Heft 89 (1929) 1192; LoTHIAN,
Proc. Linn. Soc. N.S.W. 71 (1947) 210.—W.
dehiscens (Roxs.) DC. Mon. Camp. (1830) 145;
Prod. 7 (1839) 434; LoTHIAN, Proc. Linn. Soc.
N.S.W. 71 (1947) 216.—W. agrestis DC. Mon.
Camp. (1830) 145; Prod. 7 (1839) 434; WIGHT,
le: Pl; Ind> Or 471850) 11753 HooKe ya cc an.
Proc. Linn. Soc. Lond. 2 (1858) 21; LOTHIAN,
Proc. Linn. Soc. N.S.W. 71 (1947) 215.—W.
lavandulaefolia DC. Mon. Camp. (1830) 144;
Prod. 7 (1839) 433; ZoLi_. Nat. Geneesk. Arch.
Neérl. Ind. 2 (1845) 567.—W. indica DC. Mon.
Camp. (1830) 146; Prod. 7 (1839) 434; WiGur,
Ic. Pl. Ind. Or. 4 (1850) 1176; LOTHIAN, Proc.
Linn. Soc. N.S.W. 71 (1947) 209.—W. sieberi
DC. Mon. Camp. (1830) 144; Prod. 7 (1939)
433; LoTHIAN, Proc. Linn. Soc. N.S.W. 71 (1947)
219.—W. multicaulis Bru. in Hiigel, En. PI.
Nov. Holl. (1837) 75; DC. Prod. 7 (1839) 433;
N. E. Brown, Gard. Chron. 54 (1913) 337;
LOTHIAN, Proc. Linn. Soc. N.S.W. 71 (1947) 229;
STEARN, Dict. Gard. R.H.S.4(1951) 2258.—Campa-
nula sieberi DiEtR. Syn. Pl. 1 (1839) 752.—Campa-
nula indica Dtetr. I.c. 753.—Campanula littoralis
LABILL. Pl. Nov. Holl. 1 (1844) 49, t. 70.—Cam-
panulopsis cyanea ZoLL. & Mor. Nat. Geneesk.
Arch. Neérl. Ind. 1 (1844) 484, nomen.—W.
simplicicaulis DE VRIESE in Lehm. PI. Preiss. 2
(1846) 241; LoTrnHIAN, Proc. Linn. Soc. N.S.W. 71
(1947) 209.—Lobelia dubia DE VRtESE in Lehm.
Pl. Preiss. 2 (1846) 242.—Lightfootia gracilis
Mia. FI. Ind. Bat. 2 (1856) 567, cum var. lavan-

[ser. 1; Vola

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

dulaefolia Mig.—Campanopsis marginata O.K.
Rev. Gen. Pl. 2 (1891) 378, cum var. rigida, nom.
illeg.—Cervicina gracilis Britt. Ill. Bot. Cook
2 (1901) 56, t. 182.—W. bivalvis MerR. Philip.
J.Sc. 1 (1906) Suppl. 242.—W. gracilenta LOTHIAN,
Proc. Linn. Soc. N.S.W. 71 (1947) 217.—Fig.
3-5.

Perennial. Stem 3—60 cm, glabrous or hairy, or
only pilose in the lower part. Leaves linear to
elliptic, the lower ones usually tending to be
broadest, decreasing in size upwards, 1/5—S14 cm
by 44-5 mm. Inflorescence 1- to few-flowered.
Pedicels 1-15 cm. Calyx lobes 3-5, linear-lanceo-
late, acute or + obtuse (China), usually entire,
sometimes sparsely dentate, usually about as
long as the corolla tube, sometimes shorter or up
to twice as long, 0.8-4 mm. Corolla tube 1-5 mm,
lobes 3-5, ovate or elliptic, acute or acuminate,
1.2-7(?) mm. Anthers narrow, 14-214 mm long;

basal part of the filaments about pentagonal or
obtrapezoid, abruptly narrowed, the upper por-
tion filiform, 0.8—2 mm. Ovary obconical bell-
shaped, 2~—3-celled; 1-S mm; style 1.6—5 mm;
stigma 2-3-lobed. Fruit obconical to bell-shaped,
2-3-valved, 114-10 by 1-5 mm.

Distr. Widely distributed from China and
Japan through tropical SE. continental Asia and
Malaysia to New Caledonia, Tonga, Australia,
the Kermadecs, and New Zealand; in Malaysia:
Java (from Mt Patuha eastwards), Lesser Sunda
Is (Bali, Lombok, Sumbawa, Flores, Alor, Timor),
S. Moluccas (Aru Is), SW. Celebes (Mt Bonthain),
Philippines (Luzon), and New Guinea.

MiIQueL (Sum. 1860, 234) recorded it from
Sumatra, but this seems erroneous.

CLARKE & HEMSLEY recorded W. gracilis even
from S. Africa, but I have seen no records from
Africa; this is probably in confusion with W.

CAMPANULACEAE (Moeliono & Tuyn)

117

Fig. 4. Distribution of Wahlenbergia marginata
(THuNB.) DC. with localities dotted in the area
covered by the map.

gracilis E. MEyY., a nomen nudum and a later
homonym.

Ecol. In Malaysia only in the mountains (100,
800—900—) 1000-3500 m, in light Eucalyptus and
Casuarina forest, along forest edges, among rocks,
on lava-streams, in grassfields, and along trails as
an apophyte. In India curiously recorded from
sea-level upwards (CLARKE, /.c.), in Malaysia only
found at low altitude in the Aru Is. Fi. fr. Jan.—
Dec., specially May—July.

It seems that this plant prefers regions with a
feeble to strong dry season which may explain
its absence in the wet rain-forest core of West
Malaysia (Sumatra, Malaya, Borneo). According
to DocTERS VAN LEEUWEN self-pollination is the
rule in Java.

Vern. Angkéb, kérékan lanang, patikan, télugi,
J (local), djukut riut, S; Papua: ik, Dunantina,
iki, Asaro, Kefamo, goiekul, Chimbu, Masul,
kulkal, Hagen, Togoba.

Uses. According to BACKER used against skin
diseases on Mt Diéng.

Notes. In the past many authors have already
pointed to the high degree of variability of this
species, e.g. DE CANDOLLE (1830), JUNGHUHN
(1845), Hooker f. (1860), BENTHAM (1869),
BAILEY (1900), CHEESEMAN (1906), and STEARN
(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
juvenile stage, before having attained a good,
full-grown vegetative state.

Further the woody shoots of old obviously
perennial plants may produce such thin shoots
(cf. Bro. Fl. Austr. 4, p. 137; CLARKE, FI. Br.
Ind. 3, p. 430; and BAILEY, Queensl. Fl. 3, p. 922).

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
in both the absolute and relative dimensions of
nearly all the floral parts, without an appreciable
geographical correlation (raciation). This variation
has been drawn for the shape of the filaments
in fig. 5. It must be taken into consideration that
the filaments and their margins are bent inwards
already in a very early stage (cf. DECAISNE, Rev.
Hort. 3, 1849, 41 fig.); in herbarium material it
is very difficult to flatten them out because of
their delicacy and entangled fringe-like marginal
hairs.

The length of the calyx lobes may vary as much
as 114 mm in different flowers of a single plant
during anthesis (HLB 958-307-469).

From the Philippines MERRILL has described
W. bivalvis MeRR. which would be characterized
by capsules with 2 apical valves. In the syntypes
of MERRILL 4361 (Bo, L, Ny) which I could
examine I have only found capsules with 3 valves.
In a number of other specimens ranging through
Celebes, the Lesser Sunda Is, and S. Moluccas
to New Guinea and Thursday Island I have found
indeed 2-valved capsules and all these specimens
have very slenderly branched inflorescences
(except ELBERT 1083 from Lombok): Celebes:
BUNNEMEIJER 11528, partly; Lombok: ELBERT
1332; Aru Is: BUWALDA 5354; SE. New Guinea:
ForBES 847; Thursday Island: JAHERI s.n.; Warrior
Island: Le GuiLLou Warrior 16.

But there are other less delicate-branched
specimens which have in a single plant 2- and 3-

Fig. 5. Wahlenbergia marginata (THUNB.) DC.

Geographical survey of stamen types (boiled

material). a. From Hunan (China), 6. Hongkong,

c. Hongkong, d. Central Java (Mt Sumbing),

e. Lombok, f. Timor, all < 10 (a FAN & LI 57,

b LAMONT 405, c WRIGHT s.n., d LORZING 3, e
ELBERT 1332, f Forses 4048).

118

FLORA MALESIANA

[ser. I, vol. 6!

valved capsules: Lombok: ELBert 1557; China:
Herb. Bog. sine coll. 133584; Celebes: sine coll.
136. Further there are also slenderly branched
specimens with 3-valved capsules: Celebes:
BUNNEMEIER 11528, partly, VAN STEENIS 10400.

For these reasons W. bivalvis MERR. cannot be
maintained.

Chromosomes. According to the Chromo-
some Atlas by DARLINGTON & WYLIE (1955) W.
gracilenta, W. consimilis, W. quadrifida, and W.
gracilis would have 2 n= 18, 36, 54, and 64
chromosomes respectively, with the basic numbers
9 and (?) 8 which might point to the occurrence
of chromosome races.

var. grandiflora Tuyn, nov.var.—W. consimilis
LOTHIAN, Proc. Linn. Soc. N.S.W. 71 (1947)
223; MELVILLE, Bot. Mag. 172 (1959) t. 343.—
W. gloriosa LOTHIAN, l.c. 224.

Corolla longior, c. 1Y.—2% cm longa.

Similar to the species, but the corolla (in Mal.)
134-2 cm long, the tube c. 6-10 mm, usually c.
114-2 times as long as the calyx lobes (3—5 mm).
Calyx lobes always longer than the ovary (2-214
mm). Fruit broad-obconical to bell-shaped,
414-6 by 4-5 mm.

Distr. Australia (S. New South Wales to S.
Australia and Tasmania), in Malaysia: New
Guinea.

Notes. Also in this variety the hairiness is
variable.

Var. grandiflora seemed to differ from the
descriptions of W. consimilis and W. trichogyna

mainly in the calyx lobes, which are somewhat
shorter compared with the corolla tube (cf.
LOTHIAN, /.c. and MELVILLE, /.c.). This character,
however, proved to be very variable in the
Australian material which I could examine. The
relative length of calyx lobes and corolla tube
differs sometimes considerably in different
flowers of the same specimen (e.g. HOOGLAND
3074, 3075), whereas in the typical trichogyna
forms the corolla tube is sometimes much longer
than the calyx lobes.

It seems that W. trichogyna STEARN, Gard.
Chron. III, 130 (1951) 169.—Campanula gracilis
(non Forst.) Sims, Bot. Mag. 18 (1803) t. 691.—
Campanula vincaeflora VENT. Jard. Malm. (1803)
t. 12, nom. illeg—W. gracilis var. vincaeflora DC.
Mon. Camp. (1830) 142; Prod. 7 (1839) 433.—W.
vincaeflora (VENT.) DECNE, Rey. Hort. 3 (1849) 4,
cum ic. col.; LOTHIAN, Proc. Linn. Soc. N.S.W.
71 (1947) 220, differs only in having a hairy ovary
and is better subordinated as subvar. trichogyna
(STEARN) TUYN, nov. subvar. It is native in East
Australia and has not yet been found in New
Guinea.

CRUTTWELL 776 and BRAss 22258 have the calyx
lobes clearly dentate and the broadened part of
the filaments seems longer than usual.

As there are found sometimes specimens with
a few teeth at the margin of the calyx lobes (e.g.
CLEMENS 4873, 5870) and the filaments are very
variable throughout the species, there seems no
reason to create a new variety.

3. CODONOPSIS

WALL. in Roxb. FI. Ind. 2 (1824) 103; Komarovy, Act. Hort. Petrop. 29 (1908)
98; Cupp, J. Linn. Soc. Bot. 38 (1908) 374; ANTHONY, Not. R. Bot. Gard. Edinb.
15 (1926) 173; NANNFELDT, ibid. 16 (1931) 149; Bot. Tidsskr. 34 (1940) 381.—Glo-
socomia D. Don, Prod. Fl. Nepal. (1825) 158.—Campanumoea BL. Bijdr. (1826)
726.—Cyclocodon GrirF. Not. Pl. As. 4 (1854) 279; Kurz, J. As. Soc. Beng. 46, i
(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 cylindric; 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 Cupp, /.c., and KoMAROY, /.c., among the former for
instance Hooker f. (Ill. Him. Pl. 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 ce myays = 1

by 5 small, apical valves (slits) in the first and indehiscent in the latter. When capsular, the valves are
formed in the conically protruding apical part of the fruit which is crowned by the style.

In order to find out whether this is a really ‘natural’ character I 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
Campanumoea lancifolia shows in foliage and habit more resemblance with for instance Codonopsis
purpurea WALL. in Roxs. than with other Campanumoeas; 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 different place of insertion of calyx and corolla, schematic,
largely after Cuipp. a—d. Types used for subdivision in CuIpp’s key, /.c. 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.

In merging the two genera I feel strengthened by a succinct note by DieLs, who remarked under
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.

Cupp, ANTHONY, and NANNFELDT have subdivided the genus into sections and series. I have refrained
from fitting in this scheme the Malaysian Campanumoeas. 1 do not believe that they can be main-
tained under one subgenus or section, as they differ 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 difficult 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 CuippP has illustrated four
floral types can be distinguished in Codonopsis, depending on the insertion of the calyx and corolla,
viz (1) both calyx and corolla adnate to the ovary halfway, (2) ditro 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
|. Sprawling or erect. Leaves decurrent at the base. Flowers 34-114 cm, white. 2. C. lancifolia

{. Codonopsis javanica (BL.) Hook. f. Ill. Him. panumoea javanica Bu. Bijdr. (1826) 727; DC.
Pl. (1855) 116, t. 16B; Mig. Fl. Ind. Bat. 2 (1857) Prod. 7 (1839) 423; Hassxk. 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. f. & Tu. J. Proc. Linn. Soc.
Bot. 2 (1858) 18; Kurz, J. As. Soc. Beng. 46, ii
(1877) 209; CLARKE, FI. Br. Ind. 3 (1881) 435;
Fors. & Hems_. J. Linn. Soc. Bot. 26 (1889) 8;
Diets, Bot. Jahrb. 29 (1901) 606; MAkINo, Bot.
Mag. Tokyo 22 (1908) 155, incl. var. japonica;
Koorp. Exk. FI. Java 3 (1912) 300; Fl. Tyib. 3
(1923) 54; HeEyne, Nutt. Pl. 2 (1927) 1427;
Danouy, FI. Gén. I.-C. 3 (1930) 694 & f. 78,10;
Hocur. Candollea 5 (1932) 290; Merr. Lingn.
Sc. J. 11 (1932) 60; Crais, Fl. Siam. En. 2 (1936)
307; MASAMUNE, FI. Kain. (1943) 330; BAcK.
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 183, p. 3;
Makino, Ill. Fl. Japan (1954) 81, cum var.—C.
cordata Hassk. Nat. Tid. N.I. 10 (1856) 9; Ned.
Kruidk. Arch. 4 (1856) 1; Mia. Fl. Ind. Bat. 2
(1857) 566; WALP. Ann. 5 (1858) 393; Bot. Mag.
89 (1863) t. 5372.—Campanumoea cordata Mia.
Sum. (1862) 599; Maxim. Bull. Ac. Imp. Sc.
Pétersb. 12 (1868) 68; Cras, Kew Bull. (1911) 404.
—Campanumoea japonica MAxiM. Bull. Ac. Imp.
Sc. Pétersb. 12 (1868) 67, non SIEBOLD et MORREN,
1863.—C. cordifolia KOMAROV, Act. Hort. Petrop.29
(1908) 108.—Campanumoea maximowiczii HONDA,
Bot. Mag. Tokyo 50 (1936) 389.—Fig. 7.

A sinistrorse-twining plant, up to 2 m (ac-
cording to Koorpers 6-14 m long); stem 114-4
mm diam., glabrous. Leaves opposite, higher up
often spirally arranged, ovate to oblong-ovate,
2.6-8 by 2-5 cm, base cordate, apex obtuse, acute,

Fig. 7. Codonopsis javanica (BL.) Hook. f. a.
Flower, 5. fruit sustained by the star-shaped calyx
and topped by the marcescent corolla, all «x %.

sometimes acuminate, serrulate to serrate, some-
times crenate, on both sides puberulous to hairy,
sometimes glabrous, underside often pruinose;
petiole 144-614 cm. Flowers axillary, 114-3.8
mm. Pedicels 1-5'4 cm. Calyx lobes oblong to
lanceolate, 11-23 by 3-4 mm, entire or denticulate
to serrate, glabrous or puberulous to hairy.
Corolla 12-35 mm, more than half-way lobed,
greenish-white or yellowish-white, outside some-
times hairy, tube inside with purplish streaks or
veins, lobes ovate, acuminate. Anthers c. 2-4 mm;
filaments 3-8 mm, linear, broadened towards the
base, glabrous. Style 5-10 mm, glabrous or hairy
with 3-6 ovate-acute to elliptic lobes. Ovary
(3-)4-S5(-6)-celled, the outside partly concealed
by the corolla, campanulate, visible part from the
outside 3-4 by 3-4 mm, glabrous. Fruits sub-
globular, 144-114 by 1-2'4 cm, red to dark-purple
to black-bluish, at the base sustained by the
persistent and patent or sometimes reflexed wine-
red calyx lobes and crowned by the withered
corolla; walls membranous to coriaceous, some-
times fleshy; placentas fleshy. Seeds ovoid to
ovoid-cylindrical, c. 1 mm, reticulate, light brown.
Distr. In SE. Asia (from the Deccan through
the Himalayan region, Burma, and Siam to South
and Central China), Japan, Hainan; in Malaysia:
Central to South Sumatra (Mts Kerintji and
Dempo), Java (from Mt Pangrango eastwards).
Ecol. In open forest, mostly on forest edges,
in secondary forest and thickets, sometimes in
grassy fields, 900-2200 m. Fi. fr. Jan.—Dec.
Vern. Kitjépot, srintil-srintil, susu munding,
S, gutji, indil-indil, sigerpolo, urék-urék polo, J.
Uses. The fruits are eaten and the tuberous
roots are used for drugs.

2. Codonopsis lancifolia (ROxB.) MOELIONO, nov.
comb.—Campanula lancifolia Roxs. Fl. Ind. 2
(1824) 96; DC. Prod. 7 (1839) 485.—Campanumoea
celebica BL. Bijdr. (1826) 727; DC. Prod. 7 (1839)
423; CLARKE, Fl. Br. Ind. 3 (1881) 436; BogeRL.
Handl. 2 (1891) 259; Srapr, Trans. Linn. Soc. II,
4 (1894) 188; Gamste, J. As. Soc. Beng. 74, ii
(1905) 53; Rosrnson, Philip. J. Sc. 3 (1908)
Bot. 216; Crats, Kew Bull. (1911) 404; DreLs,
Bot. Jahrb. 55 (1917) 121; Merr. En. Born.
(1921) 585; Rip. Fl. Mal. Pen. 2 (1923) 202;
Merr. En. Philip. 3 (1923) 587; DANGuy, FI.
Gén. I.-C. 3 (1930) 693; Craip, Fl. Siam. En. 2
(1936) 307; MASAMUNE, En. PI. Born. (1942) 123;
BAcK. Bekn. FI. Java (em. ed.) 8 (1949) fam. 183,
p. 4.—C. truncata WALL. [Cat. (1829) n. 1301,
nomen] ex DC. Mon. Camp. (1830) 122; Prod.
7 (1839) 423; Mia. FI. Ind. Bat. 2 (1857) 566.—
C. albiflora GrirF. Not. Pl. As. 4 (1854) 279.—
Cyclocodon adnatus GrirF. I.c. 278.—C. celebica
Mig. Fl. Ind. Bat. 2 (1857) 566.—C. leucocarpa
Mia. /.c. 565.—Cyclocodon truncatum Hook. f. &
Tu. J. Proc. Linn. Soc. Bot. 2 (1858) 18.—
Cyclocodon lancifolium Kurz, Flora 55 (1872)
303; J. As. Soc. Beng. 46, ii (1877) 210.—Cam-
panumoea axillaris Ovty. in Hook. Ic. III, 8
(1888) t. 1775; Hemsi. J. Linn. Soc. Bot. 26
(1889) 7.—Campanumoea truncata Diets, Bot.

Dec. 1960]

-CAMPANULACEAE (Moeliono & Tuyn) |

ait

Jahrb. 29 (1901) 606; MAKINo, Bot. Mag. Tokyo
8 (1904) 21; Merr. Philip. J. Sc. 7 (1912) 104.—
Campanumoea lancifolia Merr. En. Philip. 3
(i923) ros7s Lmens Se. Js 5 (1927) 185 Rap:
Mich. Ac. Sc. 20 (1935) 111; YAMAMOTO, Obs.
Fl. Form. 13 (1936) 147; MASAMUNE, FI. Kain.
(1943) 33; HarA, En. Sperm. Jap. 2 (1952) 96.

Sprawling or erect herb, with a hollow stem,
c. 3 m high, nearly glabrous or hairy, mostly
branched. Leaves opposite, petioled, lower ones
ovate to ovate-elliptic, higher ones elliptic, often
bract-like, 3-14 by 1-4 cm; acute to acuminate,
bluntly edged at the base, coarsely serrate to
serrulate, upperside glabrous, underside glabrous
or puberulous to hairy; petiole 3-15 mm, glabrous.
Flowers 7-15 mm, axillary, solitary or in cymes
of 3, through reduction of the higher leaves re-
sembling a terminal panicle. Pedicels or peduncles
(1-)3-6 cm, glabrous, with two bracteoles, 14-2
cm long, linear to elliptic, glabrous or hairy.
Buds winged, ovate. Calyx lobes 4-7, lanceolate,
4-10 by | mm, entire to dentate-serrate to pinnati-
fid, carnosulate, glabrous to puberulous. Corolla
white, pale pink, lilac, outside glabrous, corolla
tube as long as the lobes, lobes ovate to trian-
gulate, acute, entire. Stamens 4-6 mm. Anthers
as long as the filaments. Filaments linear to
broadened at the base in various degree (deltoid
to ovate), the broad forms inside with long hairs
at the base. Sty/e glabrous or scabrid. Ovary
cupular to campanulate, 2-4 mm diam., glabrous
to puberulous. Berry + globular with a flattened
apex, the persistent calyx adnate about half-way,
greenish, turning to white, c. 14-1 cm diam.
Seeds many, testa fine-reticulate.

Distr. SE. Asia (India: Himalaya and Sikkim;
Burma) to S. China, Formosa, and Hainan; in
Malaysia: Malay Peninsula (Selangor: Taiping
Hills; Perak: Semangkok Pass, ex RIDLEY, not

seen), Sumatra, E. Java (Pantjur Idjen, once
found), N. Borneo (Kinabalu), Celebes, Philip-
pines (N. Luzon; Panay; Mindanao), Moluccas
(Buru and Ambon), and New Guinea.

Ecol. In open terrain or along forest-borders
and trails, on tali and near streamlets, rarely
in forest regrowth and young secondary forest,
often in wet places, 280-1500 m. Fi. fr. Jan.—Dec.

Vern. Gordang-gordang, M (Toba), émlapagar,
Palu (W. Cel.); Philippines: /akoronbolan, Buk.

Note. There are two subspecies which largely
replace one another geographically.

ssp. lancifolia.—All synonyms except C. celebica
and C. leucocarpa.

Leaves generally large, 8!4-16!4 by 3-714 cm,
towards the apex of the stems not very much
reduced, never glaucous beneath, entirely glabrous.
Calyx lobes coarsely dentate to subpectinate.
Filaments distinctly widened downwards, either
triangular or roundish, long-hairy inside.

Distr. Continental Asia, in Malaysia: N. Su-
matra (Atjeh, Toba), Philippines, and Moluccas
(Buru, Ambon).

ssp. celebica (BL.) MOELIONO, comb. nov.—Cam-
panumoea celebica BL. I.c.—C. celebica (BL.)
MiQ.—C. leucocarpa Mia.

Leaves averagely smaller, 5!4-8(-1114) by
114-234(-334) cm, towards the apex distinctly
shorter giving the apical part of the stem the ap-
pearance of a panicle, glaucous and hairy beneath
or glabrous. Calyx lobes entire to toothed. Fila-
ments linear to triangular-broadened towards the
base, glabrous (once found short hairy inside,
OUWEHAND 215).

Distr. Siam (KERR 1217), in Malaysia: Central
and South Sumatra, SE. Java, Borneo, Celebes,
Philippines, Moluccas, and New Guinea.

6. LOBELIA

LINNE, Gen. Pl. (1754) 401; Sp. Pl. 2 (1753) 929, non sensu ADANS. Fam. Pl. 2
(1763) 157 et MILLER, Gard. Dict. 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) 408.—Pratia GAupicH. Ann. Sc. Nat. 5 (1825) 103; PResL, Prod. Mon.
Lob. (1836) 46; Wimmer, Pfl. R. Heft 106 (1943) 104.—Dortmanna ADANS. Fam.
Pl. 2 (1763) 134; SteuD. Nomencl. (1840) 526; O. K. Rev. Gen. 2 (1891) 379 (Dort-
mannia).—Rapuntium MILLER, Gard. Dict. ed. 8 (1768); PREsL, Prod. Mon. Lob.
(1836) 11.—Piddingtonia DC. Prod. 7 (1839) 341; Mia. FI. Ind. Bat. 2 (1857) 572.—
Isolobus DC. Prod. 7 (1839) 352.—Speirema Hook f. & Tu. 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, alternate 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 ~, 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 ~, 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 angulata, 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, efc.) 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, -_ sparsely long-white-pilose along the margin
and on the upper surface. Stems angular, unbranched. Cauline leaves oblanceolate, upwards
gradually smaller, 2-5 by 4-1 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, 2-414 cm long. Berry globose, 7-12 mm diam., on a reflexed pedicel,

wiolet: giinally, solack=purple 1. (7. \flsntie. hou fae SE ee es ere ie 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)

23

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

8. Free basal part of all filaments of equal width. Stem terete, under each leaf with 2 narrow ridges.

Stem,

leaves, pedicels, and calyx crisped-patent-hairy. Leaves ovate to roundish with very
blunt to subcordate base set off against a distinct petiole (1—-)3—20 mm long
7. Only the two anterior anthers bearded at their apex. Seeds ellipsoid

4. L. zeylanica
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,

Filaments glabrous
10. Leaf-base not semi- -amplexicaulous:

geass Plant glabrous.
. 12. L. conferta

11. Leaves glabrous, sessile, with acute base, subentire, oblong to elliptic, rather narrow. Testa

smooth

6. L. chinensis

11. Leaves exceptionally glabrous, at least the lower almost always petioled, mostly ovate to rounded,

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, Fei campanulate, with narrow-lanceolate

or ovate-lanceolate lobes. Hairs 1-celled. Testa finely reticulate

1. Stemless. Flowers solitary axillary,

1. Lobelia nicotianaefolia RoTH ex R. & S. Syst.
Veg. 5 (1819/?20) 47; RotH, Nov. Pl. Sp. (1821)
143; WALL. in Roxb. FI. Ind. 2 (1824) 110; PI.
As. Rar. 2 (1831) 43; G. Don, Gard. Dict. 3
(1834) 709; DC. Prod. 7 (1839) 381; WiGur,
Illustr. 2 (1850) 111, t. 135 f. 1-10; Hoox. f. &
Tu. J. Proc. Linn. Soc. Bot. 2 (1858) 29; Hook.
f. Bot. Mag. 92 (1866) t. 5587; CLARKE, FI. Br.
Ind. 3 (1881) 427, incl. var. trichandra; TRIM.
Handb. FI. Ceyl. 3 (1895) 57; PEARSON, J. Linn.
Soc. Bot. 34 (1898) 348; Merr. & MERRITT,
Philip. J. Sc. 5 (1910) Bot. 392; Merr. En. Philip.
3 (1923) 588; Sxkotrss. Medd. Géteb. Bot.
Tradg. 4 (1928) 9-13, f. 1, 8=11, 12b, incl. var.
macrostemon, l.c. 13, f. 13-14; ALSTON, FI. Ceyl.
Suppl. (1931) 175; Etmer, Leafl. 9 (1934) 3180,
incl. var. mollis; KAusik, J. Ind. Bot. Soc. 17
(1938) 161-168; Wimmer, Pfl. R. Heft 107 (1953)
643, incl. var. nicotianaefolia, bibarbata, & tri-
chandra; SANTAPAU, Rec. Bot. Surv. Ind. 16
(1953) 157.—L. pyramidalis WALL. As. Res. 13
(1820) 376; D. Don, Bot. Mag. 50 (1823) t. 2387;
WALL. in Roxb. Fl. Ind. 2 (1824) 113; D. Don,
Prod. Fl. Nepal. (1825) 57; WALL. Pl. As. Rar.
2 (1831) 42; G. Don, Gard. Dict. 3 (1834) 709;
DC. Prod. 7 (1839) 381, incl. var. 6; Hook. f.
& TH. J. Proc. Linn. Soc. Bot. 2 (1858) 29;
CLARKE, Fl. Br. Ind. 3 (1881) 426; ForBes &
HEMSLEY, J. Linn. Soc. Bot. 26 (1889) 3; SkoTTsB.
Medd. GG6teb. Bot. Tradg. 4 (1928) 17, 21, f. 12e,
25-31; DANGuy, FI. Gén. I.-C. 3 (1930) 676,
f. 76 1-4; Crats, Fl. Siam. En. 2 (1936) 304;
Wimmer, Pfl. R. Heft 107 (1953) 646.—L. rosea
WALL. in Roxb. Fl. Ind. 2 (1824) 115; Pl. As.
Rar. 2 (1831) 42, t. 152; G. Don, Gard. Dict. 3
(1858) 29; Drury, Handb. Ind. FI. 2 (1866) 108,
excl. syn.; Kurz, J. As. Soc. Beng. 46, ii (1877)
212; CLARKE, FI. Br. Ind. 3 (1881) 427; J. Linn.

11. L. angulata

c. Y% cm long. Leaves very thin, roundish, petioled.

7. L. archboldiana

Soc. Bot. 25 (1890) 41; DANGuy, FI. Gén. I.-C.
3 (1930) 675; Wimme_r, Pfl. R. Heft 107 (1953)
653.—L. excelsa LEscH. ex Roxs. Fl. Ind. 2
(1824) 114, non BonpL. (1813); WALL. Pl. As.
Rar. 2 (1832) 42; G. Don, Gard. Dict. 3 (1834)
709; DC. Prod. 7 (1839) 381; WiGurt, Ic. 4 (1850)
t. 1173-4; Tuwaites, En. Pl. Zeyl. (1860) 170;
CLARKE, Fl. Br. Ind. 3 (1881) 427.—L. stimulans
Ham. ex D. Don, Prod. Fl. Nepal. (1825) 157.—
L. purpurescens WALL. Cat. (1828) 1307, non R.BR.
(1810), nomen nudum ab auctore ipso ibidem
relictum p. 157.—L. colorata WALL. Pl. As. Rar.
2 (1831) 42; DC. Prod. 7 (1839) 380; Hook. f.
& fn J. Proc, Einn, Socs Bot. 2) @s58)ezse
Drury, Handb. Ind. FI. 2 (1866) 108; CLARKE,
Fl. Br. Ind. 3 (1881) 426; Wimmer, Pfi. R. Heft

Pres_, Prod. Mon. Lob. (1836) 23.—Rapuntium
nicotianaefolium PRESL, /.c. 24.—Rapuntium colo-
ratum PRESL, l.c. 24.—Rapuntium wallichianum
Pres_, /.c. 24.—Rapuntium roseum PRESL, l.c.
24.—Rapuntium leschenaultianum PRESL, l.c. 24.—
L. wallichii SteuD. Nomencl. 2 (1841) 62, nomen
superfl. ad L. purpurescens WALL.—L. robusta
WALL. ex VoiGcT, Hort. Suburb. Calc. (1845) 367,
non GRAHAM (1831).—L. eurostos VoicrT, l.c. 367.—
L. aromatica Moon [Cat. Pl. Ceyl. (1824) 14,
nomen] ex WiGurT, Ic. 4 (1850) 2, t. 1172; Hook.
fe & De. Je Proc. Binns Soc; Bot. 2) (1858)F 23;
ALSTON, FI. Ceyl. Suppl. (1931) 175.—L. tri-
chandra WiGur, Ic. Pl. Ind. Or. 4 (1853) t. 1171;
Sxottss. Medd. Géteb. Bot. Tradg. 4 (1928) 16,
f. 12d, 18.—L. wallichiana Hoox. f. & Tu. J.
Proc. Linn. Soc. Bot. 2 (1858) 29; Kurz, J. As.
Soc. Beng. 46, ii (1877) 211.—L. erecta Hook. f.
& Tu. J. Proc. Linn. Soc. Bot. 2 (1858) 28, non
DE VRIESE (1845); CLARKE, Fl. Br. Ind. 3 (1881)
426.—Dortmannia leschenaultiana O. K. Rev.

124

FLORA MALESIANA

(ser. 1). voli

Fig. 8. Lobelia nicotianaefolia RoTH ex R. & S. Bandarawele, Ceylon, c. 2-244 m tall, 1956.

Gen. 1 (1891) 972.—Dortmannia erecta O. K. l.c.
972.—Dortmannia nicotianaefolia O. K. l.c. 973.—
Dortmannia rosea O. K. lI.c. 973.—Dortmannia
colorata O. K. l.c. 973.—Dortmannia pyramidalis
O. K. Lc. 380.—L. seguinii LEVEILLE & VAN, in
Fedde, Rep. 12 (1913) 186; Wimmer, Pfil. R. Heft
107 (1953) 648, incl. var.—L. fossarum WIMMER,
Akad. Anz. Wien n. 14 (1924) 3.—L. eryliae
FISCHER, Kew Bull. (1928) 141; Wimmer, Pfl. R.
Heft 107 (1953) 652.—L. leschenaultiana SKOTTSB.
Medd. Géoteb. Bot. Tradg. 4 (1928) 4, f. 3-7;
Wimmer, Pfi. R. Heft 107 (1953) 659.—L. doniana
Sxottss. Medd. Goteb. Bot. Tradg. 4 (1928) 19,
f. 12, 19-24, nomen illegit.—L. philippinensis
Sxorttss. /.c. 13, f. 12c, 15-17; STEEN. Bull. Jard.
Bot. Btzg III, 13 (1934) 178; Steup, Trop. Natuur
27 (1938) 142, f. 60; Toxopeus, /.c. 109, f. 21.—
L. epilobioides WIMMER, in Fedde, Rep. 38 (1935)

79: Ann. Naturh. Mus. Wien 56 (1948) 367, incl.
var. sarasinorum; Pfl. R. Heft 107 (1953) 652.—
L. leucanthera Kerr, Kew Bull. (1936) 34; CRAIB,
Fl. Siam. En. 2 (1936) 304; Wimmer, Pfl. R. Heft
107 (1953) 641.—L. palustris KERR, Kew Bull.
(1936) 35; Craip, Fl. Siam. En. 2 (1936) 304.—
L. camptodon Wimmer, Ann. Naturh. Mus. Wien
56 (1948) 366; Pfl. R. Heft 107 (1953) 637.—L.
beddomeana Wimmer, Pfi. R. Heft 107 (1953)
645.—Fig. 8-9.

A coarse herb, up to 2(-414) m. Stem terete at
the base, angular towards the apex, hollow, simple
or apically branched. Leaves spirally arranged,
densely set in juvenile plants, oblong to narrow-
lanceolate, the lower ones sometimes obovate-
oblong, gradually decurrent towards the petiole-
like base, 50-10 by 8-4 cm to 9-3 by 3-34 cm;
acuminate towards the apex (tip acute or blunt),

Dec. 1960]

Fig. 9. Lobelia nicotianaefolia ROTH ex R. & S.
SW. Celebes, 1937.

serrate-toothed along the margin, hairy on both
sides, especially on the nerves, rarely glabrous,
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
1-214 cm, obliquely patent, ascending, terete,
hairy; bracteoles 2, minute, 3-12 mm long (in
Indian material often caducous or absent).
Flowers \14-334 cm long, variable in colour, pale,
whitish, blue, dark purple, or rose. Sepa/s linear
to lanceolate, 4-12 by | mm, entire to distinctly
toothed, acute, glabrous or hairy. Corolla 3/4,-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,
all three ovate-acute, with a slightly crenate
margin. (In immature corollas the lobes are

CAMPANULACEAE (Moeliono & Tuyn)

Fig. 10.

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, 74-1 cm by 4-6 mm, glabrous
or hairy. Seeds flattened ellipsoid, 14—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, katlabung, kany-
uong, \g., balinyungyung, balyongyong, lufgog-
lufigog, subasob, Bon.

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
many local forms but in my opinion they should
not be named.

Kausik (J. Ind. Bot. Soc. 17, 1938, 161-167),
who has studied the gametogenesis and embryo-
logy of this species, stated that the chromosome

Distribution of Lobelia nicotianaefolia
RoTtH ex R. & S., localities in Malaysia indicated
by dots.

126

FLORA MALESIANA

[ser. I, vol. 6!

number of a plant collected at Koppa (Mysore)
is n= 14.

Fig. 11. Lobelia sumatrana MeErRR. Mt Losir, Gajo
Lands (N.Sumatra), 1937.

2. Lobelia sumatrana MerR. Not. Nat. Ac. Sc.
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, 15-40 cm, with a firm
rootstock. Stem angular, glabrous to sparsely pi-
lose. Rosette /eaves narrowly spathulate to obovate-
lanceolate, tapering towards the base, shallowly
crenate-dentate, blunt, 2-5 by 14-1 cm; cauline
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 114-21% cm long. Calyx
lobes oblong, c. 5 by 114-2 mm, blunt to broadly
triangular at the apex, shallowly crenate, glabrous
or pilose. Corolla 34-2 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 114-2 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 2-44 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 \y by 14 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, Dict. Bot. 3 (1791)
588; R. & S. Syst. Veg. 5 (1819) 60; DC. Prod.
7 (1839) 378; Sonp. Fl. Cap. 3 (1865) 539,
excl. syn. 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
(non LAMK) Cav. Ic. 6 (1801) 7, t. 511, f. 1.—L.
filiformis var. luzoniensis PERS. Syn. 2 (1807) 214;
R. & S. Syst. Veg. 5 (1819) 61; G. Don, Gard.
Dict. 3 (1834) 713; DC. Prod. 7 (1839) 368;
Mia. FI. Ind. Bat. 2 (1856) 577; F.—VILL. Nov.
App. 4 (1880) 121.—L. trigona Roxs. [Hort.
Beng. (1814) 85, nomen] Fl. Ind. 2 (1824) 111;
Fl. Ind. ed. Carey (1832) 506; G. Don, Gard.
Dict. 3 (1834) 709; DC. Prod: 7 (1839) 359;
Hook. f. Ic. Pl. 4 (1841) t. 358, excl ‘syne:
trialata et L. heyneana; Wicurt, Ic. 4 (1848) t.
1170; Mig. Fl. Ind. Bat. 2 (1857) 574; Hook. f.
& Tu. J. Proc. Linn. Soc. Bot. 2 (1858) 27, p.p.;

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

127

Mia. Sum. (1861) 234; Kurz, J. As. Soc. Beng.
46, ii (1877) 211; CLARKE, FI. Br. Ind. 3 (1881)
423; Trim. Fl. Ceyl. 3 (1895) 56; Koorpb. Exk.
Fl. Java 3 (1912) 302; YAmMAmoTo, Obs. FI.
Formosa 13 (1936) 148, excl. syn. L. trialata et
L. heyneana.—L. triangulata Roxs. Hort. Beng.
(1814) 16, nomen nudum.—L. stipularis RorTH, in
Remecasusyst. Veg. 5 (1819) 9673, Nov, Pl: Sp.
(1821) 144; Wa... Pl. As. Rar. 2 (1831) 43.—
Rapuntium alsinoides Prest, Prod. Mon. Lob.
(1836) 22.— L. sp. GrirF. Not. Pl. As. 4 (1854)
281.—L. griffithii Hook. f. & Tu. J. Proc. Linn.
Soc. Bot. 2 (1858) 28; Kurz, J. As. Soc. Beng.
46, ii (1877) 211, incl. var. genuina et var. dopa-
trioides; CLARKE, Fl. Br. Ind.3 (1881) 424; RIDL.
J. Str. Br. R. As. Soc. n. 33 (1911) 124; Crats,
Kew Bull. (1911) 404; Ripi. Fl. Mal. Pen. 2
(1923) 201; DanGuy, FI. Gén. I.-C. 3 (1930)
682, f. 75, 7-9; Cras, Fl. Siam. En. 2 (1936)
303; Wimmer, Pfl. R. Heft 107 (1953) 569.—L.
dopatrioides Kurz, J. As. Soc. Beng. 39, 11 (1870)
77; Flora 55 (1872) 302; Crais, Fl. Siam. En. 2
(1936) 303.—L. microcarpa CLARKE, FI. Br. Ind.
3 (1881) 424; Koorp. Exk. FI. Java 3 (1912) 302;
Dancuy, Fl. Gén. I.-C. 3 (1930) 681; MERR. &
Perry, J. Arn. Arb. 22 (1941) 385; WIMMER,
Pfi. R. Heft 107 (1953) 574.—L. terminalis CLARKE,
Fl. Br. Ind. 3 (1881) 424; Crarp, Kew Bull.
(1904) 404; Dancuy, FI. Gén. I.-C. 3 (1930)
680; Crais, Fl. Siam. En. 2 (1936) 306; WIMMER,
Pfl. R. Heft 107 (1953) 573.—Dortmannia grif-
fithii O.K. Rev. Gen. Pl. 2 (1891) 380.—Dort-
mannia trigona O.K. l1.c., incl. var. microcarpa et
terminalis.—Dortmannia alsinoides O.K. l.c. 972.—
L. luzoniensis (PERsS.) MERR. En. Philip. 3 (1923)
588; Wimmer, Pfl. R. Heft 107 (1953) 543.—L.
chevalieri DANGuy, Bull. Mus. Paris (1929) 263;
Fl. Gén. I.-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, /.c. 2, t. 71 f. 5; DANGUy, FI. Gén. I.-C.
3 (1930) 681; Cratp, Fl. Siam. En. 2 (1936) 304,
incl. var.; WIMMER, Pfl. R. Heft 107 (1953) 574.—
L. hainanensis WIMMER, Ann. Naturh. Mus. Wien
56 (1948) 348; Pfl. R. Heft 107 (1953) 506.—L.
chinensis (non Lour.) HANcE, J. Linn. Soc. Bot.
13 (1873) 110; Forspes & HEMSLEY, J. Linn. Soc.
Bot. 26 (1889) 3; DANGuy, FI. Gén. I.-C. 3 (1930)
680, f. 75, 10-12.—L. radicans (non THUNB.)
Hosseus, Beih. Bot. Centralbl. 28, 1i (1911) 446.—
Fig. 16c.

Erect to ascending, unbranched to strongly
branched herb, 3—35(—40) cm; stem 3-angled and
winged. Leaves 4-114 by 14-2 cm, sessile or short-
petioled, contracted or narrowed to the base,
acute or blunt, entire or toothed, glabrous to
sparsely hairy; basal leaves ovate-oblong,
cordate or elliptic, upwards sometimes sublan-
ceolate, up to | by 0.3 cm. Flowers 8-12 mm,
axillary, often in the higher axils, and then
forming a lax terminal raceme. Pedicels 1-314 cm,
3-angled, glabrous or slightly patent-hairy.
Bracteoles basal, minute to 2 mm long and linear,
often caducous. Calyx lobes triangular, subulate,
244-314 by 14-1 mm, entire, sometimes ciliate,

glabrous or sparsely hairy. Corolla 4-12 mm,
varying from bright blue to violet (sometimes, in
Celebes, white), inside hairy (except for the
specimens described as L. hosseusii which are
glabrous, its var. villosa excluded), with 2 (white)
gibbosities, dorsally split to the base, dorsal
lobes connate with the lateral ones for 3-414 mm,
lateral lobes connate with the ventral one for
314-6 mm; dorsal lobes 1-3 mm, falcate-oblong
or falcate-ovate, acute or acuminate, margin entire
or subentire, lateral and ventral lobes oblong or
ovate, 1-214 mm, acute or acuminate, entire or
subentire. Filaments 3-5 mm, free to half-way up
or higher, connate, the two anterior ones + twice
as broad as the anthers, the outside patent hairy;
dorsal anthers 114, ventral ones 1 mm, each
anther at the apex with a hair tuft, otherwise
glabrous or hairy. Ovary 2-214 by 114-214 mm,
oblong, trumpet-shaped to cupular, glabrous or
sparsely hairy; style glabrous. Capsule + hemi-
spherical, 2-3 by 1144-3 mm. Seeds 14 by 1/3
mm, trigonous, brown.

Distr. SE. Asia (Ceylon and the Deccan to
S. China: Kwangtung), Hainan, and Formosa, in
Malaysia: Malaya (Perlis: Chupeng; Singapore,
Wimmer, /.c.), W. Java (Indramaju), Celebes

Fig. 12. Distribution of Lobelia alsinoides LAMK
with localities in Malaysia dotted.

(SW. & SE. peninsula), Philippines (Mindanao,
Davao), S. New Guinea (N of Merauke, Wuroi,
Oriomo R., Lake Daviumbu, Borovia). Fig. 12.

Ecol. In West Java and in SW. Celebes during
the wet season in marshy grassland often do-
minated by ‘siil’ (Sorghum nitidum) and/or
Cyperaceae; in S. Celebes also along margins
of dry rice-fields. In New Guinea this species is
found on swampy grounds, together with Erio-
caulon, Utricularia, and several Cyperaceae; on
wet places in savannahs with Melaleuca, Acacia,
Eucalyptus stands and_ scattered Antidesma
ghaesembilla trees. From sea-level up to 1000 m,
restricted to regions subject to a dry monsoon.
Fl. fr. Nov.—Aug.

Note. A polymorphic species with intergrading
forms, so we can find a range of subsequent
‘species’, beginning from the typical L. griffithii,
with its scale-like leaves to the normal broad-

128

FLORA MALESIANA

[ser: I, volo

ovate leaves of L. dopatrioides and the petiolate
leaves of L. terminalis. I cannot trace any geo-
graphical or ecological replacement of these
forms. The shape and size of the bracteoles have
also lead to superfluous names: the Indian speci-
mens have relatively long bracteoles, which in-
duced RoTH to name this form L. stipularis.

L. luzoniensis (PERS.) MERR. was based upon a
specimen in herb. NEE from Luzon near Santa
Cruz de la Lagufa. MERRILL mentioned two
recent collections from Luzon (Laguna) and
Mindanao (Davao), citing COPELAND 368 and
WEBER 1472 respectively. WIMMER quoted the last
mentioned number under L. alsinoides, Pfl. R.
Heft 107 (1953) 573. I have seen COPELAND 368,
which was distributed as a Wahlenbergia (NY);
it has all anthers bearded and trigonous seeds and
undoubtedly represents L. alsinoides. This con-
firms my opinion, made from the plate and
description, that L. /uzoniensis is conspecific with
L. alsinoides. CAVANILLES’s figure of the plant is
twice enlarged; he mentions the short seta on
top of the corolla lobes and hairy anthers.

4. Lobelia zeylanica LINNE, Sp. Pl. 2 (1753) 932;
OsBEcK, Dagbok Ostindisk Resa (1757) 241;
Bur. f. Fl. Ind. (1768) 186; R. & S. Syst. Veg. 5
(1819) 64; Roxs. Fl. Ind. 2 (1824) 113; WALL.
Pl. As. Rar. 2 (1831) 43; G. Don, Gard. Dict.
3 (1834) 709; Kurz, Flora 55 (1872) 302; J. As.
Soc. Beng. 46, ii (1877) 211, incl. var. affinis;
YAMAMOTO, Obs. FI. Form. 13 (1936) 149; MERR.
& Perry, J. Arn. Arb. 22 (1941) 386; GREEN-
woop, ibid. 30 (1949) 78.—L. succulenta BL.
Bijdr. (1826) 728; DC. Prod. 7 (1839) 373; Mia.
Fl. Ind. Bat. 2 (1856) 577; Koorpb. Exk. Fl. Java
3 (1912) 303; Ocuse, Trop. Groent. (1925) 22,
fig.; OcHsE & BAKH. Ind. Groent. (1931) 92,
f. 54; Hocur. Candollea 5 (1932) 292; MEeErR.
Pap. Mich. Ac. Sc. 20 (1935) 111; Crats, FI.
Siam. En. 2 (1936) 305; Wimmer, Ann. Naturh.
Mus. Wien 56 (1948) 361, incl. var.; BACK.
Bekn. FI. Java (em. ed.) 8 (1949) fam. 184, p. 3;
Wimmer, Pfl. R. Heft 107 (1953) 576, incl. var.
lobbiana et f. glabra.—L. affinis WALL. [Cat.
(1828) 35 et 158 no 1131] ex G. Don, Gard. Dict.
3 (1834) 709; non MirsBacu, 1805; DC. Prod. 7
(1839) 360; Mig. FI. Ind. Bat. 2 (1856) 574;
Hook. f. & Tu. J. Proc. Linn. Soc. Bot. 2 (1858)
27; Drury, Handb. Ind. FI. 2 (1866) 108; CLARKE,
Fl. Br. Ind. 3 (1881) 424, incl. var. lobbiana;
Trim. Fl. Ceyl. 3 (1895) 57; GAMBLE, J. As. Soc.
Beng. 74, ii (1905) 52; Cratp, Kew Bull. (1911)
404; Koorp. Exk. Fl. Java 3 (1912) 121; S. Moore,
Trans. Linn. Soc. Bot. II, 9 (1916) 88; DiIELs,
Bot. Jahrb. 55 (1917) 121; Ripv. Fl. Mal. Pen. 2
(1923) 200, f. 89; DaNncuy, FI. Gén. I.-C. 3
(1930) 679; Merr. Lingn. Sc. J. 6 (1930) 332.—
Rapuntium affine Pres_, Prod. Mon. Lob. (1836)
13.—Rapuntium succulentum PrReEsL, I.c. 13.—
Rapuntium zeylanicum Presv. I.c. 13.—L. sub-
cuneata Mig. Fl. Ind. Bat. 2 (1857) 574.—L.
lobbiana Hook. f. & Tu. J. Proc. Linn. Soc.
Bot. 2 (1858) 28; Drury, Handb. Ind. Fl. 2
(1866) 110.—Dortmannia succulenta O.K. Rev.

Gen. Pl. 1 (1891) 973.—Dortmannia zeylanica
O.K. /.c. 380, pro nomen, excl. sched.—Dortman-
nia subcuneata O.K. lL.c. 973.—Dortmannia
trigona O.K., pro var. affinis O.K. l.c. 380.—L.
barbata Wars. Bot. Jahrb. 13 (1891) 444; K.
Scu. & Laut. Fl. Schutzgeb. (1900) 593.—Pratia
torricellensis K. ScH. & Laut. Nachtr. (1905)
402.—Pratia ovata ELMER, Leafl. Philip. Bot. 2
(1909) 593; Merr. Philip. J. Sc. 11 (1916) Bot. 317;
En. Philip. 3 (1923) 589.—Pratia begonifolia
(non LINDL.) Hosseus, Beih. Bot. Centralbl. 28,
ii (1911) 477.

Creeping to ascending herb, 20-90 cm, stems
often rooting at the lower nodes, terete, higher up
sometimes angular, glabrous to sparsely hairy.
Leaves spirally arranged, ovate to broad ovate,
(34-)1-6 by (14-)1-344 cm; base cordate to
truncate or sometimes decurrent, apex acute,
sometimes blunt, margin subentire to subdentate
to repand-dentate; upper surface puberulous or
glabrous, underneath puberulous, especially the
nerves, or glabrous; petiole terete, 1-20 mm,
sometimes puberulous to appressed-hairy. Flowers
axillary, 7-12 mm. Pedicels terete, 1—2 cm,
ebracteolate. Calyx lobes lanceolate to oblong
triangular, acute, patent-hairy, sometimes gla-
brous, 3—5 by '4 mm, entire to subdentate, with
curved hairs to dentate-ciliate. Corolla 5—9 mm,
purplish to pale or creamy, inside glabrous to
subglabrous with two gibbosities, dorsally slit
to the base; dorsal lobes connate with the lateral
ones for 2-314 mm, lateral lobes connate with the
ventral one for 4-7 mm: dorsal lobes oblong or
elliptic, 2144-3 by 1-114 mm, falcate, entire to
wavy, acute, outside on the nerves with long
hairs, lateral and ventral lobes spathulate to ovate,
1-2 by 1-2 mm, entire, at the centre of the ventral
lobe a white spot surrounding a purple stripe.
Filaments 3-5 mm, for more than 34 free; free
parts narrowed towards the base, all of equal
width, haired; anthers from dorsal to ventral
1-114 mm to 34-1 mm, hairy; all anthers at the
apex with a hair tuft. Ovary cupular to obconical,
114-3 by 1 mm, scattered hairy to hairy (glabrous
in specimens from Hainan); style glabrous.
Capsule obconical to oblong ovate, 3-7 by 2-4
mm, with distinct nerves and a membranous
pericarp. Seeds trigonous, 0.4 by 0.3 mm, light-
brown; after withering the nerves and remains
of the placenta persistent.

Distr. SE. Asia (from Ceylon and the Deccan
through the Himalaya to SE. China: Kwangtung,
Kwangsi, Chekiang), Hainan, and Formosa, al-
most throughout Malaysia, but not yet found in
the Lesser Sunda Islands and the Moluccas; also
in the Fiji Is.

Ecol. On shaded grassy grounds and moist
places, under everwet climatic conditions. In
coffee, tea, and rubber plantations, in open places
in primary forest along streams, in shaded ravines,
etc., from the lowland up to 1500, rarely 1650-
2000 m. Fi. fr. Jan.—Dec.

Vern. Bélimbing tanah, M, rantji djadjar, J,
djukut bulu mata kérbo, ramo kujah, S.

Use. According to OcHSE the young leaves are

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

_129

eaten as ‘lalab’ (steamed vegetable) with rice.

Notes. There is in literature no unanimous
opinion about the typification of this Linnean
species and this even induced CrRaiB to suggest
to reject it as a nomen confusum (Fl. Siam. En. 2,
1936, 305-306). De CANDOLLE accepted it as
conspecific with L. affinis WALL. which is in turn
conspecific with L. succulenta BL.; he excluded
from it LINNAEUS’s reference to SEBA which he
found fit to describe as L. sebae DC., now ac-
cepted as a synonym of Monopsis simplex (L.)
Wimmer. In this interpretation he was followed
later by several others, for instance KURZ, and
recently MERRILL & PERRY.

The other interpretation started with CLARKE,
who, for his revision of the Indian Lobelias,
examined the material of L. zey/anica in the Lin-
nean Herbarium, and stated (FI. Br. Ind. 3, 1881,
425):—‘that LINNAEUus’s excellent specimen of the
species is named L. zeylanica in his own hand, but
the name has been altered (erroneously) by Sir
J. E. SmituH to L. anceps, an Australian species.”

CRAIB, /.c., re-examined the Linnean specimens,
two sheets: —‘‘on one sheet marked L. zeylanica
are 2 specimens but different plants, viz one what
we call now “‘L. succulenta’ BL. and a second
non-campanulaceous plant.” .. .“‘Pinned to this
“LL. zeylanica’’ sheet is another, on which is
written, in LINNAEUs’s hand, “*18”’, i.e. the number
of L. zeylanica in Sp. Pl. The plant on this sheet
belongs to L. zeylanica as usually understood
to-day.”

This latter is obviously the specimen which
CLARKE designated as the lectotype. It is referred
to by SAVAGE (Cat. Linn. Herb. 1945, 165) under
1015 as no 42. According to SAVAGE it is not
indicated on any of the Linnean sheets which
came from China and was collected by OsBEck.

MERRILL & PeERRY rightly considered that,
though LINNAEUs derived his specific name from
Campanula ceylanica, senecionis folio, flore
PULDULCORSEBA. “Lhes) I: 37, t 22, £ 12. 1734;
yet, at the same time, he added a question mark
to this reference; his description was based wholly
on a plant collected by Ospeck, near Canton,
in China, which represents a species totally dif-
ferent from the form Sepa illustrated (J. Arn.
Arb. 22, 1941, 386). They concluded that the
OSBECK specimen(s) are unquestionably the type
of L. zeylanica.

In the rather detailed description OsBECK gave
the following characters :—‘‘Perianthium...subtus
villosum ... filamenta... duo basi villosa... Cau-
lis teres... Folia cordata...petiolata.’ This
definition leaves no doubt that Osspeck’s plant
must agree with L. succulenta BL.

This is corroborated by an OsBECK collection
in the Bergius Herbarium, which we could examine
through the courtesy of Prof. FLorIN, Stockholm.
On the back is written:—‘‘ceylanica. Lobelia
foliis ovatis obtusis petiolatis crenatis caule dif-
fuse: In China agris aquosii oryzae legi 1751
OsBECK.”” The specimen doubtless represents L.
succulenta BL.

Through the diligence of Dr NorLInpH, Lund,

two other OsBECK specimens were located in the
Riksmuseet Stockholm, both marked ‘‘zeilanica”’
and “/8 zeylanica’”’ respectively. The latter speci-
men has possibly actually been in the hands of
LINNAEUS.

5. Lobelia heyniana R. & S. Syst. Veg. 5 (1819)
50; non SPRENG. 1825; BL. Bijdr. (1826) 728;
G. Don, Gard. Dict. 3 (1834) 709 (heyneana);
Wimme_r, Ann. Naturh. Mus. Wien 56 (1948) 344;
Pfl. R. Heft 107 (1953) 474, incl. var. div.; SAN-
TAPAU, Rec. Bot. Surv. Ind. 16 (1953) 158.—L.
decurrens RoTH, Nov. Sp. (1821) 145; non Cay.
1801.—L. micrantha Hook. Exot. Fl. 1 (1823)
t. 44; non H.B.K. 1818.—L. trialata HAM. ex
D. Don, Prod. Fl. Nep. (1825) 157; G. Don,
Gard. Dict. 3 (1834) 709; DC. Prod. 7 (1839)
360; CLARKE, FI. Br. Ind. 3 (1881) 425, incl. var.
lamiifolia; CRAtB, Kew Bull. (1911) 404; Koorp.
Exk. Fl. Java 3 (1912) 302; BoLp. Zakfl. (1916)
41; GaAmBLeE, FI. Pres. Madras 4 (1921) 736;
Craip, Fl. Siam. En. 2 (1936) 306.—L. subincisa
WALL. [Cat. (1828) n. 1320, nomen] ex DC.
Prod. 7 (1839) 367; Mia. Fl. Ind. Bat. 2 (1856)

575.—Rapuntium reinwardtianum PReEsL, Prod.
Mon. Lob. (1836) 14.—Rapuntium trialatum
Pres_, /.c. 13.—Rapuntium arenarioides PRESL,

l.c. 17.—L. arenarioides DC. Prod. 7 (1839) 367.—
L. reinwardtiana DC. I.c. 367; Mia. FI. Ind. Bat.
2 (1856) 565; Koorp. Exk. FI. Java 3 (1912) 303.—
L. inconspicua Ricu. Tent. Fl. Abyss. 2 (1851) 8.—
L. zeylanica (non L.) Moon, Cat. (1824) 14, nomen;
CLARKE, Fl. Br. Ind. 3 (1881) 425, incl. var.
walkeri; TRIM. Fl. Ceyl. 3 (1895) 56; Koorpb.
Exk. Fl. Java 3 (1912) 302; Bop. Zakfl. (1916)
41; Hanes, Bot. Bihar & Orissa 4 (1922) 501,
incl. var. aligera; BACK. & S toot. Theeonkr.
(1924) 213, fig.; DANGuy, FI. Gén. I.-C. 3 (1930)
678, incl. var. parviflora; HocHR. Candollea 5
(1932) 292; Back. Bekn. Fl. Java (em. ed.) 8
(1949) fam. 184, p. 2.—L. subracemosa Mia. FI.
Ind. Bat. 2 (1857) 576, incl. var. rigidior.—L.
dichotoma Mig. I.c. 576; WIMMER, in Fedde,
Rep. 38 (1935) 78; Ann. Naturh. Mus. Wien 56
(1948) 345, incl. var. aligera et var. pilosella;
Pfl. R. Heft 107 (1953) 476, incl. var.—L. trigona
(non Roxs.) Hook. f. & Tu. J. Proc. Linn. Soc.
Bot. 2 (1858) 27, p.p.; THwattes, En. Pl. Zeyl.
(1860) 169, p.p.; BENTH. Fl. Hongk. (1860) 197,
p.p.; Hosseus, Beih. Bot. Centralbl. 28, ii (1911)
466, p.p.—Dortmannia zeylanica O.K. Rev. Gen.
Pl. 2 (1891) 380.—Dortmannia trialata O.K. l.c.
973.—Dortmannia inconspicua O.K. I.c. 972.—
Dortmannia_ reinwardtiana O.K. l.c. 973.—L.
aligera HAINES, J. As. Soc. Beng. n.s. 15 (1920)
316.—L. bialata Merr. Philip. J. Sc. 7 (1912)
Bot. 105, ex descr.; Wimmer, Pfl. R. Heft 107
(1953) 474. Fig. 16d.

An ascending or erect herb, 5—50(—60) cm; stem
3-angled and winged. Leaves spirally arranged,
the lower ones rhomboid to broad-ovate, upwards
elliptic to linear-lanceolate, decurrent at the base,
acute at the apex, 14-4 by 14-3 cm, serrate to
serrate-dentate, glabrous or sparsely hairy,
especially the higher ones; petiole up to 3 mm.

FLORA MALESIANA

[ser. I -voliaet

Fig. 13. Distribution of Lobelia heyniana R. & S.

Flowers 414-12 mm, axillary, often forming a
terminal raceme. Pedicels 14-2 cm, 3-angled,
patent, glabrous or hairy with 2(-1), small, linear
to minutely reduced bracteoles. Calyx lobes
lanceolate-elliptic, 2-4 by 14-1 mm, acute, entire
to subdentate, glabrous to sparsely hairy. Corolla
314-10 mm, pale-purple, lilac, sometimes white,
inside with 2 gibbosities and a dark spot, nearly
glabrous to hairy, slit to the base; dorsal lobes
connate with the lateral ones for 1144-5 mm,
lateral lobes connate with the ventral one for 2—7
mm; dorsal lobes linear to lanceolate, 1-4 by
14-1 mm, often reflexed, sometimes longitudinally
folded, small compared with the others, lateral
and ventral lobes spathulate to ovate, 1-7 by
14-214 mm, 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 1-114 to 14-34 mm, glabrous or hairy,
only the 2 anterior ones with an apical hair tuft.
Ovary trumpet- to cup-shaped, 114-4 by 1-214
mm, glabrous, sometimes hairy; style glabrous.
Capsule obconical to obconical-campanulate,
3-8 by 2-214 mm. Seeds semi-ellipsoid, 4 by
14 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. bialata
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 kébo, kitombe, S, krémo,
krinjingan, kukunnang, 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,
ed. Willd. (1793) 628; R. & S. Syst. Veg. 5 (1819)
41; G. Don, Gard. Dict. 3 (1834) 709; DC.

Prod. 7 (1839) 360; ForBes & HEMSLEY, J. Linn.
Soc. Bot. 26 (1889) 2; Diets, Bot. Jahrb. 21
(1901) 607; Merr. Lingn. Sc. J. 5 (1927) 181;
Crais, Fl. Siam. En. 2 (1936) 302; YAMAMOTO,
Obs. Fl. Form. 13 (1936) 148; Wimmer, Pfi. R.
Heft 107 (1953) 609.—L. erinus (non L.) THUNB.
Fl. Jap. (1784) 325.—L. erinoides (non L.) THUNB.
lic. 325.—L. radicans THUNB. Trans. Linn. Soc.
2 (1794) 330; R. & S. Syst. Veg. 5 (1819) 60;
Roxs. FI. Ind. 2 (1824) 110; CLARKE, FI. Br. Ind.
3 (1881) 425; Forspes & Hems Ley, J. Linn. Soc.
Bot. 26 (1889) 3; Koorp. Exk. Fl. Java 3 (1912)
302; BoLp. Zakfl. (1916) 41; Koorp. FI. Tjib. 3
(1918) 55; Back. & St toor. 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.
MAKINO, Ill. Fl. Japan (1954) 80.—L. campanu-
loides THUNB. Trans. Linn. Soc. 2 (1794) 331;
Ker, Bot. Reg. 9 (1823) t. 733; G. Don, Gard.
Dict. 3 (1834) 709.—L. caespitosa BL. Bijdr.
(1826) 729; DC. Prod. 7 (1839) 366; Mia. Fl. Ind.
Bat. 2 (1856) 575.—Rapuntium caespitosum
PreEsL, Prod. Mon. Lob. (1836) 13.—Rapuntium
campanuloides PRESL, l.c. 13.—Rapuntium chinense
PresL, /.c. 13.—Rapuntium radicans PRESL, l.c.
14.—Isolobus radicans DC. Prod. 7 (1839) 353.—
Tsolobus kerii DC. 1.c. 353.—Isolobus roxburghianus
DC. L.c. 353; Hook. f. & Tu. J. Proc. Linn. Soc.
Bot. 2 (1858) 27.—Isolobus campanuloides DC.
Prod. 7 (1839) 353.—Isolobus caespitosus HASSK.

Fig. 14. Lobelia chinensis Lour. Bogor (West
Java), in dense sods in the Tji Liwung valley
(1928).

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

131

Bonpl. 7 (1859) 180.—Pratia radicans G. Don,
Gard. Dict. 3 (1834) 700.—Pratia thunbergii G.
Don, /.c. 700.—Dortmannia chinensis O.K. Rev.
Gen. Pl. 2 (1891) 380.—Dortmannia campanuloides
O.K. /.c. 380.—Dortmannia radicans O.K. |.c.—
Fig. 14.

A glabrous, 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 shallowly
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, 114-3 by
14-4, 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 4-414 mm, lateral lobes
connate with the ventral one for 414-6 mm;
dorsal lobes 414-8 mm, linear, sometimes, reflexed,
blunt, lateral and ventral lobes 3-6 mm, equal,
linear to lanceolate, acute. Filaments 5-6 mm,
for more than half-way free, all of equal width,
the 2 anterior ones patent hairy, with a seta or
hair tuft at the top; anthers from dorsal to ventral
2-1 mm, glabrous or hairy. Ovary 2-5 by 1 mm,
trumpet-shaped, glabrous; style hairy at the base.
Capsule on a recurved pedicel, 4-6 mm, obconical,
glabrous. Seeds 3/8—14 mm, ellipsoid, somewhat
compressed, dark-brown, smooth.

Distr. SE. to E. Asia (from the Deccan to
China and Japan), in Malaysia: Malay Peninsula
(Singapore) and Java (eastwards to Mt Diéng),
according to BACKER & VAN SLOOTEN since long
introduced from continental Asia.

Ecol. On moist, grassy places, along water-
courses and on cultivated land, on rice-fields
and in tea and cinchona plantations, 500-1600 m,
occasionally at lower altitudes. According to
BACKER & VAN SLOOTEN no capsules with ripe
seeds are found in Java. VAN STEENIS has found
a plant at Bogor (n. 2151) with ripe capsules and
seeds. I also saw a sheet in the Leyden Herb.
(SCHIFFNER 2728, from Tjibodas) with ripe capsules
and seeds. It remains questionable whether these
seeds are viable.

Propagation is normally vegetative. According
to BACKER & VAN SLOOTEN it descends along
water-courses to lower altitudes, even to the
lowlands, by stolons or stems dispersed by water.
However, such lowland habitats are only tem-
porary.

Vern. Djukut mata keujeup, ki tombe, S.

7. Lobelia archboldiana (MerRR. & PERRY) MOE-
LIONO, nov. comb.—Pratia archboldiana MEeERR.
& Perry, J. Arn. Arb. 30 (1949) 59; Wimmer,
Pfl. R. Heft 107 (1953) 765.—Fig. 15.
Stemless, delicate plant, sometimes with stolons,
2-4 cm. Leaves in rosette, ovate to roundish,
14-114 cm diam., subcordate or truncate at the
base, wavy to denticulate along the margin,

aavay

NS}

Fig. 15. Lobelia archboldiana (MERR. & PERRY)

MOELIONO. a. Habit,
aS iG: Open tlowers

x 2144, b. young flower,
x 3 (CLEMENS 12442).

obtuse at the apex, very thin, glabrous or sparse-
ly hairy above, glabrous underneath; petiole
3—7(—23) mm, glabrous or sparsely pilose. Flowers
5-614 mm, axillary or terminal. Pedicels 2-12 mm,
glabrous or hairy. Calyx lobes linear-lanceolate,
with 1-2 pairs of teeth, hairy, 2-3 mm. Corolla
red to dark wine-red, the tube 2-3 mm long,
outside glabrous to sparsely hairy, inside glabrous,
dorsally split to 1 mm from the base; dorsal lobes
half-way connate with the lateral ones, their free
part lanceolate, 2-3 mm, reflexed, lateral and
ventral lobes also half-way connate, their free
part lanceolate, 3—3.2 mm, acute. Stamens c.
2 mm, filaments for 934 of their length free, gla-
brous; anthers from dorsal to ventral 134-34 mm;
2 anterior ones with a short bristle and some hairs.
Ovary campanulate to trumpet-shaped, 1-214 by
1-2 mm, sparsely hairy; style glabrous. Capsule
roundish to ovoid, 3 by 3 mm, sparsely hairy,
with a thin wall. Seeds roundish to ovoid, c. 14
by 14 mm, smooth, dark-brown to black.

Distr. Malaysia: E. New Guinea (Murray
Pass, Wharton Range; Rawlinson Range, Morobe
Prov.), rare, twice collected.

Ecol. Under a rock wall on grassy bank of a
creek and in deep holes above deep water-courses,
2840-3600 m. Fl. fr. July—Aug.

8. Lobelia brachyantha Merr. & Perry, J. Arn.
Arb. 22 (1941) 385; Wimmer, Pfl. R. Heft 107
(1953) 487.—Fig. 16a-b.

Tiny, creeping, branched herb; stem terete,
rooting, sparsely pilose. Leaves alternate, orbicular-
reniform, 2!4—5 by 214-6 mm, base cordate to
truncate, margin subundulate to dentate, upper
surface sparsely pilose, beneath glabrous; petiole

132

FLORA MALESIANA

fser. I> vols?

aN

Fig. 16. Lobelia brachyantha MERR. & PERRY.

a. Habit, x 214, b. flower, x 5.—L. alsinoides

LAMK. c. Stamens, xX 5.—L. heyniana R. & S.

d. Stamens, * 5 (a—b Brass 11570, c VAN STEENIS
6705, d DE Voocp 2615).

7 BE)

1-3 mm. Flowers c. 5 mm, axillary. Pedicels
terete, 4-5 mm, hairy, ebracteolate. Calyx lobes
oblong-triangular, 2-244 by 4-34 mm, with a
distinct tooth at the base, sparsely pilose. Corolla
subcampanulate, 4 mm long, 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 2mm;
all lobes ovate-deltoid, 114 mm, acute to blunt.
Filaments 114 mm, up to 1/3 free, equal, glabrous
or hairy. Anthers from dorsal to ventral 1-34 mm,
glandular hairy; the 2 anteriors at the apex finely
hairy (according to MERRILL & PERRy ‘setigeris’).
Ovary cupular, 1 by 144 mm, long-hairy; style
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) 386.—Pratia
montana Hassk. Flora 25, 2 (1842) Beibl. 1, p. 23;
Cat. Hort. Bog. (1844) 106; CLARKE, FI. Br. Ind. 3

(1881) 423; Koorp. Exk. Fl. Java 3 (1912) 304;
Fl. Tjib. 3, 2 (1918) 56; DANGuy, FI. Gén. I.-C.
3 (1930) 677, f. 75, 1-5; Hocur. Candollea 5
(1932) 293, incl. f. variegata; HEND. Gard. Bull.
S.S. 7 (1933) 109; DOCTERS VAN LEEUWEN, Verh.
Kon. Ak. Wet. A’dam sect. 2, 31 (1933) 240;
STEEN. Bull. Jard. Bot. Btzg III, 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
Mia. FI. Ind. Bat. 2 (1857) 573; BulJSMAN, Flora
106 (1913) 127, excl. syn.—Piddingtonia patens
Mia. Fl. Ind. Bat. 2 (1857) 573.—Speirema
montanum Hook. f. & TH. J. Proc. Linn. Soc.
Bot. 2 (1858) 27; CLARKE, J. Linn. Soc. Bot. 15
(1876) 147.—Piddingtonia cyanocarpa HASssK.
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, 314-12 by 114-414 cm, decurrent at the base,
dentate to subdentate, acute to acuminate; petiole
c. %4 cm, glabrous to appressed-hairy. Flowers
axillary, solitary, secund, scentless. Pedicel 2414
cm, during anthesis erect. Calyx lobes lanceolate,
5-11 by 1-114 mm, obliquely patent, entire, acute.
Corolla 14-3 cm, purplish to lilac-blue with

Fig. 17. Lobelia montana REINW. ex BL. a. Apex
of flowering stem, x 74, b. ripe fruit, natural
poise, x %/.

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

=e

white-shaded margins, outside glabrous, inside
hairy and with two pilose gibbosities, dorsally
slit to 1-2 mm from the base; dorsal lobes connate
for 1-3 mm, connate with the lateral lobes for
7-10 mm, lateral lobes connate with the ventral
one for 9-12 mm; dorsal lobes linear-lanceolate,
9-10 mm, flat or crisped, margins glabrous, lateral
and ventral lobes oblong-lanceolate, the ventral
one broadest, 7-12 mm, crisped and ciliate,
acuminate. Filaments 4-7 mm, for 1/3 of their
length free, glabrous; anthers from dorsal to
ventral 5—3 mm long, hairy; the two anteriors
besides at their apex with a hair tuft. Ovary
campanulate to hemispherical, 3-5 by 3-614 mm,
glabrous; style glabrous. Berry globose, 34-114
cm diam. (living c. 2-214 cm), on long (over 6 cm),
patent, afterwards recurved pedicels, violet, later
dark purple. Seeds broad-ellipsoid to ovoid, c.
14 mm long, brown, smooth.

Distr. India (from the Deccan to the Himalaya)
to Indo-China (Tonkin), S. China (Yunnan), and
Malaysia: Malay Peninsula (Cameroon High-
lands), Sumatra, and Java (eastwards to Mt
Diéng.) Fig. 18.

+

Fig. 18. Distribution of Lobelia montana REINW.

ex BL. (continuous line, localities in Malaysia

dotted), and Lobelia borneensis (HEMSL.) MOE-

LIONO (dotted line, localities in Malaysia indicated
by crosses).

Ecol. Open or half-shaded places in mountain
forest, light spots in mossy forests, 1400-2600 m.
According to DocTeRs VAN LEEUWEN (1933) the
species is autogamous in Java.

Vern. Ki béwo, ki leuntja, tétéra, S, kémalon,
tjélengan, J.

Note. Kurz recorded to have found this
species on the summit of Mt Menumbing in
Banka I. (Nat. Tijd. Ned. Ind. 27, 1864, 206);
though I have not seen his collection this record
must rest On an error.

10. Lobelia borneensis (HEMSL.) MOELIONO, nov.
comb.—Pratia borneensis HrEmst. in Hook. Ic.
Pl. 16 (1886) t. 1532; Srapr, Trans. Linn. Soc.
Bot. IT, 4 (1894) 188, incl. var. grandiflora; GiBBs,

J. Linn. Soc. Bot. 42 (1914) 100; Merr. En. Born.
(1921) 585; Wimmer, Pfi. R. Heft 106 (1943)
117; ibid. Heft 107 (1953) 765; Lam, Blumea 5
(1945) 568.

Coarse, erect half-shrub, c. 1-114 m, branching,
sometimes reclining; stem glabrous, slightly to
strongly angular and furrowed. Leaves spirally
arranged, elliptic-lanceolate, 9-11 by 234-314 cm,
slightly dentate, decurrent at the base, acute to
acuminate at the top, glabrous to puberulous,
darker green at the upper side, paler beneath,
glossy on both surfaces; petiole 3-7 mm. Peduncle
angular and furrowed, inflorescence a terminal
raceme or pseudo-axillary by sympodial growth,
erect, faintly puberulous. Bracts lanceolate, 6—7
by 1-2 mm, slightly dentate. Pedicels 5-10 mm,
thin, whether or not with | or 2 tiny lanceolate
bracteoles in the basal half. Flowers c. 244 cm,
Calyx lobes lanceolate, 614-7 by 14 mm, longitudi-
nally nerved, subentire, obtuse to acute, faintly
puberulous. Corolla 134-2% cm, blue purple or
white with purple at the base of the margin of
the segments or lavender tinged, inside with 2
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!4 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 mm (ma-
ture?), globose, green. Seeds 34-1 mm, globose to
broad ellipsoid, slightly angular, reticulate.

Distr. Malaysia: Borneo (N. Borneo: Mt
Kinabalu; E. Kutei), Celebes (Central part and
SW. peninsula). Fig. 18.

Ecol. Insecondary jungle, on forest edges, along
forest paths, and in open grassfields, (SCO—)900-
1800 m. Fi. fr. Jan.—Dec.

Vern. Akar maga pawang, N. Bornew.

Note. The nearest allies of this and the foregoing
species are obviously found in remote places,
China, New Zealand, and Guatemala!

11. Lobelia angulata Forsr. Fl. Ins. Austr. Prod.
(1786) 58, n. 309; R. & S. Syst. Veg. 5 (1819) 65;
A. Ricu. Fl. Nouv.-Zél. (1832) 227; G. Don,
Gard. Dict. 3 (1834) 713; DC. Prod. 7 (1839)
366.—L. nummularia LAMK, Dict. Bot. 3 (1789)
589; R. & S. Syst. Veg. 5 (1819) 64; BL. Bijdr.
(1826) 727; G. Don, Gard. Dict. 3 (1834) 709.—
L. begonifolia WALL. Asiat. Res. 13 (1820) 377;
in Roxb. Fl. Ind. 2 (1824) 115; D. Don, Prod.
Fl. Nep. (1825) 158; WALL. Pl. As. Rar. 2 (1831)
43.—L. javanica THUNB. FI. Jav. (1825) 9, ef.
Blumea 6 (1950) 360.—L. obliqua HAm. ex D.
Don, Prod. Fl. Nep. (1825) 158.—L. pratiana
GaupicH. Ann. Sc. Nat. Bot. 5 (1825) 103.—

134

FLORA MALESIANA

[ser. I, vol. 61

Pratia repens GaAupicu. /.c. 103; in Freye. Voy.
Bot. (1826) 456, t. 79; G. Don, Gard. Dict. 3
(1834) 340, incl. var.; Hoox. f. Fl. Antarct. 1
(1844) 42; ReicHe, Fl. Chil. 5 (1910) 63.—L.
rugulosa GRAHAM, Edinb. N. Phil. J. (1829)
186.—Pratia begonifolia LINDL. Bot. Reg. (1830)
t. 1373; G. Don, Gard. Dict. 3 (1834) 699 (begoniae-
folia); PReEsL, Prod. Mon. Lob. (1836) 46; CLARKE,
Fl. Br. Ind. 3 (1881) 442; Ripv. J. Str. Br. R. As.
Soc. n. 33 (1900) 102; Merr. Philip. J. Sc. 1
(1906) Suppl. 241; BuRKILL & HoLtrum, Gard.
Bull. S. S. 3 (1923) 56 (begoniifolia); Rip. FI.
Mal. Pen. 2 (1923) 201; DANGuy, FI. Gén. I.-C.
3 (1930) 674; Craip, FI. Siam. En. 3 (1936) 302.—
L. reniformis CHAM. Linnaea 8 (1833) 210; DC.
Prod. 7 (1839) 365.—L. hederacea CHAM. Linnaea
8 (1833) 212.—Pratia hederacea G. Don, Gard.
Dict. 3 (1834) 699; Hook. f. & ARN. J. Bot. 1
(1834) 277; Prest, Prod. Mon. Lob. (1836) 46;
DC. Prod. 7 (1839) 340, incl. var. elliptica; HOOK.
f. Fl. Antarct. 1 (1884) 43; KANITz, in Mart. FI.
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.—Rapuntium angulatum
PresL, /.c. 30.—Rapuntium reniforme PRESL, l.c.
15.—Rapuntium nummularium PRESL, l.c. 30.—
Piddingtonia nummularia DC. Prod. 7 (1839) 341;
Mig. FI. Ind. Bat. 2 (1857) 572; Hook. f. & 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. 148.—Pratia zeylanica HAssk.
Flora 25, 2 (1842) Beibl. 1, p. 23; Cat. Hort.
Bog. (1844) 106.—Pratia arenaria Hook. f. FI.
Antarct. 1 (1844) 106, t. 29 in icon. errore Pratia
arenosa; Fl. Nov. Zel. 1 (1853) 157.—Pratia
angulata Hook. f. Fl. Antarct. 1 (1844) 43; FI.
Nov. Zel. 1 (1853) 157, incl. var. arenaria; Handb.
Fl. New Zeal. (1867) 172; CHEESEMAN, Man. New
Zeal. Fl. (1906) 397; Ill. New Zeal. Fl. (1908) t.
16; Winner, Pfi. R. Heft 106 (1943) 110, incl.
var.—Piddingtonia palliardii LEHM. Hamb. Gar-
tenz. 8 (1851) 337; Linnaea 25 (1852) 310; WALP.
Ann. 5 (1858) 391.—L. rotundifolia BANKS &
So. ex Hook. f. Fl. Nov. Zel. 1 (1853) 158.—
L. horsfieldiana Mia. FI. Ind. Bat. 2 (1857) 577.—
Pratia nummularia A. Br. & ASCHERS. Index
Sem. Hort. Berol. (1861) Append. 6; Kurz, J.
As. Soc. Beng. 46, ii (1877) 210; Koorp. Exk.
Fl. Java 3 (1912) 303; Botp. Zakfi. (1916) 41;
Koorpb. Fl. Tjib. 3, 2 (1918) 55; HeyNe, Nutt.
Pl. 2 (1927) 1428; Merr. Lingn. Sc. J. 1 (1927)
181; YAMAMOTO, Obs. FI. Form. 13 (1936) 150;
Wimmer, Pfi. R. Heft 106 (1943) 112; Lam,
Blumea 5 (1945) 569; BAcK. Bekn. Fl. Java (em.
ed.) 8 (1949) fam. 184, p. 3; Blumea 6 (1950) 360.
—Pratia linnaeoides Hoox. f. Handb. New Zeal.
Fl. (1867) 172.—Pratia reniformis KANITZ, in
Mart. FI. Bras. 6, 4 (1878) 136, t. 40 f. 2; WIMMER,
Rev. Sudamer. Bot. 2 (1935) 96; Pfi. R. Heft 106
(1943) 115.—L. linnaeoides PETRIE, Trans. New Zeal.
Inst. 23 (1890) 405; CHEESEMAN, Man. FI. New
Zeal. (1906) 400; Perrier, Gard, Chron, III, 47

Fig. 19. Lobelia angulata Forst. Mt Patuha, West
Java (DE VooGpD).

(1910) 99, f. 50; Wimmer, Pfi. R. Heft 107 (1953)
483, incl. var. brevipilis—Pratia papuana S.
Moore, Trans. Linn. Soc. II, 9 (1916) 88; DIELs,
Bot. Jahrb. 55 (1917) 125; ibid. 62 (1929) 493;
Wimmer, Pfi. R. Heft 106 (1943) 114.—Pratia
wollastonii S. Moore, Trans. Linn. Soc. II, 9
(1916) 89; Wimmer, Pfi. R. Heft 106 (1943) 111. —
?Pratia podenzanae S. Moore, J. Bot. 55 (1917)
306; Wimmer, Pfi. R. Heft 106 (1943) 111.—L.
arfakensis Gipps, Arfak (1917) 183; KANEHIRA &
Hatusima, Bot. Mag. Tokyo 57 (1943) 128;
Wimmer, Pfi. R. Heft 107 (1953) 483.—L. paradoxa
Wimmer, in Fedde, Rep. 26 (1929) 2; Pfl. R. Heft
107 (1953) 483.—Fig. 19-20.

Polymorphous, creeping and branching herb;
stem terete, up to 14 m long, rooting at the
nodes, glabrous or hairy. Leaves alternate, round
to ovate, broad-ovate or reniform, 2—25 by 2-35
mm, cordate, truncate, or even decurrent at the
base, crenate, dentate or subdentate to subentire

Dec. 1960] a

CAMPANULACEAE (Moeliono & Tuyn)

we

at the margin, acute or rounded at the top,
glabrous to hairy; petiole 0-25 mm, glabrous or
densely hairy. Flowers 6-18, axillary, solitary.
Pedicels 14—5(—7) cm, ebracteolate, glabrous or
hairy. Calyx lobes linear-lanceolate to triangular,
1-11 mm, entire to distinctly dentate with up to 3
teeth at either side, acute or blunt, glabrous to
puberulous. Corolla 444-15 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 114-5 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.
Filaments 3-11 mm, very variable. free for 34
of their length, equalin width, sometimes narrowed
to the base, or the two anteriors broader, some-
times the two anteriors adnate to the corolla,
glabrous or hairy or only the two anterior ones
hairy; anthers from dorsal to ventral 1-2 to 1-!4
mm, glabrous or hairy, only those of the two
anteriors with a hair tuft or with a seta and some
hairs. Ovary trumpet-shaped to obconical or
ovoid or campanulate, 1-4 by 1-3 mm; glabrous
to densely hairy; style glabrous or sparsely pilose.
Fruit an indehiscent or dehiscent capsule or a
subcarnose to baccate purple fruit, (1-)6!4-16
by (1-)5-13 mm, glabrous to hairy, ellipsoid to
globose, sometimes flattened at the apex. Seeds
flattened-ellipsoid to ellipsoid, nearly 1 mm,
brown, reticulate to fine-reticulate.

Distr. Very widely distributed, through SE.
Asia (Nepal, Sikkim, Khasia, Burma, Siam, Indo-
China) to China (Yunnan, Kwangsi, Hainan,
Formosa), throughout Malaysia, Borneo and the
Moluccas excepted, Australia, Tasmania, New
Zealand, and adjacent islands, to S. America.

Ecol. Moist open or shaded places, in mountain
forest, mossy forest, along river-banks and forest
trails, 600-3300 m.

Vern. Adntingan, ki tombe, kitomé, kuweung,
ramo keujeup, djukut mata keujeup gunung, tangkal
sutji, S, kétrus alus, manikan, sérintil, urék urék
polo, J; Philippines: gubagubai, kanapa, pisa, (g.,
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 Drets, who, in his revision of the Papuan
species, remarked:—‘‘Diese drei Arten (Pratia
papuana, P. nummularia, und P. angulata) sowie
die P. wollastonii von der Nassaukette, und mehrere
andere bei Pratia diagnostisierte Spezies, sind

iibrigens so geringfiigig, dass nach genauem
Studium der Formenkreis, wahrscheinlich um-
fassender Arten angenommen werden miissen.”
(Bot. Jahrb. 55, 1917, 125; ibid. 62, 1929, 493).

Fig. 20. Various fruits of Lobelia angulata Forst.

a—c. Fleshy berry to capsule, from Celebes,
d-e. capsular New Guinea specimens, all « 21/4
(a—c EyMA 1161, d Brass 10622, e NGF 4789).

Also WIMMER himself, in identifying a Philip-
pine specimen which was astonishingly resembling
Pratia nummularia but possessed fruits with valves
on top, came obviously reluctantly to the conclu-
sion that this character settled it as a Lobelia;
he named it L. paradoxa!

BEUZENBERG & Hair recently found the fol-
lowing chromosome numbers: for L. linnaeoides
(Hook. f.) PETRIE n = 7 and for Pratia angulata
(Borst) Hooke fz ni — 35nCNeZealee Seaee G59:
32 Si):

12. Lobelia conferta Merr. & PERRY, J. Arn.
Arb. 59 (1949) 59.—Pratia conferta WIMMER,
Pfl. R. Heft 107 (1953) 764.—Fig. 21.

Tiny, prostrate, fleshy, glabrous herb, with
short stems. Leaves distichous, conferted, semi-
amplexicaulous, sessile, oblong-lanceolate, entire,
4-5 mm by 1-2 mm, blunt to obtuse, smooth and
shining. Flowers 5 mm long, in the upper leaf
axils. Pedicels 2-5 mm, ebracteolate. Calyx lobes
linear to lanceolate, 114-214 mm by | mm, entire,
blunt to slightly acute. Corolla 44144 mm,
glabrous, very light purple, inside with two
gibbosities, dorsally split to the base; dorsal lobes

WA
LS

(|

{ @s9

Fig. 21. Lobelia conferta MERR. & PERRY. a.
Habit, x 214, b. flower, x 5, anther tube opened
in front (BRAss 4417).

136

FLORA MALESIANA

{ser: T;-voli6t

connate with the lateral ones for 114-2 mm; lateral
lobes connate with the ventral one for 2-24 mm;
dorsal lobes lanceolate, 14-214 mm, reflexed,
sometimes of unequal length; lateral and ventral
lobes equal, lanceolate to ovate, 114—214(-3) mm
by 34-1 mm, reflexed. Filaments 114-3 mm, half-
way free, linear, glabrous; anthers from dorsal to
ventral 114-1 mm, glabrous, the two anterior
anthers finely haired at the apex. Ovary trumpet-
shaped to cupular, 14—34mm long. Fruit not seen.

Distr. Malaysia: New Guinea (Owen Stanley
Range, SW. slope of Mt Albert Edward), once
found.

Ecol. Wet grasslands, 3680 m.

KEY TO SOME
CULTIVATED SPECIES

(in all spp. both anterior anthers with bearded tip)

1. Corolla tube less than 7 mm long. Pedicels

glabrous. Leaves petiole-like decurrent.

2. Corolla tube 5-7 mm long. Flowers axillary.
Pedicels 114-5 cm. Calyx tube 2-4 mm, lobes
314-11 mm. Corolla blue or violet, rarely
white. Stem triangular. Leaves 1!4—5 by 1/3-1
cm, oblong lanceolate, entire to serrate.
Annual, native in S. Africa. L. erinus L.

2. Corolla tube 3-314 mm, inside hairy. Pedicels
less than 114 cm. Calyx tube 2-3 by 2 mm,
lobes 2-214 by 14 mm, acute, entire. Corolla
blue or blue-violet. Stem quadrangular,
fistulose. Leaves broad-ovate, rather coarsely
sinuate-dentate, 114-314 by 1-2 cm. Native
of Central America, early introduced as an
ornamental . . . . L. cliffortiana L.

1. Corolla tube longer than 10 mm. Pedicels

hairy. Leaves sessile, the upper ones longer
than 5 cm. Perennials.

3. Flowers all axillary. Pedicels 2!4-8 cm,
shorter than the sustaining leaf. Sepals erect,
214-314 mm. Corolla tube bright red, 18-21
mm; lobes yellow to orange, 12-14 mm, lower
lip short-3-lobed, posterior lobes linear.
Leaves ovate-lanceolate, very shallowly den-
tate, 6-12 by 134-3 cm. Erect shrub 114-214
m. Native in Mexico. L. laxiflora H.B.K.

3. Upper flowers in bracteate, terminal racemes.
Pedicels shorter than 3 cm. Sepals longer than
5 mm. Lower lip over halfway 3-split. Herbs.

4. Flowers purple, rarely white. Pedicels 6-8
mm. Racemes 5—20 cm, lower bracts folia-
ceous. Sepals erect, 8-10 mm, with recurved
ciliate margins. Corolla tube 12-15 mm,
lower lip 10-12 mm, at the base with 2 white
convexities. Leaves oblong, 6-15 by 2—5S cm.
Native in N. America L. syphilitica L.

4. Flowers dark-red. Pedicels 8-28 mm.
Racemes 8-30 cm. Bracts narrow. Sepals
recurved, glabrous, 10-15 mm. Corolla tube
15-20 mm, lobes 20-35 mm long, posterior
ones much narrower than the anteriors.
Leaves linear-lanceolate, glabrous, 8-20 by

1-214 cm. Native in America, mostly cited
as L. fulgens WILLD. L. cordigera Cay.

Excluded

Lobelia anceps THuns. Prod. (1794) 40, non
LINN. f. 1781; Mia. Fl. Ind. Bat. 2 (1856) 578 =
Lobelia alata LABILL. Noy. Holl. Pl. 1 (1804) 51,
t. 72; Wimmer, Pfl. R. Heft 107 (1953) 469.

MIQUEL, /.c., quoted this species as possibly
occurring in the Sunda Islands. According to
WIMMER it is an extra-Malaysian species with a
distribution in S. Africa, Australia, and Chili.

Lobelia pumila Burm. f. Fl. Ind. (1768) 186,
t. 60, f. 3 (type in G) = Mazus pumilus (BURM.
f.) STEEN. Nova Guinea n.s. 9 (1958) 31 (Scro-
Phulariaceae).

Lobelia tetragona Bu. Bijdr. (1826) 729; HAssk.
Cat. Hort. Bog. (1844) 105 = Lobelia cliffortiana
LINNE, Sp. Pl. 2 (1753) 931; Miq. Fl. Ind. Bat. 2
(1857) 577; Wimmer, Pfi. R. Heft 107 (1953) 526.

BLUME described this species as an endemic
from Mt Gedeh, in West Java, but his specimens
have doubtless been erroneously localized and
were derived from specimens cultivated in the
Botanic Gardens at Bogor or possibly naturalized
in their vicinity. It is a native of Central America
and has been introduced as an ornamental plant
into several other countries at an early date and
seems to have frequently naturalized.

Rapuntium longifolium Pres_, Prod. Mon. Lob.
(1836) 26.—L. longifolia DC. Prod. 7 (1839) 382;
Mia. FI. Ind. Bat. 2 (1856) 578; F.—VILL. Nov.
App. 4 (1880) 121; Merr. En. Philip. 3 (1923)
588 = Lobelia graminea LAMK, Dict. Bot. 3 (1791)
583; Wimmer, Pfl. R. Heft 107 (1953) 413.

PrESL based this species on a specimen from
the Malaspina Expedition collected by HAENKE.
According to MERRILL it was erroneously localized
in Luzon and came really from Central America.
WIMMER has examined the type specimen (in
Mus. Prague) and has referred it to L. graminea
LAMK, which is a native of Panama and Mexico.

Rapuntium haenkeanum PrReEst, Prod. Mon.
Lob. (1836) 26.—L. haenkeana DC. Prod. 7
(1839) 382; Mia. Fl. Ind. Bat. 2 (1856) 578;
F.-VILL. Nov. App. 4 (1880) 121; Merr. En.
Philip. 3 (1923) 588; Wimmer, Pfl. R. Heft 107
(1953) 685.—Dortmannia haenkeana O.K. Rev.
Gen. Pl. 2 (1891) 972 = Lobelia nelsonii FERNALD,
Proc. Am. Ac. Arts Sc. 36 (1901) 503 = Lobelia
laxiflora var. nelsonii MCVAuUGH, North Am. FI.
32A (1943) 97.

According to PRESL this species was based upon
a specimen from Luzon. MERRILL, /.c., already
suspected that this record was based ona Malaspina
Expedition plant from tropical America er-
roneously localized as Philippine. WIMMER, /.c.,
saw HAENKE’s type specimen (Mus. Prague) and
stated that it is conspecific with “Lobelia laxiflora
var. nelsonii MCVAUGH”’ from Guatemala, cor-
roborating MERRILL’s statement about its native
country.

Dec. 1960] CAMPANULACEAE (Moeliono & Tuyn) 137

7. PHYLLOCHARIS

Diets, 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

a, 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 from Lobelia in the epiphyllous flowers, with 2 much prolonged
upper lobes. A third differential character mentioned by DIELS, viz esetose anther tips, does not hold
for P. subcordata. 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 differential characters appear satisfactory it remains to
be seen whether they will prove to be constant.

Sincere thanks are due to Dr VAN RoyeEN 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 . : Rate 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

1. Phyllocharis oblongifolia Die_s, Bot. Jahrb. 55
(1917) 124, fig. 1 L-N; Wimmer, Pfl. R. Heft 107
(1953) 724, fig. 108 c.—Fig. 22g.

Almost glabrous herb, stems 15-30 cm. Leaves
6-11 by 214-3 cm, oblong-elliptic, + acuminate,
base cuneate, entire but the veins protruding as
small callose-fimbriate teeth; nerves c. 8 pairs,
arching; petioles 1-314 cm. Flowers inserted at
about the middle of the midrib. Pedicels c. 5 mm.
Calyx tube 114-2 by 2 mm, lobes lanceolate,
acute, 2-3 mm, both warty. Corolla tube 14-1
mm; posterior lobes 9 by 14 mm, at least twice as
long as the anterior ones, narrow-spathulate,
broadened at the base, acute; anterior lobes
314-4 by 2-214 mm, ovate, acute, connate up to
the middle, at the inside near the posterior lobes
somewhat pilose, further glabrous. Androecium
214-34 mm long; anthers 1-114 mm, posterior
ones dorsally + hairy. Seeds verruculose.

Distr. Malaysia: NE. New Guinea (Udu:
SCHLECHTER 17445).

Ecol. Rain-forest, 300 m. Fi. fr. March.

2. Phyllocharis schlechteri Die_s, Bot. Jahrb. 55

4. P. saxicola

(1917) 124, fig. 1 A-K; Wimmer, Pfi. R. Heft
107 (1953) 725.—Fig. 22h.

Stems 10-20 cm, glabrous below, the upper part
slightly puberulous. Leaves 2-5 by 1'4-2'4 cm,
ovate to ovate-elliptic, acute, base cuneate, very
sparsely hairy above, puberulous on the nerves
beneath, margin dentate, ends of the veins cal-
lously protruding at the margin; nerves c. 6—7
pairs; petioles glabrous, 4-114 cm. Flowers
inserted in the basal quarter of the midrib.
Pedicels 5—6 mm, glabrous. Ca/yx tube c. 144 mm
long, hairy, 10-ribbed; lobes 4 by 14 mm, linear,
acute, with scattered hairs along the margin.
Corolla tube 1 mm; posterior lobes 7-8 by 14 mm,
1, to 4 longer than the anterior ones, linear,
acute, glabrous; anterior lobes 5-6 by 114-214
mm, oblong-ovate, acute, connate for about
24 of their length, inside papillose. Androecium
3 mm long; anthers | mm, posterior ones dorsally
hairy. Capsule 6-7 by 3 mm. Seeds smooth.

Distr. Malaysia: NE. New Guinea (Bismarck
Mts: SCHLECHTER 18620).

Ecol. In humous rain-forest, 1200 m. Fi. fr. Nov.

FLORA MALESIANA [ser. I, vol. 6!

Fig. 23. Laurentia longiflora (L.) PETERMANN. Java (ROEPKE).

Dec. 1960]

CAMPANULACEAE (Moeliono & Tuyn)

139

Fig. 22. Phyllocharis subcordata MERR. & PERRY.

a. Habit, x %4, b. flower, x 4, c. fruit, x 4.—

Phyllocharis saxicola VAN ROYEN. d. Habit, « 74,

e. fruit, x 4, f. seed x 13.—Phyllocharis oblongi-

folia Diets. g. Flower, x 4.—Phyllocharis schlech-

teri Diets. h. Flower, * 4 (a—c CLEMENS 4404,
d-f Carr 14083, g—h after DIELS).

3. Phyllocharis subcordata Merr. & PERRY, J.
Arn. Arb. 22 (1941) 387; Wimmer, Pfi. R. Heft
107 (1953) 725, fig. 108b.—P. lamiifolia WIMMER,
Ann. Naturh. Mus. Wien 56 (1948) 372; Pfi.
R. Heft 106 (1943) 11, fig. 8e.—Fig. 22a-c.

Plant almost glabrous, 5-20 cm. Leaves
1-314 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; merves c. 4 pairs;
petioles c. 1-114 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,
1-114 mm wide at the base, twice as long as the
anterior ones, acute, glabrous, anterior lobes 3
by 114 mm, ovate, acute, connate at least up to
the middle, inside papillose. Androecium 2—3 mm;
anthers 1-114 mm, posterior ones dorsally pilose,
anterior ones with a few setae on top. Capsule
4-414 by 2-3 mm. Seeds about 14 mm, 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. Fi. fr. June, Nov.—Dec.

4. Phyllocharis saxicola VAN ROYEN, roy. sp.—
Fig. 22d-f.

Caules, folia et calyx pilis crispis albis pubes-
centes. Folia ovata ad elliptica, obtusiuscula, in-
tegra vel propter venulas ultra marginem in cal-
lositate breviter producta denticulata, basi rotun-
data ad truncata. Flores in costae parte quarta
inferiore inserti. Semina verruculosa. Typus: CARR
14083, L.

Stems 5-12 cm, short crisped-hairy. Leaves
sparsely hairy on both sides, 1-2!4 by 14-2 cm,
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 1/,-1 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
2-214 mm, linear, + acute, sparsely whitish
hairy. Posterior corolla lobes green with a crimson
median line, anterior lobes crimson, whitish pilose
near-the apex. Androecium c. 2 mm long; posterior
anthers with a few white hairs. Capsule 4-6 by
2-3 mm, very obliquely oblong-obovoid, 10-
nerved, pilose. Seeds verruculose, 4 mm.

Distr. Malaysia: Papua (Lala River).

Ecol. On rocks in forest, 1650 m. Fi. fr. Dec.

8. LAURENTIA

MIcu. ex ADANS. Fam. 2 (1763) 134, 568; ENDL. Gen. PI. 1 (1838) 511; DC. Prod.
7 (1839) 409; McVauGu, Bull. Torr. Bot. Cl. 67 (1940) 778; Wimmer, Pfi. R. Heft
107 (1953) 386.—Isotoma LINDL. Bot. Reg. 10 (1824) t. 964; PREsL, Prod. Mon.
Lob. (1836) 42; DC. Prod. 7 (1839) 412; BENTH. FI. Austr. 4 (1869) 134.—Hip-

140 FLORA MALESIANA

pobroma G. Don, Gard. Dict. 3 (1834) 717; McVauGu, Bull. Torr. Bot. Cl. 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 anthers, free from the corolla,
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 o, 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. Cl. 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, /.c. 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 /sotoma as representing
a monotypic genus, although it is manifestly allied to its Australian congeners. GLEASON on the other
hand (Bull. Torr. Bot. Cl. 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 Micn. Gen. 18, t. 14 = Lobelia laurentia L.
1753 = Laurentia gasparrinii (TINEO) STROBL.. As he reduced Lobelia L. to Laurentia a formal conser-
vation of Laurentia would seem necessary.

[ser. I, vol. 6%

1. Section Isotoma

(R. Br.) ENDL. Gen. Pl. 1 (1838) 512; Wimmer, Ann. Naturh. Mus. Wien 56 (1948)
335; Pfl. R. Heft 107 (1953) 398.—Lobelia sect. Isotoma R. BR. Prod. (1810) 564;
McVauGH, Bull. Torr. Bot. Cl. 67 (1940) 794.—Jsotoma LINDL. Bot. Reg. 10 (1824)
t. 964.—Hippobroma G. Don, Gard. Dict. 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-
reich (1845) 444, t. 118 fig. 665; ed. 2 (1847) id.;
Wimmer, in McBride Fl. Peru 6? (1937) 474;
Ann. Naturh. Mus. Wien 56 (1948) 337, cum
var. runcinata (HASSK.) WIMMER; Pfl. R. Heft
107 (1953) 405.—Lobelia longiflora LINNE, Sp.
Pl. (1753) 930; LINDL. Bot. Reg. 14 (1828) t. 1200.
—Rapuntium longiflorum Miu. Dict. ed. 8 (1768)
n. 7.—Hippobroma longiflora G. Don, Gard. Dict.
3 (1834) 717; McVAuGH, Bull. Torr. Bot. Cl. 67
(1940) 783; N. Am. FI. 32A, pt 1 (1943) 99.—

Isotoma longiflora PresL, Prod. Mon. Lob. (1836)
42; DC. Prod. 7 (1839) 413; HEYNeE, Nutt. PI.
(1927) 1428; Back. Trop. Natuur 9 (1920) 129,
fig. 1-3; Fawc. & RENDLE, Fl. Jam. 7° (1936)
138; Back. Bekn. Fl. Java (em. ed.) 8 (1949)
fam. 184, p. 5; STEEN. FI. Sch. Indon. (1949)
376; Quis. Med. Pl. Philip. (1951) 955.—Isotoma
runcinata HaAssk. Bonplandia 7 (1859) 189, ex
descr.—Fig. 23.

Perennial (always?), erect or ascending, 10-30
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
running from the leaves downwards. Leaves
sessile, elongate-obovate, attenuated at the base,
acute or obtuse, mostly sparsely hairy, coarsely
sinuate-dentate, 3-16 by 1—3 cm. Flowers solitary,
axillary, almost regular. Pedicels densely short-
hirsute, 3-15 mm. Bracteoles 2, at the base of
the pedicels, narrow, acute, up to 5 mm. Calyx
lobes inequal, narrow-lanceolate or linear, ir-
regularly dentate, acute, hairy, 8-20 mm. Corolla
white, hairy, persistent, tube funnel-shaped,
5—9 cm, lobes spreading, almost equal, lanceolate,
acute or obtuse, 10-25 mm. Filaments inserted
above the middle of the corolla tube, more or less
united, as long as or somewhat exceeding the
corolla tube, anthers curved forward, witha bearded
top, 4-6 mm, the 2 in front 2-3 mm. Ovary
obconic to bell-shaped, (mostly 10-)ribbed,
densely hairy, especially along the ribs, 3-7 mm;
style equalling the anther tops or somewhat
longer: stigma broad, flat, -- 2-lobed, bearing
long slender hairs beneath. Capsule nodding,
ellipsoid. Seeds c, ellipsoid, foveolate-reticulate.

Distr. Native in the West Indies (the type from

Jamaica), in the lowlands from Florida and
Sonora to Brazil and Peru; introduced into Penn-
sylvania (U.S.A.), Hawaii, the Marianas, and the
Old World tropics, almost throughout Malaysia.

Ecol. In damp places, along ditches and
streams, against walls, from sea-level up to 300
(-1200) m. Fi. Jan.-Dec.

This is the sole species which does not possess a
trigger-hair on the two lower anthers, as described
in Isotoma by HILDEBRAND (Bot. Zeit. 27, 1869,
476, f. 8-12) and MELVILLE (Kew Bull. 14, 1960,
277-279, f. 1). In several American Laurentias the
two lower anthers have each 2 setae.

Vern. Daun tolod, S, daun kéndali, J, tambakis,
N. Born., Philip.: estrélla, Sp., Tag., lagrimao de
San Diego, revienta caballos, Sp.

Note. The milky juice is said to be very
poisonous, in particular for horses. According to
BoorsMA (HeEyYNE, Quis., //. cc.) its alcaloids
have a paralysing effect. The frequent occurrence
along streams seems to point to seed dispersal
by water; that against walls to seed dispersal by
ants, but there is no elaiosome.

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 (JACQ.) DC.

JUGLANDACEAE (M. Jacobs, Leyden)

Juglandaceae 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 Jug/ans 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.. Platycarya and Pterocarya are
Asiatic, Alfaroa is American, while Engelhardia, Carya, and Juglans occur in both the Old
and the New World. As for the disputed monotypic genus Annamocarya CHEV. (synonyms:
Caryojuglans KircHH., Juglandicarya Rem & CHANDLER, Rhamphocarya KUANG), described
in 1941 from S. China and Tonkin, we refer to LeRoy, Mém. Mus. Hist. Nat. n.s. Bot. 6
(1955) 66.

The family is a coherent one. Engelhardia and Al/faroa 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. Kjobenhavn (1870) 159-174. The second is by
KoImpzumMI, 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, /.c. 1-246. The three systems have
in common, that they always keep Engelhardia together with Pterocarya, and Carya together
with Juglans.

The affinity of the Juglandaceae within the Amentiferae seems to be closest with the Myricaceae.

Palynology. According to ERDTMAN (Pollen Morph. & Pl. Tax. 1952, 216) the pollen
grains have several characters incommon with those in Betulaceae, 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 Juglans, Carya, and Pterocarya. 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 of Juglandaceae; fresh barks and leaves of Juglans
rupestris ENGELM., Carya illinoensis (WANG) K. Kocu, 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%
of flavonoid glycosides (engelitin, isoengelitin, afzelin, astilbin, quercitrin). Similar compounds
were isolated from leaves of Juglans 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 Platycarya strobilacea.

(143)

144 FLORA MALESIANA [ser. I, volicg?

Fig. 1. Engelhardia rigida BL. a. Fruiting twig, x 1, d-f. variation in the 3 flowers, all x 6, g. 3 flower

after removal of the stamens, with 2 perianth lobes and 3 bract lobes, fh. 3 flower from above, with

the bract lobes (b) and 4 perianth lobes.—E. serrata BL. (?) b. Seedling, x 14.—E. spicata LECHEN.

ex BL. c. 2 Flower, < 6 (a Brass 11061, b 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. Hamamelidaceae, Fagaceae, etc.—R. HEGNAUER.

1. ENGELHARDIA

LECHEN. ex BLUME, Bijdr. 10 (1825) 528; Fl. Jav. Jugl. (1829) 5, ‘Engelhardtia’ ;
C.DC. Ann. Sc. Nat. IV, 18 (1862) 35; Prod. 16, 2 (1864) 140; NAGEL, Bot. Jahrb.
50 (1914) 475.—Pterilema REINW. Syll. 2 (1826) 13.—Oreomunnea OrrstT. Vid.
Medd. Nat. For. Kjobenh. (1856) 52.—Fig. 1-9.

Trees, mostly lepidote with golden yellow, glandular scales 0.1-0.2 mm diam.
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.—< Flowers: perianth often reduced and irregular; stamens
4-13 (in Asiatic members), (sub)sessile-—Q Flowers: perianth lobes in 2 whorls
partly connate with the ovary, the median pair exterior; ovary 2-carpellate, the
carpel sutures median, |-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 |
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 filling 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 + 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 Engelhardia 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 Eupatorium inulifolium 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. rigida 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-anat. BEEKMAN, Meded. Proefstat. Boschw. 5 (1920) 96; DEN BERGER, Meded. Proefstat.
Boschw. 13 (1926) 13, Determinatietabel van Malesié, Veenman, Wageningen (1949) 62 (hand lens.)
HermscH, Lilloa 8 (1942) 164; Kriss, Trop. Woods 12 (1927) 16; METCALFE & CHALK, Anat. Dic. 2
(1950) 1287; MoLL & JANssonius, Mikr. Holzes 6 (1936) 308; Trepo, Bot. Gaz. 100 (1938) 42. For
various opinions about the affinities of the family see METCALFE & CHALK, /.c.—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 Myricaceae, 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 ¢ 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 9 flowers, ibid. 35 (1948) 606-621 on the ¢ flowers, Bull. Torrey Bot. Club
86 (1959) 190-198 on Engelhardia and Alfaroa in America; HJELMQvisT, Bot. Notis. Suppl. 2! (1948)
5-171; Leroy, Mém. 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

JUGLANDACEAE (Jacobs) 147

and also of N. America in the Tertiary. Dr W. H. ZAGWUN 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 ZAGwisN, Aspects of the Pliocene and early Pleistocene
Vegetation in the Netherlands, thesis Leiden 1960. An endocarp of +E£. nucifera (LUDWIG) 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 2 catkin or the few 3 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 individual 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 2 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: 1; voles

in dry material the exact moment when the stigmas are receptive; material with 9 flowers is, moreover,
very rare in herbaria. The collecting of 2 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. Trichotocarpeae 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. rigida BL., E. serrata BL., and E. apoensis ELMER
ex NAGEL.

Sect. 2. Psilocarpeae NAGEL emend. LeRoy. Leaves spirally arranged. Inflorescences terminal, sometimes
also lateral. Male flowers: stalked; bract very obscurely 3-lobed; receptacle orbicular; perianth regular,
4-lobed, lobes cucullate, 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 (OeERsST.) 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 + 14 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 Oreamunoa, 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 all names given to Asiatic Engelhardias. | 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 ITO THE SPECIES

1. Nut hispid, (sub)sessile, with distinct style and stigmas. Bract of the ¢ flowers with 3 distinct narrow
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 mm long, 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 . . el Aa WE As) OG ee aes
4. Leaflets (more or less distinctly) serrate or crenate ; . 2. E. serrata
3. Nut elongate, ovoid, 6-8 mm long, wing 514—7!, cm long. ‘Leaflets ovate, the base subcordate

to rounded. (Flowers unknown). . ‘ 3. E. apoensis

2. Leafiets 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. 2.—E. subsimplicifolia MeERR. Govt Lab.
Fl. Jay. Jugl. (1829) 13, t. 3; Mig. Fl. 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; Nace, Bot. Jahrb. 50 (1914) 477;
K. & V. Bijdr. 5 (1900) 175; Koorp. Exk. Fl. Merr. En. Philip. 2 (1923) 24.—E. 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; Koorp. Atlas 4 (1916) t. 698; BAkH. f. in (1923) Suppl. 53.—E. 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)

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!4—)3-6(-21) cm, mostly
glabrous or scaly, rarely puberulous, (1—)2-3
(-5)-jugate; leaflets rather thick-coriaceous, ses-
sile to 5 mm stalked, blade 2!4—314(—4) times as
long as wide, widest at the middle or rarely above,
514-1014(-1614) by 2!4-4(-7'4) 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 !4-1 cm long stalk, 114-5
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, (!4—)1 mm, more or less hispid,
connective not or shortly protruding; if there is a
single proximal stamen, this has sometimes a
filament up to | 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 mm diam., hispid, wing
(2-)214-3(-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 mm long.

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
and Numfoor). In New Guinea rather common,
elsewhere rare.

Ecol. Primary forest, only once reported from
secondary forest, c. 1000—1800(—2300) m.

Vern. Ki hudjan, S, sibiri, Papua.

Uses. The wood is used for canoes, native
buildings, etc.

Notes. Only in Borneo, the Philippines, and
Celebes occur specimens with 1l-jugate leaves.

Closely related to E. serrata, and arguments
could be advanced to regard the two as subspecies.
Separation based on reproductive characters is
practically impossible. However, apart from the
different distribution, either species has its own
particular pattern of variability, as has been
pointed out in the descriptions. For separation
two characters have been found, notably the leaf
margin, which is entire or serrate, and the decreas-
ing in size of the leaflets towards the leaf base;
this feature is generally far more pronounced in
E. serrata, where the basal leaflets are about half
as long as the largest ones. These characters
mostly concur. In the cases where they do not, I
am inclined to take the leaf margin character as
decisive, since in a few E. rigida specimens from
New Guinea and elsewhere such small basal

Fig. 4. Fruits of Engelhardia. a. E. roxburghiana

WALL., ventrally, nat. size, 6. the stigmas,
shown by removal of 2 perianth lobes, x 4.—E.
rigida BL. c. Fruit dorsally, * 2, d-f, fruits
ventrally, showing variation in perianth and
adaxial lobe, x 2.—E. serrata BL. g. Longi-
tudinal section of fruit, showing ovulum on
columella (c) and partial septa (s), = 4 (ab
CLEMENS 27250, c-d WINCKEL 1540/8, e BRAss
11051, f KALKMAN 4363, g JACOBS 4677).

leaflets were also found. Engelhardia zambalensis,
described on a specimen with entire leaflets but
much reduced towards the leaf base, is therefore
placed by me under E. rigida, but if one accepts
the narrowing leaf base as the key character, it
would come under E. serrata.

Two sterile specimens from Japen and Numfoor
are believed to belong here, notwithstanding they
lack scales on the underside of the leaves.

In ENpeRT 3833 and KOSTERMANS 7648, both
from E. Borneo, the fruit bears no long hairs,
only a soft pubescenceand scales. From E. roxburg-
hiana these specimens are distinguished by well-
developed style and stigmas. In RosBBINs 569
from New Guinea even the anthers bear scales.

150

FLORA MALESIANA

[ser. I) volongs

2. Engelhardia serrata BL. FI. Jav. Jugl. (1829)
14, t. 4, Sc; Mia. Fl. Ind. Bat. 1, 1 (1856) 843;
C. DC. Prod. 16, 2 (1864) 141; Hook. f. FI. Br.
Ind. 5 (1888) 596; K. & V. Bijdr. 5 (1900) 172,
174; Koorp. Exk. Fl. Java 2 (1912) 53, f. 7;
NAGEL, Bot. Jahrb. 50 (1914) 477; Koorp.
Atlas 4 (1916) t. 699; Dope, Fl. Gén. I.-C. 5
(1930) 928; Quis. Philip. J. Sc. 76 (1944) 37;
BAKH. f. in Back. Bekn. Fl. Java (em. ed.) 7A
(1948) fam. 155 p. 3.—E. palembanica Mig. Sum.
(1861) 346, 139.—E. 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.—E. nudiflora Hook. f. Ic.
Pl. 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 Hoox. f. ex GAMBLE; RIDL.
Fl. Mal. Pen. 3 (1924) 369, f. 158; CoRNER,
Ways. Trees (1940) 333, f. 117.—E. permicrophylla
Eimer, Leafl. Philip. Bot. 9 (1934) 3194.—Fig.
1b, 4g, 5.

Tree (2—)20—25(-42) m, sometimes with but-
tresses up to 3-4 m high and 1!4 m 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, 114-3 times as long as
wide, widest at the middle or above, (1—)2-13!4
by (14-)1+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
stalked, 3-10 cm long; perianth completely
reduced or almost so; stamens (3—)5-—7, sessile to
1, mm stalked, anthers sometimes unequal,
14-1 mm, sparsely hirsute, connective not or
hardly protruding. Fruiting catkins 9-14 cm long
in all, 2-5 cm stalked, rachis as thick as that of
the leaves, shallowly grooved. Nut (sub)sessile,
+ 3 mm diam., hispid, wing 114-314(-334) cm
long with the nut, 5-10 mm wide: perianth lobes
rather equal, often not adnate to the style, style
about as long as the stigmas, the whole 2-4 mm
long.

Distr. Upper Burma, N. Siam, Indo-China
(Laos, Cambodia), in Malaysia: Malay Peninsula,
Sumatra (S of Lake Toba, also Bengkalis, Banka,
Billiton), W. Java, Borneo, Philippines (Luzon to
N. Mindanao), Celebes, Moluccas (W. Ceram).

Ecol. Primary dryland forest (rarely secondary
forest) on sandy or clayey soil, from sea-level to
2200 m. In Malaya the leaves are shed at the end
of a dry spell without fading into autumn tints
(CORNER).

Vern. Ki hudjan, (lang)kédi, (bé)béri, M, ki
keper, S.

Fig. 5. Variation in leaflets of E. serrata BL.,

the acroscopic side on the left, a-d. « 7%, e. leaf

base densely set with glandular scales (a Hout-

soorten Gedeh 126, b Suit 21595, c, e HOLTTUM
14857, d CLEMENS 16888).

Notes. In the Philippines several collections
possess very small leaflets, glabrous, viz 2-3 cm,
the fruit wing being + 15-21 mm long. In Celebes
specimens occur in which the leaflets are slightly
larger and 2-3-jugate, the fruit wing being
slightly smaller. In Malaya, Sumatra, Banka, and
Billiton the leaflets are generally 5-7 cm and
glabrous. In Java the leaflets are thin in texture,
up to 9-jugate, 8-1314 cm long, serrate from the
base, and hirsute, especially on the nerves. Inter-
grading specimens occur throughout the area.

PIERRE 3301, from Cambodia, is a very bad
specimen with entire, acuminate leaves glandular-
scaly on both sides. Most probably it belongs here.

JAcoBS 4677, from Mt Sago in W. Sumatra,
in fruit, is the only specimen with entire leaflets.
The leaf rachis is sparsely hairy, the leaflets are
4 times as long as wide, the perianth lobes are
not connate with the style. It tallies with a

Dec. 1960]

JUGLANDACEAE (Jacobs)

151

specimen from the same locality that has serrate
leaves.

Most of the extra-Malaysian specimens, es-
pecially those from N. Siam, are very densely
brown-pubescent on the twigs, the rachises of
leaves and catkins, on the leaflets underneath and
the nerves above. The largest leaflet seen was 1714
by 514 cm. The leaflets are sometimes nearly
entire, with their indument resembling E. spicata
var. colebrookeana, but have always yellow scales
below.

3. Engelhardia apoensis ELMER ex NAGEL, Bot.
Jahrb. 50 (1914) 477; Emer, Leafl. Philip. Bot. 7
(1915) 2693.—Engelhardtia sp. CORNER, Ways.
Trees (1940) 333, f. 117.—Fig. 6.

Leaf 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
-+- 21%4-3 times as long as wide, ovate, (7—)9-13
by 214-3(—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 24 (CF 24424 SymincrTon).

some minute tomentum on the nerves. Flowers
not known; inflorescences presumably lateral.
Fruiting catkins 12-30 cm long in all, stalk 4-6
cm. Nut -++ 6-8 by 5-6 mm, the body ovoid,
elongate, wing (5!4-)6-7!14 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
the type specimen from Mindanao, ELMER 11744,
very well, the Malayan plants having somewhat
longer catkins and larger fruits. In RICHARDS 1521
from N. Sarawak the leaves are entire.

4. Engelhardia spicata LECHEN. ex BL. Bijdr. 10
(1825) 528; Fl. Jav. Jugl. (1829) 8, t. 1, 5A; Mia.
Fl. Ind. Bat. 1, 1 (1856) 842; C. DC. Prod. 16,
2 (1864) 140; Kurz, For. Fl. Burma 2 (1877)
491; VIDAL, Sin. Atlas (1883) t. 90A; Hook. f.
Fl. Br. Ind. 5 (1888) 595; K. & V. Bijdr. 5 (1900)
165, 170, incl. var. « genuina et var. B aceriflora,
var. y colebrookiana quoad specim. tantum;
Koorp. Exk. Fl. Java 2 (1912) 51, f. 6; NAGEL,
Bot. Jahrb. 50 (1914) 476, incl. var. aceriflora;
LreEFM. Trop. Natuur 3 (1914) 46 with fig., 77;

Fig. 7. Engelhardia spicata LECHEN. ex BL. Large

tree, over 40 m tall, in primary forest near

Tjikahuripan, Mt Gedeh, W. Java; stem base

8.84 m in girth, dbh 2.8 m (F. KRAMER, who is
standing in front).

152

FLORA MALESIANA

[ser.. 5, -volieG:

GAMBLE, J. As. Soc. Beng. 75, ii (1915) 400;
Koorp. Atlas 4 (1916) t. 700; Merr. Sp. Blanc.
(1918) 120; En. Philip. 2 (1923) 24; Koorp. FI.
Tjibodas 7 (1923) 10, var. aceriflora (REINW.)
K. & V.; Rip_. Fl. Mal. Pen. 3 (1924) 368;
DocTEeRS VAN LEEUWEN, Zoocecidia (1926) 103;
Dope, FI. Gén. I.-C. 5 (1930) 930, f. 107, 3-7;
CuHuNn, Sunyatsenia 4 (1940) 245; CorRNER, Ways.
Trees (1940) 333, f. 117; KANJ. c.s. Fl. Assam 4
(1940) 229; p—E VooGp, Trop. Natuur 30 (1941)
103net 56.1125) BAKE: o/2in) Backes Bekne nl:
Java (em. ed.) 7A (1948) fam. 155 p. 2.—Prerilema
aceriflorum RetNw. Sylloge 2 (1826) 13.—E.
aceriflora Bu. Fl. Jav. Jugl. (1829) 11, t. 2, 5B;
Mig. Fl. Ind. Bat. 1, 1 (1856) 842; C. DC. Prod.
16, 2 (1864) 141; Hook. f. Fl. Br. Ind. 5 (1888)
596, ‘acerifolia’; FoRBES & HeEMSL. J. Linn. Soc.
Bot. 26 (1899) 495; von MALM, Fedde Rep. 34
(1934) 271.—E. roxburghiana LINDL. ex WALL.
Pl. As. Rar. 2 (1831) 85, t. 199 f. 7, quoad fruct.—
Juglans pterococca Roxs. Fl. Ind. (ed. Carey) 3
(1832) 631, quoad fruct., ‘plerococca’.—Giro-
carpus pendulos BLANCO, Fl. Filip. ed. 2 (1845)
55, ed. 3 (1877) 104, t. 387.—E. philippinensis

Fig. 8. Engelhardia spicata LECHEN. ex BL. in
clearing of coffee estate Kalisat, Mt Idjen, E.
Java (J. H. CoermT).

GDE. Anns Sc: Nat) IV; 18)\(1862)" 345 sta
15; Prod. 16, 2 (1864) 141.—Fig. 1c, 3, 7-9.

Tree (5—)12—30(-36) m, sometimes with small
buttresses. Leaf rachis vigorous, blackish, (51—)
10-30(—40) cm, glabrous and/or scaly to hirsute
and/or tomentose, (2—)4-5(—7)-jugate; leaflets
firmly herbaceous to coriaceous, sessile to 5(—15)
mm stalked, blade (1.7—)2.4-3.0(—3.4) times as
long as wide, widest about the middle or rarely
above, (6—)814-16(—30) by (3-)5'4-6(—8) cm; base
very unequal, acute to subcordate, top shortly
acuminate, the tip obtuse to acute, rarely the top
rounded to acutish; margin entire, rarely serrate;
surfaces glabrous to hirsute, especially underneath
and on the nerves, sometimes underneath with
domatia and/or thin flat greyish (never yellow)
scales. In juvenile specimens and suckers leaves
large, the leaflets serrate, hirsute. Inflorescences
mostly bisexual, paniculate, lateral on leafy and
on slightly older twigs, with i(—0) central 2 catkin
and 2-5 basal ¢ catkins subtended by a caducous
subulate bract 3-5 mm, rarely the basal catkins
also 9. Male catkins (5—)10-18 cm long in all,
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—12(—13), anthers (sub)-
sessile, equal or unequal, + 1 mm long, hirsute,
connective 14-14 mm pointed. Fruiting catkins
(12-)21-40(—60) cm long in all, peduncle 2-1014
cm, vigorous, angular, glabrous and/or greyish-
scaly. Nut (sub)sessile, 3-4 mm diam., hispid,
wing (2-)234-334(-614) cm long with the nut,
(3-)7-9 mm wide, 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
+ 34%-9 mm long.

Distr. India (in the montane Himalayas
between about 75° and 90° E; Assam: Khasia,
Cachar, and Naga Hills, Manipur; E. Bengal:
Tripura, Chittagong), Tibet (MANNING, in /ift.),
China (Yunnan, Kwangsi, n.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 Vooap, /.c., on Mt Rindjani in Lombok;
he also observed it pioneering in mountain
savannahs. The dominance is due to succession
and seral; with Pittosporum, Homalanthus gigan-
teus, Vernonia arborea, Dodonaea, Wendlandia,
etc. belonging to the pioneers of the rain-forest

Dec.

JUGLANDACEAE (Jacobs)

153

Fig. 9. Engelhardia spicata LECHEN. ex BL. in pyrogenous mountain savannah near Sembalun, Mt
Rindjani, Lombok, c. 2000 m (DE VooGp).

which try to invade the pyrogenous Casuarina
junghuhniana stands.

Often deciduous for a short time and then
flowering, not in definite periods.

Vern. Ki hudjan, S, sowo, donglu, J, with various
prefixes and variations.

Note. Easily distinguished from other species
by lacking the golden-yellow glandular scales
on the lower surface of the leaves. If scales are
present, these are flat, greyish and never golden-
yellow.

var. colebrookeana (LINDL. ex WALL.) O. KUNTZE,
Rev. Gen. Pl. 1 (1891) 637, non K. & V. Bijdr.
5 (1900) 169, 172.—E. colebrookeana LINDL. ex
WALL. Pl. As. Rar. 3 (1832) 4, t. 208; BRANDIS,
For. Fl. NW. & C. India (1874) 499; Hook. f.
Fl. Br. Ind. 5 (1888) 596; Dope, FI. Gén. I.-C.
5 (1930) 929; Merr. En. Philip. 2 (1923) 23.—E.
villosa KURz, For. Fl. Burma 2 (1877) 491, incl.
var. integra (according to MANNING in /itt.).—
E. esquirolii Lév. in Fedde Rep. 12 (1913) 507,
cf. REHDER, J. Arn. Arb. 10 (1929) 118.—Fig.
4a-b.

Differs from E. spicata var. spicata in the
following characters: tree mostly comparatively

small; twigs, leaf rachis, and leaflets underneath
densely hirsute, leaflets with rounded top, often
grey-greenish when dry. Fruiting catkins 16—26
cm longin all, 2!4-8 cm stalked, the rachis
densely hirsute.

Distr. India (largely the same area as var.
spicata, but not known from E. Bengal), S. China
(Kwangsi), Hainan, Burma (Kurz), N. and E.
Siam, Indo-China (Annam, Laos); in Malaysia:
Philippines (Luzon, see below).

Notes. The material on which the Malaysian
record is based, PARAISO 25483 and RAMos &
EpDANOo 37939, was identified by Dr MANNING;
I have not seen it.

E. spicata is a very variable species and the
variation of its characters seems to occur more or
less at random and not bound to a geographical
pattern. The reason that the combination of char-
acters described above has received taxonomic
recognition is solely because the specimens re-
presenting this combination are conspicuous. All
these characters, however, were also found to
occur separately, and in other combinations as
well. Therefore I adhere no more taxonomic value
to this variety than as a random assemblage of
marked paramorphs.

154

FLORA MALESIANA

[ser. I, vol. 6!

In East Java, apart from the common form, a
form dominates with densely hairy leaves, which
has incorrectly been referred to var. colebrookeana
by KoorpDEeRS & VALETON. In my opinion this
form represents a local tendency in the population
that originated independently from the paramorphs
on the Asiatic continent. Moreover, a complete
series of intergrades exists between the glabrous
and the hairy form in Java.

5. Engelhardia roxburghiana WALL. Pl. As. Rar.
2 (1831) 85, t. 199, excl. fruct.; BRANDIS, For.
Fl. NW. & C. India (1874) 500.—Juglans
pterococca Roxs. Hort. Beng. (1814) 68, nomen;
Fl. Ind. (ed. Carey) 3 (1832) 631, ‘plerococca’,
descr., excl. fruct.—E. chrysolepis HANCE, Ann.
Sc. Nat. IV, 15 (1861) 227; NAGEL, Bot. Jahrb.
50 (1914) 475; CorNER, Ways. Trees (1940) 333,
f. 117; MeTCALFE, Fl. Fukien 1 (1942) 45; P’EI,
Bot. Bull. Acad. Sin. 1 (1947) 208.—E. wallichiana
[LINDL. ex WALL. Cat. (1831/2) no 4942] C. DC.
Prod. 16, 2 (1864) 141, incl. 6 chrysolepis; HooK.
f. Fl. Br. Ind. 5 (1888) 596; ForRBEs & HEMSL.
J. Linn. Soc. Bot. 26 (1899) 195; NAGEL, Bot.
Jahrb. 50 (1914) 475; GAMBLE, J. As. Soc. Bengal
75, ii (1915) 402; RipL. Fl. Mal. Pen. 3 (1924)
370; Dope, Fl. Gén. I.-C. 5 (1930) 931.—E.
polystachya RADLK. Sitz. Ber. Bayer. Ak. Wiss.
Math.-Phys. KI. 8 (1878) 385, ex descr.; BRANDIS,
Ind. Trees (1906) 620; KANJ. c.s. Fl. Assam 4
(1940) 301.—E. pterococca O.K. Rev. Gen. Pl. 2
(1891) 637, quoad nomen et var. « roxburghiana
O.K.—E. spicata var. formosana Hay. Fl. Mont.
Form. (1908) 199.—E. formosana Hay. Ic. PI.
Form. 6 (1916) 61; KANEH. Form. Trees (1936)
80, f. 35.—E. fenzelii MERR. Lingn. Sc. J. 7 (1931)
300, ex descr.; METCALFE, FI. Fukien 1 (1942) 45.

Tree -+- 5-20 m, sometimes with low, rounded
buttresses. Innovations bronze-coloured with
golden yellow glandular scales which become
brown when older. Leaf rachis 514-14 cm, as the
twig glabrous to sometimes densely brown
puberulous, 2—4-jugate; leaflets coriaceous, on a
stalk 6-12 mm forming mostly a distinct joint
with the rachis; blade (2.4—)2.8-3.6(-4.0) times
as long as wide, widest mostly below the middle,
sometimes halfway, (8—)10—-16(—23) by (2-)314-5
(-714) cm; base very unequal, acute, often basis-
copically somewhat decurrent, top narrowed,
gradually bluntly acuminate; midrib narrowly
raised above; margin entire, surfaces in dry state
never even-coloured, but with dull shades of
greenish, greyish, brownish, or blackish, glabrous
but underneath yellow to brown glandular-scaly.
Probably sometimes dioecious, but mostly the
inflorescences bisexual, paniculate, terminal on

normal or slightly weaker lateral twigs, 2-4 cm
peduncled, with central 2 catkin (which may be
wanting) and 2-6 basal ¢ catkins subtended by
early caducuous bracts; rarely the 2 catkin
solitary. Flowers scaly. Male catkins 8-12 cm
long in all, 14—2 cm stalked; rachis slender, scaly,
not hairy; bract + 2 by 1 mm, obscurely 3-
lobed at the top, perianth lobes 4, not reduced,
strongly cucullate, 1-114 cm diam., with mem-
branous margin; stamens (8—)12, inserted with
(2-)3 at the base of each perianth lobe, filament
and anther both 144 mm long, in some stamens
reduced to 14 mm, glabrous. Fruiting catkins
12-22 cm long in all, peduncle 2-514 cm, slender,
rachis tomentose, sometimes also scaly. Nut 4-8
mm stalked, transversely ellipsoid, 4-5 mm diam.,
scaly, not hairy; wing (3—)5—5'4 cm long with the
nut, 0.7—1.4(-1.8) cm wide, hardly connate with
the nut, adaxial lobe almost none; perianth lobes
very equal and regular, 1-2 mm long, on top of
the nut together enclosing the 4 sessile stigmas.

Distr. India (Assam), S. China (all provinces
S of about 30° N lat.), Hainan, Formosa, Indo-
China (Tonkin, Laos, Annam), in Malaysia:
Sumatra (in the N. part especially, S to Bencoolen),
Malay Peninsula (Kedah, Perak, Pahang, Penang),
Borneo (Brunei and Kinabalu region). Everywhere
more or less rare.

Ecol. Primary forests in hilly country between
about 700 and 1500 m alt. Flowers and fruits
not at definite times.

Uses. In Sumatra said to be a fish-poison.

Note. WALLICH’s treatment of this species com-
prises three elements: a two-line diagnosis from
LINDLEY’s MS, an extensive analysis after Rox-
BURGH’S MS (the latter was to be published again,
under the name Juglans plerococca, in 1832), and
a plate which is a copy of ROXBURGH’s drawing.
The diagnosis that WALLICH took from LINDLEY
refers, as HooxkerR has already pointed out, to
E. spicata and the same is true for that part of
ROXBURGH’s plate and description where the fruit
is given as hairy. The remaining part of Rox-
BURGH’s plate and description clearly refer to
E. roxburghiana in the present sense. Since only a
minor part of WALLICH’s mixtum belongs to E.
spicata, it is thought correct to accept WALLICH’s
E. roxburghiana pro majore parte.

Excluded

Engelhardia selanica Bu. Fl. Jav. Jugl. (1829)
8 = Shorea selanica BL. Mus. Bot. 2 (1852) 33
(Dipterocarpaceae).

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FLORA MALESIANA

UNDER THE AUSPICES

OF

THE BOTANIC GARDENS OF INDONESIA, BOGOR,

AND THE RIJKSHERBARIUM, LEYDEN,

SPONSORED BY THE FOUNDATION
FOR PURE SCIENTIFIC RESEARCH

*

SCIENTIFIC COMMUNICATIONS
concerning Flora Malesiana should be addressed to the General Editor,
Dr C. G. G. J. VAN STEENIS, Rijksherbarium, Leyden, Holland

OK

SUBSCRIPTIONS

should exclusively be directed to N.V. DE ERVEN P. NOORDHOFF,
P.O. Box 39, Groningen, Holland

Phytolaccaceae
Pittosporaceae
Plumbaginaceae
Podostemaceae
Polemoniaceae
Pontederiaceae
Proteaceae
Punicaceae
Restionaceae
Rhizophoraceae
Salicaceae
Salvadoraceae .

Sarcospermaceae .

Saururaceae

Scyphostegiaceae . ;
4: cay 513

Sonneratiaceae
Sparganiaceae
Sphenocleaceae
Stackhousiaceae
Staphyleaceae
Stylidiaceae
Styracaceae
Thymelaeaceae
Trigoniaceae .
Turneraceae
Typhaceae
Umbelliferae .
Valerianaceae .
Xyridaceae
Zygophyllaceae

Volume 1. Cyclopaedia of collectors & collections. 1950. pp. clii. + 639.
Volume 2. Malaysian vegetation is Hebe, Sen eaiges
Volume 3. Malaysian plant geography
Volume 4. General chapters and revisions. 1948 - 1954. pp. ‘ccix + 631 ;
Volume 5. Bibliography, specific delimitation, & revisions. 1955-1958 ~
For sale only , Not for exchange
INDEX TO REVISED FAMILIES
Aceraceae . 4: 3, 592 Dioscoreaceae 4: 293
Actinidiaceae s.str. . 4: 37 Dipsacaceae 4: 290
Aizoaceae . . » 4: 267 Droseraceae 4: 377
Alismataceae. . . 5: 317 Elatinaceae : 4: 203
Amaranthaceae . 4: 69, 593 Erythroxylaceae . 5: 543
Ancistrocladaceae . 4: 8 Ficoidaceae 4: 267
Aponogetonaceae . 4: 11 Filacourtiaceae eee
Basellaceae . . . 5: 300 Filagellariaceae 4: 245
Batidaceae . . . 5: 414 Gmetaceae. 4: 336
Betulaceae . . . 5: 207 #Gonystylaceae ; 4: 349
Bixaceae . . . . 42239 Goodeniacede |... ~5: 335
Burmanniaceae 4: 13, 592 Haemodoraceae . 5: 111
Burseraceae . . . 5: 209 #Hamamelidaceae. . 5: 363
Butomaceae . . . 5: 118 # Hydrocaryaceae. . 4: 43
Callitrichaceae . . 4: 251 ##Hydrocharitaceae 5: 381
Campanulaceae . 6:.107 | Hydrophyllaceae 4: 207
Cannabinaceae 4: 223 Juglandaceae . 6: 143
Capparidaceae . 6: | 61 Juncaceae 4: 210
Caprifoliaceae . 4: W715, 598 Juncaginaceae 4:* 57
Centrolepidaceae . ,5: 421 Malpighiaceae §3,125
Ceratophyllaceae. . | 4:: 41 Martyniaceae 4: 216
Chenopodiaceae . 4: 99, 594 Molluginaceae 4: 267
Cochlospermaceae 4: :* 61 Moringaceae . 4: 45
Combretaceae “4: ~ 533 Myoporaceae 4: 265
Connaraceae . - 5:495 Myricaceae _4: 277
Convolvulaceae . 4: 338,599 Nyssaceae . . . 4: 29
Corynocarpaceae . 4:'262 Papaveraceae she a See
Crassulaceae . 4: 197 Pedaliaceae . . .* [433216
Datiscaceae . . ... 4: 1382 Pentaphragmataceae . =, 4: 517
Dichapetalaceae . . '5:'305 Pentaphylacaceae .* 5: 121
Dilleniaceae 4: 141 Philydraceae . . . 4: ..5

PRINTED IN THE NETHERLANDS BY
N.V. DISKSTRA’S DRUKKERIJ V.H. BOEKDRUKKERIJ GEBR. HOITSEMA - GRONINGEN

4: 229
5: 345
4: 107
4: 65
4: 195
4: 255
5: 147
4: 226
5: 416
5: 429
5: 107
4: 225
4: 32
4: 47
5: 297

4: 233
4AP327

4: 35

6: 49

4: 529

4: 49 —
“4: 349, 6: 1
ee a
4: 235

. 4: 243
4: 113, 595
4: 253

| 4: 366, 598

4: 64