LAV Ran SA qn af 5
alla MAY 10 1963

mm FLORA
MALESIANA

UNDER THE AUSPICES OF
THE BOTANIC GARDENS OF INDONESIA, BOGOR,
AND THE RIJKSHERBARIUM, LEYDEN,
SPONSORED BY THE FOUNDATION
FOR PURE SCIENTIFIC RESEARCH

NAJADACEAE (W.J.J.O. de Wilde, Leyden)

Within the Helobieae there has been a great deal of controversial opinion about the evaluation
of the genera belonging to the Potamogetonaceae, among which Najas finds by almost unanimous
opinion its closest relatives. Generally Najas has been accepted to represent a separate monotypic
family on account of the basal ovule and the structure of the anther (with a thin, tight, 2-lipped
envelope and apically escaping pollen). The closest allied genus among Potamogetonaceae seems
to be Zannichellia, which is by HUTCHINSON (1934) accepted as a separate family, Zannichellia-
ceae, put together with Najadaceae in his order Najadales.

Within the Helobieae some authors accept the structure of Najadaceae as primitive, notably
CAMPBELL (1897) and RENDLE (1930), but others find it a derived, advanced state within the
order, cf. HUTCHINSON (1934) and LAWRENCE (1951).

An excellent concise account of opinions is given by LAWRENCE (Taxonomy of Vascular
Plants, 1951, 375-378).

NAJAS

LINnNE, Gen. Pl. ed. 5 (1754) 445; Sp. Pl. (1753) 1015; WitLp. Mém. Ac. R. Sc.
Berl. (1798) 85; A. BRAUN, J. Bot. 2 (1864) 274-279, 8 fig.; P. MAGNUS, Beitr. z.
Kenntn. Gatt. Najas (1870) 1-63, t. 1-8; C. BAtLey, J. Bot. 22 (1884) 305, fig.
1-89; P. MAGNus in E. & P. Pfl. Fam. 2, 1 (1889) 214-218; Ber. Deut. Bot. Ges.
12 (1894) 214, t. 11; K. Scu. in Mart. Fl. Bras. 3, 3 (1894) 715-734, t. 123-124;
RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 379-444, t. 39-42; Pfl. Reich Heft 7
(1901); DE WiLpe, Act. Bot. Neerl. 10 (1961) 164, fig. 1-9.—Caulinia WILLD.
Mem. Ac. R. Sc. Berl. (1798) 87, t. i; cf. DE WILDE, Willdenowia 2 (1960) 595—
597._Fig. 1-8.

(Mostly) fragile, slender, submersed annuals, rooting from the base and
lower nodes. Stems often forked, terete, sometimes provided with teeth (subg.
Najas). Leaves sessile, in pseudowhorls of 3 or more, linear, sometimes more or
less subulate or linguiform, l-nerved; midrib bordered on both sides, especially
clearly visible in the lower part of fullgrown leaves, by a mostly distinctly trans-
versely septated intercellular space, here called ‘cavity’. Sheath more or less auri-
cled, or truncate, or decurrent; upper part of the sheath and margin of the blade,
sometimes also the dorsal surface of the midrib, spiny-dentate or spiny to various
degree; spine cells yellowish-brown, ascending. Sheath with 2 axillary intravaginal
scales (see fig. 2g-1). Plants mostly monoecious. Flowers small, unisexual, soli-
tary or sometimes 2-4 together, at the very base of an (sometimes very short)
axillary shoot, each often enclosed in a membranous (sub)sessile spathe (9 fl. in
Mal. spp. espathaceous). Spathe tapering to the top, or constricted into a cylindri-
cal neck, the edge mostly with some erect spines. 3 Flowers consisting of one
subsessile or stalked anther, which is tightly enveloped by a membranous ‘perianth’
apically mostly produced into 2 more or less distinct ‘lobes’; anther l- or 4-
(rarely 2-)celled, basifixed, + pore-like dehiscing at the top; ‘pedicel’ mostly
considerably elongating just before anthesis. 2° Flowers (in Mal.): ovary subsessile,
naked, elliptical to oblong, |-celled, style cylindrical, with (1—)2(-4) linear, often
unequal stigmas. Ovule one, basal, erect, anatropous; integuments two. Fruits
elliptical-oblong (to ovate); style and stigmas persistent; pericarp very thin in
the herbarium, not dehiscent, tearing away at the base or dwindling by rotting.

(157)

158 _ FLORA MALESIANA [sere Ty vole?

Seeds oblong (to ovate), with a conspicuous raphe and a hard, (in Mal. spp.)
distinctly areolate, mostly brownish testa; embryo straight; hypocotyl and radicle

large, plumule well developed; cotyledon terminal, blunt. No endosperm.

Distr. About 35 spp., from the tropical to the temperate parts of the world.

Ecol. Mostly in shallow, stagnant or sluggish waters, ditches, rice-fields, small streams in forest,
pools, ponds, swamps, mostly in freshwater, but also in brackish water near coasts and inland, a few
species even preferring this habitat. N. malesiana is the only species which is preferring streams and
ponds in forests. The others are all found in eutroph waters. According to SUNIER (Treubia 2, 1922, 190)
N. indica is usually found in freshwater sites but occurs in the brackish fish-ponds near Djakarta with
a salinity of up to 32/59. N. browniana seems almost confined to coastal brackish water and is found
inland only in the saline water of the Kuwu mudwells, SE of Semarang, as the only aquatic represented
there. Further N. marina var. zollingeri is confined to the alkaline waters of Batur Lake, in Bali, with a
rather high content of dissolved minerals. Stems and leaves of Najas are not seldom found encrusted
with calcium which adds to their brittle nature. Some species are resistant to rather hot water, e.g.
N. tenuifolia ssp. pseudograminea var. pseudograminea which is found in the craterlake of Mt Kelud,
Central Java, on black mud in water of 60° C; this water contains the usual large amount of minerals
in solution common in craterlakes, further sulphates, sulphides, Ca and Mg compounds, efc. BRANDIS
recorded N. graminea from a hot spring in Burma in water of 92° F, and the first record of it in England
was from hot water emitted by a factory (J. Bot. 22, 1884, 326). HERMANN recorded N. marina from
hot springs in America (Leafl. West. Bot. 1, 1935, 182).

As to depth Najas species are adapted to shallow waters; during the German Limnological Sunda
Expedition the greatest depth at which Najas occurred was fixed at c. 5 m, but these specimens proved
all to be sterile (cf. VAN STEENIS, Arch. Hydrobiol. Suppl 11, 1932, 240, 271).

The relatively rare occurrence of N. marina in Malaysiaand in many parts of its almost ubiquitous range
as well, points to a sensitive and selective ecology rather than to random chance dispersal by aquatic
birds, especially in connection with its raciation into geographical varieties of sometimes restricted
areas. The phenomenon is also found in Hydrocharitaceae and Alismataceae (cf. Fl. Mal. I, 5, 1957, 317).
In checking localities it has appeared that generally one lake seems to have only one species; N. indica
in Toba, Ranau, Situ Bagendit, Singkang, and Gorontalo lakes, N. graminea var. graminea in Situ Gu-
nung, Matana & Towuti lakes, N. tenuifolia ssp. pseudograminea var. pseudograminea in Kelud and
Rawa Bening (Kediri) lakes, and the same holds for N. browniana and the three varieties of N. marina
which are all found in one lake only, with no other species present. The only exception is Tondano
lake, in NE. Celebes in which according to the data on the labels three species have been collected,
viz N. graminea var. graminea, N. indica, and N. tenuifolia ssp. pseudograminea var. celebica.

As to altitude most species and localities prefer the lowland areas, but N. marina goes up to 1000 m,
and several others are found in Toba Lake, 900 m, Tondano Lake, 700 m, Kelud Lake, 1000 m, the
highest altitude recorded being 1400 m. In the Himalayas Malaysian species are recorded to over
2000 m.

As to climate none of the species shuns the seasonal areas, and some seem often to be more abundant
in them than in everwet areas which may again point to preference for eutroph water.

Flower biology. Pollination takes place in submerged condition, but there are no detailed actual
observations in Malaysia. The stalk of the anther elongates at maturity pushing the anther, either la-
terally or terminally, beyond the spathe (if there is any), then sometimes recurving. The pollen grains
are either globular or ellipsoid and contain much starch; an exine is absent; they escape from the top
of the anther through a hole between the two lobes which diverge. MAGNus (in E. & P. Pfl. Fam. 2,
1, p. 216) stated that the pollen of N. marina germinates before leaving the anther. This has also been
observed in other species by MIKI (Tokyo Bot. Mag. 49, 1935, 774, fig. 6 G, P). The pollen tube facilitates
of course floating and will increase the chance to fasten to a stigmatic arm of the 9 flower. Through this
mechanical way, without aid of water animals, pollination must take place. As Najas mostly grows gre-
gariously the distance between ¢ and 9 flowers is not large, and small movements in the water may effect
contact of pollen and stigmas.

Though in a few instances two species are growing in the same locality and are found mixed in collec-
tions they retain their specific characters well and I have not found any trace of hybridization.

Morph. The leaves of Najas are essentially subopposite, but appear to be placed in pseudowhorls
of 3 or more. This phyllotaxis originates as follows: The sheath of the not fully amplexicaulous lower
leaf of each pair imbricately overlaps the fully amplexicaulous sheath of the upper one. Only the lower
leaf bears an axillary bud growing into a lateral branch. The first internode of this lateral branch is ex-
tremely short and its first node becomes in this way almost sessile; it bears a leaf pair, but of this pair
only the upper one is developed as a real leaf which forms together with the 2 leaves of the main stem the
pseudowhorl of three. The lower leaf of the first node is very much reduced; according to MAGNUS
(1870, 1894) it is, in vegetative shoots, represented by a very small scale bearing a vegetative bud in its
axil, whereas in fertile lateral shoots a flower occupies the place of this scale. Fig. 1b-c.

The flower would, therefore, be homologous with a scale plus its axillary bud. It has appeared that
a fully developed flower of many Najas species is surrounded by a spathe (fig. 2d, 2f) and from the

Dec. 1962] NAJADACEAE (de Wilde) 159

Fig. 1. Najas tenuifolia ssp. pseudograminea (W. Kocu) DE WILDE var. pseudograminea. a. Habit (arrows
pointing to an open and 2 closed ¢ flowers respectively), * 5, b. situation at a node in which the lowest
leaf has been recurved and one flower removed, 15, c. diagram with identical signatures as in b:—1.
main axis of stem, 2. lower leaf of a pair, 3. upper leaf of the pair, 6. axis of shoot in axil of lower leaf,
bearing: 4. flower (removed here) either j and enveloped by spathal bract or 9 and naked, representing
lower fertile leaf of lateral shoot, and 5. leaf opposite 4, representing upper leaf of the lowest pair of
lateral shoot, 6—7—8-9-10 a repetition on the next node of the lateral shoot of 1-2-3-4—5 on the node
of the main axis. This drawing represents an average situation; it can be more complicated if the lower
leaf pairs on axillary shoots are reduced to mere bracts. Sometimes 2, 3, and 5 form a pseudowhorl of
three leaves which is always represented on sterile nodes; in this figure 2, 3, 5, 7, 8, and 10 form a pseudo-
whorl of six leaves (BACKER 7903).

homology it would appear that the spathe is not representing a perianth but a tubular reduced leaf, and
must be understood as homologous with the scale found in its place in vegetative shoots. I have been
able to find an almost complete series of transitions between an open scale and a fully closed bottle-
shaped spathe (pe WILDE, 1961) and believe to have proved this homology.

In a few cases there are more flowers together with one whorl at a node. As these flowers are always
different in size, hence in age, the situation can be explained in that more than one node of the lateral
shoot is reduced in a similar way as the first one. This follows also from the fact that the number of
leaves of the pseudowhorl may be up to 5 or more in exceptional cases.

In the 3 flower the anther is tightly surrounded by a very thin membrane which, in N. marina, consists
of only 2 cell layers. According to MAGNus (1870, p. 23; 1894, p. 216) this envelope is practically ad-

160 FLORA MALESIANA [ser: 1) volisG2

nate to the anther and ends in two more or less thickened lobes. Whether this would represent a ‘pe-
rianth’ seems doubtful and it is not clear whether the stalk under the anther is a pedicel or a filament.
Fig. 2c and 2e.

The pericarp of the fruit is almost pellucid and very thin in dried specimens, clinging very closely
to the seed. According to BAILEY (1884, p. 322) and RENDLE (1899, p. 385) it would in fresh specimens
be “‘succulent” in N. marina and N. graminea. It seems gradually to decay on the plant or separates from
the seed by breaking irregularly away at the base.

Anat. As usual in water plants the structure of stem and leaf is very simple. In most species the
epidermal cells of the stem closely resemble those of the underlying cells of the cortex, only in N. marina
the epidermal cells are distinctly smaller. In a cross section of the stem the cortex appears to consist
of two layers of parenchymatic cells; the outer one is always more cell layers thick, the inner one is
also mostly a few cell layers thick but may consist of only one cell layer (the endodermis). These two
cortical layers are connected by radial septa mostly one cell thick, which separate large intercellular
cavities. The stele is small-celled and has a central cavity. Fig. 2a—b.

Fig. 2. Details of Najas. a. Partial cross-section of stem of N. marina var. zollingeri RENDLE, b. ditto
of N. kurziana RENDLE, c. naked ¢ flower, anther 1-celled, of N. malesiana DE Wi bE, d. spathed 3
flower, anther |-celled, of N. browniana RENDLE, e. naked 3 flower, anther 4-celled, of N. graminea DEL.
var. graminea, ‘perianth lobes’ extraordinarily conspicuous, f. spathed 3 flower, anther 4-celled, of
N. tenuifolia ssp. pseudograminea (W. KOCH) DE WILDE var. pseudograminea, g. half a sheath of N.
marina var. sumatrana DE WILDE, no auricle, squamulae intravaginalis irregularly quadrangular, A. leaf
base of N. marina L. var. marina, ditto, i. half a sheath of N. tenuifolia ssp. pseudograminea var. cele-
bica, (KOORD.) DE WILDE, arrows indicate how and where length and width of auricles and sheath have
been measured, squamulae intravaginalis linear, j. ditto of N. graminea DEL. var. graminea, k. ditto
of N. malesiana DE WILDE, I. ditto of N. indica (a-—b X 50, c-f X 25, g X 10,h X 5, i-l X 10; a RUTTNER
313, b VAN STEENIS 18226, c MEIJER 5772 (type), d HORSFIELD s.n., e EYMA 3329, f THUNG s.n. (isotype),

g JACOBSON s.n. (type), A JENSEN 298, i KoorDERS 17347 (type), / SUNIER S.n., k CLEMENS 9699, / LORZING
11067).

Dec. 1962] NAJADACEAE (de Wilde) ial

The leaf shows in cross section a midrib consisting of small cells covered above and beneath by one
to two layers of larger cells and is on both lateral sides accompanied by an intercellular cavity. The
blade proper consists of only two cell layers (upper and under epidermis), but has additional subepider-
mal cell layers in N. marina.

Taxon. The genus is very coherent in structure and I have followed A. BRAUN (1864) and K. ScHu-
MANN (1894) in not accepting subgenera, but merely two sections; sect. Najas contains only one species,
N. marina; the other species are accommodated in sect. Caulinia. RENDLE found it useful to subdivide
his subgenus Caulinia into 4 ‘sections’ which were mostly defined according to the presence or absence
of the spathe in one or both sexes. I have found (1961) that this character is variable and I have con-

Uses. According to SUNIER (Treubia 2, 1922, 222) Najas, which is profusely found in brackish fish-
ponds (tambaks) near Djakarta, may be important as a source of food for fish. From the Toba Lake it
is reported that Najas can be utilized as pig food, a use it shares with many other water plants (BACKER,
Teysmannia 1911, p. 514).

Notes. As the vegetative characters are very uniform and show besides a rather large variability
within one species, it has been impossible to frame a key to identify sterile material. Collectors should
try to select fertile material either recognizable by the presence of fruit in the lower axils, or the presence
of a yellowish or reddish tinged anther in the higher axils. In some species the sexual organs are very
small. Dissection and examination of boiled herbarium specimens is a delicate, time-consuming work.

Attention is called to the fact that sometimes more than one species is represented in a single habitat.

Measurements of leaves as found in descriptions have been made as is shown in fig. 21.

Thanks are due to Dr. P. W. LEENHOUTS whose help and advice I have appreciated.

KEY TO THE SPECIES

1. Seed (314-)4-414(-714) by 2-214(-3) mm. Intravaginal scales up to 114 times as long as broad,
fleshy (fig. 2g-h). Cells of epidermis distinctly smaller than underlying cortex cells. Neck of the
spathe (only ind fl.!) up to twice as long as broad, the margin without or with a few very minute spines.
Internodes sometimes dentate. Sect. NAJAS . . . .1, N. marina

1. Seed less than 314 mm long, if nearly 344 mm then only. @ 2/3(-3 4) mm broad. Intravaginal scales
lanceolate to filiform, thin (fig. 2i-l). Cells of epidermis not distinctly smaller than underlying cortex
cells. Neck of the spathe more than twice as long as broad, the margin with 3-9 distinct spines. Inter-
nodes smooth. Sect. CAULINIA.

2. d Flower enclosed in a spathe.

3. Mature anther 1I-celled, 1/3-134 mm. Spathe up to 2 mm.
4. Seeds 214-3 mm “On areoles of testa much broader than long (ladder-like). Leaves often coarsely
dentate 4 oe 2eaNs minor.
4. Seeds up to 2 mm long; areoles of testa ‘about ‘isodiametrical. Leaves not coarsely dentate.
5. Seeds 1-1.2 by 1/3-0.4 mm. Spathe 0.9-1.2 mm. Blade with (30—)50—60 spines on each side.

Auricles mostly long-triangular, 4,-0.8 mm long . . 3. N. kurziana
5. Seeds 114-2 by 14-34 mm. Spathe 114-2 mm long. Blade with 12 zy spines on each side. Auri-
cles mostly short triangular, up to 14 mm long. . . 4.N. browniana

3. Anther 4-celled, when mature 1'14—2'4(-2%) mm. Spathe 2 33, mm long.

6. Auricles (0—)0.2-0.6(—1) mm long, mostly broader than long, the inner edge without spines.
Leaf-blade flat to nearly terete or triangular in cross-section, with 1—-30(-40) spines on each side,
sometimes moreover with one or more dorsal spines. Fruits solitary. Seeds ¥4—-0.9 mm broad.
Cavities at least half as wide as the leaf-halves, their outer aes not brown. Marginal spines well
visible with the naked eye . ; 5. N. indica

6. Auricles 0.6-1.6(-3) mm long, longer than broad, the i inner edge with (0-)1- 4 spines. Leaf-
blade flat, with 15—60(—70) spines on each side, dorsally always spineless. Fruits solitary or
2(—3) together. Seeds up to 34 mm broad; cavities variable in width, from very narrow to nearly
reaching the margin. Marginal spines (mostly) just visible with the naked eye . 6. N. tenuifolia

2. § Flower not enclosed in a spathe.

7. Auricles less than twice as long as broad, up to 1.1 mm long. Anther 1-celled, when mature 0.6-1
by 0.15-0.3 mm. Seeds 1-1.8 by (0.35—)0.4—0.6 mm; testa with 16—22(—26) longitudinal rows of
24-30 areoles. Leaves 114-214 cm, with Ce ae ace spines along each side. Plant slender, up
to’ DS emi high) =... . 7. N. malesiana

7. Auricles at least twice as long as ; broad, 1- 5h mm long. Anther 2: or ‘4. celled, when mature 1.2—2
by 0.4-1 mm. Seeds 1.8-2.7 by 0.6-0.9 mm (if less than 2 mm long then broader than 0.6 mm and
with more than 25 longitudinal rows of areoles). Testa with 20-34 longitudinal rows of 30-45
areoles. Leaves 134—6 cm, with 40-185 spines along each side. Plant slender to rather coarse, up
to 50(—75) cm high Pair ee Pre anythin es sro ets: Gre Metisia, Gok or) sy) Ae ee SOP New eraminea

162 FLORA MALESIANA [ser. I, vol. 6?

1. Section Najas

Sect. Eunajas A. BRAUN, J. Bot. 2 (1864) 275; Ascuers. FI. Prov. Brand. 1 (1864)
669 (status not mentioned).—Subg. Eunajas ASCHERS. ex RENDLE, Trans. Linn.
Soc. II, Bot. 5 (1899) 389; Pfl. Reich Heft 7 (1901) 7.

Mostly coarse plants. Epidermis distinctly differentiated by smaller cells from
the underlying cortex. Internodes often spiny. Leaves up to 12(-15, very rarely
more) times as long as broad, mostly fleshy, coarsely dentate along the margin
and mostly on (or near) either side of the lower portion of the midrib; marginal
teeth up to c. 15 (very rarely more) on each side. Sheath with or without incon-
spicuous auricles. Intravaginal scales less than twice as long as broad, blunt, often
asymmetrical, fleshy (fig. 2g-h). Dioecious. 3 Flowers: spathe constricted into a
short (up to twice as long as broad), (sub)cylindrical neck, the margin without,
or with a few, very minute spines; anther 4-celled. 2 Flowers espathaceous, bearing
(2-)3(-4) stigmas. Seed 3-714 by 2-3 mm; testa consisting of more than three
layers of hardened cells. Areoles often unequal in size, not distinctly arranged in

longitudinal rows.
Distr. Monotypic, ubiquist.

1. Najas marina Linng, Sp. Pl. (1753) 1015, incl.
var. B ety; K. Scu. in Mart. Fl. Bras. 3, 3 (1894)
723, t. 123 f. 2; RENDLE, Trans. Linn. Soc. II, Bot.
5 (1899) 389, t. 39 f. 1-30; ibid. (1900) 437; Pfl.
Reich Heft 7 (1907) 7, f. 1 A-C, 3 A-Q.— N. major
ALL. Fl. Ped. 2 (1785) 221; BENTH. FI. Austr. 7
(1878) 181; Hoox. f. FL. Br. Ind. 6 (1893) 569;
Koorp. Exk. Fl. 1 (1911) 91; Miki, Bot. Mag.
Tokyo 49 (1935) t. 8 f. E-I.— Fig. 2a, g-h, 3.

Plant up to c. 70 cm, lower internodes 3-8 cm
by 0.9-2.6 mm, glabrous to very spiny. Leaves
441, cm by 114-314 mm, flat, sometimes
triangular in section, almost always fleshy; tip
acute to blunt (blade c. 3 mm below the tip 1.8-3
mm wide); margin on either side with 5—12(—-40)
spiny teeth; dorsal surface sometimes without,
often with 1—6(—40) spines; teeth up to long trian-
gular or conical; up to as long as the width of the
blade; spine cells 0.05—0.15 mm, brownish; midrib
c. 0.1 as wide as the blade, cavities occupying up
to 34 of the leaf-width, septa indistinct. Sheath
on either side (without or) with 1—5(—10), mostly
inconspicuous spines, 2-614 by 3-9 mm; auricles
absent or up to c. 0.2 mm long. Flowers solitary.
Spathe of 3 flower (in Mal. unknown) c. 444 mm
long (Kashmir specimens); neck about cylindrical,
0.3—-0.5 mm, edge somewhat lobed, sometimes with
(23) spines; anther 4-celled, c. 344 by 214 mm; ‘pe-
rianth lobes’ rather indistinct; ‘pedicel’ c. 0.5 mm,
in anthesis longer. Spathe in 2 flower absent; & fl.
1.9-3.8 mm, ovary 0.8—1.2 by 0.6-0.8 mm, style
0.2-0.8 mm, with 2-4 stigmas 0.9-2 mm. Seeds
(asymmetrical) elliptical to ovate, narrowing to the
tip, somewhat compressed, (314—)4-4.4(-7) by
(2—)2.4-2.6 mm; testa shiny with rather irregularly
arranged + isodiametrical, 5-6 angular, unequal-
sized areoles.

This ubiquist species has been split up into a
large number of varieties, three of which occur in
Malaysia.

KEY TO THE VARIETIES
1. Leaves 15—25 times as long as broad, flat and
thin, (when dry) more or less pellucid. Stem
densely dentate; leaf margin with 30-40 slender
spiny teeth, 34-1 mm long, on either side. Style
narrow, longer than 14 mm var. sumatrana
1. Leaves less than 15 times as long as broad,
somewhat fleshy, opaque. Stem smooth; leaf
margin with up to 12 broad triangular (conical)
spiny teeth on either side. Style thick, up to

Y mm long.

2. Leaves 3-414 cm by 3-314 mm; widest in the
middle of the blade. Leaf margins each with

(5—)8-10(-12) teeth, on the dorsal surface

rarely with 1 tooth. Leaf sheath (4-)5-6% by

7-9 mm. SU ele Ps var. zollingeri
. Leaves up to 3 cm by 3 mm, mostly broadest

above the middle. Leaf margins each with

(2-)4-7 teeth; dorsal surface with (0—)2-6

teeth. Leaf sheath up to 3.7 by 5.6 mm.

var. Marina
var. marina.—N. major B augustifolia et e interme-
dia A. Braun, J. Bot. 2 (1864) 275, 276.—N.
marina yar. angustifolia et intermedia RENDLE,
Trans. Linn. Soc. II, Bot. 5 (1899) 395, 439,
t. 39 f. 13, 28, 14 et 29; ibid. (1900) 439; Pfi.
Reich Heft 7 (1901) 8, f. 3 E, O, F, P.—N. inter-
media Gorski, Eichw. Nat. Skizze Lithauen
(1830) 126, non vidi.—Fig. 2h.

Plant up to 30 cm, lower internodes up to 6 cm
by (1.1—)114-2(-2'4) mm. Leaves linear or slightly
spathulate, (14—)1—214(-3) cm by 114-2% mm, c.
3 mm below the tip 2-3 mm wide, flat to triangular
in section, somewhat fleshy; apex blunt, rarely
acute; margin on either side with (2—)4~—7 coarse
spiny teeth, up to the lower 7 mm sometimes with-
out teeth; dorsal surface mostly with 1-6 spiny
teeth, rarely unarmed; teeth (long) triangular to
conical, up to as long as the width of the blade;
spine-cells c. 0.05 mm or smaller, brown; cavities

tN

Dec. 1962]

NAJADACEAE (de Wilde)

163

often indistinct, occupying 14-34 of the leaf-
width. Sheath on either side with 1-2 incon-
spicuous (< 0.05 mm) spines, 2—3.7 by 3—5.6 mm;
auricles absent. Spathe of 3 flower c. 1.8 mm, tip
with c. 3 small hyaline spines c. 0.1 mm; anther
4-celled, elliptic c. 1 by 0.7 mm; ‘pedicel’ c. 0.5
mm. 2 Flower 1.9-2.7 mm; ovary 0.7—1 by 0.7-0.8
mm; style 0.2-0.5 mm by 0.25-0.3 mm, with
2-3 stigmas 0.9-1.3 mm. No seeds present.

Distr. According to RENDLE this variety is
very widely distributed, from NW. Europe to West
Australia, Japan, and the Sandwich Is, in Malay-
sia: South Moluccas (Kei Is). In this area there are
wide gaps. Fig. 4.

Note. The anther I have observed in this
material, the only one I saw of N. marina in Malay-
sia, is far too small for the species and is in my
opinion not normally developed. This deserves
further field study.

Fig. 3. Najas marina L. var. zollingeri RENDLE.
a. Habit, = 3/5, b. 2 flower, c. fruits, d. seeds, * 3.

var. zollingeri RENDLE, Trans. Linn. Soc. II, Bot. 5
(1900) 438-439; Pfl. Reich Heft 7 (1901) 7;
Steen. Arch. Hydrobiol. Suppl. 11 (1932) 271,
272, f. 2; RUTTNER, /.c. 374, tab. II.—Fig. 2a, 3.

Up to c. 70 cm, lower internodes up to c. 8 cm
by 1.2-2.6 mm, glabrous. Leaves linear-lan-
ceolate, 314-414 cm by 2.9-3.6 mm, c. 6 mm below

the tip c. 2.5 mm wide, flat, fleshy; apex acute,
rarely blunt; margin on either side with (S—)8-10
(-12) coarse, spiny teeth, up to the lower 3 mm
sometimes without teeth; dorsal surface without
or with 1 tooth; teeth triangular, up to as long
as 1/3 of the width of the blade; all spine-cells
c. 0.1 mm, brown; cavities as wide as to twice as
broad as the midrib. Sheath on either side with
2-5 inconspicuous spines, 5—6(—61) by 7-9 mm;
auricles absent or up to 0.2 mm long. 3 Flowers
not seen. 2 Flowers 2.5—3.8 mm; ovary 0.8—1.2
by 0.6-0.8 mm; style 0.2-0.5 by 0.2-0.4 mm, with
2-4 stigmas 1.6—2 mm. Seeds 44.3 by 2.4-2.6 mm.

Distr. Malaysia: Lesser Sunda Islands (Bali:
Batur Lake). Fig. 4.

Ecol. Gregarious along the sandy shore, at c.
1030 m. The water of Lake Batur is alkaline and
contains a high percentage of dissolved minerals.

a a ; |

ere n

Fig. 4. Malaysian localities of Najas marina var.

marina (QO), var. sumatrana DE WILDE (+),

var. zollingeri RENDLE (x), N. browniana RENDLE
(e), N. kurziana RENDLE (#9).

var. sumatrana DE WILDE, Act. Bot. Neerl. 10
(1961) 169.—Fig. 2g.

Up to 70 cm, lower internodes 3-7 cm, by
0.9-1.4 mm, densely set with prickles (15-20 per
2 mm stem), more densely on (and just below)
the nodes. Leaves 3144-444 cm by 2-214 mm,
flat and thin, apex acute (to slightly obtuse), c.
3 mm below the tip c. 1.8 mm wide. Margin on
either side with 30-40 conspicuous spiny teeth,
often up to the lower 2 mm unarmed; dorsal
surface with as much spines as along one margin,
mostly on the midrib; spine-cells c. 0.15 mm,
situated on several brown cells, all together
14-0.4 mm; all spines (dark) brown; the whole
tooth c. conical, 744-1 mm, 14-14 as wide as the
blade; cavities often indistinct, as wide as the mid-
rib. Sheath 3-314 by 4-414 mm, on either side
of the margin with 7-10, on the upper 1/3 of the
dorsal surface with numerous spines; auricles
absent. 3 Flowers not seen. 2 Flowers 244-3 mm;
ovary c. 1 by 0.6 mm; style 0.6—-0.8 by 0.1 mm,
with 2-3 stigmas c. 1.2 mm. Seeds greyish-brown,
4414 by 2144 mm.

Distr. Malaysia: West Central Sumatra (Lake
of Manindjau), c. 500 m. Fig. 4.

164 FLORA MALESIANA [ser. I, vol. 6?

2. Section Caulinia

A. BRAUN, J. Bot. 2 (1864) 276; Ascuers. Fl. Prov. Brand. | (1864) 67 (status not
mentioned ).—Caulinia WILLD. Mém. Ac. R. Sc. Berl. (1798) 87.—Subg. Caulinia
ASCHERS. ex RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 398; Pfl. Reich Heft 7
(1901) 10.

Mostly slender plants. Epidermis not distinctly differentiated from the underly-
ing cortex. Internodes smooth. Leaves usually much more than 15 times as long
as broad, mostly thin, (mostly) finely dentate along the margin and rarely so on
the lower portion of the midrib, with (0O—)10-80(—180) spiny teeth on either side.
Sheath mostly with distinct auricles. Intravaginal scales lanceolate to filiform,
thin (fig. 2i-1). Monoecious. 3 Flowers with or without spathe. Anther either | or
(2-, or) 4-celled. 9 Flowers mostly without spathe, stigmas 2(—3); spathe tapering
to the top, or constricted into a long cylindrical neck (more than twice as long as
broad), the margin with 3-9 distinct spines. Seed less than 314 mm long, if nearly
314 mm then only c. 0.6—0.8 mm broad; testa consisting of 3 layers of hardened
cells; areoles all of about the same size, except on the raphe, usually arranged in

length-rows.

Distr.. About 35 spp. throughout the warm and temperate regions of the globe.

2. Najas minor ALL. Fl. Ped. 2 (1785) 221; A.
Braun, J. Bot. 2 (1864) 277, excl. var. indica
A. Br.; Hook. f. Fl. Br. Ind. 6 (1893) 569, excl.
Caulinia indica et N. indica; RENDLE, Trans.
Linn. Soc. II, Bot. 5 (1899) 413, excl. stirp.
GRIFFITH 5609/5 & 5609/7; non RipDL. Mat. FI.
Mal. Pen. 2 (1907) 129; non Koorp. Junghuhn
Gedenkb. (1910) 160, quae est N. indica.

Up to c. 25 cm, lower internodes 2—S cm by
14-1 mm. Leaves (1—)1144—-2(-3) cm by 0.4-0.6
(0.7) mm, flat, sometimes subterete or triangular
in section; apex acute to slightly obtuse (blade
c. 3 mm below the tip 0.3—0.35 mm wide); margin
on either side with (5—)7—15(—17) mostly con-
spicuous spiny teeth, up to the lower 3 mm often
unarmed; dorsal surface sometimes with some
spines .(cf. RENDLE); all spines (dark) brown;
spine-célls (0.075—)0.15(—0.2) mm; teeth up to
as long as half (in the upper portion of the blade
as long as) the width of the blade; midrib c.
0.1 as wide as the blade; cavities occupying
half to almost the entire width of the blade, often
with distinct septa. Sheath 11/-3(-3.5) by 144-3
mm, on either side with 5—10(—15?) spines, the
inner side of the auricle unarmed; auricles (excl.
spines) broad to very broad triangular, truncate
to rounded, (0.1—)0.2-0.5 by 0.3-0.7 mm, often
shallowly lobed or lacerate. ¢ Flowers (mostly?)
solitary, enclosed in a spathe 114-2 mm; above
the anther a conical(-cylindrical) neck 0.3(—0.4)
mm, tip almost truncate or (slightly) oblique,
often somewhat lacerate; anther 1-celled (?, see
notes), oblong, often somewhat constricted towards
the top, 0.8-1.3 by 0.3—0.4 mm; ‘perianth lobes’
inconspicuous; ‘pedicel’ 1/3—2/5 mm, in anthesis
114-2 mm. 2 Flowers without spathe, often 2(—3)
together, in different stages; 1.9—2.3(—2.5) mm;
ovary 0.7-0.8 by c. 0.35 mm; style 0.4-0.6 mm,
with 2 stigmas 0.6-1 mm. Seeds narrow-oblong,

the top sometimes slightly bent, 2.4-3 by ec.
0.6 mm; (sometimes greyish); testa with (12-)
14-16(-19), ladder-like, length-rows of many
(100-150) areoles, which are much wider than
high.

Distr. According to RENDLE widely distribut-
ed in Europe, tropical and North Africa, tropical
and temperate Asia to Japan and Pegu (KURZ),
in Malaysia: ?Java (Bogor), see below.

Notes. N. minor has repeatedly been reported
from Malaysia. The two GRIFFITH numbers 5609/5
and 5609/7 cited by RENDLE and RipLey I have
seen in Herb. Kew; they have no fruit; the latter
number is sterile, the first has ¢ flowers with a
4-celled anther which excludes N. minor, but points
to either N. indica or N. oguraensis Miki. The
Koorpers record from Java is based on a speci-
men of N. indica with which this species has
sometimes been confused, specially by those who
referred N. indica as a variety to N. minor.

The only specimens from Malaysia which repre-
sent with certainty N. minor were raised at
Ziirich from ‘“‘Schlammproben von Buitenzorg
mit Hydrilla verticillata, im Gewachshaus yon
Prof. Ernst in Ziirich gekeimt 1933; 1934 von mir
im Warmhaus weitergezogen; fruchtend eingelegt
am 3. Febr. 1935. WALo Kocu. leg. M. ERNST-
SCHWARZENBACH”, in Herb. E. T. H., Ziirich.

It is most remarkable that nobody has collected
N. minor in Buitenzorg or vicinity where so
many collections have been made. If these ERNST
samples were extracted from the small concrete
water-plant tanks in the Botanic Gardens, there
is every reason to assume contamination with
imported water plants. Contamination with for-
eign seed could possibly also have happened at
Ziirich. For the present N. minor seems very
doubtfully native in Malaysia.

A closely related species is N. oguraensis MIKI,

Dec. 1962]

NAJADACEAE (de Wilde)

165

Bot. Mag. Tokyo 44 (1935) 775, t. 7 A-L, which
was described from Japan as a rigid plant up to
1 m long with seeds c. 31 by 0.6 mm and 4-celled
anthers c. 134 mm long. I have seen plants from
India with a slender habit, seeds (2—)3-334 mm
and large 4-celled anthers 114-134 by 34 mm which
thus come very close to the Japanese species and
could well belong to it (Herb. Persoon, in L;
SAXTON 491, in K; STEWART 3347, in K).

3. Najas kurziana RENDLE, Trans. Linn. Soc. I,
Bot. 5 (1899) 413, t. 41 f. 116-121; Pfl. Reich
Heft 7 (1901) 15, f. 4 U; Horn AF RANTZIEN,
Kew Bull. (1952) 37.—Fig. 2b.

Up to 15 cm, lower internodes 1-2 cm by
0.5-0.7(-0.8) mm. Leaves 114-2 cm by 0.45—
0.95 mm, flat (and thin), apex rounded to obtuse
(blade c. 3 mm below the tip 0.4-0.7 mm in width);
margin on either side with 30-60 inconspicuous
teeth, mainly consisting of the spine-cell, dorsal
surface unarmed; spine-cells 0.05-0.1 mm, all
(dark) brown; midrib 0.07-0.05 as wide as the
blade, cavities occupying 0.1—'4 of the width of
the blade, often with distinct septa. Sheath on
either side with (3—)5—12 spines, often 1-3 on
the inner edge of the auricle, 1.7—2.4 by 1.3-2.2
mm; auricles (broad-)triangular to long-triangular,
subentire, tip obtuse or more or less rounded,
(0.2-)0.5-0.8 by 0.3-0.5 mm. Flowers mostly up
to 5 together, in different stages, often with 1 male,
or male solitary. Spathe of ¢ flower 0.9-1(-1.2)
mm, neck mostly cylindrical, 0.3—-0.4(-0.5) mm,
tip almost truncate, entire to slightly lobed;
anther 1-celled, oblong, (0.3—)0.4—0.7(—1) by (0.1—)
0.15-0.25 mm; ‘perianth lobes’ inconspicuous.
*Pedicel’ 0.1-0.2(—0.3) mm, in anthesis c. 0.8 mm.
9 Flowers espathaceous, c. (1—)1.4 mm; ovary
c. (0.4-)0.5 by 0.2 mm; style 0.3-0.5 mm, with
2(-1) stigmas 0.3-0.4 mm. Seeds 1-1.2 mm by
0.35(—0.4) mm; testa with 16-18 length-rows of
18-26 subquadrate or 5—6-angular areoles.

Distr. India (North Bengal between Kishen-
ganj and Oolabena, Kurz s.n. in CALC & BM,
once found), in Malaysia: Lesser Sunda Islands
(Port. Timor: in Irabére R., near Uato Carabao,
c. 400 m alt., VAN STEENIS 18226), once found.
Fig. 4.

Notes. The Timorese collection differs from
the holotype only in having c. 50-60 spiny teeth
on each leaf margin (in the type there are 30-34).

According to HORN AF RANTZIEN (Kew Bull.
1952, 35-37, t. 3) the African N. hagerupi H. a. R.
is very closely allied, the only difference being
in the anther which is 1.2-1.4 mm long; its fruit is
unknown.

4. Najas browniana RENDLE, Trans. Linn. Soc. II,
Bot. 5 (1899) 420, t. 42, f. 163-167; Pfl. Reich
Heft 7 (1901) 17, f. 5 J.—Fig. 2d.

Up to 15 cm, lower internodes 1—3(—6) cm by
0.5—0.8(-0.9) mm. Leaves 114-214 cm by (0.35-)
0.4-0.7(-0.8) mm, flat, sometimes slightly fleshy,
apex obtuse to rounded, sometimes acute (c.
3 mm below the tip 0.25—0.6 mm wide); margin

on either side with 12-21 spiny teeth largely in the
upper portion of the blade; no spines on the dorsal
surface; spine-cells < 0.05-0.15 mm, (dark)
brown; teeth up to as long as 0.2 of the width of
the blade; midrib c. 0.07 as wide as the blade;
cavities occuping 14-34 of the width of the blade,
septa often indistinct. Sheath on either side with
(1—)3-7(-10) spines, no (?) spines on the inner
edge of the auricle, (1.5—)2(—2.3) by 1.5—2.9 mm;
auricles mostly broad, seldom long-triangular,
truncate-rounded to acute, sometimes acuminate,
rarely falcate, (0.1—)0.2-0.3(-0.5) by 0.25-0.75
mm, entire to lobed or lacerate (incisions up to
halfway (or more) the auricle). Flowers mostly up
to 3 together, in different stages, together with one
male'!, or the male solitary. ¢ Flowers: spathe
1.3-2 mm, neck almost cylindrical, 0.3-0.5 mm,
the tip almost truncate, entire to lacerate; anther
l-celled, ovate to oblong(-lanceolate), 0.6—0.8
(-0.9) by 0.2-0.3 mm; ‘perianth lobes’ indistinct;
‘pedicel’ 0.3-0.6 mm, at anthesis 1-14, mm.
2 Flowers: spathe absent; 9 fl. 114-2 mm; ovary
0.5-0.8 by 0.2-0.3 mm; style (0.2—)0.5—0.7 mm,
with 2 stigmas (0.3—)0.5—0.7(-0.9) mm. Seeds
(1.5— and narrow) 1.6-2 by (0.45—)0.5—0.7
(0.75) mm, testa with 18-30 length-rows of
27-37 subquadrate or 5—6-angular areoles.

Distr. India (once found) and Northern Terri-
tory of Australia (Cavern I.), in Malaysia: Java
(SE of Semarang: Kuwu mudwells; Surabaja:
Keputih tambaks; Madura I.), S. New Guinea
(coast between Oriomo and Fly R.). Fig. 4.

Ecol. In the Kuwu saline mudwell area the only
aquatic water plant, near Surabaja in brackish
water of fish ponds, in Madura in pools near the
sea, in S. New Guinea in open Eleocharis swamp,
all localities as far as known saline and at very
low altitude.

Notes. Already found by HorsFiELD at Kuwu,
but that specimen was identified by RENDLE as N.
falciculata (I.c. 1899, p. 418). In his key he placed
N. browniana along with species possessing 4-celled
anthers, although the anther depicted (/.c. t. 42
f. 165) is l-celled. I have examined the holotype
and this proves to be correct, they are mature and
1-celled.

The only specimen from continental tropical
Asia is a specimen in the Kew Herbarium bearing
three labels, reading: “1217/1830 Tutichorin;
N. rigidula Herb. Wight prop.; N. graminea var.
minor Rendle, Flora of Madras, det. C. Binker
19.4.1929”’.

I am not entirely satisfied that this species is
specifically distinct from N. gracillima (A. BRAUN)
Maanus (N. indica var. gracillima A. BR. ex
ENGELM. in A. GRrAy) which was originally des-
cribed from eastern North America. The North
American material is very distinct from the ma-
terial of N. browniana in having much longer seeds
(c. 2.6-3.3 mm) with elongated areoles and large
anthers (c. 114 mm). As RENDLE supposed that
N. browniana would have 4-celled anthers, they
appear in his key wide apart. This, however,

(1) Male flower often appressed to the female one.

166

FLORA MALESIANA

[ser. I, vol. 6?

not being correct, we are faced with the problem of
their discrimination through the statements of
Miki who recorded WN. gracillima from Japan
(Tokyo Bot. Mag. 49, 1935, 773, f. 6 A—L)
supplemented by many sheets collected by FAURIE
in Japan and Formosa which I had on loan from
the Geneva Herbarium. Through Mrkxr’s_ de-
scription and FAurRIe’s material it appears that
the plants from Japan and Formosa do not exactly
match the North American plants, as they have
small anthers (c. 0.7 mm) and smaller seeds (2 by
14 mm), to which should be added that in the
Faurie material I observed seeds up to c. 3 mm
long. The areoles of the Japanese plants are about
twice as long as wide. The Japanese-Formosan
plants are, therefore, more or less intermediate
between true gracillima and browniana. It was
beyond the scope of the present revision to go
much deeper into this delicate situation, although
it should be realized that if these species are merged
into one (as three replacing subspecies) which
would then possess an enormous distributional
area, N. gracillima is the oldest binomial for it.
The fact that the Japanese plants are largely from
rice-fields and that N. browniana has, in Malaysia,
an exacting natural habitat has added to my provi-
sional decision to keep it apart.

5. Najas indica (WILLD.) CHAM. Linnaea 4 (1829)
501; KUNTH, En. (1841) 113; HAssk. Pl. Jav. Rar.
(1848) 142; ZoLL. Syst. Verz. 1 (1854) 74; Mia.
Ill. Fl. Arch. Ind. (1871) 44-45, pro parte, incl.
var. macrodictya et var. rigida A. BR., nom. nuda;
MAGNnus, Ber. Deut. Bot. Ges. 2 (1894) 218-219;
RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 399,
t. 39 f. 31-45; Pfl. Reich Heft 7 (1901) 10, f. 3
R-T; Merr. En. Philip. 1 (1923) 25; FiscHer,
in Gamble, Fl. Madras (1931) 1603; DE WILDE,
Willdenowia 2 (1960) 595; Act. Bot. Neerl. 10
(1961) 167, fig. 6—9.—Caulinia indica WILLD.
Mém. Ac. R. Sc. Berl. (1801) 89, t. 1 f. 3; Sp. Pl. 4
(1805) 182; cf. DE WILDE (1960) /.c.—N. palustris
BLANCO, FI. Filip. (1837) 660; KuNtTH, En. 3
(1841) 590; RENDLE, Trans. Linn. Soc. II, Bot. 5
(1899) 427; Pfl. Reich Heft 7 (1901) 18; Merr.
Sp. Blanc. (1918) 57; En. Philip. 1 (1923) 25;
VENKATESH, Bot. Notis. 109 (1956) 75-82, f. 1,
3-6, 8-18, 24-27, 29-36.—N. lobata BLANCO, FI.
Filip. ed. 2 (1845) 459; Naves, ibid. ed. 3, 3 (1879)
65; Nov. App. (1880) 298; RENDLE, Trans. Linn.
Soc. II, Bot. 5 (1899) 427; Pfl. Reich Heft 7
(1901) 18.—N. tenuifolia (non R. Br.) Mia. FI.
Ind. Bat. 3 (1856) 224, pro specim. sumbaw.
(ZOLL.) et tranqueb. (KLEIN); NAves, Noy. App.
(1880) 297.—N. falciculata A. BRAUN, J. Bot. 2
(1864) 278, f. 4; MARTENS, Preuss. Exp. Ost-As.
Bot. Teil Tange (1866) 143; Hoox. f. Fl. Br.
Ind. 6 (1893) 569; RENDLE, Trans. Linn. Soc. II,
Bot. 5 (1899) 417, t. 42, f. 145-151; Pfl. Reich
Heft 7 (1901) 15; SunierR, Treubia 2 (1922)
190-195, f. 6; Merr. En. Philip. 1 (1923) 25;
Back. Handb. Fl. Java 1 (1925) 51; HEYNE,
Nutt. Pl. (1927) 139; Back. Onkruid. Suik.
(1928) 20; FiscHER in Gamble, Fl. Madras (1931)
1604; Steen. Arch. Hydrob. Suppl. 11 (1932)

271, t. ii; BuRK. Dict. (1935) 1533; BACK. Bekn.
Fl. Java (em. ed.) 10 (1949) fam. 210, p. 1.—
N. minor var. indica A. Br. J. Bot. 2 (1864) 278,
excl. specim. maurit.; MARTENS, Preuss. Exp. Ost-
As. Bot. Teil Tange (1866) 143.—N. foveolata
A. Br. ex (MAGNUS, Beitr. 1870, vii, 43, nomen)
RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 416,
t. 41, f. 139-144; ibid. (1900) 443; Pfl. Reich
Heft 7 (1901) 15, f. 4 W; Camus in FI. Gén. I.-C.
6 (1942) 1213.—N. minor (non ALL.) Hook. f.
Fl. Br. Ind. 6 (1893) 569, pro majore parte, pro
syn. indica; Koorp. Jungh. Gedenkb. (1910)
160.—N. kingii RENDLE, Trans. Linn. Soc. II,
Bot. 5 (1899) 415, t. 41 f. 126-131: ibid. (1900)
442; Pfl. Reich Heft 7 (1901) 15 f. 5 A-C; Rip.
Mat. Fl. Mal. Pen. 2 (1907) 129; Fl. Mal. Pen. 4
(1924) 366; Camus in FI. Gén. I.-C. 6 (1942)
1213, t. 115 f. 13-19; Hemne, Aquar. & Terr. Z.
(Datz) 11 (1958) 375, fig. 1-2.—N. lacerata
RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 416,
t. 41 f. 132-138; Pfi. Reich Heft 7 (1901) 15, f. 5
E-F; FIscHer in Gamble, Fl. Madras (1931) 1603,
1604.—Fig. 2 1.

Up to c. 40 cm, lower internodes 2-10 cm by
(0.6—)0.75—-1.2(-1.7) mm. Leaves (114-)24%,4'4
(—6) cm by (0.3—)0.8—1.2(—1.6) mm, flat to subterete
or triangular in section, apex acute to obtuse
(blade c. 3 mm below the tip 0.2-0.4(-0.5) mm
wide); margin on either side with (1—)5—30(-40)
conspicuous spiny teeth, sometimes up to the
lower 15 mm of the blade unarmed; dorsal surface
sometimes with 1-—10(—30) spines; spine-cells
0.1—0.25 mm, all (dark) brown; teeth up to as long
as half (in the upper part of the blade as long as)
the width of the blade; midrib 0.12-0.1 times as
wide as the blade; cavities occupying half or almost
the entire width of the blade, often with distinct
septa. Sheath (2-)2.5—4(-S) by (1.5-)24(-6) mm,
on either side with (2—)5—12(—20) spines, but un-
armed on the inner edge of the auricles; auricles
absent to (long-, especially in Indian specimens)
triangular, truncate to rounded or almost acute,
rarely falcate, (0.1—)0.3-0.6(-1) by (0.3—)0.5-1
(-1.5) mm, mostly entire, sometimes toothed or
lobed, seldom lacerate up to halfway. Flowers
mostly solitary. ¢ Flowers: spathe 2.2-3.6 mm,
neck mostly cylindrical, 0.3-1 mm, tip almost
truncate to very oblique, entire to up to halfway
lacerate; anther subsessile, 4-celled, about ellipti-
cal to oblong, (1.2—)1.5—2.2(—2.5) by 0.7-1.2 mm;
‘perianth’ lobes mostly distinct, (+) nipple-
shaped. ‘Pedicel’ c. 0.2 mm, in anthesis 2.7—3.5
mm. 2 Flowers: spathe absent (except in some
Indian specimens), (1.7—)2-3(-3.4) mm; ovary
(0.6—-)0.7-0.9(—1.2) by (0.25—)0.3—-0.4(—0.45) mm,
style (0.4-)0.5—0.8(-1.2) mm, with 2(-1) stigmas
(0.4-)0.8-1.6(-1.8) mm. Seeds (1.5—)1.7-2.3 by
0.75—-0.9 mm; testa with (14—-)16—20(—25) length-
rows of (17—)22-25(-30) subquadrate or 5-6-
angular areoles, which are mostly slightly wider

than long. :
Distr. ?Africa, tropical continental Asia
(Kashmir, Deccan, Bengal, Andamans, Siam,

Tonkin) northward to Japan, throughout Ma-
laysia (not yet coJlected in Borneo and Moluccas).

Dec. 1962]

NAJADACEAE (de Wilde)

167

Fig. 5.

eee}

ae
ar)

Fig. 5. Malaysian and some adjacent continental
Asian localities of Najas indica (WILLD.) CHAM.

Ecol. Found in a great variety of habitats,
in various biotopes of freshwater, mostly shallow
but up to 5 m depth, and c. 1000 m altitude, also
found in brackish water of fish-ponds near
Djakarta with a salinity of up to 30°/,, (SUNIER,
l.c.), both in everwet regions and in those subject
to a dry season. One of the commonest species in
Malaysia.

Vern. Lumut-siarang, Toba, ganggéng, Ba-
tavia; Philippines: aragdn, Ilk., bangbafgi, Bon.,
labui, Ig.

Notes. A variable species, the variability
obviously mostly due to external conditions. Spe-
cimens from India not seldom deviate to forms with
thin stems and narrow leaves and rather large
seeds (up to 2.3 by 0.9 mm) with many (up to 25)
longitudinal rows of many (up to c. 30) rather
small areoles of the seed coat.

In Asiatic specimens the 2 flowers are often en-
closed by a spathe, but these may be found together
with naked ones on a single individual (cf. DE
WILDE, 1961). In Malaysia the 2 flowers are as far
as known always espathaceous.

It was judged beyond the scope of this revision
to make a thorough study of African material des-
cribed under other various names which may pos-
sibly be referred to N. indica as circumscribed here.
I have not the impression that this species occurs
in Australia.

6. Najas tenuifolia R. Br. Prod. 1(1810) 345;
BenTH. FI. Austr. 7 (1878) 181; RENDLE, Trans.
Linn. Soc. II, Bot. 5 (1899) 419, t. 42, f. 158-162;
Pfl. Reich Heft 7 (1901) 17, f.5 G-H.—N. graminea
DEL. var. tenuifolia (R. BR.) A. BRAUN, J. Bot. 2
(1864) 278.—N. celebica Koorp. Minah. (1898)
270, 637.—N. pseudograminea W. Kocu, Ber.
Schweiz. Bot. Ges. 44 (1935) 339.—Fig. 1, 2f, i.

Up to c. 40 cm, lower internodes 2-6 cm, by
(0.5—)0.7-0.85(-1.2) mm. Leaves (114-)134-4
cm by 0.6—-1.2 mm, flat, tip blunt (rounded) to
acute (blade c. 3 mm below the tip 0.3-0.9 mm

wide); margin on either side with 14—60(—70)
spiny teeth; dorsal surface without spines; spine-
cells (0.05-)0.1-0.2 mm, on the auricles up to
0.3 mm, on the tip of the spathe rarely up to
0.4 mm long, brownish, rarely yellowish green;
teeth shallow, up to c. 0.16, near the tip up to
c. 1/3 as long as the width of the blade; midrib
0.1-0.07 as wide as the blade; cavities variable
in width, septa sometimes distinct. Sheath (1.5—)
2-4.7(-5S) by (1.5—)2-3(—3.5) mm, on either side
with 5—25 spines, (0O—)1-3(-4) on the inner edge
of the auricle; auricles long-triangular, mostly
114-2 times as long as broad, with an obtuse
(rounded) to acute apex, rarely falcate, (0.4—)0.6—
1.9(-3) by (0.25—)0.6-1 mm, entire, sometimes
shallowly lobed (or somewhat lacerate). Flowers
solitary or 2-3 together. ¢ Flowers: spathe 2-3.8
mm, subsessile (very rarely up to 0.8 mm stalked),
neck mostly cylindrical, (0.4-)0.5-1 mm, tip
almost truncate to oblique, entire to lobed; anther
subsessile, 4-celled, ovate-elliptical to slightly
oblong, 1.2-2.5 by 0.75-1.1 mm, ‘perianth’ lo-
bes distinct, (shallowly) rounded. ‘Pedicel’ c. 0.2
mm, in anthesis 1.5—2.3(-2.7) mm. 2 Flowers
espathaceous, 1.5—3.5 mm; ovary 0.5-0.9(-1) by
0.2—0.35 mm, style 0.4—1(—1.2) mm, with 2 stigmas
0.3-0.8(—1.7) mm. Seeds (1.1—)1.2-2.3 by 0.4-0.75
mm; testa with 16-36 length-rows of (15—)20(—25)
or 35-45 subquadrate or 5-6-angular areoles.

Distr. Australia and Malaysia.

Notes. The anther of this species has origi-
nally been described by R. BRown as 1-celled and
so it was defined by BENTHAM (1878); RENDLE
already observed, however, that this was an error
and he is correct as I could verify on the holotype
(in BM).

I have divided this species into two subspecies,
ssp. tenuifolia which occurs both in Australia and
Malaysia and ssp. pseudograminea which is confined
to Malaysia. In order to facilitate identification
the two subspecies are opposed here and I have
added the characteristics of a closely allied species
known only from continental Asia, viz. N. bre-
vistyla, which is most closely allied to the Malay-
sian ssp. pseudograminea.

KEY TO THE SUBSPECIES

1. Leaves small, 14-24 by 0.4-0.6 mm, often some-
what fleshy, on either side with 17-33 rigid
spiny teeth up to as long as the width of the
blade. Spathe of ¢ fl. 1.9-2.2 mm; anther
small, 1-1.3 by 0.5—0.6 mm. 2 Flower c. 1.5
mm. Seeds small, 1-1.2 by 0.4—0.45 mm.

N. brevistyla RENDLE

1. Leaves (15—)17—40 by 0.6-1.2 mm, not fleshy,
on either side with 14~-60(—70) rather shallow
spiny teeth, up to 1/3 as long as the width of the
blade. Spathe of 3 fi. 2-3.8 mm; anther 1.2
—2.5 by 0.75-1.1 mm. 2 Flower 1.5—3.5 mm.
Seeds (1.1—)1.2-2.3 by 0.4-0.75 mm.

2. Leaves 21-40 mm, apex acute (or slightly
obtuse), c. 3 mm below the tip 0.3—0.4 mm in
width; margin on either side with 14—-35(-
42) spiny teeth (spine-cells dark brown);

168

FLORA MALESIANA

[ser. I, vol. 6?

cavities nearly reaching the margin (their
outer edge not brown-coloured); septa often
distinct. Flowers solitary. Mature anther
2-2.5 by 0.8-1.1 mm. Seeds 2—2.3 by 0.7(-0.75)
mm. N. tenuifolia ssp. tenuifolia
. Leaves (15—)17—28 mm, apex blunt (round-
ed), rarely slightly acute, c. 3 mm below the
tip (0.5-)0.7-1 mm in width; margin on
either side with 28—60(—70) spiny teeth (spine-
cells light brown, rarely yellowish green);
cavities occupying 0.12-0.5 of the width of
the blade, their outer edge often dark-brown;
septa indistinct. Flowers mostly 2-3 together.
Anther 1.2—2 by 0.75-—0.9 mm. Seeds (1.1—)

1.2-2 by 0.4— 0.65 mm.
N. tenuifolia ssp. pseudograminea

Nm

ssp. tenuifolia.—WN. tenuifolia R. Br. Prod. (1810)
345; BENTH. FI. Austr. 7 (1878) 181; RENDLE,
Trans. Linn. Soc. II, Bot. 5 (1899) 419, t. 42 f. 158-
162; ibid. (1900) 443; Pfl. Reich Heft 7 (1901) 17,
f. 5 G-H.—N. graminea var. tenuifolia (R. Br.)
A. BRAuN, J. Bot. 2 (1864) 278.

Leaves 2-334 cm by 0.6—1.3 mm, on either side
with 14-40 (dark) brown spiny teeth. Sheaths on
either side with 6-17 spines. Spathe of 3 flowers
2.8-3.8 mm, neck 0.7—0.9 mm, bearing up to c.
0.3 mm long spine-cells; anther 2—2.5 by 0.8-1.1
mm. & Flowers 2.3—3(—3.5) mm; ovary 0.8 by
(0.25—)0.3 mm; stigmas 0.6—0.8(-1.2) mm. Seeds
2-2.3 by 0.7-0.75 mm; testa with 26-36 length-
rows of 35—40 subquadrate or 5—6-angular areoles.

Distr. Australia, and 7New Caledonia (REND-
Le, 1900, p. 443), in Malaysia: SE. Celebes
(Lepo-lepo near Kendari, BEccARI sh. no. 11810
Fir), once found.

ssp. pseudograminea (W. KOCH) DE WILDE, comb.
nov.—N. celebica Koorb. Minah. (1898) 270, 637.
—N. pseudograminea W. Kocn, Ber. Schweiz.
Bot. Ges. 44 (1935) 339.—Fig. 1, 2f, i.

KEY TO THE VARIETIES

1. Spathe (of ¢ fi.) 2-2.8 mm; anther 1.2-1.85
mm. & Flower 1.5—2 mm, stigmas 0.3-0.7 mm.
Seeds small, (1.1—)1.2—-1.3(—1.4) by 0.4—0.5 mm.

var. pseudograminea

1. Spathe (of 3 fl.) 3-3.5 mm; anther 1.6-2 mm.

2 Flower 2.7-3.5 mm, stigmas 1-1.7 mm.

Seeds 1.9-2.05 by 0.5-0.65 mm. var. celebica

var. pseudograminea.—N. graminea (non DEL.)
Mia. Illustr. (1871) 45, pro specim. javan.—N.
tenuifolia (non R. Br.) Auct.: BACK. Teysm. 22
(1911) 514, excl. specim. tobaic.; Handb. Fl. Jav.
1 (1925) 51; Bekn. FI. Java (em. ed.) 10 (1949)
fam. 210, p. 1.—N. falciculata (non A. Br.)
CoeERT, Trop. Natuur 23 (1934) 27, f. 11.—N.
pseudograminea W. Koc, Ber. Schweiz. Bot. Ges.
44 (1935) 339; Horn AF RANTZIEN, Kew Bull.
(1952) 37.—Fig. 1, 2f.

Leaves (1 14—)2(—234) cm by (0.6—)0.9(—1.1) mm,
green, margin of the blade on either side with
28-53(—63, Buru) spiny teeth, brown; sheaths

on either side with 5—15(—23, Buru) spines. ¢
Flowers: spathe 2-234 mm, neck (0.4—)0.5-0.6
(-0.9) mm, bearing up to 0.25 mm long spine-
cells; anther 1.2-1.85 mm; ‘pedicel’ at anthesis up
to 2.3 mm. 2 Flower 114-2 mm; ovary 0.5—0.8 by
0.2-0.3 mm; stigmas 0.3-0.7 mm. Seeds (1.1—)
1.2-1.3(-1.4) by 0.4-0.5 mm; testa with 16-18
length-rows of (15—)20—23(—25) (sub)quadrate or
5—6-angular areoles.

Distr. Malaysia: Java (also Madura I.), Les-
ser Sunda Islands (Bali), Philippines (Luzon; Pam-
panga Prov.), and Moluccas (Buru). Fig. 6.

a -
f\
5 aa

\
— \

= A eae An

”
fe

2S
ae

Fig. 6. Localities of Najas tenuifolia R. BR. ssp.
pseudograminea (W. KOCH) DE WILDE var.
pseudograminea.

Ecol. Common in Java, at low altitude, mostly
in freshwater but found in brackish water in Bali
(O. JAAG), twice found in the mountains, viz
Rana Lake (Buru) at c. 700 m and in the
rivulet and craterlake of Mt Kelud in Central
Java at 1200 m abundant in sulfurous hot water
(up to 60° C) on blackish, muddy soil (CLASON &
COERT).

Notes. The identification of the Buru col-
lection is somewhat doubtful.

One plant from Java possessed among numerous
naked 2 flowers and fruits a single one which was
enveloped by a spathe (cf. DE WiLpe, Act. Bot.
Neerl. 10, 1961, 167).

var. celebica (KOORD.) DE WILDE, comb. nov. —
N. celebica Koorp. Minah. (= Med. Lands Pl.
Tuin) 19 (1898) 270, 637; RENDLE, Pfil. Reich
Heft 7 (1901) 18; Koorp.-ScHum. Syst. Verz.
(Cel.) (1914) 9.—Fig. 2i.

Leaves 134-234 cm by 0.95-1.2 mm, often
bluish-green (when dry), margin of the blade on
either side with 40-60(—70) spiny teeth, yellowish
green. Sheaths on either side with 10-25 spines.
3 Flowers: spathe 3-314 mm, neck 0.8-0.9 mm,
bearing up to c. 0.4 mm long spine-cells; anther
1.6-2 mm; ‘pedicel’ at anthesis up to 2.7 mm.
Q Flowers 2.7-3.5 mm; ovary 0.7-0.9(-1) by
0.2-0.35 mm; stigmas 1-1.7 mm. Seeds 1.9-2.05
by 0.5—0.65 mm; testa with 16-24 length-rows of
38-45 areoles, which are often somewhat broader
than high.

Dec. 1962]

NAJADACEAE (de Wilde)

169

Distr. Malaysia: NE. Celebes (Minahassa: Ton-
dano Lake), c. 700 m altitude.
Vern. Sengit, tl.

7. Najas malesiana DE WILDE, Act. Bot. Neerl. 10
(1961) 168.—N. graminea var. minor RENDLE,
Trans. Linn. Soc. II, Bot. 5 (1899) 426; Pfl. Reich
Heft 7 (1901) 18; Camus, in Fl. Gén. I.-C. 6
(1942) 1212.—N. graminea var. angustifolia
RENDLE, Trans. Linn. Soc. II, Bot. 5 (1899) 427,
t. 42 f. 202; Pfl. Reich Heft 7 (1901) 18, f. 5 V;
Ript. Mat. Fl. Mal. Pen. 2 (1907) 128; GrsBs,
J. Linn. Soc. Bot. 42 (1914) 172; Merr. En. Born.
(1921) 37; Rpt. Fl. Mal. Pen. 4 (1924) 366;
Disp. (1930) 180.—N. graminea (non Dev.) RIDL.
Mat. FI. Mal. Pen. 2 (1907) 128; Fl. Mal. Pen. 4
(1924) 365.—N. bengalensis HORN AF RANTZIEN,
ad int. Act. Hort. Gotob. 18 (1950) 192, 193;
Kew Bull. (1952) 39, in clay. angl.—Fig. 2c, k.

Up to 15 cm, lower internodes 1-3 cm by 0.5—
0.8 mm. Leaves (114-)2(-2144) cm by (0.4-)
0.5—0.7(-0.9) mm, flat, apex obtuse or slightly
acute (blade c. 3 mm below the tip 0.25-0.5 mm
wide); margin on either side with (20—)25-30
(—50-60) inconspicuous teeth, mainly consisting
of the spine-cell; dorsal surface without spines;
spine-cells (0.05—)0.1(—0.25) mm, (dark) brown;
midrib c. 0.05 times as wide as the blade, cavities
occupying up to half the width of the blade,
often with distinct septa. Sheaths (1.5—)2—2.5(-3)
by 1—2.5 mm, on either side with (2—)6—14 spines,
0-4 spines on the inner edge of the auricle;
auricles long-triangular or linguiform, sometimes
falcate, 0.5-0.8(-1.1) by 0.3-0.8 mm, entire
(rarely shallowly lobed). 2 Flowers often up to
3 together, in different stages, together with one
male (often appressed to the 2), or male solitary.
3 Flowers without spathe; anther 1-celled, ellipti-
cal-oblong, apically sometimes narrowed, 0.6—1
by 0.15—0.3 mm; ‘perianth lobes’ often indistinct;
‘pedicel’ 0.2-0.4, at anthesis c. | mm. 2 Flowers:
spathe absent; rarely up to 0.3 mm stalked;
(1.2-)1.5(-1.8) mm; ovary 0.4—0.8 by 0.2—0.3 mm;
style 0.25-0.5 mm, with 2(—1) stigmas 0.3-0.6 mm.

FE ms

Fig. 7. Malaysian Jocalities of Najas malesiana
DE WILDE.

Seeds (0.9-)1-1.5 by (0.35—)0.4—0.5 to 1.5-1.8 by
0.5-0.6 mm; testa with 16-26 length-rows of
24-30 subquadrate or 5—6-angular areoles.

Distr. India, E. Bengal, Burma, Pegu, Lower
Siam, Indo-China, throughout Malaysia (not yet
found in New Guinea, Lesser Sunda Islands, and
the larger parts of Java and Borneo), on the whole
rare. Fig. 7.

Ecol. In streams and pools often in the forest,
in swamp forest (Tjitjadas), more rarely in rice-
fields (Setul, Padang), at low and medium altitude,
once at 900-1000 m (Padang).

Vern. Ganggeéng, S.

8. Najas graminea DEL. Descr. Egypt. Hist. Nat. 2
(1813) 282, t. 50 f. 3; Fl. Egypt. (1813) 138, t. 50 f.
3; Kuntu, En. (1841) 115; A. BRAuN, J. Bot. 2
(1864) 278 f. 5, pro parte, excl. specim. javan. et
celeb., excl. var. B tenuifolia A. BRAUN; MuiQ.
Illustr. (1871) 45, excl. specim. javan.; NAVES, Nov.
App. (1880) 298; BaiLey, J. Bot. 22 (1884) 305,
fig. 1-89, pro maj. p. incl. var. delilei MAGNUS;
Hook. f. Fl. Br. Ind. 6 (1893) 569; K. Scu. in Mart.
Fl. Bras. 3, 3 (1894) 730, t. 124 f. 2; RENDLE,
Trans. Linn. Soc. II, Bot. 5 (1899) 424, t. 42
f. 192-201, excl. var. minor et angustifolia REND-
LE; ibid. (1900) 443; Pfl. Reich Heft 7 (1901) 18,
f. 5 Q-V; Koorp. Exk. Fl. Java 1 (1911) 91;
Me_rr. FI. Manila (1912) 68; En. Philip. 1 (1923)
24; FiscHer, in Gamble Fl. Madras (1931) 1603,
1604; W. Kocu, Ber. Schweiz. Bot. Ges. 44 (1935)
340; Miki, Bot. Mag. Tokyo (1935) 774, t. 6
M-—R; GuILLAUMIN, Bull. Soc. Bot. Fr. 84 (1937)
256; Camus, Fl. Gén. I.-C. 6 (1942) 1211; Horn
AF RANTZIEN, Act. Hort. Gotob. 18 (1950) 191-193,
map; Kew Bull. (1952) 33, 39; VENKATESH, Bot.
Not. 109 (1956) 75-82, f. 2, 7, 19-23, 28; DE
Wipe, Act. Bot. Neerl. 10 (1961) 167, fig. 1—-S.-
N. seminuda GrirF. ex Voict, Hort. Suburb.
Calc. (1845) 694; GrirF. Not. 3 (1851) 184;
Ic. Pl. As. t. 251, f. 2, t. 253-254.—Fig. 2e, j.
Up to 50(—75?) cm, lower internodes 1-10 cm,
by (0.6-)0.7—-1.5(-2.25) mm. Leaves (134-)214-
314(-6 )cm by (0.6 -) 0.7—1(—-4) mm, flat, seldom
canaliculate, apex obtuse (-rounded), rarely some-
what acute (blade c. 3 mm below the tip 0.4-1.2
mm wide); margin on either side with (40—)60
(-185) inconspicuous spiny teeth, mainly con-
sisting of the spine-cell; dorsal surface without
spines; spine-cells 0.05—0.15 mm, yellowish brown;
midrib c. 0.07-0.05 times as wide as the blade,
cavities occupying up to '4 of the width of the
blade, septa (-_) indistinct. Sheath (3—)4—S(-10.5)
by (2-)3-4(-8.5) mm, on either side with (10-)
15—30(—50) spines; 3—8 spines on the inner edge of
the auricle; auricles long-triangular, seldom slight-
ly faleate; apex acute, rarely obtuse or acuminate,
(1-)2(-5.5) by 0.5-1(-1.5) mm, entire, rarely
(shallowly) lobed. Flowers often up to 3 together,
in different stages, together with one male, or
male solitary (in the var. robusta all flowers prob-
ably solitary). gd Flowers: spathe 0; anther
4(-2)-celled, elliptical, rarely ovate or oblong,
(1-)1.5(-2) by (0.4-)0.7(-1) mm; ‘perianth lobes’
distinct, rounded to globular. ‘Pedicel’ 0.2-0.5 mm,

170

FLORA MALESIANA

[ser. I, vol. 6?

at anthesis (0.7—)1—2.5 mm. 2 Flowers: no spathe;
(1.6-)2(—3.5) mm; ovary (0.6—)0.7(-1) by (0.2—)
0.35(—0.4) mm; style (0.4-)0.7(-1.2) mm, with
(1-)2(—3) stigmas (0.4—-)0.8(-1.6) mm. Seeds (in
the var. robusta unknown) (1.6-, broad seeds)
2.3(-2.7) by (0.6-, long seeds) 0.8(—0.9) mm;
testa with 20-34 length-rows of 30-45 subquadrate
to 5-6-angular areoles (often slightly higher than
broad).

Distr. Widely distributed from North Africa
and the Middle East, to tropical Asia (Punjab,
Deccan, Bengal, Assam, Burma, Cochinchina,
ex lit.), northwards to Formosa and Japan,
southwards to Malaysia (not yet found in Borneo
and Malaya, rare in Sumatra and Java), New
Caledonia, and Australia. Introduced in the rice-
fields in North Italy, England (BaAILey, /.c.), and
probably in Brazil (K. SCHUMANN, 1894, RENDLE
1899, HoRN AF RANTZIEN 1950, Il.cc). Fig. 8.

ie A fA Ls

Fig. 8. Malaysian localities of Najas graminea
DEL.

Ecol. Chiefly in the lowland in rice-fields,
ditches, small streams, in both everwet and seasonal
areas, up to 400 m, rarely at 1000 m (W. Java),
once at 1400 m (Wéetar), often together with
Nitella, Salvinia, Marsilea, Pistia, and other
aquatics. By BRANDIS noted to occur in a hot
spring in Burma at 92° F.

Note. As an exception, part of the 3 fiowers,
seldom all on one individual, are enclosed by a
more or less developed spathe; this seems to be a
teratological deviation (DE WILDE, 1961).

Vern. Ganggéng-leutik, S, ira-walaki, Wetar,
welak, amila, Alor; Philippines: aragdan-tamndi,
Ilk., rigmdu, Tag.

KEY TO THE VARIETIES

1. Leaves up to c. 4 cm by 114 mm, on either side
with up to 100 spiny teeth. Stem up to 144 mm
thick. Leaf sheaths up to 6 mm, auricles up to
2144 mm long. 2 Flowers up to 244 mm long.

var. graminea

1. Leaves 5-6 cm by 3-4 mm, on either side with
160-185 spiny teeth. Stem (1.6-)2-2144 mm
thick. Leaf sheaths (4-)7144(-104%4) mm,
auricles (2—)4-514 mm long. 2 Flowers c. 314
mm long. Robust plant var. robusta

var. graminea.—Fig. 2e, j.

Up to 50(—75) cm, lower internodes 1-10 cm, by
(0.6—)0.7-1.5 mm. Leaves (134-)2'44-3 4(-4)cm by
(0.6—)0.7-1(-1.4) mm, flat, rarely canaliculate;
apex obtuse (-rounded), rarely somewhat acute
(blade c. 3 mm below the tip 0.4-1 mm wide);
margin on either side with (40—-)60(—100) spiny
teeth; midrib c. 0.07—0.05 times as wide as the
blade; cavities narrow, occupying up to 14 of the
blade; septa indistinct. Sheaths (3—)4—-5(-6) by
(2-)3-4(—5) mm, on either side with (10—)15—20
(-30) spiny teeth; 3-8 teeth on the inner edge of
the auricle; auricles about straight, (l—)2(-2.6) by
0.5-1 mm, entire, rarely shallowly lobed. Flowers
often up to 3 together, in different stages, together
with a male, or male solitary. ¢ Flowers: anther
4(-2)-celled, elliptical, rarely ovate or oblong,
(1-)1.5(-2) hy (0.4-)0.7(-1) mm; ‘perianth lobes’
distinct, rounded to globular; ‘pedicel’ 0.2-0.5
mm, at anthesis 1-2.5 mm. 2 Flowers (1.6-)
2(-2.5) mm; ovary (0.6—)0.7(-1) by (0.2—)0.35
(—0.4) mm; style (0.4-)0.7(-1.2) mm, with 2(—3)
stigmas (0.4—-)0.8(-1.6) mm. Seeds, see under the
species.

Distr. efc. as under the species.

Note. The two collections made in the large
Matana and Towuti Lakes in Central Celebes
deviate slightly in having a condensed habit,
short (34-134 cm) rather coarsely dentate leaves with
few spines (20-35), a rather short (2- or 4-celled)
anther (c. 1 mm) and short stigmas only 0.3 mm
long (KJELLBERG 3820 and sine coll. 4, 31-10-09
respectively).

var. robusta DE WILDE, Act. Bot. Neerl. 10
(1961) 169.

Up to c. 50 cm, lower internodes c. 5 cm, by
(1.6-)2-2.25 mm. Leaves 5-6 cm by 3-4 mm, flat,
apex rounded to slightly acute (blade c. 3 mm
below the tip c. 1.2 mm wide); margin on either
side with 160-185 spiny teeth; midrib 0.07—-0.05
times as wide as the blade, cavities narrower than
the midrib; septa + distinct. Sheaths (4-)7'4
(-1014) by (4-)6.2(-8!14) mm, on either side with
(15—)30(—S0) spiny teeth, c. 7 teeth on the inner
edge of the auricle; auricles slightly falcate,
2-)4(-5.5) by (0.8—)1(-1.5) mm, rarely lobed.
Flowers solitary. 3 Flowers: anther 4-celled,
elliptic(-oblong), (1.5—)1.7-1.8(-2) by 0.8-1 mm;
‘perianth lobes’ distinct, rounded; ‘pedicel’ c.
0.2 mm, in anthesis 0.7-? mm. 2 Flowers c.
3.5 mm, ovary c. 0.9 by 0.35 mm; style c. 1 mm,
with 2(-1) stigmas c. 1.6 mm. Seeds unknown.

Distr. Malaysia: Lesser Sunda Islands (Wetar:
Lake Tihu), once found.

Ecol. The lake lies in the Eucalyptus forest at
c. 485-500 m.

Note. In a transverse section of the stem it
appears that the septa between the cavities are 2-4
cell layers thick; the outer cortex layer is also
well developed and consists of c. 5 cell layers.
This cortex structure is also found in N. marina,
some specimens of N. indica, and the African
N. horrida.

Dec. 1962] NAJADACEAE (de Wilde) 171

Excluded 332 is according to MERRILL, Sp. Blanc. (1918)
49 = Ceratopteris thalictroides BRONGN. (Pterido-
Najas obyoluta BLANco, Fl. Filip. ed. 2 (1845) = phyta).
460; ed. 3, 3 (1879) 66; NAves, Nov. App. (1880)

Excluded

Lemnopsis Zou. Syst. Verz. 1 (1854) 74 is according to BACKER and VAN STEENIS, cf. Fl. Mal. I, 4
(1949) 68 = Halophila (Hydrocharitaceae).

Najadeae comprised with Mique, FI. Ind. Bat. 3 (1856) 223 a large number of genera now partly
referred to Potamogetonaceae and partly to Hydrocharitaceae.

Ruppia has been referred to Najadaceae by ZOLL. Syst. Verz. 1 (1854) 74 = Potamogetonaceae or
Ruppiaceae.

—)

PRIMULACEAE (P. A. J. Bentvelzen, Leyden)

Annual or perennial herbs, erect, ascending or prostrate, less than 114 m high.
Leaves spirally arranged or alternate (often various in one plant), or opposite,
often in a basal rosette, exstipular, simple, sometimes lobed, penninerved. In-
florescences racemose, terminal (sometimes axillary) racemes or umbels, or
flowers in whorls, or solitary axillary. Bracts small or leafy. No bracteoles.
Flowers bisexual, actinomorphic (rarely zygomorphic), isomerous, in Mal. always
5-merous, often dimorphous in sexual organs. Ca/yx dentate or cleft, persistent,
sometimes leafy, rarely coloured (G/aux). Corolla connate, shallowly to deeply
cleft (free in Pelletiera), in bud often quincuncial or contorted, variously coloured
(absent in Glaux). Stamens inserted on the corolla, epipeta/ous, rarely alternating
with staminodes or their vestiges; anthers dorsifixed or versatile, sometimes
basifixed; cells opening with apical pores or latrorse, filaments free or connate.
Disk absent. Ovary superior (in Samolus semi-inferior), 1-celled with ~ ovules
on a free central placenta; style simple. Capsule mostly 5-valved (valves epi- or
alternisepalous) or 10-valved, sometimes irregularly bursting, or circumsciss.
Seeds mostly ~, often angular, small; embryo straight, endosperm present;

integuments 2.

Distribution. Genera 21 with approximately 900 spp., all over the world, but mainly developed
in the temperate and cold regions of the northern hemisphere; in the tropics mostly on the mountains.
The largest genera, Primula (incl. Androsace) with c. 500 spp. and Lysimachia with c. 150 spp. are almost
confined to the northern hemisphere and centre in the Sino-Himalayan region. In Malaysia and Melanesia
Primula extends across the equator and finds its southernmost stations in the Old World. Lysimachia
and Anagallis have a worldwide area. It is remarkable that the almost cosmopolitan species Samolus
valerandi L., which occurs in the surrounding continents of Asia and Australia and is widely distributed
in the Pacific (New Caledonia, Loyalty Is., Norfolk I., Chatham, Auckland Is., Kermadec, New Zealand,
and Easter I.), has never been found in Malaysia.

Ecology. Primulaceae have generally a temperate thermo-ecology, hence are montane to alpine in
the tropics. They are also generally heliophilous, preferring open habitats, dry or boggy. Few data are
available; the dimorphous Primulas are apparently adapted to cross-pollination. Monomorphic (i.e.
homostylous) Primulas are capable of effective self-pollination. In P. prolifera DOCTERS VAN LEEUWEN
(Verh. Kon. Ned. Ak. Wet. 31, 1933, 215) found only twice a Bombus on the flowers on Mt Pangrango;
self-pollination seems there the rule.

Dispersal. No special means of dispersal are found in the Malaysian Primulaceae and the only
positive record is the descent of seeds of Primula prolifera by rainwash along the trail on Mt Gedeh.
All representatives have capsules with many small seeds.

Phytochemistry. Many characteristic chemical compounds have been found in this family. Most
of them, nowever, were traced hitherto only in a few genera or species.

One remarkable exception can be made to this statement. It is very probable that saponins occur in
practically all species of Primulaceae (G. SCHNEIDER, Diss. Berlin, 1930). As in the case of most other
Dicotyledones, the sapogenins belong to the triterpenes. This was established for the saponins of the
roots and rhizomes of Primula veris L. and P. elatior (L.) Hii; both species contain primulagenin A,
C,)H;,O03, as an aglycone of the saponins. The saponins of the rhizomes and roots of Primulaceae
(e.g. Primula, Cyclamen) probably replace carbohydrates partially as carbohydrate reserve.

Concerning storage of reserve substances, Primulaceae are distinguished by other pecularities. In sub-
terranean organs some species accumulate beside saponins, reducing sugars, saccharose and starch a
variable amount of fructosanes. There exists a tendency in the family to replace accumulation of starch
by accumulation of fructosanes (J. Lys, Rév. Gén. Bot. 61, 1954, 154, 226, 300). Furthermore the
genus Primula is distinguished by the replacement of part of the mono- and disaccharides of the sub-
terranean organs by heptitols (volemitol) and heptoses (sedoheptulose) (J. BouGAULT & G. ALLARD,
C. R. Paris 135, 1902, 796; A. NorpAL & D. OklseTH, Acta Chim. Scand. 5, 1951, 1289). In this respect
Primulaceae are similar to Crassulaceae and Saxifragaceae. The seeds contain fatty oil and hemicelluloses
but no starch. The hemicelluloses (deposited in the membranes of the cells of the endosperm) belong
to the so-called amyloid type (in all species investigated). Amyloid is a polysaccharide, which stains
blue with iodine like starch; chemically amyloid is a galactoxyloglucan (P. Kooiman, Diss. T. H. Delft,
1959). Amyloid is also present in the seeds of Plumbaginaceae and Myrsinaceae.

(173)

174 FLORA MALESIANA [ser. I, vol. 6?

The secondary products of metabolism of Primulaceae are mostly phenolic in nature. Phenolics
are very common in the family. At the moment we may discern 4 different types of phenolic com-
pounds in the family.

(i) Leucoanthocyanins occur frequently (E. C. BATE-SmitH & N. H. LERNER, Biochem. J. 58, 1954,
156). It is probable that the so-called “‘Inklusen”’ (tannin-containing idioblasts with a solidified vacuol
content giving a red colour with HCI and vanilline), which were observed in some members of the
family (i.e. roots and rhizomes of Primula), contain polymeric leucoanthocyanins.

(ii) Many Primulaceae contain in their subterranean parts diglycosides of volatile, odorous phenols.
The sugar part of these heterosides is the disaccharide, primverose. There exists also an enzyme, prim-
verase, which is able to split the heterosides in primverose and the aglycones. The latter are mainly
derivatives of salicylic acid or of acetophenon. The presence of the heterosides and the corresponding
enzyme, explains the fact that the non-odorous fresh roots of many Primulaceae become odorous
gradually on drying or after injury. Heterosides of this type have been found in species of the genera
Primula, Dodecatheon, Lysimachia, and Anagallis (A. Goris, Industrie de la parfumérie 5, 1950, 121, 177).

(iii) The leaves and flowers seem to contain frequently flavonol glycosides. Rutin was isolated from
Lysimachia vulgaris L. and a compound called primulaflavonoloside (probably a dirhamnosid of kaemp-
ferol) was extracted from Primula veris L. In red, pink and blue flowers anthocyanins are common
(compare J. B. HARBORNE & H. S. A. SHERRATT, Nature 181, 1958, 25-27).

(iv) The glandular hairs of many Primulaceae excreet oily or granular, wax-like substances, termed
farina in the latter case. In most instances flavone, C,;H,)O., is the main constituent of farina (W. C.
BLASEDALE, J. Am. Chem. Soc. 67, 1945, 491; J. R. Hort. Soc. 72, 1947, 240; H. BRUNSwWIK, Sitz. Ber. Ak.
Wiss. Wien, M.-N. KI., Abt. 1, 131, 1922, 221). In species with yellow coloured farina the latter con-
tains oxyflavones beside flavone (primetin = 5,8-dioxyflavone: W. BAKER, J. Chem. Soc. 1939, 956;
5-oxyflavone: P. KARRER & G. SCHWAB, Helv. Chim. Acta 24, 1941, 297). Flavones have been demon-
strated to be present in the secretions of many species of Primula and in some species of Cortusa and
Dionysia.

A few species (Primula obconica HANCE, P. cortusioides L., P. sieboldii MORREN, P. mollis HooK.,
and Cortusa matthioli L.) contain allergenic substances in the secretion of their glandular hairs. The
latter (“Primeltoxin’’, primin) produce skin irritations in persons sensitive to the allergenes. Primin
is a fairly volatile, well crystallizing substance with a very high toxicity (skin irritation may be produced
by 1/50 to 1/100 of a milligram) (Br. BLock & P. KaArrer, Vierteljahrsschr. Naturf. Ges. Ziirich 72,
Beibl. No. 13, 1927, 1-26).

Concluding it may be stated amyloid and fatty oil in seeds and saponins in all organs are common
in Primulaceae. The family is furthermore characterized by the accumulation of different types of phenolic
substances; the distribution and chemistry of many of these compounds, however, have not yet been
studied intensively. If we search for chemotaxonomical relationships we find that similarities exist between
Primulaceae and Saxifragaceae (leucoanthocyanins, which are rare in herbaceous families, saponins,
volemit, sedoheptulose), between Primulaceae and Myrsinaceae (saponins, amyloid) and between
Primulaceae and some families of Centrospermae (leucoanthocyanins, saponins).—R. HEGNAUER.

Taxonomy. Primulaceae are affiliated with some other sympetalous families with epipetalous
stamens, superior ovary with free basal placenta, and 2 integuments; for example HANDEL—MAZZETTI
(Not. R. Bot. Gard. Edinb. 16, 1928, 70) postulated Lysimachia solanoides HM. to represent a possible
link with Myrsinaceae. The latter are, however, almost always woody plants and are (often pellucid-)
gland-dotted which rarely occurs in Primulaceae (Lysimachia, Anagallis spp., Primula spp.). According
to MetTcALFE & CHALK these substances are not yet analyzed. An other family, which agrees more
in habit, is Plumbaginaceae; the relation is sustained by anatomical characters brought forward by VAN
TIEGHEM (Bull. Mus. Hist. Nat. Paris 16, 1900, 131-135) but here is only one ovule, 5 styles, and an
obviously cymose inflorescence.

Subdivision. According to VALENTINE (Progress Study British Flora 1957, 80) five subfamilies can be
distinguished, viz:

(i) Androsaceae. Almost confined to the northern hemisphere comprising inter alia Primula and
Dedecatheon.

(ii) Cyclamineae. Central Europe and Mediterranean; only genus Cyclamen.

(iii) Lysimachieae. Worldwide, inter alia Lysimachia, Anagallis, and Trientalis.

(iv) Samoleae. Temperate worldwide. Only genus Samolus with c. 10 spp.

(v) Corideae. Mediterranean and Somaliland. Only genus Coris with 2 spp.

Generic delimitation. Far from the centre of the greatest development genera are often represented
by few species of distant alliance; these may appear so sharply separable that they are sometimes
assumed to deserve generic status in local or regional floras. In monographical studies it often appears,
however, that such ‘genera’ which are for example readily distinguishable in Europe are connected by
transitional species in Asia and can therefore not be maintained. Thus, for instance, KLATT and
HANDEL—MAZzzeTTI merged the genus Naumburgia MOENCH with Lysimachia in their monographs of
the latter genus. For similar reasons P. TAYLOR, in his revision of the tropical East African species of
Anagallis, came to the conclusion that Centunculus can not be maintained as a separate genus, although
in Europe one would conclude from the species growing there that they are distinct genera. A similar

Dec. 1962] PRIMULACEAE (Bentvelzen) 175

case is found with the pair Primula and Androsace which are in Europe and America separated by hav-
ing a distinctly salver-shaped or an almost rotate corolla respectively. This single character hasno great
taxonomic importance in itself as for example in Lysimachia the length of the corolla tube is as variable
as it is in Primula and Androsace. Besides transitional species are found in the Sino-Himalayan region
according to FRANCHET (1886) and Pax (Bot. Jahrb. 10, 1888, 133-136). PAx did not unite them because
he assumed that in the further evolution of the group the transitional species will disappear by which
the two segregates will further ‘diverge’. Such theoretical considerations are of course inadmissable
in phytography; we have to deal with the present situation and this is such that these two genera are
only separated by one character which does not hold. And although the consequence must be rather
impopular and nomenclaturally inconvenient because of the large number of species described in
Androsace, the two genera should be united from the scientific point of view. Although authors have
shrunk from putting the reduction into practice we should not yield to such inertia. If Androsace had
only a few species, hesitation would have been overcome more easily, as shown for example by the
uniting of Anagallis and Centunculus, Lysimachia and Naumburgia, etc.. O. KUNTZE (1891) reduced
Androsace to Primula.

Cytotaxonomy. According to DARLINGTON & WyLIe’s ‘Chromosome Atlas’ the basic chromosome
numbers do not yield a clear-cut picture for the whole family, although for instance in the genus Primula,
Bruun (Symb. Bot. Ups. 1, 1932, 1-239) could find a correlation between karyotypes and sections
based on morphological characters. In the genus Cyclamen the basic numbers are 5, 11, 12, and 17
(cf. DE HAAN & Doorensos, Med. Landbouwhogeschool Wagen. 51, 1951, 151). In Primula these
numbers are 8, 9, 10, 11, 12, and 13; in P. auricula L. the haploid number is surprisingly n = 31. Many
polyploids occur in this genus; P. kewensis W. WATS. is a classic example of an artificially obtained
good allotetraploid species (cf. W. F. Newton & C. PELLEWw, J. Genetics 20, 1929, 405-467).

Genetics. Much experimental research has been performed on species of Primula and Cyclamen.
Many Mendelian factors have been found responsible for characters of flower morphology and colour;
especially P. praenitens KER GAWL. (P. sinensis LINDL. 1821, non Lour. 1790) has intensively been
examined.

A. Ernst has devoted many decades of his life to study the problem of heterostyly in Primula, on
which further comment is given under the genus Primula.

From his work it appears that many interspecific crosses yield an entirely fertile progeny in which
differential characters appear often to be of Mendelian value only. Therefore, we can conclude that
such ‘species’ do not deserve specific status.

Cultivated. BACKER (Bekn. FI. Java (em. ed.) 8, 1949, fam. 180) mentioned, as cultivated in Java,
Cyclamen persicum L. and several Primulas (see there).

Note. By exception some genetical remarks are inserted because genetical work with Primula in-
volved taxonomical conclusions.—v. ST.

KEY TO THE GENERA

1. Leaves in a basal rosette. Flowers in one or more superposed whorls. Corolla lobes imbricate (quin-
cuncial) . . Srohonin Ge tao ey ast deca
. Leaves cauline. Flowers not i in " whorls. Corolla lobes contorted.
3 Corolla mostly yellow, pate white. Plants ae gland-dotted. eapsie dehiscing with valves or
irregularly bursting . . i . . . 2. Lysimachia
2. Corolla white (in Mal.). Plant ‘not ‘gland: dotted. Capsule circumsciss. Very small plant.
1. Anagallis

176 FLORA MALESIANA [Secs TevoliiG?

1. ANAGALLIS

LINNE, Gen. Pl. ed. 5 (1754) 189; Sp. Pl. (1753) 148; KNurn, Pfl.R. Heft 22
(1905) 321; P. TAyLor, Kew Bull. (1955) 321. — Centunculus LINNE, Gen. PI.
ed. 5 (1754) 135; Sp. Pl. (1753) 116; KNuru, /.c. 334. — Micropyxis DuBy in DC.
Prod. 8 (1844) 71. — Fig. 1.

Annual or perennial, prostrate or decumbent, small herbs. Stem terete or
angled, not rarely winged. Leaves many, entire, opposite at least at the base
of the stem, spirally arranged, or whorled. Flowers solitary axillary, sessile or
pedicelled. Calyx 5-cleft, campanulate. Corolla rotate to urceolate, the lobes
large or small. Stamens 5, free from each other; filaments often villous. Capsule

circumsciss. Seeds co, planoconvex.

Distr. About 25 spp., all over the world, 14 in tropical-montane Africa, a cosmopolitan species once
found in Malaysia (Timor).

Ecol. A characteristic temperate to warm-temperate genus; according to P. TAYLor, /.c., it occurs
in tropical East Africa between 2000-4000 m in bogs and by streamsides and in the Rhodesia-Angola
region in southern tropical Africa between 1000-1500 m in marshes and on seasonally flooded ground.

Taxon. P. TAyLor, /.c., found no reason to maintain the genus Centunculus. He distinguished three
subgenera, viz subg. Anagallis, subg. Centunculus (L.) P. TAYLOR, and subg. Jirasekia (SCHMIDT) P.

Taytor. His new conspectus makes a sound impression.

1. Anagallis pumila Sw. Prod. Veg. Ind. Occ. 1
(1788) 40; Bru. Fl. Austr. 4 (1869) 270; BAILEy,
Queensl. FI. 3 (1900) 946; KNuTH in Pfl. R. Heft
22 (1905) 331, f. 71; Ewart & Davies, Fl. North.
Terr. (1917) 217; GAmBLE, FI. Madras 4 (1921)
747; HutcH. & Daz. Fl. W. Trop. Afr. 2
(1931) 184; SreeNn. Bull. Jard. Bot. Btzg III, 13
(1934) 236; P. TayLor, Kew Bull. (1955) 342,
345.—Centunculus pentandrus R. Br. Prod.( 1810)
427; Mig. Fl. Ind. Bat. 2 (1859) 1004.—Centun-
culus indicus Roy er, Ul. Bot. Him. (1839) 310,
nomen.—Micropyxis pumila Dusy in DC. Prod.
8 (1844) 71.—Centunculus tenellus Dusy, I.c. 72;
Hook. f. Fl. Br. Ind. 3 (1882) 506.—Micropyxis
tenella WiGHT, Ic. 4 (1850) t. 1585.—Centunculus
pumilus O.K. Rev. Gen. Pl. 3 (1891) 193.— A.
nana SCHINZ, Vierteljahrschr. Naturf. Ges. Ziirich
55 (1910) 244.—Fig. 1.

Slender nearly erect herb up to 25 cm, not
rooting at the nodes. Stem terete, scarcely winged.
Leaves \ess than twice as long as wide, more or
less ovate, a few mm long. Flowers towards the
apex in the leaf axils, short-pedicelled. Calyx
cleft to the base, lobes lanceolate-oblong, acute,
mucronate. Corolla white, as long as the calyx
or exceeding it, subpersistent, deeply S-cleft; tube
wide, c. 2/5 mm high; lobes acuminate, c. 2 mm
long, the insertions separated by a sinus. Stamens
as long as the corolla; filaments glabrous, c. 1 mm;
anthers c. 1/8 mm. Pistil c. 2 mm long, style
somewhat shorter than the ovary, with a flattened
stigma. Ovary with granular glands. Capsule as
high as the calyx.

Distr. Africa, India, N. Australia, South and
Central America; in Malaysia: Timor (once col-
lected, WALSH 448a).

Ecol. Damp grounds, swamps, efc., up to 2100

Fig. 1. Anagallis pumila Sw. a. Habit, nat. size,

b. flower, * 5,c.corollalaid open, < 5, d. dehiscing
fruit, * 5 (TH. THOMSON s.n. in L).

m. The only Malaysian material was collected as
a mixture with Lobelia heyniana R. & S. but un-
fortunately no altitude was indicated.

Note. The Timor material belongs to var.
pumila; two other varieties occur in Africa (P.
TAYLor, /.c.).

Excluded

Anagallis arvensis LINNE, Sp. Pl. (1753) 211;
Mia. FI. Ind. Bat. 2 (1859) 1004, almost ubiquitous,
has as yet not been found introduced as a weed
in Malaysia.

Dec. 1962] PRIMULACEAE (Bentvelzen) _ 177

2. LYSIMACHIA

Tourn. ex LINNE, Gen. Pl. ed. 5 (1754) 72; Sp. Pl. (1753) 146; Kiatr, Abh.
Naturw. Ver. Hamb. 4, 4 (1866) 1-45, t. 1-24; KNuTH, Pfl. R. Heft 22 (1905)
256; HAND.—Mazz. Not. R. Bot. Gard. Edinb. 16 (1928) 51-122; Pfl. Areale 2,
5 (1929) 39-41, maps 44-49; Ray, Illinois Biol. Monogr. 24, pts 3-4 (1956)
13.— Cerium Lour. Fl. Coch. (1790) 136; ed. Willd. (1793) 167, cf. MERR. Comm.
Lour. (1935) 300.—Lubinia VENT. Hort. Cels. (1800) 96; Pax in E. & P. Pfl.
Fam. 4, 1 (1897) 112.—Orescia Retnw. Syll. Pl. 2 (1825) 15, cf. STEEN. Bull.
Bot. Gard. Btzg III, 17 (1948) 458.—Bernadina BAubo, Ann. Sc. Nat. IH, 20,
Bot. (1843) 349, nomen. —Fig. 2.

Erect ascending or prostrate herbs. Leaves cauline, opposite, spiral, alternate,
or whorled (sometimes various in one specimen), often glandular-punctate,
margin entire or not. Flowers in terminal or subterminal racemes, or solitary
axillary. Calyx deeply 5-fid. Corolla contorted in bud, S-cleft, yellow or white
(sometimes with purple background). Stamens sometimes largely connate with the
corolla lobes; anthers basifixed or versatile, opening with an apical pore or with
lateral slits. Ovary globose, style-tip as high as the anthers. Capsule about as high

as the calyx, 5—valved or irregularly bursting. Seeds numerous, testa crustaceous.

Distr. About 150 spp., all over the world save in northern Siberia, Greenland, northern Canada,
Alaska, and New Zealand. According to HANDEL—MAzzZETTI there is a distinct centre of development
in the Sino-Himalaya where c. 80 spp. occur. With regard to these numbers it should be remembered
that HANDEL—MAzzeTTI employed a rather narrow specific concept. Of the 8 spp. which occur in Malaysia
four are only found along its northern border: L. peduncularis WALL. ex Kurz in the Langkawi Is.,
L. mauritiana LAMK in the Batan Is. (between Luzon and Formosa), L. capillipes HEMSL. in Luzon and
L. sikokiana Mia. in the Philippines. All occur also in East or SE. Asia. The tropical-montane species
have followed either the Sumatra track (L. laxa, L. decurrens, L. montana, L. japonica var. japonica)
or the Luzon track (L. capillipes, L. sikokiana, L. japonica var. papuana, L. decurrens). Large-distance
disjunctions occur in the distributional areas of L. decurrens (fig. 10) and L. mauritiana (fig. 9), the
first a montane species, the second largely occurring on the seashore.

Ecol. Largely a genus of temperate to warm-temperate thermo-ecology, but some sections contain
tropical lowland species and must be defined as eurytherm. All species occur in Malaysia above 1000 m,
except L. mauritiana LamK and L. peduncularis WALL. ex Kurz. The first one is in Malaysia and the
Pacific almost bound to coastal limestone rocks; it is surprising that it ascends the coastal hills in East
China to 70—400(—?700) m.

Taxon. Kiatr, KNuTH, and HANDEL—MAzzetTI, who all monographed Lysimachia, are unanimous
about a broad concept, including for example segregate genera as Lubinia VENT., Coxia ENDL., Steironema
RAFIN., and Naumburgia MOENCH.

HANDEL—MAzzeTTI has subdivided the genus into five subgenera; the Malaysian species belong to
subg. Lysimachia (Eu-lysimachia) and subg. Palladia (MoENCH) H.—M. (L. mauritiana and L. decurrens).
Within the subgenera he further distinguished sections, and series. The sections seem to me acceptable
but the finer division into series is not satisfactory: for instance in sect. Apodanthera subsect. Ramosae
there are two series, viz Valvatae and Evalves based on the dehiscence of the capsule by valves and
irregular bursting respectively. However, this character may vary within species. Another example
showing that the distinction of series is going too far is found in sect. Nummularia in which L. japonica
is placed in ser. Japonicae and L. debilis in ser. Debiles, though there is no doubt in my opinion that
they are conspecific.

Note. In Lysimachia the stamens are always, in degree, adnate to the corolla tube. Moreover they
are mutually connate in a ring or low tube, or even in a high tube; in some flowers of several species
I have found them occasionally entirely free from each other. Such free stamens have a basally dilated
filament.

178 FLORA MALESIANA [ser. I, vol. 6?

KEY TO THE SPECIES

1. Leaves never spiral, opposite or subopposite, the apical ones not unfrequently alternate, in Mal.
broad-ovate to orbicular, often abruptly very short-apiculate. Flowers solitary axillary. Pedicels shorter
than their sustaining leaf. Anthers versatile, shorter than the filaments. Prostrate herb. 6. L. japonica

1. Leaves spiral, rarely a few subopposite.

2. Flowers in terminal or leaf-opposed, 1—5-flowered, stunted, often pseudo- or subumbellate racemes.

Leaves usually ovate-acute . Rake te ae ae 4. L. montana
2. Flowers either solitary axillary or in | many- flowered, ferminal, ‘bracteate racemes.
3. Bracts almost filiform (except sometimes the lowest). Corolla white or pale purple, in Mal. with
prominent, elliptic red glands in the tissue. Stamens conspicuously exceeding the corolla.
8. L. decurrens
3. Flowers subtended by normal leaves or leafy bracts. Stamens not exceeding the corolla.
4. Filaments halfway adnate to the corolla. Leaves ae ee blunt or bluntish. Flowers white,
yellowish in the dried state. 3 .. . 7. L. mauritiana
4. Filaments adnate to the corolla at the base only. ‘Leaves acute or acuminate, not spathulate.

Flowers yellow.

5. Corolla deeply cleft, distinctly exceeding the calyx. Anthers as long as or longer than the filaments.
6. Leaf venation between the nerves, especially beneath, distinctly prominent-reticulate in the

dried state

6. Venation not distinctly prominent- -reticulate.

1. L. sikokiana

7. Stamens as long as the corolla. Anther cells opening with an apical pore. Flowers confined to
the upper part of the stem. Leaves often truncately cut away at the base then decurrent on

one or both sides. Capsule shorter than the calyx. Lax decumbent herb.

2. L. capillipes

7. Stamens shorter than the corolla. Anther cells opening with lateral slits. Flowers present almost
along the entire stem. Leaves lanceolate. oe somewhat exceeding the calyx. Erect rather

robust plant.

3. L. laxa

5. Corolla cleft halfway, about as slong: as or shorter than the calyx. Anthers shorter than the

filaments .

1. Lysimachia sikokiana Mig. Ann. Mus. Bot.
Lugd. Bat. 3 (1867) 121; FRANcH. & Sav. En
Pl. Jap. 1 (1875) 302; ENGL. Bot. Jahrb. 6 (1885)
64; HemsL. J. Linn. Soc. Bot. 26 (1889) 51;
KNuTH, Pfl. R. Heft 22 (1905) 270; HAND.-MAzz.
Not. R. Bot. Gard. Edinb. 16 (1928) 74; SASAKI,
Cat. Govt. Herb. (1930) 402; MASAmMuUNE, FI.
Stud. Yakush. (1934) 360; HARA, En. Sperm. Jap.
1 (1948) 88; Makino, Ill. Fl. Jap. (1954) 233.—
L. simulans Hemsv. J. Linn. Soc. Bot. 26 (1889)
57; SASAKI, Cat. Govt. Herb. (1930) 402.—L.
ramosa var. typica KNUTH, Pfl. R. Heft 22 (1905)
271, p.p., excl. typ.; MERR. Philip. J. Sc. 2 (1907)
Bot. 298; ibid. 5 (1910) Bot. 377; En. Philip. 3
(1923) 275; Merr. & CHUN, Sunyatsenia | (1930)
77; STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 238,
p.p.—L. fragrans HAyaTA, J. Coll. Se. Univ.
Tokyo 30 (1911) 175, ex descr.; HAND.—MAzz.
Not. R. Bot. Gard. Edinb. 16 (1928) 74; ibid. 16
(1931) 167; Merr. Sunyatsenia 1 (1934) 203; J.
Arn. Arb. 19 (1938) 61; Merr. & CHUN, Sunyat-
senia 5 (1940) 156, syn. nov.—L. ardisioides
MASAMUNE, J. Soc. Trop. Agr. Taiwan 4 (1932)
302, ex descr., syn. nov.—L. garrettii FLETCH.
Kew Bull. (1936) 41.—Fig. 2a—d.

Erect or decumbent herb, rooting at the base,
stem terete at the base, 30-60 cm. Leaves spirally
arranged, elliptic-lanceolate, acute, sometimes
shortly acuminate, base acute to rounded, decur-
rent along the stem, 214-10 by 1—3'4 cm, both
surfaces glabrous, no glands; upper side very dark
brown when dry, midrib and nerves depressed,

5. L. peduncularis

veins more or less prominent; underside glaucous,
midrib, nerves, and veins prominent, nerves
ascending, passing into a marginal vein; margin
entire or undulate-crenate; lower leaves scale-like
reduced; petiole 44-1'4 cm. Flowers axillary,
solitary, confined to the upper part of the plant.
Pedicels as long as the sustaining leaves, drooping
or + recurved during anthesis, in fruit obliquely
erect. Calyx lobes ovate, acuminate, 3-7 mm, the
margins sometimes with stalked glands. Corolla
yellow, deeply cleft, 10-15 mm long, lobes elliptic-
oblong, acute at the apex. Stamens only adnate at
the base, whether or not free from each other,
4-8 mm; anthers oblong, basifixed, 3-5 mm long,
the cells opening with an apical pore. Style-tip
as high as the anthers. Capsule 5-valved to the
base, but often irregularly bursting.

Distr. S. Japan, Ryukyu Is., southern con-
tinental China (Yunnan, Kwang-tung), N. Siam,
Hainan, Formosa; in Malaysia: Philippines
(Luzon: Bontoc; Mindoro; Negros; Camiguin de
Misamis; Mindanao). Fig. 3.

Ecol. Mossy forests, damp ravines, 1200-2300 m.

Vern. Philip.: /upo-lupo, C. Bis., tasig-tauig, Bag.

Notes. This species has been confused with L.
laxa Baupo (L. ramosa WALL. ex DuBy) by
KNUTH and by MERRILL.

It differs from WALLICH’s type in many charac-
ters: less flowers and these confined to the upper
part of the plant; stamens with apical pores; ovate
leaves discoloured in the herbarium with visible
veins and not glandular-punctate.

Dec. 1962]

PRIMULACEAE (Bentvelzen)

Fig. 2. Lysimachia sikokiana Mig. a. Habit, x lA,
b. flower, nat size, c. androecium, 2, d. flower,
stamens and petals removed.—L. /axa BAuDo.
e. Flower, nat. size, f. part of inside of corolla
showing insertion of stamens, * 2, g. dehiscing
fruit, 2 (a-d MERRILL BS 4383, e-g VAN
STEENIS 9109).

Fig. 3. Distribution of Lysimachia sikokiana Mia.

After HANDEL—MAZzzETTI’s revision had been
published Merritt identified the Philippine
specimens to belong to L. fragrans HAYATA
which HANDEL—MAzzeTTI already pointed out
may run into L. sikokiana Miq. I have seen
MIQUEL’s type and MERRILL’s specimens, though
less robust than the Japanese ones, are doubtless
conspecific. I have not seen HAyYATA’s type, but
material identified by HANDEL—-MAzzeTTI as L.
fragrans proved to belong to L. sikokiana. A
topotype of L. ardisioides MASAMUNE (TANAKA
13561) has the same habit as the Philippine
plants.

2. Lysimachia capillipes Hemsv. J. Linn. Soc. Bot.
26 (1889) 48; KNuTH, Pfl. R. Heft 22 (1905) 270;
PETITMENGIN, Bull. Ac. Géogr. Bot. 17 (1907) 223;
Merr. Philip. J. Sc. 7 (1912) Bot. 93; En. Philip.
8 (1923) 274; HAND.-MaAzz. Not. R. Bot. Gard.
Edinb. 16 (1928) 74; J. Arn. Arb. 15 (1934) 294;
STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 237.—
Andrachne cavalerieri LévL. in Fedde, Rep. 12
(1913) 187.—L. cuspidata (non BL.) HAND.—
Mazz. Not. R. Bot. Gard. Edinb. 16 (1931) 167,
pro specim. philip.

Ascending or prostrate herb, branched only at
the base, 20-60 cm; stem thin, angular or ridged.
Leaves alternate and spirally arranged, mem-
branous, ovate-lanceolate, acute, sometimes a-
cuminate, mucronate, 114-4 by 4-2'4 cm; base
unequal, acute, obtuse or truncate, decurrent
along the petiole, margin entire or undulate-
crenate; midrib and nerves on the upper surface
slightly depressed; prominent below, veins hardly
visible; no marginal vein. Flowers solitary, axil-
lary, confined to the upper part of the stem.
Pedicels 114-414 cm, obliquely erect during an-
thesis then drooping. Ca/yx lobes lanceolate,
acuminate, 3-5 mm. Corolla yellow, deeply cleft,
7-10 mm; lobes oblong; broad-triangular at apex.
Stamens as long as the corolla, filaments very
short, at the base adnate with the corolla tube,
also connate with each other in a basal ring.
Style-tip as high as the stamens. Capsule S-valved,
shorter than the calyx.

Distr. China (Yunnan, Kweichow, Szechuan,

FLORA MALESIANA

[ser. I, vol. 6?

eX Tybee

Sak

s\ ~
ste

: Pa) ze cae (2 :
Poe eens

Fig. 4. Distribution of Lysimachia capillipes
HEMSL.

\
» S| “
%&
QU:
\a +

Hupeh) and Formosa; in Malaysia: Philippines
(North Luzon). Fig. 4.

Ecol. In damp thickets, along streams, 1300—
1500 m, in China 1200-2000 m.

Vern. Philip.: osiak, Ig.

Notes. Merritt (1912) determined some
Philippine specimens with KNUTH’s key, which
is rather useless, especially for sect. Alternifoliae
KNutTH, under which this species was arranged.
By elimination MERRILL reached (ex descr.) the
conclusion that his material belonged to L.
capillipes. 1 have now seen the type of L. capillipes
and can confirm MERRILL’s deduction.

Later HANDEL—MAZzZETTI tentatively identified
some other Philippine specimens collected by
CLEMENS (16264a, 16459, 18783) as L. cuspidata
BL. (= L. montana), but they really represent L.
capillipes.

3. Lysimachia laxa BAUDOo, Ann. Sc. Nat. II, 20,
Bot. (1843) 347.—L. ramosa WALL. [Cat. (1828)
n. 1490, nomen] ex DuBy in DC. Prod. 8 (1844)
65; De VrieseE in Miq. Pl. Jungh. (1851) 88;
Tuwaites, En. Pl. Zeyl. (1860) 172; Miq. Sum.
(1861) 246; KLatr, Abh. Naturw. Ver. Hamb. 4,
4 (1866) 31, t. 17; Hook. f. Fl. Br. Ind. 3 (1882)
503, incl. var. zeylanica; Trim. Fl. Ceyl. 3 (1895)
65; Koorp. Nat. Tijd. Ned. Ind. 60 (1901) 273;
KNuTH, Pfl. R. Heft 22 (1905) 271, typ. incl. sed,
excl. yar. typica KNUTH pro parte et var. grandi-
flora FRANCH.; Koorp. Jungh. Gedenkb. (1910)
186; Exk. Fl. Java 3 (1912) 35; Fl. Tjib. 3 (1918)
30; HAND.-Mazz. Not. R. Bot. Gard. Edinb. 16
(1928) 75; STEEN. Bull. Jard. Bot. Btzg Ill, 13
(1934) 238, p.p.; Merr. J. Arn. Arb. 19 (1938)
61; Backer, Bekn. FI. Java (em. ed.) 8 (1949)
fam. 180, p. 3; INGRAM, Baileya 8 (1960) 95.—L.
floribunda Zo... & Mor. Nat. & Geneesk. Arch.
Neérl. Ind. 2 (1845) 575; Hassk. Flora 30 (1847)
600; WALp. Ann. Bot. Syst. 1 (1848) 494; ZoLL.
Syst. Verz. 3 (1855) 59.—Fig. 2e—g.

Erect herb, sometimes decumbent at the base,
to a 44 m high; stem angular, often strongly
branched. Leaves spirally arranged, lanceolate,
acute at both ends, sometimes acuminate at
apex, 3-11 by 14-2 cm; upper surface often with
lax thin hairs, midrib depressed, further venation
hardly visible; underside glabrous, midrib pro-

minent, nerves slightly so, ascending, veins in-
visible; both sides provided with scattered flat
brown glands. Petiole 0-1 cm, decurrent along the
stem. Flowers axillary, solitary, all along the stem.
Pedicels nearly as long as the leaves, obliquely erect.
Flowers during anthesis drooping; in fruit erect
ascending. Calyx lobes elliptic-obovate, short-
acuminate, 4-8 mm; margin sometimes provided
with stalked glands. Corolla yellow, deeply cleft,
somewhat longer than calyx, 5-10 mm, lobes
elliptic-oblong, acute at the apex or rounded.
Stamens 2-5 mm; filaments adnate to the corolla
only at the base, further connate with each other
in a small basal ring (in some Indian specimens
the filaments are further free); anthers as long as
the filaments incl. the ring, basifixed, oblong,
sagittate at the base, opening with lateral slits.
Style nearly as high as the corolla. Capsule ex
ceeding the calyx, 6-10 mm, 5-valved to the base,
often irregularly bursting.

Distr. Ceylon, Eastern Himalaya, Southern
China, Indo-China, and Thailand; in Malaysia:
Sumatra, Java, and Lesser Sunda Is. (Bali, Lom
bok, and Sumbawa). Fig. 5.

Fig. 5. Distribution of Lysimachia laxa BAuDo.

Ecol. In forests in sunny spots along trails, on
volcanic walls, 1000-3000 m.

Notes. WALLICH’s type of L. ramosa WALL.
was first validated by BAupo with a succinct
description in a precursory study for a revision
of the Primulaceae intended for DE CANDOLLE’s
Prodromus under the name L. /axa; this name was
not taken up by Dupy a year later in DC. Prod.
vol. 8. BAuDo’s paper has been neglected by later
monographers.

Hooker f. distinguished a var. zeylanica to
which Koorpers (1912) referred the specimens
from Java. HANpEL—MAzzeTri has correctly
pointed out, however, that it is impossible to up-
hold this variety because the differences concerned
(size of the flowers, shape of the corolla lobes,
etc.) have proved to be highly variable.

KNUTH (1905) and MERRILL (1923) have refer-
red some Philippine specimens to L. ramosa which
species does not occur in the Philippine Is.; all
these sheets belong to L. sikokiana.

Dec. 1962]

PRIMULACEAE (Bentvelzen)

181

4. Lysimachia montana (REINW.) BAKH. f. comb.
nov.—Orescia montana REINW. Syll. Pl. 2 (1825)
15.—L. cuspidata BL. Bijdr. 14 (1826) 737; DuBy
in DC. Prod. 8 (1844) 67; Mor. Syst. Verz.
(1846) 45, incl. var. glabra; DE VRIESsE in Miq.
Pl. Jungh. (1851) 88; Mie. Sum. (1861) 245, non
Kiatt, Abh. Naturw. Ver. Hamb. 4, 4 (1866) 36,
quae est L. klattiana HANcE, J. Bot. 16 (1878)
236; Mig. Ann. Mus. Bot. Lugd. Bat. 4 (1868)
144; O. K. Rev. Gen. Pl. 2 (1891) 397; KNUTH,
Pfl. R. Heft 22 (1905) 272, incl. var. glabrescens
et hispida; Koorp. Exk. Fl. Java 3 (1912) 35; Fl.
Tjib. 3 (1918) 31; ReENDLE, J. Bot. 63 Suppl.
(1925) 58; HANp.-Mazz. Not. R. Bot. Gard.
Edinb. 16 (1928) 75; ibid. 16 (1931) 167, excl.
specim. philip.; STEEN. Bull. Jard. Bot. Btzg III,
13 (1934) 237; ibid. III, 17 (1948) 458; BACKER,
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 180, p. 3.
L. uliginosa (non Bu.) KLatt, Abh. Naturw. Ver.
Hamb. 4, 4 (1866) 36, t. 20.—L. platyphylla
Merr. Contr. Arn. Arb. 8 (1934) 134, t. 12.—L.
chapaensis Merr. J. Arn. Arb. 20 (1939) 350,
syn. nov.

Fig. 6. Distribution of Lysimachia montana
(REINW.) BAKH. f., dotted line is demarcation in
Sumatra between the two varieties.

var. montana.—Orescia montana REINW.—L.
cuspidata BL.—L. uliginosa (non BL.) KLATT.
Decumbent or erect herb up to 14 m high;
stem angular, twisted. Leaves alternate, spirally
arranged or sometimes opposite, narrow to broad
elliptic, acute, mucronate, sometimes slightly
acuminate, base acute to rounded, 1-6 by 0.5—2.5
cm, margin entire or slightly dentate; whether or
not hairy at one or both sides; midrib depressed
above; prominent underneath as are the ascending
nerves, the latter passing into a marginal vein;
petiole c. 1 cm. Flowers in terminal or subterminal
1—5-flowered racemes, exceeding the leaves;
peduncle c. 1 cm. Pedicels 1-3 cm, each in the
axil of a small leafy bract. Calyx lobes 4-7 mm,
lanceolate, acute-acuminate, sometimes hairy on
midrib, whether or not glandular-punctate.
Corolla yellow, cleft halfway or deeper, 6-13 mm
high, lobes obtuse or rounded, with stalked glands
on the margin. Stamens c. 5 mm; filaments as long
as anthers, adnate with the corolla only at the

base, free from each other; anthers oblong,
sagittate at the base, basifixed, opening with
lateral slits. Style c. 7 mm. Capsule somewhat
smaller than the calyx, 5-valved.

Distr. Indo-China (Laos: Kerr 21026, 21038)
and Malaysia: Sumatra and Java (Diéng to
Tengger Mts); Flores (not seen.) Fig. 6, partly.

Ecol. Primary forest, along trails, in shaded
places, on old lava-streams, 1000-2500 m.

Notes. Dr. BAKHUIZEN VAN DEN BRINK Jr sug-
gested that the epithet montana has priority over
cuspidata.

HANDEL—MAzzeTTI (1931) referred some Philip-
pine specimens collected by CLEMENS to L. cus-
pidata; they rightly belong to L. capillipes HEMSL.

MIQuEL (1868) pointed out that Kiatr had
made an erroneous interpretation of BLUME’s
species: what KLAtTr described as L. uliginosa BL.
(= L. japonica THUNB.) is L. montana, and that
the Asiatic specimens which he referred to L.
cuspidata belonged to another species (in 1878
described as L. klattiana HANCE). KLATT was
curiously sensitive to MIQUEL’s correct criticism
(cf. Linnaea 37, 1873, 502).

Hitherto this species was thought to be confined
to Malaysia, although KNUTH suggested that the
Chinese L. trichopoda FRANCH. might represent
only a variety of it; according to HANDEL—MAz-
ZETTI, however, this species has stamens opening
by apical pores. Also var. platyphylla to which L.
chapaensis is here reduced occurs in continental
Southeast Asia.

I see no reason to distinguish varieties based
on hairiness of the leaves. Within specimens from
a local population some were glabrous, others had
leaves hairy to different degree, and of different
nature, sometimes only at the underside, sometimes
only at the upper side. Often the hairiness was
strigose on the upper surface but villous beneath.

var. platyphylla (MERR.) BENTVELZEN, Stat. novy.—
L. platyphylla Merr.—L. chapaensis MERR.

Differs from the type variety by larger leaves,
5-10 by 3-6 cm, larger corolla (12-16 mm), anthers
much longer (5 mm) than filaments (2 mm),
which are shortly connate at the base.

Distr. Indo-China (Tonkin) and Malaysia:
N. Sumatra southwards to the boundary between
East Coast and Tapanuli. Fig. 6, partly.

Ecol. Flat forest ridges along stream margins or
on rocks, 1000-1500 m.

Vern. Dukut-dukut lutu, dukut lohut, Toba.

Note. MERRILL distinguished L. platyphylla by
the very much larger, broader leaves, shorter
pedicels, rounded petals, and glandular-punctate
sepals. The latter two characters are, however,
also found in Javanese specimens of L. montana,
in which besides the length of the pedicels is very
variable. But indeed the North Sumatran plants
have distinctly larger leaves and also very large
anthers, a character not mentioned by MERRILL,
and I have accepted it as a distinct variety. To
this variety I have also referred L. chapaensis
Merr. from Tonkin (PETELOT 6347). MERRILL
thought this species to be allied to L. ramosa

182

FLORA MALESIANA

[ser. I, vol. 62

WALL. but in my opinion the type is clearly the
same as L. montana var. platyphylla.

5. Lysimachia peduncularis WALL. [Cat. (1828) n.
1489, nomen] ex Kurz, J. As. Soc. Beng. 46, ii
(1877) 219; Hook. f. Fl. Br. Ind. 3 (1882) 504;
KNuTH, Pfl. R. Heft 22 (1905) 271; Cras, Kew
Bull. (1911) 406; Haines, Bot. Bihar & Orissa 4
(1922) 506; HAND.-Mazz. Not. R. Bot. Gard.
Edinb. 16 (1928) 77; Bonati, Fl. Gén. 1.-C. 3
(1930) 760; FLETCHER, FI. Siam. En. 2 (1938) 324;
HENDERSON, J. Mal. Br. R. As. Soc. 17 (1939)
53.—L. pierrei PETITMENGIN, Bull. Ac. Géogr.
Bot. 18 (1908) 337; HAND._MAzz. Not. R. Bot.
Gard. Edinb. 16 (1928) 77; Merr. J. Arn. Arb.
19 (1938) 61, ex descr. syn. nov.

Decumbent herb, branched above the middle,
rooting at the base. Stem up to 30 cm, angular,
thin, narrowly winged. Leaves spirally arranged,
lanceolate, acute at both ends, 114%4-5 by %-1%
cm, strigillose, faintly gland-dotted, midrib at the
underside slightly prominent, nerves ascending,
veins invisible; petiole 14-1 cm. Flowers yellow,
axillary, solitary, confined to the upper part of the
plant, during anthesis drooping. Pedicels somewhat
longer than the leaves, in fruit erect ascending.
Calyx lobes oblong, shortly acuminate, 3—S mm.
Corolla cleft halfway, 244-4 mm high, lobes ellip-
tic, acute. Stamens free from each other, as long as
the corolla, only at the base adnate to it; filaments
2-3 mm; anthers c. 14 mm, oblong, sagittate at
the base, basifixed, cells latrorse. Style 2-3 mm.
Capsule shorter than the calyx, 5-valved, often
irregularly bursting.

Distr. India (Bihar), S. Burma, Siam, South
Indo-China; in Malaysia: Malay Peninsula (Lang-
kawi Is.), once collected (HENDERSON SF 21371).
Fig. 7.

t

Fig. 7. Distribution of Lysimachia peduncularis
WALL. ex Kurz.

Ecol. In coastal limestone vegetation; in Burma
to 420 m; according to HAINes in Bihar in damp
places in bamboo forests.

Note. PETITMENGIN compared his L. pierrei
with L. ramosa WALL. ex Dusy (= L. laxa
Baubo), L. floribunda ZoLL. & Mor. (= L. laxa
Baupbo), and L. capillipes HemsL. and found it
to be distinct. According to HANDEL—MAZZETTI
and MERRILL it will come very near to L. pedun-

cularis. The only difference is that L. pierrei is
glabrous but as the indument in this genus is very
variable this cannot warrant specific separation.

6. Lysimachia japonica THUNB. FI. Jap. (1784) 83;
LAMK, Tabl. Enc. Bot. 1 (1792) 440; THuns. Ic.
Pl. Jap. (1794) t. 16; DuBy in DC. Prod. 8 (1844)
67; Step. & Zucc. Fl. Jap. 2 (1846) 139; Kvarr,
Abh. Naturw. Ver. Hamb. 4, 4 (1866) 34, t. 19:
Mia. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 121;
Bru. Fl. Austr. 4 (1869) 269; FRANCH. & SAV.
En. Pl. Jap. 1 (1875) 303; Hook. f. Fl. Br. Ind. 3
(1882) 505; Hems-. J. Linn. Soc. Bot. 26 (1889)
53; CouteTT, FI. Siml. 1 (1902) 302; KNutH, Pfl.
R. Heft 22 (1905) 262; Mats. & HayatTa, J. Coll.
Sc. Univ. Tokyo 22 (1906) 222; Crais, Kew Bull.
(1911) 406; Koorp. Exk. Fl. Java 3 (1912) 34;
Fl. Tjib. 3 (1918) 30; Merr. En. Philip. 3 (1923)
275; HAND.-Mazz. Not. R. Bot. Gard. Edinb.
16 (1928) 96; BonatTi, FI. Gén. I.-C. 3 (1930) 759;
STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 237;
H. J. Lam, Blumea 5 (1945) 585; Hara, En.
Sperm. Jap. 1 (1948) 86, incl. var. thunbergiana et
var. Subsessilis; BACKER, Bekn. FI. Java (em. ed.)
8 (1949) fam. 180, p. 3.—L. maculata R. Br.
Prod. (1810) 428; SpreNG. Syst. Veg. 1 (1825)
572; Dusy in DC. Prod. 8 (1844) 66.—L. debilis
WALL. in Roxb. FI. Ind. 2 (ed. Wall.) (1824) 25;
Don, Prod. Fl. Nepal. (1825) 83; BAupo, Ann. Sc.
Nat. II, 20, Bot. (1843) 348, incl. var. minor et
vulgaris; DuBy in DC. Prod. 8 (1844) 66; Mor.
Syst. Verz. (1846) 44; Mia. FI. Ind. Bat. 2 (1859)
1002; Hanp.-Mazz. Acta Hort. Gothob. 2
(1926) 118; Not. R. Bot. Gard. Edinb. 16 (1928)
92; FLETCHER, FI. Siam. En. 2 (1938) 323.—L.
uliginosa BL. Bijdr. 14 (1826) 737; HAssk. Tijd.
Nat. Gesch. & Phys. 10 (1843) 130; Dusy in DC.
Prod. 8 (1844) 67; Mig. Fl. Ind. Bat. 2 (1859)
1002.—L. microphylla Merr. Philip. J. Sc. 1
(1906) Suppl. 221.—L. siamensis BONATI, Bull. Soc.
Bot. Genéve II, 5 (1913) 304, f. 62.—L. sub-
orbicularis WENT f. Nova Guinea 14 (1924) 113.—
L. minutissima (MASAM.) MASAM. Bull. Soc. Bot.
Fr. 83 (1937) 695.

var. japonica.—L. maculata R. Br.—L. debilis
WALL. in Roxb.—L. uliginosa BL.—L. siamensis
BONATI.

Slender, prostrate or decumbent, little-branched
herb, 10-40 cm long. Stem angular, brown-hairy;
internodes 2-5 cm. Leaves in opposite or sub-
opposite pairs, 2—5 cm spaced, orbicular apiculate
to ovate acute or subacuminate, 1-244 by 4-2
cm, base unequal, truncate, decurrent along the
petiole (0-1 cm long), upper side scanty appressed-
hairy, midrib and ascending nerves slightly distinct,
glandular punctate; underside more hairy, midrib
and nerves slightly prominent, veins subdistinct,
glands depressed. Flowers axillary, solitary, found
all along the stem, drooping then obliquely erect.
Pedicels 0-1 cm, always shorter than leaves.
Calyx 4-8 mm long, lobes lanceolate, acuminate,
undersurface scantily provided with glands,
hirsute-hairy, inner surface with distinct midrib.
Corolla yellow, shorter to a little longer than the
calyx, deeply cleft, lobes broad-elliptic, at the apex

Dec. | 962]

PRIMULACEAE (Bentvelzen)

183

broad-triangular, apiculate, whether or not
provided with glands. Stamens 3-4 mm, filaments
connate in a short tube or free from each other and
then dilated at the base; anthers | mm, versatile,
opening by lateral slits. Ovary hirsute. Style 3-4
mm. Capsule shorter than the calyx, 5-valved.

Distr. Kashmir to Yunnan, Siam, Eastern
China, Formosa, Ryukyu Is., Japan, also in
Australia (N. S. Wales, introduced?); in Malaysia:
Sumatra, Java (Preanger, Mt Diéng, and Mt
Tengger). Fig. 8.

= Ay
=

L So:
L

Fig. 8. Distribution of Lysimachia japonica
THUNB. var. japonica (dots), var. papuana S.

Moore (triangles).

Ecol. Damp banks, roadsides, abandoned fields
in montane rain-forest areas, 1100-2000 m.

Notes. According to HANDEL—MAzzetTTi L.
debilis WALL. would not be synonymous with L.
japonica THUNB. He arranged the first in the ser.
Debiles, the latter in ser. Japonicae. L. debilis
would be distinct by the glandular and twice as
large corolla and acute leaves. VAN STEENIS (1934,
237) pointed out that these characters do not hold
and I can fully confirm this: I have seen specimens
with orbicular leaves and a glandular-punctate,
small corolla but there are also eglandular small-
flowered ones. Also the hairiness of the calyx and
length of the pedicel is very variable (2-10 mm)
as is the length of the blade (19-30 by 18-22 mm).
In my mind the Chinese species of sect. Num-
mularia are too finely split and a new critical
synthesis with a wide species concept is necessary.

Kiatr described the staminal tube of L.
japonica as to be high and provided with glands.
BonatTI said the same of the specimen KERR 1146,
which he called L. siamensis. I have seen the type
and found a thickish or fleshy membrane. The
‘glands’ actually represent fruit bodies of a fungus
and hyphae can be observed in the tissue of the
corolla, and I assume the fleshy character of the
staminal tube to be due to the same cause.

In Japanese specimens the filaments are often
partially connate in a basal ring; in Sumatran
and Javan specimens they are free.

Tuwaites (En. Pl. Zeyl. 1860, 172) united L.
deltoides WiGur from Ceylon and India with L.
japonica THunB. Although they look very similar,

I think they are specifically distinct. In L. deltoides
the pedicels are longer than the leaves and it
has a much larger corolla. In passing it may be
remarked that the name L. deltoides WiGHT is
obviously a synonym of L. procumbens BAubo,
succinctly described(Ann. Sc. Nat. II, 20, Bot.. 1843,
348): ““Subprostrata rigide hirsuta. Cor. calyce
major. praecedenti (L. debilis Wall.) affinis.” In my
opinion there is no doubt that he had material
of L. deltoides WiGHT.

The name L. procumbens BAuDo, 1843, has
priority over L. deltoides WiGuT, 1851.

var. papuana S. Moore, Trans. Linn. Soc. II, 9,
Bot. (1916) 102; Went f. Nova Guinea 14 (1924)
113; Merr. & Perry, J. Arn. Arb. 29 (1948)
162.—L. microphylla Merr.—L. suborbicularis
WENT f.—L. japonica var. minutissima MASAM.
Prel. Rep. Veg. Yakus. (1929) 108; J. Soc. Trop.
Agr. Taiwan 4 (1932) 195; Hara, En. Sperm. Jap.
1 (1948) 86.—L. minutissima (MASAM.) MASAM.
Small prostrate herb, 5-10 cm long. Leaves
opposite, the pairs 0.3-2 cm spaced, scantily to
dense (Papua) strigose-hairy, broad-elliptic-orbi-
cular, 4-7 by 3-5 mm. Corolla c. 3-5 mm long.
Distr. Japan (Yakushima I.), Indo-China
(Annam); in Malaysia: Philippines (Luzon),
Celebes, and New Guinea. Fig. 8.
Ecol. In valleys, along river-banks, in grassland,
and in dense forests, also in fields, 1300-2900 m.
Vern. Jamp, Enga lang., Yogos, New Guinea.
Notes. According to Moore the pubescence
would be the only reliable character to separate
this from the type variety. This does not hold,
however, as many Javanese and Sumatran
specimens of var. japonica, including also THUN-
BERG’S type, are very hairy. The small, condensed
habit, the small leaves, small flowers efc. are,
however, very characteristic and for that reason I
have accepted this variety. Its occurrence over a
wide area geographical and altitudinal, and its
almost replacing geographical distribution as
compared with var. japonica justify a separate
status and plead against an environmental variant.
It is not endemic in New Guinea as supposed by
some authors; for that reason epithets as micro-
phylla and minutissima would be more appropriate,
but papuana has priority.
In the Papuan specimens the filaments are
whether or not connate in a basal ring.

7. Lysimachia mauritiana LAMK, Encycl. 3, Bot.
(1791) 572; Tabl. Encycl. 1 (1792) 440; Mia,
Ann. Mus. Bot. Lugd. Bat. 4 (1868) 147; Cor-
pEMoy, Fl. Ile Réunion (1895) 441; KNuTH,
Pfl. R. Heft 22 (1905) 273, f. 58; Merr. Philip
J. Sc. 3 (1908) Bot. 425; En. Philip. 3 (1923) 275;
HaAND.—MaAzz. Not. R. Bot. Gard. Edinb. 16
(1928) 106; SASAKI, Cat. Govt. Herb. (1930) 402;
Hosokawa, Trans. Nat. Hist. Soc. Form. 25
(1935) 34; KANEHIRA, En. Micron. Pl. (1935) 387;
Brown, Bull. Bish. Mus. 130 (1935) 222; GulL-
LAUMIN, Fl. Nouv. Caléd. (1948) 270; Hara, En.
Sperm. Jap. 1 (1948) 87; WALKER & RODIN,
Contr. U. S. Nat. Herb. 30 (1949) 463; MAKINO,
Ill. Fl. Jap. (1954) 232; INGRAM, Baileya 8 (1960)

184

FLORA MALESIANA

[Sera ieevolenGe

Mauritius

20
Réunion

Fig. 9. Distribution of Lysimachia mauritiana LAMK.

97.—Lubinia spathulata VENT. Hort. Cels. (1800)
t. 96; Dusy in DC. Prod. 8 (1844) 60; BAKER,
Fl. Maur. (1877) 188.—Lubinia mauritiana
SPRENG. Syst. Veg. 1 (1825) 572.—L. lineariloba
Hook. & ARN. Bot. Beech. Voy. (1841) 268;
Dusy in DC. Prod. 8 (1844) 61; Stes. & Zucc.
Abh. M.—Ph. KI. K. Ak. Wiss. Bayern 4, 3 (1846)
140; Kiatr, Abh. Naturw. Ver. Hamb. 4, 4
(1866) 31; Mig. Ann. Mus. Bot. Lugd. Bat. 3
(1867) 121; B. & H. Gen. Pl. 2 (1873) 635; FRANCH.
& SAv. En. Pl. Jap. 2 (1879) 431; Hest. J. Linn.
Soc. Bot. 26 (1889) 53; Mats. & Hayarta, J. Coll.
Sc. Univ. Tokyo 22 (1906) 221.—L. glaucophylla
Hook. & ARN. Bot. Beech. Voy. (1841) 306, t.
68; KLATr, Abh. Naturw. Ver. Hamb. 4, 4 (1866)
32, t. 18; KNuTH, Pfl. R. Heft 22 (1905) 275;
HAND.—-MAzz. Not. R. Bot. Gard. Edinb. 16
(1928) 107, syn. nov.—Bernadina laurina BAUDO
var. stenia BAUDO, Ann. Sc. Nat. Il, 20, Bot.
(1843) 349, nom. illeg.—Bernadina mauritiensis
Baupo, Ann. Sc. Nat. II, 20, Bot. (1843) 349,
nom. illeg—L. lubinoides Step. & Zucc. Abh.
M.-Ph. KI. K. Ak. Wiss. Bayern 4, 3 (1846) 140;
Mia. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 121;
FRANCH. & SAv. En. Pl. Jap. 1 (1875) 302.—L.
spathulata’ ScHouw, Linnaea 24 (1851) 160;
Katt, Abh. Naturw. Ver. Hamb. 4, 4 (1866) 32,
t. 18, ‘spatulata KLATT’; HILLespR. Fl. Hawaii
(1888) 285.—Lubinia lubinoides PAx in E. & P.
Pfl. Fam. 4, 1 (1897) 112.—Lubinia lineariloba
Pax, /.c.—L. nebeliana GitG, Bot. Jahrb. 24,
Beibl. 75 (1905) 57.—L. rubida Kotnz. Bot. Mag.
Tokyo 33 (1919) 110; HARA, En. Sperm. Jap. 1
(1948) 87.

Robust ascending or erect herb, to 50 cm; stem
at the base with two lateral branches, terete, at
the end branched again. Leaves alternate or
spirally arranged, blunt, spathulate, decurrent
along the petiole (0-1 cm), 244-6 by 14-2'4 cm,

coriaceous, at the underside with scattered black
glands, midrib prominent, venation invisible.
Upper leaves sessile. F/owers in head-like congested,
terminal racemes, after anthesis elongated and
3-12 cm. Bracts leafy, blunt and spathulate,
upwards decreasing in length. Pedicels longer than
the sustaining leaves. Calyx campanulate, lobes
lanceolate, broad-triangular or elliptic, apex
acute, obtuse, or rounded, 4-7 mm, outside a
prominent midrib, black-punctate, but not at the
scarious margin. Corolla white (in the dry state
yellow!), deeply cleft, lobes elliptic, obtuse, 12-16
mm. Stamens 9-12 mm, filaments halfway adnate
to the corolla, dilated at the base, not connate
with each other, anthers | mm, oblong, versatile,
acuminate, rounded at the base, opening with
lateral slits. Style 8-10 mm. Capsule pear-shaped,
exceeding the calyx, irregularly bursting.

Distr. Mauritius and Réunion, East Asia (E.
China: rare; Korean Is.; Japan; Ryukyus; For-
mosa), Bonin Is., Marianas, Hawaiian Is., New
Caledonia, and Loyalty Is.; in Malaysia: Philip-
pines (Batan Is.). Fig. 9.

Ecol. On coastal coral rocks and beaches; in
China also occurring in the coastal hills to
400(—700?) m, remarkably absent in the equatorial
zone between 13° NL and 18° SL.

Notes. This species has often been regarded as
representing a separate genus Lubinia VENT. but
I agree with later authors that it should be kept
in Lysimachia. BAUDO took L. mauritiana, the
type species of Lubinia, together with L. decurrens
to represent a separate genus Bernadina, but he
gave no generic description. It is the equivalent of
Lysimachia subg. Palladia (MorENCcH) HAND.—
MAzz.

L. lineariloba Hook. & Arn. and L. lubinoides
Stes. & Zucc. have been kept distinct from L.
mauritiana, but only on the strength of very

Dec. 1962]

PRIMULACEAE (Bentvelzen)

185

variable characters, such as the length of the in-
florescence; I agree with HANDEL—MAZZETTI to
reduce them.

Both HANDEL—MAZZETTI and KNUTH suggested
that L. glaucophylla Hook. & ARN. might belong
to L. mauritiana.

According to the first author it could be a local
form, possibly native in the Bonin Is. I have
examined the type which consists of a single
specimen (in K); it bears only a few flowers of
which the lowest is still in anthesis. The habit and
floral characters of the specimen are exactly those
of L. mauritiana, but the calyx differs in being
more or less fleshy, with + twisted lobes which
lack the scarious margin of typical L. mauritiana
and measure 9 mm in length (the longest in L.
mauritiana being 7 mm). In the Rijksherbarium
there is, however, a specimen (908. 176-1390) of
L. mauritiana, cultivated in the Botanic Garden
at Groningen (27—6—-1860) of which the lowest
flower has a calyx of exactly the same structure
as that in L. glaucophylla, but of which the upper
flowers have the calyx structure as in typical L.
mauritiana.

Another species which may be not different is
L. rapensis F. B. H. BRown (Bull. Bern. P. Bish.
Mus. 130, 1935, 22, f. 32), described from Rapa I.
in Polynesia; it differs by an acute leaf apex, short
pedicels, 4-merous flowers, and small size; this
also may be a local form. If so it would extend
the distribution considerably. I have not seen the
material.

8. Lysimachia decurrens Forst. f. Prod. (1786) 12;
LAMK, Tabl. Encycl. Bot. 1 (1793) 441; Dusy in
DC. Prod. 8 (1844) 67; SEEM. FI. Vit. (1866) 147;
F. y. M. Contr. Phytogr. New Hebr. (1874) repr.
p. 17; BritTEN in Forbes, Wand. (1885) 509;
Hems_L. J. Linn. Soc. Bot. 26 (1889) 51; Mats.
Tokyo Bot. Mag. 14 (1900) 71, incl. var. recurvata;
KnutTH, Pfi. R. Heft 22 (1905) 296; Merr. Philip.
J. Sc. 7 (1912) Bot. 93; Koorp. Exk. Fl. Java 3
(1912) 93; Fl. Tjib. 3 (1918) 31; Merr. En.
Philip. 3 (1923) 274; Lingn. Sc. J. 5 (1927) 144;
HAND.-Mazz. Not. R. Bot. Gard. Edinb. 16
(1928) 114; Merr. Sunyatsenia 1 (1930) 762;
Bonatl, Fl. Gén. I.-C. 3 (1930) 762; STEEN. Bull.
Jard. Bot. Btzg III, 13 (1934) 238; MAsam. FI.
Stud. Isl. Yakush. (1934) 358; FLeTcHer FI.
Siam. En. 2 (1938) 323; HAND.—MaAzz. Acta
Hort. Gothob. 13 (1939) 222; Hara, En. Sperm.
Jap. 1 (1948) 86; Backer, Bekn. FI. Java (em. ed.)
8 (1949) fam. 180, p. 4; MAKINO, IIl. Fl. Jap.
(1954) 231, incl. var. acroadenia; INGRAM, Baileya
8 (1960) 97.—Cerium spicatum Lour. Fl. Coch. 1
(1790) 136; ed. Willd. (1793) 168, cf. MERR.
Comm. Lour. (1935) 300.—L. javanica BL. Bijdr.
14 (1826) 736; Dusy in DC. Prod. 8 (1844) 62;
ZOLL. Syst. Verz. 3 (1855) 59; Mia. FI. Ind. Bat.
2 (1859) 1002; Kiatr, Abh. Naturw. Ver. Hamb.
4, 4 (1866) 16, t. 5; Hook. f. Fl. Br. Ind. 3 (1882)
502.—Bernadina parviflora BAUDO, Ann. Sc. Nat.
II, 20, Bot (1843) 349, nom. illeg., based on L.
multiflora —L. multiflora WALL. [Cat. (1828) n.
1487, nomen] ex Dusy in DC. Prod. 8 (1844) 63;

Kxatt, Abh. Naturw. Ver. Hamb. 4, 4 (1866) 14,
t. 4; Maxim. Bull. Ac. St. Pétersb. 12 (1868) 70;
Kurz, J. As. Soc. Beng. 46, ii (1877) 219.—L.
sinica Mia. J. Bot. Néerl. 1 (1861) 110; Hemsv.
J. Linn. Soc. Bot. 26 (1889) 57.—L. consobrina
Hance, Ann. Sc. Nat. V, 5 (1866) 224.—L.
keiskeana Mia. Ann. Mus. Bot. Lugd. Bat. 3
(1867) 120; HAND.-MAzz. Not. R. Bot. Gard.
Edinb. 16 (1928) 113.—L. acroadenia Maxim.
Bull. Ac. St. Pétersb. 12 (1868) 70; FRANCH. &
Sav. En. Pl. Jap. 1 (1875) 302; ibid. 2 (1879) 431;
KnuTH, Pfi. R. Heft 22 (1905) 296, f. 61B;
PETITMENGIN, Bull. Herb. Boiss. 7 (1907) 534;
Bonatl, Fl. Gén. I.-C. 3 (1930) 763; Hara, En.
Sperm. Jap. 1 (1948) 84.—L. recurvata (Mats.)
Masam. J. Soc. Trop. Agr. Taiwan 4 (1932) 302.

Erect, robust, little branched herb, up to 50 cm;
stem angular, provided with lenticels. Leaves
usually opposite, sometimes subopposite or alter-
nate, membranous, glabrous, narrowly ovate-
lanceolate, 3-7 by 1-214 cm, subacuminate, acute
at both ends, decurrent along the stem; upper
surface with point-like glands often confined to
the margin; midrib prominent beneath; nerves
slightly visible, veins obscure. Flowers in 1-5
terminal, sometimes leafy racemes 10—25 cm long.
Pedicels obliquely, erect or recurved, provided
with stalked glands, during anthesis 2-7 mm,
lengthening to 12-18 mm. The upper flowers are
sterile and monstruous. Bracts linear. Calyx lobes
lanceolate, acuminate, 3-4 mm, inside with
longitudinal black glands. Corolla white or purple,
as long as the calyx or smaller, deeply cleft; lobes
oblong, rounded, with prominent oblong glands;
inside with scattered granular red _ glands.
Stamens free from each other, 4-5 mm; filaments
with small sessile glands; anthers | mm, rounded,
versatile. Style 5 mm, persistent. Capsule longer
than calyx, bursting irregularly.

v

Fig. 10. Distribution of Lysimachia decurrens
Forst. f.

Distr. Eastern Himalaya, Burma, Siam, Indo-
China, Southern China, Japan, and Formosa,
through Malaysia to New Hebrides, and New
Caledonia. In Malaysia: Philippines (Luzon:
Benguet), West Java (from Tugu to Tyjikurai), and
Lesser Sunda Is. (Timor). Fig. 10.

Ecol. In pastures and fields, along roadsides

186

FLORA MALESIANA

[ser. I, vol. 6?

and trails, in brushwood and hedges, 1000-1500
m (in the Himalayas to 2000 m).

Notes. HANDEL—Mazzettl, /.c., had L. acroa-
denia MAXIM. as a synonym under L. keiskeana
Mia., which he kept as a separate species. Because
it was not included in his key, it is difficult to
verify his reason for doing so. I deduce from the
place he gave to L. keiskeana in relation to that
of L. decurrens that he assumed L. keiskeana Mia.
to have smaller flowers and stamens which do not
exceed the corolla. I examined MIQUEL’s type but
could find no difference in corolla between it and
that of L. decurrens. In many flowers the stamens
exceed the corolla but this character is variable in
L. decurrens in Java. The leaves of MIQUEL’s type
are not glandular-punctate but this also varies in
L. decurrens. 1 have consequently reduced L.
keiskeana to L. decurrens.

THENEN (Zur Phylog. der Primul. Bliite 1911,
99) stated that L. acroadenia MAxim. has branched
vascular bundles in the calyx which do not occur
in L. decurrens, but I cannot see how this would
warrant a separate taxonomic status.

After examining a topotype of L. recurvata
MasaAM. from Formosa (TANAKA 13596) I agree
to reduce it to L. decurrens. Recurved pedicels are
not rare in that species, even within one raceme
one can find drooping and ascending ones.

Excluded

Lysimachia lobelioides WALL. in Roxb. Fl. Ind.
ed. Carey & Wall. 2 (1824) 29; Dusy in DC. Prod.
8 (1844) 61; LINDL. Bot. Reg. (1842) t. 6; KLATT,
Abh. Naturw. Ver. Hamb. 4, 4 (1866) 16, t. 2;
Hook. f. Fl. Br. Ind. 3 (1882) 502 was mentioned
for Java by Hooker f. but I have seen no specimen
to justify this record. Dr. HusBarRD kindly inform-
ed us that there are in the Kew Herbarium no
specimens to support the record from Java.

Lysimachia obovata Ham. ex Hook. f. Fl. Br. Ind.
3 (1882) 502 was mentioned for Java but I haveseen
no specimen justifying this record. Dr. HUBBARD
kindly informed us that there are in the Kew
Herbarium no specimens to support the record
from Java.

3. PRIMULA

TouURN. ex LINNE, Gen. Pl. ed. 5 (1754) 70; Sp. Pl. ed. 1 (1753) 142; Dusy in
DC. Prod. 8 (1844) 34; Pax, Bot. Jahrb. 10 (1888) 75-241; O.K. Rev. Gen.
Pl. 2 (1891) 400; Pax, Pfl. R. Heft 22 (1905) 17; Bar. f. J. R. Hort. Soc. 39
(1913) 128-183; SmirH & Forrest, Not. R. Bot. Gard. Edinb. 16 (1928) 1-50;
SMITH & FLETCHER, Trans. Bot. Soc. Edinb. 33 (1941) 168.—Androsace TouRN.
ex! ILINNE; ‘Gen. PI. ed: 5 (1754) 179: Sp. Pl, (11753) 141; PAX. Pill Resblerm22
(1905) 172.—Cankrienia DE Vrirse, Jaarb. Kon. Ned. Mij. Tuinb. (1850) 29;
ZOLL. Syst. Verz. (1855) 59.—Fig. 11—12, 14—15.

Erect, sometimes robust, perennial, rarely annual herbs, up to c. 1 m high.
Leaves radical rosulate. Flowers 5-merous in | or more superposed whorls or
umbels, seldom solitary, sometimes dimorphous. ca/yx dentate or deeply
cleft. Corolla with non-recurved lobes. Stamens inserted in the tube; filaments
short. Ovary globose; style variable in length, stigma capitate. Capsule shortly

5-10-valved. Seeds numerous.

Distr. About 535 spp. throughout the temperate zone of the northern hemisphere where the largest
number of species and sections is found in the Sino-Himalayan region; secondary centres are found
elsewhere in Asia and Europe; no section is endemic in North America.

Ecol. The thermo-ecology of the genus is cool to frigid; though at higher latitudes occurring in
the lowland, the richest development is in the mountains, in shaded or open, moist or dry places.

Only in Malaysia and America the genus has extended its area over the mountains across the equatorial
belt, viz in Sumatra and Java by P. prolifera WALL., in New Guinea by P. umbellata (LouR.) BENTY.,
while in the Americas races of P. farinosa L. are found in Southern Chile and the Falklands (cf. BAKER,
Am. Natur. 93, 1959, 225-272), for some obscure reason widely disjunct from the North American
part of the population. It is significant that these southern-hemisphere populations of P. prolifera and
of P. farinosa are both homostylous and have an inbreeding system which is obviously due to genetic
recombination; this gives apparently a better adaptation for progressive dispersal and establishment to
the marginal part of the populations.

Taxon. Pax (Bot. Jahrb. 10, 1888, 76) subdivided Primula sens. str. into c. 20 sections, W. W. SMITH
& Forrest (1928) had even 32 sections and often of different circumscription. It is not clear whether
for the nomenclature typification was always strictly followed, for example sect. Candelabra BALF. f.
1913 contained amongst others P. prolifera and P. imperialis, here accepted to be conspecific. In 1905
Pax had already included these species in his sect. Cankrienia (DE VrRigEsE) Pax, but still earlier, in 1889,
Pax (/.c. p. 217 and in clay. p. 162) had included them in sect. Proliferae PAX based on P. prolifera

PRIMULACEAE (Bentvelzen)

ae

Fig. 11. Primula prolifera WALL. Group of plants along a brooklet on the Jang Plateau, East Java,
c. 2000 m altitude, in pyrogenous mountain savannah of Casuarina junghuhniana MiqQ., in
front of a stool of Preris wallichiana AGARDH (1938).

188 FLORA MALESIANA [ser. I, vol. 6?

WALL. As taxa exert priority in their own rank, it is clear that for this group sect. Proliferae Pax is the
correct name.

Generic delimitation. Hooker f. (in B. & H. Gen. Pl. 2, 1876, 631) could find no good generic dif-
ferences between Primula and Androsace (strong rhizome and vividly coloured flowers in Primula);
at least five species are intermediate in these two respects.

FRANCHET (Bull. Soc. Bot. Fr. 33, 1886, 63) has distinctly shown that the single systematic difference
between Primula and Androsace, viz a long or short tube of the corolla, is untenable because of many
intermediate Asiatic species showing this character in degree. PAx (Bot. Jahrb. 10, 1888, 133-136)
agrees with the presence of not a few intermediary links, but he avoided to take the consequence and
merge the genera, in arguing that Androsace would then have to be the accepted generic name — an
erroneous assumption because only based on page priority — and furthermore because he assumed
that the two genera were proportionally young and therefore still connected by transitions. O. KUNTZE
(Rey. Gen. Pl. 2, 1891, 399) rightly observed that the later assumption is a mere hypothesis, *“‘ein Wechsel
auf die Zukunft, der vielleicht erst nach Jahrmillionen fallig wird, den ich deshalb als vorsichtiger
Mann nicht in Zahlung nehmen kann ...”. In phytography it is of course inadmissable to enter such mere
theoretical considerations in practical research work. This is an illustration of a case in which two
genera appeared well distinct in early time when only few species were known, but which appeared
later to grade into one another when the centre of species development was gradually explored. From
the standpoint of scientific taxonomy there is of course no sense in retaining such a fictitious distinction;
but genera prove to have often a long life mostly due to the uneasiness of botanists connected with the
disappearance of a familiar generic name.

Cytotaxonomy. BRUUN (Symb. Bot. Ups. 1, 1932, 1-239) made very extensive cytotaxonomical
research of Primula. He concluded, that the species could be arranged into natural groups possessing
the same karyotype and that this division would coincide with that of W. W. SmitH & Forresr (Not.
R. Bot. Gard. Edinb. 16, 1928, 76), based on morphological criteria. Although the species within a
section would rarely have quite identical nuclei, in general the size and shape of the chromosomes
are rather stable.

These two characters together with the occurrence of constrictions he used for cytotaxonomical
demarcation.

It is remarkable that in cases where the systematics are rather difficult, their are also difficulties in
the distinction of the karyotypes.

For cytological evolution in Primula polyploidisation, accompanied by aneuploidisation, has presumably
been important for speciation, but the most important has been the regrouping of chromosome parts
(translocations).

Genetics. One of the most interesting problems of this genus is that of heterostyly, causing the flower
dimorphism. Of the same species there are plants bearing flowers with a short style and highly inserted
anthers (thrum type), and other individuals with a reverse situation (pin type). Thrum type styles have
to be pollinated with pollen from low-inserted stamens and the reverse. It will be clear that this is
genetically an outbreeding mechanism.

Already in the early days of the rediscovery of MENDEL’s laws, BATESON and GreGoryY (1905) found
in Primula sinensis LINDL. (= P. praenitens KER—GAWL.) heterostyly was controlled by a single pair
of alleles. The dominant one giving thrum, the recessive one pin type. The thrum type corresponded
normally with a heterozygous genotype; the pin type with the homozygous recessive one. In fact the
legitimate crossings thrum x pin and reciprocally were back-crossings (like sex-mechanism of fly, mouse
and man), producing constantly the same genotypes.

Ernst (Arch. Klaus Stift. 1, 1925, 13-62) detected in P. ‘hortensis’ and P. hirsuta ALL. the existence
of homostylous ‘‘Sippen’’. Crossing in all directions were possible: so homost. x thrum and reciprocally
and homost. x pin and reciprocally. The results of these crossings he interpreted by accepting two
separate, but narrowly coupled pairs of alleles. d—a controlling style length, B—b determining the
place of insertion of the anthers. A would give a short style, a a long one, B high-inserted anthers,
£ low-inserted ones. AB phaenotype was thrum, ab pin. The normal crossing is AB/ab = ab/ab, in fact
a scheme not much different from that of BATESON and GreGory. The phaenotype Ab corresponds
with a short homostyle, aB with a long one.

Ernst (Z. Vererb. 71, 1936, 156-230) recognized later on differences in size of pollen grains, again
controlled genetically independently of the other characters. These he explained not by means of separate
genes, but erroneously by multiple alleles. According to the fashion of this time Ernst (Z. Vererb.
88, 1957, 517-599) postulated that all characters connected with heterostyly are located within one
gene, which is subdivided into several subgenes.

Because of the absence of any position effect and besides the independent dominance of the factors,
it is my contention that they must be separate narrowly coupled genes. According to MATHER (Eyolu-
tion 4, 1950, 340-352) probably the genesis of this coupling has been promoted by natural selection.
In case of free recombination namely the advantageous heterostyly will easily break down and give
rise to homostyles.

According to Lewis (Biol. Rey. 24, 1949, 472-496) the size of the stigmatic papillae is also a character
tied up with heterostyly. But the essential thing of heterostyly is the selfincompatibility. Therefore he

Dec. 1962] _PRIMULACEAE (Bentvelzen) 189

postulated (sub)genes for incompatibility reactions of pollen tube and style. Nevertheless the situation
is not so clear as Lewis suggested. The results of ERNst’s laborious experiments do not fit entirely
with Lewis's hypothesis. Possibly there exists a biochemical interaction between the four ‘morphological
genes’ with relation to the incompatibility.

MATHER demonstrated convincingly the great pleiotropical effects of some mutants on heterostyly.
Furthermore he found some influence of the genetical background (‘polygenic systems’ in his ter-
minology) on the style-length ere. His finding that a heterozygous background produced a greater
variation seems important from a general genetic point of view.

According to A. Ernst (Arch. Klaus Stift. 28, 1953, 1-159) and H. G. BAKER (Recent Advances
in Botany, Toronto, 1961, 882) there might exist a correlation between polyploidy and homostyly,
as found for instance in P. farinosa L. sens. lat. The latter author has two explanations for this phenom-
enon. First: if a plant has a tendency to produce unreduced gametes, it will only produce polyploids
if it is selfcompatible. Second: polyploidy would enhance crossing-over and induce in this way a break-
down of heterostyly. Without experimental evidence it will be difficult to judge whether we are dealing
here with polyploidy as cause or as consequence of homostyly.

KEY TO THE SPECIES

1. Robust. Flowers yellow in 1-6 superposed whorls. Corolla tube well-developed. Leaves obovate
or spathulate-oblong, 10-50 by 4-10 cm... : 1. P.. prolifera
1. Small. White flowers in 1 whorl or umbel. Corolla tube short. Leaves broad- elliptic, 1-114 by 44-1 cm.
2. P. umbellata

1. Primula prolifera WALL. As. Res. 13 (1820)
372A, tab.; in Roxb. FI. Ind. ed. Carey & Wall. 2
(1824) 18; Don, Prod. Fl. Nepal. (1825) 81;
Zoi. & Mor. Nat. & Geneesk. Arch. Ned. Ind.
2 (1845) 8; Dusy in DC. Prod. 8 (1844) 34; ZoLL.
ex Hassk. Flora 30 (1847) 600; Mia. Ann. Mus.
Bot. Lugd. Bat. 4 (1868) 143; Hook. f. Fl. Br.
Ind. 3 (1882) 489; in Curtis, Bot. Mag. 110
(1884) t. 6732; Pax, Bot. Jahrb. 10 (1889) 217;
in E. & P. Pfl. Fam. 4, 1 (1897) 107; BLANc &
Decrock, Bull. Herb. Boiss. 6 (1898) 686; WaTT,
J. R. Hort. Soc. 29 (1904) 315; Pax, Pfl. R. Heft
22 (1905) 124; Bar. f. J. R. Hort. Soc. 39 (1913)
166; Cras, J. R. Hort. Soc. 39 (1913) 189;
SmitH & Forrest, Not. R. Bot. Gard. Edinb. 16
(1928) 17; STEEN. Trop. Natuur 19 (1930) 51,
77-84 f. 1-3; Bull. Jard. Bot. Btzg III, 13 (1934)
238; Tijd. Kon. Ned. Aardr. Gen. II, 55 (1938)
729, f. 1; Ernst, Ann. Jard. Bot. Btzg 49 (1940)
150; LooGEN, Trop. Natuur 29 (1940) 151, pho-
togr.; ibid. 30 (1941) 81; SmMitH & FLETCHER,
Trans. Bot. Soc. Edinb. 33 (1941) 168; STEEN,
Proc. Kon. Ned. Ak. Wet. C, 64 (1961) 435—442.—
P. imperialis JUNGH. Tijd. Nat. Gesch. & Phys. 7
(1840) 298; Hassk. Flora 25, Beibl. | (1842) 29;
Dusy in DC. Prod. 8 (1844) 668; Mia. Fl. Ind.
Bat. 2 (1859) 1001; WaALLAcgE, Mal. Arch. ed. 2
(1869) 114, f. 183; WIGMAN, Teysmannia 2 (1891)
450; Hook. f. in Curtis, Bot. Mag. 118 (1892) t.
7217; Koorp. Teysmannia 10 (1899) 446, 453,
456; Watt, J. R. Hort. Soc. 19 (1904) 315; Pax,
Pfil. R. Heft 22 (1905) 124, incl. var. gracilis;
UsterI, Vierteljahrschr. Naturf. Ges. Ziirich 50
(1905) 397, 465; VALETON, Jaarb. Dep. Landb.
Ned. Ind. (1907) 1; Ernst, Vegetationsbilder 7
(1909) t. 3a; Koorpb. in Jungh. Gedenkb. (1910)
231, f. 1-2; Exk. Fl. Java 3 (1912) 33, f. 33; BALF.
f. J. R. Hort. Soc. 39 (1913) 166; Koorp. FI.
Tjib. 3 (1918) 28; Costerus & SmitH, Ann. Jard.
Bot. Btzg 34 (1925) 47; SmitH & Forrest, Not.
R. Bot. Gard. Edinb. 16 (1928) 17; BopEGom, Fig. 12. Primula ee WALL. on Mt Sumbing,
Trop. Natuur 21 (1932) 43-47; DoctTers vAN along brook in forest, c. 2500 m altitude, immature
LEEUWEN, Proc. Kon. Ned. Ak. Wet. II, 31 (1933) Specimen, background leaflets of Thalictrum
javanicum BL. (LOOGEN, 1940).

190 |

FLORA MALESIANA

[ser. I, vol. 6?

215-218; Ernst, Ann. Jard. Bot. Btzg 49 (1940)
99-161; SmitH & FLETCHER, Trans. Bot. Soc.
Edinb. 33 (1941) 153; Backer, Bekn. Fl. Java
(em. ed.) 8 (1949) fam. 180, p. 1.—P. kuhlii BL.
Jaarb. Kon. Ned. Mij. Tuinb. (1844) 70.—
Cankrienia chrysantha DE VRiEsE, Jaarb. Kon. Ned.
Mij. Tuinb. (1850) 55 & tab.; Fl. des Serres 7
(1851) 53 & f. p. 58; ZoLL. Syst. Verz. 3 (1855)
$9.—Cankrienia farinosa ZOLL. Syst. Verz. 3
(1855) 59.—P. smithiana Crais, J. R. Hort. Soc.
39 (1913) 190; BAF. f. lc. 166, nomen; FARRER,
Gard. Chron. III, 68 (1920) 20; SMirH & ForREsT,
Not. R. Bot. Gard. Edinb. 16 (1928) 17; SmirH &
FLETCHER, Trans. Bot. Soc. Edinb. 33 (1941) 175;
Lownbes, Quart. Bull. Alp. Gard. Soc. 12 (1944)
35, t. 29.—P. khasiana BAF. f. & W. W. SMITH,
Not. R. Bot. Gard. Edinb. 9 (1916) 176; SmiTH &
Forrest, ibid. 16 (1928) 17; SmirH & FLETCHER,
Trans. Bot. Soc. Edinb. 33 (1941) 157.—P.
helodoxa Baur. f. Not. R. Bot. Gard. Edinb. 9
(1916) 171; Forr. J. R. Hort. Soc. 41 (1915) 201,
nomen; The Garden 80 (1916) 367 & 2 f.; Gard.
Chron. III, 59 (1916) 291, f. 123, 124; KiINGDON
Warp, J. R. Hort. Soc. 49 (1924) 152; SmitH &
Forrest, Not. R. Bot. Gard. Edinb. 16 (1928)17;
DARWELL, Gard. Chron. III, 84 (1928) 473;
STAPF in Curtis, Bot. Mag. 147 (1938) t. 8899;
SMITH & FLETCHER, Trans. Bot. Soc. Edinb. 33
(1941) 150.—P. sumatrana Merr. Not. Nat. Ac.
Nat. Sc. Philad. n. 47 (1940) 6; SmirH & FLET-
CHER, Trans. Bot. Soc. Edinb. 33 (1941) 150.—
Fig. 11—12, 14.

Robust perennial herb, 25-100 cm. Leaves
lanceolate-obovate, apex rounded, base acute,
decurrent along the petiole, midrib passing into a
thick petiole, 10-50 by 4-10 cm, margin regular-
dentate; nervation more or less reticulate. Flowers
in 1-7 whorls, whether or not pendulous, pedicels
2-4 cm. Bracts linear with whether or not folia-
ceous broadened apex. Calyx 4-8 mm, with
shorter or longer toothed teeth sometimes short
or long-acuminate, whether or not yellow waxy.
Corolla pale, bright or golden yellow; tube 7-15
mm, limb 10-20 mm g. Anthers inserted halfway
the corolla tube or near the limb, in Mal. specimens
high-inserted (except on Mt Jang) and homosty-
lous. Style 4-14 mm half as long as the tube or of

0

(ean

Fig. 13. Distribution of Primula prolifera WALL.

the same length. Capsule globose, shortly 5-valved,
erect. Seeds co, angular, brown.

Distr. Sino-Himalaya (Assam; Khasya and
Chumbi Hills; Upper Burma; S. Yunnan); in
Malaysia: Sumatra (Gajo Lands, e.g. Mts Losir
and Kemiri; West Coast Res.: Mts Singalang,
Kerintji, and Masurai) and Java (Mts Pangrango,
Papandajan, Sumbing, and Jang). Fig. 13.

Ecol. Rather open mossy forest, on damp soil
along brooks and in exposed marshy places,
2000-3250 m (in the Himalayas even higher), only
on mountains of which the summit reaches at
least 2650 m. The small angular seeds which are
produced in great quantity can be dispersed by
rain-wash and by streams, as has been observed
on Mts Pangrango and Papandajan in Jaya.

In JUNGHUHN’s time this beautiful plant was
on the summit zone of the extinct forest-clad

Fig. 14. Seedling of Primula prolifera WALL. from
seed washed down along the brook in the dense
forest at c. 2200 m near the Volcanological Ob-
servatory on Mt Papandajan. Specimens flower
profusely at 2500 m on Tegal Alun®, but no
flowering is ever observed in the present shaded
locality.

Pangrango cone confined between 2900 and 3000
m; in the course of time it has succeeded to descend
along the trail to the saddle between Gedeh and
Pangrango at c. 2500 m and has been grown suc-
cessfully near the shelter Lebak-saaét at 2450 m.
Attempts to introduce it on the adjacent Gedeh
Peak, either by seed or plants, by TEYSMANN,
DoctTers VAN LEEUWEN, and VAN STEENIS have
failed, obviously through the unsuitable soil
conditions, not through seed inaccessibility.

Vern. Babakuan, kémbang konéng, S, sawi utan,
Gajo M.

Notes. Hooker f. (1882) reduced P. imperialis
JUNGH. to P. prolifera, but later he changed his
opinion when he observed some specimens from
West Java and the Khasya Hills growing side by
side (1892); the first were more robust, had larger
leaves with thicker texture, broader midrib, closer
reticulate venation and bullate parenchyma, and
deeper tinged flowers. These criteria are, however,
derived from a few specimens, and are besides

Dec. 1962]

PRIMULACEAE (Bentvelzen)

ee

vague and merely quantitative. They do not hold
even for the Malaysian area as the size of the leaves
and plants and the colour of the flower varies
considerably. This has amply been demonstrated
by VAN STEENIS (1930) who could show that in
Java each population had its own facies. Specimens
from North Sumatra and East Java have the
closest similarity with those of the Khasya Hills.
Specimens from the Papandajan are transitional
between those of Mt Gedeh and Mt Jang in
length of calyx, flower colour, amount of yellow
wax on calyx, efc. It is precisely on such minor
characters that the Scotch botanists have recog-
nized further segregates of the prolifera complex,
by describing P. khasiana BALF. f., P. helodoxa
BALF. f., and P. smithiana Crats. It must of course
be kept in mind that the range of a mountain plant
is broken up into separate partial populations on
separated mountains, and that in cultivating a few
plants from each population they will differ in
details, and will never be ‘identical’. From this
it does not follow, however, that such non-identical
plants are also non-conspecific. As a matter of
fact Ernst kept all Malaysian specimens apart
from the continental Asian ones by the homostyly
of the former, notwithstanding their variability in
these details.

P. sumatrana was placed by MERRILL in sect.
Callianthae Pax, but SmirH & FLETCHER classified
it in sect. Candelabra (= Proliferae Pax) and
suggested already its affinity with P. imperialis.

Genetics. The only obvious difference between
this Primula in Malaysia and India is the homostyly
of the first one. Under the genus I have already
discussed the genetics of heterostyly. If we restrict
a further discussion to the two characters style
and height of insertion of the anthers a rare
crossing-over between A and B in a thrum type
will give the gametes 4b and aB. A combination
of these gametes with ab gametes of the pin type
will give the genotype I Ab/ab (short homostyle)
and II aB/ab (long homostyle).

As the homozygous homostyles will perpetuate
and the heterozygous homostyles will also yield
partly homozygous homostyles the number of
homostyles will steadily increase and after a few
tens of generations consist entirely of homostylous
individuals.

In the centre of the area the majority of the
individuals are heterostylous; this causes hetero-
zygosity which, as generally accepted, will be ad-
vantageous because of the greater fitness to with-
stand fluctuating environmental conditions. The
homostylous condition is here disadvantageous.

In marginal populations the gene pool is always
depauperated, hence the adaptive value of the
homostylous individuals will be greater because
of more abundant seed production which will lead
to more rapid expansion of the range, as postulated
by BAKER (Am. Natur. 93, 1959, 255-272). I
assume this reasoning to hold for P. prolifera.
Unfortunately Ernst (Arch. J. Klaus Stift. 26,
1951, 1-96; Genetica 27, 1955, 391-448) did
not succeed in crossing the ‘Javanese’ P. prolifera
with P. smithiana, but he also did not succeed in

intraspecific crossings of P. smithiana.

ErNst considered homostyly as a_ primary
character of P. ‘imperialis’ but in my opinion its
derivation from the continental-Asiatic population
of P. prolifera would point to its secondary nature.

2. Primula umbellata (Lour.) BENTV. comb. nov.—
Drosera umbellata Lour. Fl. Coch. ed. 1 (1790)
186; ed. Willd. (1793) 232; PLANcH. Ann. Sc. Nat.
III, 9, Bot. (1848) 304.—Androsace saxifragifolia
Bunce, Mém. Ac. St. Pétersb. 2 (1835) 127;
Dusy in DC. Prod. 8 (1844) 53; HANce, J. Bot.
20 (1882) 6; Hook. f. Fl. Br. Ind. 3 (1882) 496;
Forses & Hems-L. J. Linn. Soc. Bot. 26 (1889)
45; KNnuTH, Pfl. R. Heft 22 (1905) 179; Mats. &
Hayarta, J. Coll. Sc. Univ. Tokyo 22 (1906) 221;
GAMBLE, FI. Madras 4 (1921) 745; HAND.—MAzz.
Not. R. Bot. Gard. Edinb. 15 (1927) 271; Bonati,
Fl. Gén. I.-C. 3 (1930) 757; HAND.—MAzz.
Symb. Sin. 7 (1936) 753.—P. saxifragifolia O. K.
Rev. Gen. Pl. 2 (1891) 400.—P. minutiflora
Forrest, Not. R. Bot. Gard. Edinb. 4 (1908) 219,
t. 29A.—Androsace umbellata Merr. Philip. J.
Sc. 15 (1919) 237; En. Philip. 3 (1923) 274;
HAND.-MaAzz. Not. R. Bot. Gard. Edinb. 16

(1928) 162; Sasaki, Cat. Govt. Herb. (1930) 401;
STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 237;
Merr. Comm. Lour. (1935) 300; NAKAl, Rep.
Exp. Manchouko sect. IV, 4 (1936) 38; Bull. Nat.

Fig. 15. Primula umbellata (Lour.) BENTV. a.
Habit, 2/3, b. flower, * 4, c. corolla laid open.
x 4, d. stamen, 12, (VAN ROYEN, 4443).

192

FLORA MALESIANA

[ser. I, vol. 6?

Sc. Mus. Tokyo 31 (1952) 89; STEEN. Nova Guinea
n.s. 6 (1955) 279; STEWARD, Pl. Lower Yangtsze
(1958) 299.—Fig. 15.

Slender erect herb, 4-12 cm. Leaves hairy,
broad-elliptic, decurrent along the petiole, trun-
cate at the base, 8-15 by 4-10 mm, margin-crenate,
ciliate; midrib slightly prominent at the underside;
nerves and veins indistinct; petiole 5-15 mm.
Umbels 2-6-flowered, peduncle 3-9 cm, hairy;
pedicels 1-4 cm, hairy. Bracts 1 mm linear.
Flowers 2-5 mm long. Calyx cleft more than half-
way, campanulate, lobes oblanceolate acute,
hairy outside and on the margin. Corolla white,
slightly exceeding the calyx; lobes oblong, rounded
at the apex. Stamens not surpassing the style;
filaments longer than the anthers, to the middle
adnate with corolla. Capsule as high as or higher
than calyx, shallowly 5-valved or irregularly
bursting.

Distr. In India, Pakistan, and Burma widely
distributed (from the Punjab to Manipur),
northern Indo-China, China (Szechuan, Yunnan,
Kweichow, Kwangtung, Manchuria), Korea,
Japan, Ryukyu Is., and Formosa; in Malaysia:
Philippines (N. Luzon: Mountain Proy.) and
East New Guinea. Fig. 16.

E ai eerie sy

Fig. 16. Distribution of Primula umbellata (Lour.)
BENTV.

Ecol. Shaded moist places in mountain forest,
1300-1500 m.

Notes. This species has, according to KNUTH,
sometimes been confused with Androsace rotun-
difolia HARDW. which is well characterized by its
leafy, crenate bracts.

Already PLANCHON (1848) suggested that
Drosera umbellata Lour. would be an Androsace
and MERRILL (1919, 1935) was convinced of this
reduction, basing himself in the interpretation of
LouREIRO’s very meagre description on his method
of ‘elimination’; in the flora of Kwangtung, from
where LoureEIRO obtained his material, no other
plant fits the combination of characters mentioned
by Loureiro and, besides, Androsace saxifragifolia
is there a common plant. HANDEL—MAZZETTI
(1936) wanted to have more certainty before
accepting MERRILL’s conclusion, but I assume
MERRILL is right.

Cultivated species

Primula malacoides FRANCH.; cf. BACKER, Bekn.
Fl. Java (em. ed.) 8 (1949) fam. 180, p. 2: calyx
not inflated, cleft halfway, outside white-waxy,
lobes 5, recurved. A Chinese species cultivated as
an ornamental in the mountains.

Primula obconica HANCE; cf. BACKER, /.c.: calyx
not inflated, cleft much less than halfway, without
wax, lobes 5, short and broad, erect. A Tibetan
species, cultivated as an ornamental in the
mountains.

Primula praenitens KER GAWL.; syn. P. sinensis
LINDL. 1821, non Lour. 1790; cf. BACKER, /.c.:
calyx inflated, at apex with 14 or more erect
laciniae, without wax. A Chinese species occa-
sionally cultivated as an ornamental in the moun-
tains of Java.

Primula yeris L. var. elatior L. This species is
cultivated in the Mountain Garden of Tjibodas,
W. Java, where it does not flower at 1450 m.
Specimens planted near Lebak Saat, at c. 2450 m,
flower abundantly. Cf. STEEN. Bull. Jard. Bot.
Btzg Ill, 13 (1934) 337.

Excluded

Samolus valerandi LINNE, Sp. Pl. (1753) 243; Mia. FI. Ind. Bat. 2 (1859) 1004, though almost ubiquitous,

has as yet not been found in Malaysia.

SIMAROUBACEAE (H. P. Nooteboom, Leyden)

Trees or shrubs, in Mal. evergreen or almost so, usually containing very bitter
substances. Twigs pithy. Hairs mostly simple and 1-cellular, sometimes glandular-
capitate. Leaves (in Mal.) spirally arranged, simple or 1-pinnate, often articu-
lated, often provided beneath (rarely also above) with pitted, concave, or flattish
glands (in Mal. in Ai/lanthus, Brucea, Samadera, and Soulamea). Stipules usually
absent, (in Mal.) present in Jrvingia and Picrasma. Inflorescences usually com-
pound, axillary, rarely terminal; plants monoecious, rarely dioecious. Flowers
usually small, actinomorphic, uni- or bisexual, or functionally unisexual. Sepals
3-5, almost always partly connate, valvate to slightly imbricate. Petals 3-5, free,
imbricate or valvate, rarely absent (in extra-Mal.), or united into a tube (extra-
Mal.). Stamens inserted at the base of the disk, isomerous or dimerous, rarely
numerous (extra-Mal.), mostly obdiplostemonous, rarely the stamens of the outer
whorl doubled, not rarely with a scale at the inner base; anthers 2-celled, opening
lengthwise, introrse to latrorse, versatile. Disk intrastaminal, often gynophorous,
sometimes rather inconspicuous, at least when dry. Ovary often 2—S-lobed,
1—5-celled, or with free carpels; styles 1-5. Ovules 1-2 (in Mal. 2 only in Suriana),
axile, anatropous (in Harrisonia and Suriana amphitropous). Fruit(s) usually
indehiscent, often drupaceous, sometimes a samara, some carpels sometimes
aborted. Seed: endosperm 0 or scant; cotyledons planoconvex; embryo straight

or curved; no aril.

Distribution. About 30 genera, with c. 200 spp. with the main centre in tropical America, and a
second centre in tropical West Africa. With the exception of Picrasma quassioides (D. DON) BENN., ranging
as far north as North Japan and Korea, and of Ailanthus altissima (MILL.) SWINGLE, which is endemic
in subtropical to temperate China, the species of this family are native in the tropics. The monotypic
genus Suriana has the widest range and is almost pantropical along sandy beach, being absent only on
the west coast of Africa.

A remarkable distribution is exhibited by Soulamea, with 6 endemic species in New Caledonia, one
in Fiji, one widely ranging sandy beach species from Polynesia westwards to Borneo and one endemic
species in Mahé I. (Seychelles).

Another remarkable type of distribution is displayed by Picrasma, which is disjunct tropical transpa-
cific.

Ecology. Most of the Malaysian Simaroubaceae are inhabitants of the lowland forest; only Brucea
mollis WALL. ex Kurz is recorded as high as 1800 m (in the Philippines)

Climatically defined areas in Malaysia are occupied by both species of Harrisonia and to a less degree
by Ailanthus triphysa, all of which distinctly avoid the everwet forest belt of West Malaysia (in Sumatra,
Malaya, Borneo, and West Java), and prefer regions subject to a dry monsoon. Fig. 10, 11, and 20.

Brucea, Picrasma, and Ailanthus integrifolia behave rather indifferent to climate. As to soil /rvingia
and Samadera are indifferent but Eurycoma has a distinct preference for acid, leached sandy soils and is
classified as silicicolous. Quassia §Samadera shows preference for temporarily inundated places.

Suriana and Soulamea both belong to the Barringtonia formation and obviously prefer calcareous or
rocky beaches.

Pollination. Is probably performed by insects, as the flowers are often reported to be fragrant.
They are either unisexual or functionally so, or bisexual.

Dispersal. Unfortunately little is known on this subject.The fruits of Suriana maritima and Soulamea
amara are certainly dispersed by seawater and this seems to have been very effective. RIDLEY (Disp.
1930, 264) assumes dispersal by fresh-water for Quassia § Samadera which has large fruits and is
frequent in alluvial forests and swamp forest. In all cases buoyancy is gained by the seed not entirely
filling the fruit and leaving a cavity. The thin-winged samaras of Ai/anthus will be wind-dispersed over
small distances. See further under the genera.

Phytochemistry. All Simaroubaceae (with the exception of the Irvingioideae and Surianioideae)
contain bitter principles which seem to be closely related to each other. Unfortunately their chemical
structure has not yet been fully elucidated. The best known bitter principle of the family is quassin, which
has been isolated from the woods and barks of Quassia amara AuBL. and Aeschrion excelsa (Sw.) O.
Kuntze (Picrasma). According to recent investigations of ROBERTSON and collaborators (J. Chem. Soc.

(193)

194 FLORA MALESIANA [Sen UeavolaGs

1950, 3431; 1954, 3672, 4238) quassin is a mixture of two closely related compounds which were called
quassin and neo-quassin. Both bitter principles contain a carbon skeleton built up from 20 C-atoms and
belong probably to the diterpenoid compounds. Quassin, C.)H ,0,(OCHs), has the following functional
groups: 2 methoxyl, a lacton, a tertiary hydroxyl and a carbonyl group. Another bitter principle of the
family was isolated from the seeds of Simarouba glauca (E. A. HAM etal., J. Am. Chem. Soc. 76, 1954, 6066)
and termed glaucarubin. Glaucarubin like quassin seems to be a derivative of a C,)-compound; it is,
however, an ester of «-methyl-x-hydroxybutyric acid with the diterpenoid hexahydroxylacton, glaucaru-
bol. A third bitter principle examinated recently is cedrin from the seeds of Simarouba cedron PLANCHON.
According to Kress and RUser (Arzneimittelforschung 10, 1960, 500) cedrin is a sesquiterpenelacton of
the santonin-type. The common features of all bitter principles of Simaroubaceae seem to be the lactonic
function and the isoprenoid structure (sesquiterpenes or diterpenes). In this respect these constituents are
related to the bitter principles of Rutaceae (limonin nomilin, obacunon); the latter, however, contain a
carbon skeleton, which is related to triterpenes rather than to sesquiterpenes or diterpenes.

Many Simaroubaceae are used locally as therapeutic agents, especially as tonics, antidysenterics and
anthelmintics. The bitter principles are believed to be the therapeutically most important constituents of
the members of the family.

Another feature rather characteristic for the family is the common occurrence of small amounts of essential
oils and large amounts of resins. These excretions are located in perimedullar resin canals said to be of
schizolysigenous origin and in idioblasts occurring in the pith, phloem and cortex and in the leaves of some
species. Practically nothing is known about the precise chemical nature of these essential oils and resins.

Mucilages too are wide spread in the family. They are deposited in mucilage cells in the epidermis of the
leaves and in the subfamily of Irvingioideae in lysigenous cavities in the pith.

In Simarouboideae and Irvingioideae the epidermal cells of the leaves are heavily silicified as demonstrat-
ed by EDMAN (Svensk Bot. Tidskr. 30, 1936, 493).

There are some indications that tannins, coumarines and alkaloids are not rare in the family, but no
detailed informations are available, besides the statement of ALTMAN (Bol. tecn. inst. agron. do Norte,
Belém, no 31, 1956, 27) that the twigs of Picrolemma pseudocoffea DUCKE contain quinine. It would be
very interesting to investigate Simaroubaceae for alkaloids and coumarines because the closely related
Rutaceae are so rich in highly characteristic alkaloids and coumarines.

A very interesting chemical feature of the family is to be found in the composition of the fatty oils of
the seeds. As a rule the seed fats of the members of a family are rather uniform in composition. In the family
of Simaroubaceae, however, four different types of seed fats have been found up to now. The fats of the
genus /rvingia are similar to those of Lauraceae and Myristicaceae by the high amounts of lauric and my-
ristic acid. The genus Picramnia seems to be characterized by fats with the acetylenic tariric acid related
by the position of unsaturation to petroselinic acid. The latter one (characteristic for the families Um-
belliferae and Araliaceae) has been demonstrated to be a major fatty acid in the seed fat of Picrasma
quassioides (D. DON) BENN. The rest of the seed oils of the family investigated (of the genera Ai/anthus,
Brucea, Perriera, Samadera, Simarouba) belong to the common and very wide spread type characterized
by palmitic, oleic and linoleic acids as major fatty acids. There exist many striking phytochemical re-
semblances between Rutaceae and Umbelliferae. Therefore the occurrence of petroselinic acid and tariric
acid as major fatty acids in the seeds of Simaroubaceae which seem to be closely related to Rutaceae, may
be more than purely chemical convergence.—R. HEGNAUER.

Wood anatomy. DEN BERGER, Determinatietabel van Malesié, Veenman, Wageningen (1949)
several pp., due to the occurrence of gum ducts, storied structure, the distribution of the wood pa-
renchyma or the structure of the rays in the various genera (hand lens). Hetmscu, Lilloa 8 (1942) 117;
METCALFE & CHALK, Anat. Dic. 1 (1950) 321; MoLt & JANssonius, Mikr. Holzes 2 (1908) 72;
Saya, Ann. Accad. ital. Sci. for. 4 (1955) 315; WEBBER, Am. J. Bot, 23 (1936) 577; Lilloa 6 (1941)
441. According to HEImscH (/.c. 176) the Burseraceae and Anacardiaceae are highly similar in wood
structure and together are distinct from the Rutaceae, Simaroubaceae, Meliaceae, and Sapindaceae, and
in general the Simaroubaceae simulate the Rutaceae rather closely (/.c. 177); because of the large
variation in wood structure the members of the family most probably do not represent a natural group
(1c. 189, cf. also MeTCALFE & CHALK, /.c. 325). Note that the term septate fibre tracheids (WEBBER,
/.c., HeImscu, /.c.) has to be replaced by septate (libriform wood) fibres (REINDERS, Trop. Woods 44,
1935, 30; Handl. Plantenanatomie ed. 4, 1951, 147; REcorp, Trop. Woods 78, 1944, 36). According
to the definitions of SANIO-JANSSONIUS-REINDERS (REINDERS, /.c.) all species of this family possess libri-
form fibres and none of them fibre-tracheids. — C.A.R.-G.

Taxonomy. The Simaroubaceae are doubtless closest related with the Rutaceae, followed by the Melia-
ceae and Burseraceae, in that order. They lack homogeneity and there is no single character common
to all genera and not present in the other families. The bitter substances are commonly assumed to be
characteristic of the quassi family, but they are absent in the /rvingioideae (Klaineodoxa and Irvingia)
and in the Surianoideae (Cadellia and Suriana), and, besides, occur also in some Rutaceae and Melia-
ceae (Trichilia). Another very common character is the occurrence of concave or flattish but sunken spot-
glands on the underside of the leaves which are also found in the meliaceous genus Trichilia. Both vege-
tatively and in sexual organs Simaroubaceae are diverse in character: leaves are simple or compound,
stipules are present or absent, carpels are free or connate, stamens are isomerous or dimerous, and the

Dec. 1962] SIMAROUBACEAE (Nooteboom) 195

fruit structure displays a great variaton. From this follows that delimitation against the allied families
must remain arbitrary. In 1874 ENGLER (Abh. Naturf. Ges. Halle 13, 2, p. 140) considered the Sima-
roubaceae as a residue of the Geraniales: ‘Wir sind genéthigt alle diejenigen Formen aus der Reihe
der Geraniales, welche sich dusserlich an eine der verschiedenen Rutaceen-Gruppen anschliessen, in
ihrem anatomischen Verhalten aber in der angegebenen Weise von denselben sich unterscheiden, zu den
Simarubaceae zu rechnen.’

It must be remembered, however, that Simaroubaceae sometimes (Irvingia) possess lysigenous cavities
containing mucilage, METCALFE & CHALK (Anat. Dic. 1950, 317-326) recently concluded from the ana-
tomy: ‘There are very few characters common to the whole of the Simaroubaceae. This lack of homo-
geneity, which also occurs in the external morphological characters, seems to indicate that the family is
unnatural, but consists of a number of groups which are themselves relatively uniform.’ This diversity
is also expressed in the large number of subfamilies. WEBBER (Am. J. Bot. 23, 1936, 577-587) concluded
from the wood anatomy that the subfamilies Kirkioideae, Irvingioideae, Picramnioideae, and Alvaradoi-
deae might rank as distinct families or as components of other families if these were suggested on other
morphological grounds.

In phytochemical aspect it is interesting that EDMAN (Svensk Bot. Tidskr. 30, 1936, 493-514) found
lack of homogeneity in the amount of silica in the leaves, a character which is generally characteristic
for taxa of higher rank. In the Simaroubaceae it appears that the Irvingioideae and the subtr. Simarou-
binae are the only groups which are highly silicified. He assumes that this is a primitive character; it is
also found in the primitive members of the Rutaceae and Burseraceae; the ability to store a large amount
of silica has obviously been lost by some groups of all three families.

Palynologically the family also shows a certain lack of homogeneity, according to ERDTMAN (Pollen
Morph. & Tax. 1, 1952, 406-409), although a relation of Jrvingia and Suriana with the true Simarou-
baceae is probable.

The genera without bitter substances, viz Irvingia and Suriana, have often been discussed.

Irvingia, together with Klainedoxa, was by ENGLER (Pfl. Fam.3, 4, 1896, 227) distinguished as a sub-
tribe, and later by Boas (Beih. Bot. Centr. 29 i, 1913, 348) and ENGLER (Pfl. Fam. ed. 2, 19a, 1931, 396)
raised to subfamily rank. Pierre (Fl. For. Coch. 4, 263) has already given it family rank in 1892, mainly
because of presence of stipules and lysigenous mucilage cavities in the cortex and pith of branches and
petioles. He placed the Irvingiacées next to Anacardiaceae. VAN T1iEGHEM (Ann. Sc. Nat. IX, Bot. 1,
1905, 247-320) considered the Irvingiaceae close to but distinct from the Simaroubaceae. HALLIER f.
(Beih. Bot. Centr. 39, ii, 1923, 62-68) classified Irvingia next to the Linaceae-Erythroxyleae, a disposi-
tion which has recently partly been accepted by HUTCHINSON (Fam. FI. PI. ed. 2, 1959, 261) in placing
the Irvingiaceae in the order Malpighiales next to the Linaceae. As EDMAN’s phytochemical results are
in favour of affinity between Irvingia with Simarouboideae and as the leaves of Irvingia are extremely
similar to those of Quassia § Samadera (though without glands), and stipules also occur in Picrasma, I
think it is reasonable to maintain /Jrvingia within the Simaroubaceae. It is true that the stipules in Jrvingia
leave an annular scar, which is not the case in Picrasma, but in other families, e.g. Hamamelidaceae and
Rubiaceae their insertion also varies in degree.

Suriana has also a chequered taxonomical history, and was successively classified in the Crassulaceae
by LInng, in the Spiraeaceae by ENDLICHER and PLANCHON, and in the Geraniaceae by LINDLEY. ARNOTT
(W. & A. Prod. 1 (1834) 360) raised Suriana to family rank. J. G. AGARDH (Theor. Syst. Pl. 1858, 169)
placed the family next to the Geraniaceae with the following argumentation: lack of bitter substances,
thin and brightly coloured clawed petals, and an ovary with 2 amphitropous ovules per cell. Amphitro-
pous ovules occur, however, also in Harrisonia. AGARDH’S opinion was sustained by JADIN (Ann. Sc. Nat.
VIII, Bot. 13, 1901, 303) on — somewhat inadequate — anatomical characters. According to WEBBER
(1936, /.c.) the anatomical structure of Suriana supports SOLEREDER’S opinion (Syst.Anat. Dic. 1899,
207-213) of suppressing a monotypic family Surianaceae and classifying it with the Simaroubaceae as
proposed by BENTHAM & Hooker (Gen. Pl. 1, 1862, 307) and accepted by ENGLER (Pfl. Fam. ed. 2,
19a, 1931, 365) and Cronauist (Brittonia 5, 1944, 129), which seems still the most acceptable disposition.

Uses. All the bitter tasting genera are highly in demand for medicinal purposes by the people, and
are used against a wide variety of illnesses. It seems that they have sometimes some healing properties
indeed, for example the nuts of Brucea javanica. These are known under the name of ‘Makassaarse pit-
jes’ (Dutch) or ‘Kho-Sam (Chin.). They were about 1900 imported into Europe and came highly in de-
mand as a drug. According to MoussALLt (Contr. a l’étude des Simarubacées, 1939) an unidentified,
much less bitter tasting drupe was often handled by crooked merchants to adulterate true ‘Kho-Sam’.

Some of the toxic constituents, after having been extracted, are occasionally used as an insecticide.

None of the genera furnishes timber of general commercial importance, though some, e.g. Ailanthus,
produces a timber that is used locally, chiefly for packing-cases and also for house building. In none of
the genera the wood is very durable.

196 ue j FLORA MALESIANA [ser. I, vol. 6?

KEY TO THE GENERA
1. Leaves simple.

2, SOUS jose, IGA Aue SES 45 6 oo ob oo 6 oO ne bow en ee 9. Irvingia
2. Stipules absent.
3. Leaves sessile or nearly so, linear-spathulate, up to 5 mm wide. Plant not bitter 1. Suriana

3. Leaves larger, distinctly petioled. Bitter substances present.
4. Leaves obovate. Branchlets thick. Stamens without an adaxial scale. Carpels connate. Fruits

(Oloretos eNO OM oS EEDA CRICMENS LakerinE OAC MAM amceraNn Atoka rice ceases Gy SOUTED
4. Leaves subelliptic, with concave glands, usually on the undersurface. Branchlets not thick. Sta-
mens without an adaxial scale. Carpels free. Fruit + semicircular... ... . . . 2. Quassia

1. Leaves compound.
5. Leaf-rachis winged or leaves ternate. Stamens with an adaxial scale.
6. Branches with stipular thorns. Leaf-rachis narrowly winged or leaves ternate. Carpels united.
4. Harrisonia

6. Branches without thorns. Leaf-rachis broadly winged and articulated. Carpels free. . . 2. Quassia
5. Leaf-rachis not winged. Leaves pinnate. Stamens either with an adaxial scale or not.
7. Stipules present, caducous. Stamens without an adaxial scale . .°. . . . .. . . 6. Picrasma

7. Stipules absent. Stamens either with an adaxial scale or not.

8. Leaflets sessile or nearly so, attached to the rachis with a conspicuous (constricted) articulation.
Stamens with an adaxial scale, the same number as the petals, alternating with staminodal scales
3. Eurycoma
8. Leaflets distinctly stalked, not conspicuously articulated. Androecium haplo- or obdiplostemonous.

Stamens either with an adaxial scale or not.
9. Stamens twice the number of petals, either with an adaxial scale or not. Inflorescence a panicle.
10. Stamens without an adaxial scale. Large trees. Branches thick, with large, crowded leaf scars.
Fruitvavsamaraaviy 5 fis a AS Se See eh see oe cei eal tanya
10. Stamens with an adaxial scale. Branches not very thick, not with large, crowded leaf scars
2. Quassia
9. Stamens the same number as petals, without an adaxial scale. Thyrse narrow . . 5. Brucea

1. SURIANA

LINNE, Gen. Pl. ed. 5 (1754) 137; Sp. Pl. (1753) 284—Fig. 1.

Shrubs or small trees; innovations hairy, partly glandular-capitate; without a
bitter taste. Leaves sessile, simple. Stipules 0. Flowers 5-merous, bisexual, in
pauciflorous, axillary cymes, rarely solitary; stalks articulated at the base. Bracts
persistent, foliaceous. Sepals persistent, connate at the base, imbricate in bud, as
large as the imbricate petals. Stamens 10, sometimes 5 barren in 2 distinct rows,
with latrorse, versatile anthers. Disk not developed. Carpels 5, free, each with a
free, filiform, basally attached erect style; stigmas free, small, inconspicuous;
ovules 2 in each carpel, collateral, basal, amphitropous, micropyle directed to
the base. Fruits drupaceous, 3-5 together, enclosed by the calyx. Seed 1 in each

carpel; embryo curved, albumen 0.

Distr. Monotypic, pantropical. Fig. 2.

Notes. JADIN (Ann. Sc. Nat. VIII, Bot. 8, 1901, 224-226) considered Suriana as representative of a
monotypic family, Surianaceae, on behalf of the occurrence of glandular hairs (a character which he
overlooked in some other genera), the number and basal attachment of the ovules, the lack of bitter
substances, and some other minor characters. SOLEREDER (Verh. Bot. Ver. Brandenb. 47, 1905, 35-62)
referred the genus to the Simaroubaceae, which was agreed to by later authors.

1. Suriana maritima LINNE, Sp. Pl. (1753) 284; Java (1911) 193; Merr. Philip. J. Se. 7 (1912)
DC. Prod. 2 (1825) 91; SpRENG. Gen. Pl. ed. 9 Bot. 274; Guppy, Seeds & Curr. (1917) 239-242;
(1830) 383; W. & A. Prod. (1834) 361; DECNE, Laut. Bot. Jahrb. 56 (1920) 342; E. G. BAker, J.
Herb. Tim. Descr. (1835) 121; BENTH. Fl. Austr. 1 Linn. Soc. Bot. 45 (1921) 285; Merr. En. Philip. 2
(1863) 375; BENN. in FI. Br. Ind. 1 (1875) 522; (1923) 345; C. T. Wuite, J. Arn. Arb. 10 (1929)
HeEMsL. Bot. Chall. 1 (1885) 131; Trimen, FI. 227; Ripi. Disp. (1930) 264; DANIKER, Viert.
Ceyl. 1 (1893) 222; Wars. Bot. Jahrb. 18 (1893) Jahrschr. Naturf. Ges. Ziirich 77 (1932) 204;
194; BaiLey, Queensl. Fl. 1 (1899) 220; Guppy, FF. B. H. Brown, Bull. Bern. P. Bish. Mus. 130
Observ. Natur. Pac. 2 (1906) 105; BAck. Schoolfl. (1935) 131; GuILLAUMIN, Bull. Soc.’ Bot. Fr. 85

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

197

(1938) 20; Fl. Nouv. Cal. (1948) 170; Perr. DE
LA BATuiE, FI. Madag. fam. 105 (1950) 7; W. R.
Tay or, Pl. Bikini (1950) 183; YuNcKeErR, Bull.
Bern. P. Bish. Mus. 220 (1959) 154.—Fig. 1.

Shrub or small tree, up to 3(—8?) m, rather
densely pubescent in all the younger parts; hairs
partly glandular-capitate. Wood very hard. Leaves
somewhat fleshy, linear-spathulate, up to 314 by
V4 cm, crowded at the end of the branchlets, leaving
tuberculate scars; midrib, nerves, and veins in-
conspicuous. Inflorescences 2—4-flowered. Bracts
lanceolate, 4-9 by 1-114 mm. Pedicels up to c.
1 cm. Sepals ovate-lanceolate to ovate-oblong,
5-10 by 2-4 mm. Petals yellow, + obovate-
oblong to orbicular, shortly clawed, about as long
as the sepals. Filaments sericeous at the base,
up to 5 mm; anthers with emarginate top and base,
1 mm 9. Carpels hairy, obovoid, in anthesis up
to c. 1 mm long; styles glabrous, except at the very
base, up to 5 mm. Fruits hairy, subovoid, c.
314 mm long.

Distr. Pantropical (but not in West Africa,
not on the Asiatic and Australian! continents,
and not in Hawaii), in the Pacific-Indian Oceans
usually on small isolated islands or coral islets
and atolls, in Malaysia: very scarce, only found
in the Philippines (Lumbucan, Sulu Sea), Timor,
and the Tanimbar Is (unlocalized), and East
New Guinea (Kelana, Port Moresby, Misima I.).
Fig: 2:

| + ~
| °
If 5 °
i" : te
| + *
| ee
°
° * ree A Oe
2 oe
~.} “e
Ee ‘0 cm

Fig. 2. Distribution of Suriana maritima L. in
Malaysia and adjacent countries; material seen
(e), from literature (0), unlocalized (+).

Ecol. A coastal shrub of the Barringtonia
formation, usually rare, but locally often very
common and forming thickets along the sandy
beach and along coral coasts, often associated
with Messerschmidia, Scaevola, Guettarda, etc.,
from sea-level up to 10 m. Fi. fr. Jan.—Dec.

In SE. Polynesia F. B. H. BRown found that
the dense branching causes it to act occasionally
as a sand binder initiating the formation of small
dunes. In Bikini especially common on the wind-
ward side of the islet. In the Tuamotus one of the
most common littoral woody plants.

It is remarkable that almost all localities are

b

Fig. 1. Suriana maritima L. a. Flowering twig,

2/,, b. flower, X 2, c. petal, 3, d. stamen,
< 6, e. gynoecium, scars of other flower parts
visible, * 6, f. fruit, < % (a FosBerG 26890,
b-e McKee 4997, f after BRITTON).

situated in the small islands or islets in the Ma-
laysian Pacific area, except some in East New
Guinea. No specimen is known from the Aus-
tralian and Asiatic continents proper, though
BENNETT? (/.c.) recorded it from the ‘shores of the
western Peninsula’, i.e. the Deccan. Among shore
plants it shares this peculiarity with Pisonia
grandis for which it has been accepted that these
localities coincide with bird colonies, and con-
sequently guano accumulation leading to calcium
phosphate coral rock; Pisonia would then also
be dispersed by these sea birds. It is, however,
unlikely that the fruits of Suriana are dispersed in
this way.

Mr L. S. Situ (in /itt.) suggested that Suriana
and other rare littoral plants avoiding continental
shores (Messerschmidia argentea, Soulamea amara)
might prefer highly calcareous beaches which, of
course, are much more prevalent on islands than
on the mainland where rivers silt the coral by
outflow of freshwater and sediments.

Ripiey (/.c.) dwelt at length on the peculiar
distribution pattern and I agree that Guppy’s
assumption (Plants, Seeds, efc. 1917, 240) that
it would have been destroyed in spots as fire-
wood by wandering islanders is unlikely.

ScHIMpeR and Guppy (Obsery. 529) have
established that the buoyancy power of the nuts
is very great, up to at least 5 months; the floating
power being due to an unfilled space in its cavity

(1) Except two localities, near Yirrkalla, at the NE. corner of Arnhem Land and Look-Out Point in

Queensland.

198 FLORA MALESIANA [ser. I, vol. 6?

It is therefore certain that they are dispersed by In the herbaria Suriana is sometimes confused
sea-water, but ‘they could also be carried in with Pemphis acidula Forst. (Lythraceae) which
floating logs and pumice’; attachment in mud to can immediately be distinguished by a lengthwise
birds feet is possible though unlikely as its habitat sulcate, campanulate calyx tube and 6-merous
is too sandy. flowers.

2. QUASSIA

LINNE, Sp. Pl. ed. 2 (1762) 553,, app. (1763) 1679; Gen. Pl. ed. 6 (1764) 212;
Pierre, Bull. mens. Soc. Linn. Paris n. 156 (1896) 1236; NooreBoom, Blumea
11 (1962) in press. — For generic synonyms see under the sections. — Fig. 3-5.

Trees, shrubs, or suffrutices. Leaves pari- or imparipinnate, rarely simple;
leaflets usually with pitted glands in the upper surface along the margin, especially
at the apex; nerves and veins usually immersed or obscure, sometimes (in sect.
Quassia and some African and American spp.) prominent; rachis distinctly jointed
and winged in Q. amara, with 2 narrow ribs or terete and not jointed, or only so
in the apical part in other spp.. Inflorescence a simple or branched raceme, a
panicle, or an umbel; bracts usually spathulate, more or less succulent, or tri-
angular; bracteoles nearly opposite, tiny, triangular, ciliate. Flowers 4—-6-merous,
mono- or bisexual, or polygamous; pedicels jointed about the middle in Q. amara,
either jointed at the base or not in the other spp.. Calyx more or less lobed, rarely
(§ Simaba, p.p.) closed in bud and irregularly rupturing. Peta/s imbricate or con-
torted in bud, longer than the calyx, sometimes very long. Stamens obdiplostemo-
nous (in a single African sp. the outer whorl doubled), with a shorter or longer,
hairy, adaxial scale with a shorter or longer free apex. Disk -+ cylindrical or
subglobose, highly varying in size. Ovaries free or coherent, 4-6, on top of the
disk, often more or less immersed in it, the abortive ovaries of the 3 flowers in
some monoecious spp. surrounded by a barrel-like disk; style 1, but the parts of
each carpel discernible, and with as many style canals as there are carpels; stigmas
more or less stellately spreading, or one slightly lobed or capitate stigma. Fruits
1-6, drupaceous or woody, often compressed (laterally, or in one sp. dorsoven-

trally), either bicarinate or not, sometimes very large.

Distr. Pantropical, c. 25 species in tropical and subtropical America, 5-10 spp. in Africa, 2 spp. in
lower Burma and Cambodia, one of which also almost throughout Malaysia to the Bismarcks and Solo-
mons, 1 endemic in Borneo & Sumatra, and 2 in Queensland.

Ecol. In Malaysia in rain-forests at low altitude.

Notes. The new North Bornean species, which could be described thanks to the generous co-operation
of Mr ForMAn, Kew, necessitated a reconsideration of the trib. Simaroubeae. Mr FORMAN assumed it to
belong to the American genus Simaba, but though the similarity is striking indeed, Simaba is defined in
having bisexual flowers, and the flowers of the new species are male, with clearly reduced ovaries. Uni-
sexual flowers occur in the likewise American genus Simarouba. The new species showed, however, also
similarity to some African genera for example Hannoa and Odyendyea. I have come to the conclusion
that all these genera, including also Samadera and the African genus Pierreodendron, cannot be separated
generically and should be arranged in an enlarged genus Quassia, a point of view already suggested by
PIERRE in 1896. This emended genus Quassia comprises four sections, three of which occur in Malaysia.

KEY TO THE SPECIES
1. Leaves compound.
?. Leaf-rachis winged, conspicuously articulated. Flowers in racemes . «=; oy sage, Olvamara'
2. Leaf-rachis not winged, not articulated. Flowers in panicles . . ..... .. . 3. Q. borneensis
1. Leaves simple. Flowers in pseudo-umbels. 2. Q. indica

1. Section Quassia

Leaves pinnate, with a more or less winged and conspicuously articulated
rachis. Racemes terminal, either branched or not. Pedicels articulated about the

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

middle, with 2 tiny bracteoles below the joint. Flowers bisexual. Petals 5, con-
torted, oblong, erect, much longer than the calyx. Disk large, nearly as high as
broad. Styles long, with a small, slightly 5-lobed stigma.

Distr. One sp. native in Brazil and introduced in all tropical countries for medicinal and ornamental

purposes.

1. Quassia amara LINNE, Sp. Pl. ed. 2 (1762) 553,
app. (1763) 1679; Back. Fl. Bat. 1 (1907) 256;
Schoolfl. Java (1911) 190; Lecomre, Fl. Gén.
I—-C. (1911) 689; Merr. FI. Manila (1912) 272;
En. Philip. 2 (1923) 346; Crats, Fl. Siam. En. 1
(1926) 239; Herne, Nutt. Pl. (1927) 870; BAck.
Bekn. Fl. Java (em. ed.) 6 (1948) fam. 146, p. 3.

Very bitter, erect shrub, 2-3 m high. Leaves
with broadly winged rachis; rachis + petiole c.
5-16 cm; leaflets usually 5, apical ones reduced to
3-1; flush purple; almost sessile, obovate-oblong.
Racemes 10-25 cm long, often branched. Pedicels

most sometimes foliaceous, 3-14 mm long. Calyx
patent, bright red, 7-8 mm. Petals bright red out-
side, whitish inside, 27-32 by 5-6 mm. Stamens
longer than the petals, slightly unequal, 314-4 cm.
Drupes \—5, purple-black, 12-13 mm long.

Distr. Native of Brazil, in Malaysia cultivated,
occasionally naturalized.

Uses. The Quassi-wood is used as a tonic in
case of stomach diseases and as an insecticide
to destroy for instance plant lice. The active
constituent of the wood consists of a number
of bitter substances (HEYNE, /.c.).

8-14 mm, accrescent. Bracts spathulate, the lower-

2. Section Samadera

(GAERTN.) NOOTEBOOM, oy. stat.—Locandi ADANS. Fam. Pl. 2 (1763) 449, based
on RHEEDE, Hort. Mal. 6 (1686) t. 18, nom. gen. rejic.; O.K. Rev. Gen. Pl. 1
(1891) 104.—Samadera GAERTN. Fruct. 2 (1791) 352, t. 156, ‘f. 3’, nom. gen.
cons.; BOERL. Ned. Kruidk. Arch. IT, 5 (1890) 520-524.—Vitmannia VAHL, Symb.
Bot. 3 (1794) 51, t. 60.—Niota [Porr. in Lamk, Tabl. Enc. Méth. (1792) t. 299]
Lamk, Enc. Méth. 4 (1797) 490.—Biporeia Petit-THOUARS, Gen. Noy. Madag.
(1806) 14, nom. illeg —Mauduita Comm. ex DC. Prod. 1 (1824) 592, nom. inval.—
Manungala BLANCO, FI. Filip. (1837) 306.—Samandura LINNE [Fl. Zeyl. (1747)
202, pro specim. Herm., excl. RHEEDE t. 21] ex BAILLON, Hist. Pl. 4 (1873) 491,
nom. illeg.; Bot. Méd. 2 (1884) 845, 874; PrerreE in De Laness. PI. Utiles Col. Fr.
(1886) 305; Batti. Dict. Bot. 4 (1892) 11.

Leaves simple, with more or less scattered concave glands, usually on the
undersurface. Flowers bisexual, in axillary or terminal, peduncled pseudo-
umbels or in racemes. Calyx lobes 3—5, imbricate in bud, obtuse, in the centre
with a concave gland. Petals 3-5, contorted, much longer than the calyx, usually
hairy on the back. Disk large, as high as broad, gynophore-like. Style with a
terminal inconspicuous stigma. Fruits rather large, (in Mal.) laterally compressed
with a narrow unilateral sharp-edged thinner part in the apical half (in the Indo-

Chinese sp. very large and dorsoventrally compressed).
Distr. Two spp., Madagascar and from lower Burma and Cambodia throughout Malaysia (except
Java and the Lesser Sunda Islands) to the Bismarcks and Solomons. Q. indica is cultivated in Java.
Ecol. Usually at low altitude under everwet climate conditions.
Note. BACKER (1907) defined the flowers as 3—5-merous. In Q. indica I have only seen 4-merous ones.

2. Quassia indica (GAERTN.) NooTEBOOM, comb.
nov.—Samadera indica GAERTN. Fruct. 2 (1791)
352, t. 156, f. 3; W. & A. Prod. (1834) 151; Hook.
Ic. Pl. 1 (1837) t. 7; GrAH. Cat. Bomb. Pl. (1839)
37; PLANCH. in Hook. Lond. J. Bot. 5 (1846) 562;
Tuwaites, En. (1858) 70; Mia. Fl. Ind. Bat. 1, 2
(1859) 677; BENN. in Hook. f. Fl. Br. Ind. 1
(1875) 519; Kurz, For. Fl. Burma 1 (1877) 200;
Bianco, FI. Filip. ed. 3, 4 (1880) 38; VIDAL, Sin.
Atlas (1883) 19, t. 26, f. c.; Phan. Cuming. (1885)
101; Rev. Pl. Vasc. Filip. (1886) 78; TRIMEN,
Fl. Ceyl. 1 (1893) 231; GresHorr, Schetsen (1894)

17-19, t.; MeRR. Gov. Lab. Publ. Philip. 7. 27
(1905) 29; Back. Fl. Bat. (1907) 258, incl. var.
brevipetala (SCHEFFER) BAcK.; Schoolfl. Java
(1911) 191; Laur. Bot. Jahrb. 56 (1920) 342,
incl. var. papuana LAUT.; MERR. Sp. Blanc. (1918)
206; En. Born. (1921) 315; RipLey, Fl. Mal. Pen.
1 (1922) 363; Merr. En. Philip. 2 (1923) 345;
Back. Bekn. FI. Java (em. ed.) 4 (1948) fam. 146,
p. 2; Capuron, Adans. 1 (1961) 83.—Karin-
Njoti Rueepe, Hort. Mal. 6 (1686) t. 18.—
Vitmannia elliptica VAHL, Symb. Bot. 3 (1794)
51, t. 60.—Niota pentapetala Poir. in Lamk,

200

FLORA MALESIANA

[ser. I, vol. 62

Fig. 3. Quassia indica (GAERTN.) NOOTEBOOM. a. Twig with flowers and fruit, x 24, 6. bud, with glands on

calyx,

Encycl. 4 (1797) 490; DC. Prod. 1 (1824) 592;
BLANco, Fl. Filip. ed. 2 (1845) 213.—WNiota
tetrapetala Por. in Lamk, Tabl. Encycl. Méth.
(1792) t. 299; in Lamk, Encyel. 4 (1797) 490; DC.
Prod. 1 (1824) 592; BLANco, FI. Filip. ed. 2
(1845) 213.—Niota commersonii Pers. Syn. 1
(1805) 416, nom. inval—Mauduita penduliflora

< 3 (a after GreESHOFF, 6 [BoET 48).

Comm. ex DC. Prod. 1 (1824) 592, nom. inval.—
Samadera madagascariensis Juss. Mém. Mus.
Hist. Nat. Paris 12 (1825) 516, t. 27, n. 46, nom.
illeg—Niota lamarckiana Bu. Bijdr. 5 (1825)
251, nom. illeg.—Niota lucida WALL. Pl. As. Rar.
2 (1831) 54, t. 168.—Samadera tetrapetala G.
Don, Gard. Dict. 1 (1831) 811.—Samadera

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

pentapetala G. Don, /.c.—Samadera glandulifera
PRESL, Symb. Bot. 2 (1833) 1, t. 51.—Manungala
pendula BLANco, FI. Filip. (1837) 306.—Vit-
mannia lucida SteuD. Nomencl. ed. 2 (1841) 779.—
Samadera brevipetala ScHEFF. Nat. Tijd. N.I. 32
(1871) 410.—Samandura indica BAILL. Bot. Méd.
2 (1884) 874; PrerRE in De Laness. Pl. Utiles Col.
Fr. (1886) 305.—Locandia indica O.K. Rev. Gen.
Pl. 1 (1891) 104.—Locandia O.K. |.c.—Locandia
madagascariensis O.K. l.c—Samandura_ mekon-
gensis PIERRE, Fl. For. Coch. 4 (1892) t. 262, t.;
Lecomte, Fl. Gén. I.-C. 1 (1911) 694.—Locandia
glandulifera PreRRE, Fl. For. Coch. 4 (1892) sub
t. 262, text.—Locandia mekongensis PIERRE, /.c.
t. 262, text.—Locandia merguensis PIERRE, l.c.
sub t. 262, text, nomen.—Locandia pendula
Pierre, /.c. sub t. 262, text.—Samadera mekongen-
sis ENGL. in E. & P. Pfl. Fam. 3, 4 (1896) 210.—
Samandura madagascariensis PERRIER DE LA BATHIE,
Fl. Madag. Fam. 105 (1950) 6, t. 2.—Fig. 3.
Glabrous evergreen shrub or tree, up to 20 m.
Branchlets with a small pith, the base of each
shoot provided with some stiff persistent scales.
Leaves elliptic-oblong to lanceolate, rather acute
or sometimes rounded or even subcordate at the
base, blunt, more or less acuminate or sometimes
rounded at the apex, 12-30 by 4-12 cm, midrib,
nerves, and distinctly reticulated veins conspicu-
ously prominent at either surface, always with
internerval veins, usually with 2 pitted glands at
the base beneath and similar ones scattered on the
surface, less so above; petiole 1-214 cm. Flowers
up to 20 or more in an umbelliform, glabrous or
puberulous inflorescence. Peduncle more or less
flattened, thickened at apex, terminal or axillary,
sometimes on old wood, 1-30 cm. Pedicels jointed
in the lower half, 14-2'4 cm, growing during
anthesis, in fruit to more than 3 cm. Bracts minute.
Calyx 4-lobed, 2-3 mm high, lobes about as long
as the tube or longer, -_ semiorbicular, puberu-
lous outside. Petals 4, free, dorsally puberulous,
obtuse, usually narrowed to the base, growing
during anthesis, creamy-green to violet, purplish
or brownish, up to 3 by | cm. Filaments puberu-
lous, hairy except towards the apex, up to 2! cm,
inserted at the base of the disk; anthers lanceolate
to oblong, c. 4 by 1-2 mm. Disk glabrous or
nearly so, c. 2 by 1144 mm. Carpels 4, free, more
or less puberulous, c. 2 by 2 mm; styles up to 2
cm. Fruits 1-4 together, flattened, with ++ straight
inner and + semicircular outer margin, which is
sharp and thinner in the upper half, the apex more
or less overtopping the subapical stylar scar,
4-9 by 214-5 cm; pericarp with similar glands as
the leaf. Seed with an apical plumule and an
adaxial conspicuous chalaza; testa thin; endosperm
none; cotyledons planoconvex, up to 344 by 214
cm.
Distr. Madagascar, Ceylon, S. Concan, Ma-
labar, Lower Burma (Martaban, Tenasserim),

Fig. 4. Distribution of Quassia indica (GAERTN.)

Nootesoom (delineated and dots) and Q. har-

mandiana (PIERRE) NooTeBoom (dotted delineation
with localities in triangles).

Andamans, and Cochinchina, throughout Malay-
sia (not in Sumatra, Java, and the Lesser Sunda
Islands) to the Bismarcks and Solomons; cul-
tivated in Java and elsewhere. Fig. 4.

Ecol. Usually very rare, but locally rather
common in the eastern part of its area, preferably
in wet places in lowland, forests below 150 m,
sometimes in localities which are periodically
inundated by fresh or by salt water, for example
on edge of the mangrove, in East North Borneo
common in young swamp forests back of the
mangrove. Fi. fr. Jan.—Dec.

Uses. In Sarawak the wood is used for making
handles of knives, in the Solomons the macerated
leaves, mixed with coconut oil, are applied to hair
for cleansing purposes.

The seeds are given as an emetic and purgative,
and sometimes in bilious fevers. In the Philip-
pines chips of wood are put in coconut oil which
is drunk as a purgative. The same oil is used as a
liniment for rheumatism and bruises. The plant
is also used against malignant fevers, as a tonic,
and as insecticide, specially against ants.

The seeds contain oil to the extent of one third
of their weight, but by difficulty of getting a
sufficient supply it is not commercial (GRESHOFF,
Schets. 1, 1894, 19; HEYNe, Nutt. Pl. 1927, 869;

Burk. Dict. 2, 1935, 1945; Quis. Medic. PI.
Philip. 1951, 475).
Vern. Philip.: daraput, linatog-anat, linton-

gamai, mabingdato, palagarium, palagium, ponoan,
Bis., maluiggal, mdfgal, Tag., manunggdl, Tag.,
Bik., Pamp., P. Bis., Lan., Ibn., palo santo,
Spanish, rapus (tree), kélépis, klipis (fruit), Banka,
kaju pait, Borneo, gatép pait, Java, onne, Ternate.
Notes. According to CAPURON /.c. the species
is doubtless native in Madagascar and not rare
in the substage of swampy forests along the
east coast, rarely ascending on crests to 400-600 m.
The leaves show a resemblance to those of
Irvingia and Inocarpus but are distinguished by the
occurrence of scattered concave glands.

3. Section Simaba

Pierre, Bull. mens. Soc. Linn. Paris n. 156 (1896) 1236.—Quassia sect. Odyendyea

202 ; FLORA MALESIANA [ser. I, vol. 6?

Fig. 5. Quassia borneensis NOOTEBOOM. a. Habit, x %, b. 3 flower, x 5, c. ditto, from above, stamens
and petals removed, the disk seemingly consists of two rings, the star-shaped structure in the centre
represents the 5 vestigial carpels, x 10, d. frontal and dorsal view of stamen (a—d MEYER San 20499).

Pierre, /.c. 1238.—Simaba Aust. Hist. Pl. Guian. (1775) 400, t. 153.—Aruba
AuBL. /.c. 293, t. 115.—Hannoa PLANcH. in Hook. f. Lond. J. Bot. 5 (1846) 566.—
Mannia Hook. f. in B. & H. Gen. Pl. 1 (1862) 309.—Hyptiandra Hook. f. I.c. 293,
990.—Odyendyea (PIERRE) ENGL. in E. & P. 3, 4 (1896) 215.—Pierreodendron
ENGL. Bot. Jahrb. 39 (1906) 575.—Simarubopsis ENGL. Bot. Jahrb. 46 (1911) 276.

Dec. 1962] SIMAROUBACEAE (Nooteboom) 203

Leaves pinnate or simple, if simple more than thrice as long as broad. Flowers
bisexual or trees polygamous, in terminal or axillary panicles, which are sometimes
reduced to few-flowered, axillary, umbel-like clusters, or to pseudo-umbels with
forked peduncles. Petals imbricate or contorted. Scales of the stamens sometimes
nearly as long as the filament and somewhat coherent. Stigmas short, or only one
4-5-lobed or punctate stigma.

Distr. Pantropical, c. 20 spp. in tropical South and Central America, c. 5-10 spp. in Africa, 1 sp. in

Malaysia, and 2 spp. in Australia.

3. Quassia borneensis NOOTEBOOM, nov. spec.—
Fig. 5.

Arbor mediocris, foliis paripinnatis vel impari-
pinnatis, 2—3-jugatis. Foliola elliptico-oblonga vel
obovato-oblonga, abrupte acuminata, glabra, mar-
gine superne glandulis parvis munita, 8-12 cm
longa, 4-414 cm lata. Flores 3 sepalis puberulis
basi coalitis 1 mm longis; petalis imbricatis vel
contortis, elliptico-oblongis vel ovato-oblongis,
glabris, 3 mm longis 2 mm latis, staminibus squama
pilosa apice emarginata \4—1 mm longa semiadnata
instructis; disco insigni, basi 2 mm lato, apice 1 mm
lato, 4% mm alto; vestigiis carpellorum % mm
longis, stylo carpellis longitudine aequanti.—
Typus MEUER SAN 20499, in L, isotypes K, SAN.

Tree, 14 m by 25 cm 9g; outer bark densely
fissured, brittle and corky. Leaves spirally arrang-
ed, pari- or imparipinnate; leaflets 2-3 pairs,
glabrous, elliptic to obovate-oblong, shortly
rounded-acuminate, 8-12 by 4-4!4 cm; upper
surface shining, lower surface opaque; very small
pitted glands along the margins and in the acumen
on the upper surface; nerves sunken in both upper
and lower surface, or obscure, ending in a mar-
ginal vein; veins obscure; petiole c. 5 cm, as the ra-
chis + terete; petiolules 1-114 cm, articulated at
the base. Panicle puberulous in all its parts, not
quite as long as the leaves. Bracts spathulate,
succulent in the apical part, up to 24 mm long.
3 Flowers 4-5-merous. Pedicels up to 7 mm.
Calyx c. 1 mm high, outside puberulous, lobes
ovate to triangular, longer than the tube. Petals
contorted or imbricate in bud, glabrous, elliptic
to ovate-oblong, c. 3-4 by 2 mm. Stamens slightly
shorter than the petals; filaments sigmoid-folded
in bud, with a hairy adaxial scale at the base; scale

free for 14 of its length, more or less emarginate,
c. Y%-1 mm long; anthers oblong, latrorse, c.
14-1 mm long. Disk c. 4% mm high, at the basis
c. 2 and at the apex c. 1 mm wide, the upper half
distinct from the lower half and folded around the
barren ovaries. Carpels free, c. 4 mm high; style
as long as the carpels, with a small 4~S-lobed
stigma. 2 Flowers unknown. Fruits drupaceous,
1-5 from each flower, if more than one diverging
radially from a thickened torus, prune-shaped,
dark purple-red when ripe sec. coll.; in dry state
slightly flattened-ellipsoid, with a faint dorsal and
ventral ridge, c. 2-3 by 114 cm; pericarp thin but
hard. Seed with a thin testa; plumule short; cotyle-
dons large, green, plano-convex; no endosperm
(BurGEss 2849).

Distr. Malaysia: Central Sumatra (Indragiri),
Borneo.

Ecol. Primary rain-forest at low altitude,
often in peat-swamp forest, also on mineral soil.

Notes. Obviously closely allied to the African
species described by PIERRE in Quassia_ sect.
Odyendyea. All these species have the filaments
sinuously folded in bud, a condition not observed
in other species of Quassia.

Excluded

Niota globosa BLANCO, FI. Filip. ed. 2 (1845) 214
is, according to Mere. Sp. Blanc. (1918) 225 =
Cleidion spiciflorum (BURM. f.) MERR. (Euphor-
biaceae).

Niota polyandra BucH. Ham. ex W. & A. Prod.
(1834) 63, nomen subnudum; Vitmannia polyandra
SteuD. Nomencl. ed. 2, 4 (1841) 779 = Brown-
lowia tersa (L.) KosTeRM. (Tiliaceae).

3. EURYCOMA

Jack, Mal. Misc. 2 (1822) 45; Roxs. Fl. Ind. 2 (1824) 307; Pierre, Fl. For.
Coch. 4 (1892) t. 292, t. 293; ENGL. in E. & P. Nat. Pfl. Fam. ed. 2, 19a (1931)
380.—Picroxylon WaARB. in Fedde, Rep. 16 (1919) 256.—Fig. 6, 7a—e.

Treelets or rarely shrubs, up to c. 10 m high, monoecious or dioecious. Leaves
imparipinnate, usually multijugate, long and numerous, crowded at the tips of the
rather thick, pithy branches, leaving large scars. Leaflets opposite or subopposite,
slightly oblique, ovate-lanceolate to obovate-lanceolate, rarely ovate-oblong (or
linear extra-Mal.), sessile or nearly so, attached to the rachis with a conspicuous
articulation; midrib slightly prominent on the upper surface, prominent beneath;
nerves inconspicuous above and below, or slightly sulcate beneath, straight, ending

FLORA MALESIANA [sera If vole?

r* ‘
a> a one od

—

Fig. 6. Habit of Eurycoma longifolia Jack (Photogr. W. Mewer, Sandakan, June 1960).

Dec. 1962] SIMAROUBACEAE (Nooteboom) 205

in an intramarginal, looped vein. Panicles axillary, mostly large and lax, puberu-
lous, usually also with thickish, short, stiff, capitate-glandular hairs. Flowers
bisexual, 2 or 3; 2 flowers always with rather large but sterile stamens, 3 flowers
always with a sterile pistil. Calyx small, 5(—6) lobed, lobes ovate to triangular,
acute or bluntish, longer than the tube. Petals 5(—6), induplicate-valvate in bud,
lanceolate or ovate- to obovate-oblong. Stamens 5(-6), episepalous, filaments
narrowing to the top, usually with a very small (c. !/; mm long) adaxial ligule at
the base, alternating with 5(-6) small entire, emarginate or cleft staminodes,
which are usually connate with the abaxial and lateral sides of the base of the
filaments; sometimes there is a second row of still smaller entire staminodes out-
side the stamens; stamens and staminodes sometimes connate with the base of
the petals; filaments glabrous or sparsely hairy. Disk inconspicuous. Carpels
5(—6), free, the style attached adaxially near the top and mutually connate or cohe-
rent; stigma peltate, 5(—6)-lobed. Each ovary with | anatropous ovule with adaxial
placenta. Fruits up to 5, c. 3 mm stalked, spreading, ellipsoid or ovoid, slightly
bicarinate nuts with very thin exocarp and hard endocarp. Seed exalbuminous with

2 planoconvex cotyledons and a short plumule.

Distr. Three spp. in tropical SE. Asia (Lower Burma, Siam, Indo-China), Sumatra, the Malay Pe-
ninsula, Borneo, and the S. Philippines. Fig. 8, 14.

Ecol. Preferably on sandy soils below 1200 m, sometimes flowering at an early age.

Uses. The roots, and particularly the bark of the rcots, are used as a febrifuge. The Malays give it
also as a tonic, e.g. after childbirth. In Borneo a decoction of the bark is drunk to relieve pain in the bones
and a decoction of the leaves is used for washing itches. The Malayan name bédara /aut is also used for
Strychnos, which has the same uses (cf. BURKILL, Dict. 1, p. 984).

Vern. (for both Mal. spp.). Malay Peninsula: bédara mérah, b. puteh, b. pahit, bumi, lémpédu pahit,
muntah, payong ali, pénawar pahit, pétala bumi, tongkit ali, t. baginda, M; akar jangat sémang, duak,
jélas, Sakai; Sumatra: bégu-gad-jan, bésan, béséng, Karo; bidara laut, b. putih, M; kaju pétimah, k. porhis
potala, Alas; njatu suria, Taram; kaju pulae, mémpoleh, Banka; parie potala, Padang; Borneo: bina,
kabal kabal bérang, sérirama, tongkat ali; babi kurus, J for the drug.

Note. Though in current classifications Eurycoma is referred to the subtribe Eurycominae and the
genus Quassia to the subtribe Simaroubinae, of the tribe Simaroubeae, I find it unsatisfactory to di-
vorce Eurycoma from Quassia which show an astonishing similarity in both vegetative and genera-
tive aspects, save that whereas Eurycoma has 5 stamens and 5 staminodes, valvate petals and a less deve-
loped disk, there are in Quassia 10 stamens, contorted or imbricate petals, and a well-developed disk.
In my opinion they represent a couple of closely related genera of one tribe.

KEY TO THE SPECIES

1. Leaflets ovate-lanceolate to obovate-lanceolate, rarely ovate-oblong, more than 14 cm wide.

2. Leaflets not or slightly acuminate, but the apex rather acutish as compared with next species. Petals
about twice as long as wide. Anthers c. 14mm long. Styles rather aay stigma c. 1 mm above the
ovaries : Sts 1. E. longifolia

2. Leaflets usually rather abruptly very blunt- acuminate: Petals linear 4 or more times as long as wide.
Anthers c. 34 mm long. Styles very short, stigma sessile mene 5 2. E. apiculata

1. Leaflets linear, up to 7 by c. 44 cm, hard coriaceous, HES 8-18 cm n long. Dwarf tree or shrub, up to
1 m high. Petals pubescent on both sides. Anthers c. 34 mm long. Panicles without glandular hairs.
(Siam and Cambodia.) Saewicy 600 aa 3. E. harmandiana

1. Eurycoma longifolia Jack, Mal. Misc. 2 (1822) (1911) 695; Back. Schoolfl. Java (1911) 193;

10 (1915) Bot. 190; En.

45; Roxs. Fl. Ind. 2 (1824) 307; DC. Prod. 2
(1825) 86; Mig. Fl. Ind. Bat. 1, 2 (1859) 681;
BENN. in Hook. f. Fl. Br. Ind. 1 (1875) 521; F.-
Vitt. Noy. App. (1880) 39; VipAL, Rey. Pl.
Vasc. Filip. (1886) 78; Pierre, Fl. For. Coch. 4
(1892) t. 292, t. 293, incl. var. merguensis and
var. cochinchinensis; KiNG, J. As. Soc. Beng. 62, ii
(1893) 229; Rip. J. Str. Br. R. As. Soc. 30 (1897)
205, err. latifolia; J. Str. Br. Med. Assoc. n. 5
(1897) 127, ditto; Lecomte, Fl. Gén. I.-C. 1

Me_rr. Philip. J. Sc.
Born. (1921) 316; RipLey, Fl. Mal. Pen. 1 (1922)
362; Merr. En. Philip. 2 (1923) 346; Crats, FI.
Siam. En. 1 (1926) 242; Merr. Pl. Elm. Born.
(1929) 116.—E. merguensis PLANCH. in Hook.
Lond. J. Bot. 5 (1846) 584,—E. tavoyana WALL.
Cat. (1847) n. 8523, nomen.—Picroxylon siamense
Wars. in Fedde, Rep. 16 (1919) 256.—Manotes
asiatica GAGN. Bull. Soc. Bot. Fr. 98 (1951) 207
(VIDAL, in litt.).

FLORA MALESIANA

[ser. I, vol. 6?

Fig. 7. Eurycoma longifolia Jack. a. 3 Flower,
< 4, b. ditto, petals and sepals removed, * 8,
c. stamen, * 16, d. 2 flower, petals removed,
x 8, e. fruits, nat. size.—Ailanthus integrifolia
LaMK. f. Flower, X 2, g. ovary, < 4 (a-d MEIER
6712, e KOSTERMANS 6654, f—g C.H.B. III—E-2).

ssp. longifolia.—Fig. 6, 7a-e.

Leaves up to c. 1 m long. Leaflets lanceolate to
obovate-lanceolate, rarely ovate-oblong, some-
times slightly acuminate with a bluntish or acute
apex, c. 5-20 by 114-6 cm. Panicles, pedicels,
sepals, and calyx puberulous and with capitate-
glandular hairs. Flowers reddish. Bracts triangu-
lar, very small, up to c. 1 mm, caducous. Pedicels
rather thick, up to c. 7 mm. Calyx small, lobes
c. 1 mm long. Peta/s puberulous on both surfaces,
lanceolate to ovate- or obovate-oblong, c.
414-54 by 2-3 mm. Stamens usually longer than
the calyx, c. 144-214 mm long, anthers c. 44 mm
long. Staminodes from 14 mm in & flowers to c. 2

Fig. 8. Distribution of Eurycoma longifolia JACK
(delineated, dots) and its var. eglandulosa (MERR)
Nooresoom (2 lined dots).

mm in ¢ flowers. Styles rather long, with a peltate
5(—6)-lobed stigma elevated c. 1 mm above the
ovaries. Fruits 10-17(—20) by 5-12 mm.

Distr. Lower Burma, Siam, Laos, Cambodia,
Indo-China; in Malaysia: Sumatra, Malay
Peninsula, and Borneo. Fig. 8.

Ecol. Frequent at low altitude in beach forests
on sandy soil, in primary and secondary forest
as an understorey treelet, according to RICHARDS
(J. Ecol. 24, 1936, 22) in mixed dipterocarp rain-
forest, in heath forest and in low forest on ridge
crests in Sarawak on Mt Dulit, on sandstone
and dry kerangas, a characteristic silicicolous
species, locally frequent, generally at low altitude,
rarely up to 500 or even 1000 m. Fi. fr. Jan.—
Dec.

Note. Eurycoma longifolia JACK ssp. longifolia
has been recorded from the Philippines by F.-
VILLAR and VIDAL, but this record rested on an
erroneously localized specimen (Loss 486) which
according to MERRILL (1915) probably came from
Borneo or Malaya.

ssp. eglandulosa (MERR.) NOOTEBOOM, nov. stat.
E. eglandulosa MER. Philip. J. Sc. 17 (1920) 266;
En. Philip. 2 (1923) 346.

Panicles, pedicels, and flowers puberulous,
without glandular hairs. Petals 5-614 mm long.

Distr. Malaysia: Philippines (Mindanao: Su-
rigao; Dinagat), twice collected. Fig. 8.

Ecol. In forests at low altitudes.

2. Eurycoma apiculata BENN. in Hook. f. Fl. Br.
Ind. 1 (1875) 522; Kina, J. As. Soc. Beng. 62, ii
(1893) 230; Riot. Fl. Mal. Pen. 1 (1922) 363.

Leaves c. 40 cm long. Leaflets usually rather
abruptly very blunt-acuminate, c. 8-14 by 24 cm.
Panicles, pedicels, calyx, and petals with thick,
stiff, capitate-glandular hairs. Bracts small, linear,
up to c. 1 mm long. Pedicels rather slender, up
to c. 7 mm long. Calyx lobes c. 1Y%—2 mm long.
Petals puberulous, with glandular hairs outside,
glabrous within, linear, rarely lanceolate, c.
4-9 by 1-114 mm. Stamens usually as long as the
calyx or shorter, c. 2 mm long, anthers c. 34 mm
long; ligule usually absent; staminodes in 1 or 2
rows, up to c. 44 mm, small or absent. Styles
very short, with a 5-lobed sessile stigma. Fruit as
in former species.

Distr. Malaysia: Sumatra, Malay Peninsula.
Fig. 14.

Ecol. Similar as in E. longifolia, but usually at
higher altitude, up to c. 1200 m. Fi. fr. Jan.—Dec.

3. Eurycoma harmandiana Pierre, Fl. For. Coch.
4 (1892) t. 292 B; Lecomte, FI. Gén. I.-C. 1 (1911)
696; Crais, Fl. Siam. En. 1 (1926) 242.

Excluded

Eurycoma dubia Emer, Leafl. Philip. Bot. 2
(1908) 481 is according to HALLIER, Rec. Tray.
Bot. Néerl. 15 (1918) 55 and MERRILL, En. Philip.
2 (1923) 329 = Evodia meliaefolia (HANCE)
BENTH. (Rutaceae).

Dec. 1962] SIMAROUBACEAE (Nooteboom) 207

4. HARRISONIA

R. BROWN ex A. Juss. Mém. Mus. Hist. Nat. Paris 12 (1825) 517, nom. gen.
cons. prop., non ADANS. ex LEMAN, 1821; cf. Taxon 10 (1961) 243.—Ebelingia
Reus. Consp. (1828) 199.—Lasiolepis BENN. Pl. Jav. Rar. (1844) 202.—Fig. 9.

Thorny, erect or sprawling shrubs, rarely small trees, up to 12 m. Branches
pithy, older ones glabrous, lenticellate; stipular thorns accrescent, conical, finally
caducous, slightly recurved, up to 7 mm; annual shoots at the base with small
persistent bud-scales and sometimes spines. Leaves imparipinnate or ternate;
rachis narrowly winged; leaflets subentire to lobed, + sessile, the apical one
whether or not with a longer petiole than the lateral ones but without articulation,
rhomboid to ovate-lanceolate, blunt. Flowers bisexual, 4—-5-merous, in bracteate
axillary cymes or terminal, rarely axillary thyrses. Calyx small, lobes acutish-
triangular, about as long as the tube or longer. Peta/s much longer than the calyx,
slightly imbricate in bud. Stamens attached at the base of the disk, twice as many
as petals; filaments with an adnate 2-lobed cr emarginate hairy ligule free at its
top; anthers latrorse, cells diverging in lower half; filament attached between the
cells. Ovary 4—5-celled, slightly lobed, seated on a rather thick disk; ovules | per
cell, amphitropous, pendent from the adaxial side near the top; styles 4-5, connate
or sometimes free at the very base; stigma knob-shaped, slightly 4-S-lobed. Drupe
depressed-globose, sometimes + lobed; exocarp fleshy or coriaceous; endocarp
hard; fertile cells 2-5, each with a perforation of the hard endocarp at the base
of the stylar canal. Seed with a thin testa, endosperm present; cotyledons horse-

shoe-shaped, radicle pointing upwards.

Distr. About 3-4 spp. in tropical Africa and from SE. Asia through Malaysia (2 spp.) to North Austra-
lia.

Ecol. The Malaysian spp. usually on dry, open, hot places, often on limestone rocks, under distinctly
seasonal conditions, usually at low altitude, up to 700 m, locally sometimes extremely common in thickets,
less common in open monsoon forests.

Uses. In some parts of Malaysia the shoots are used as a drug against diarrhoea. In the Philippines
a decoction of the bark and roots is used against diarrhoea and dysentery, and apparently also against
cholera (HEYNE, Nutt. Pl. 1927, 871; Burk. Dict. 1935, 1128).

Note. The leaves resemble some species of Zanthoxylum (Fagara) but these are gland-dotted.

KEY TO THE SPECIES

1. Leaves ternate. Flowers usually 4-merous, c. 4mm long. Fruits 4-5 by 7-9 mm . 1. H. brownii
1. Leaves imparipinnate, (1—) 3-8 (—13)-jugate. Flowers usually 5-merous, 6-10 mm long. Fruits 4-9 by
11-15 mm. Ana ce) oaks, A 9 2. H. perforata

1. Harrisonia brownii A. Juss. Mém. Mus. Hist. 5(-8) cm; lateral leaflets usually oblique, cuneate

Nat. Paris 12 (1825) 540, pl. 28, n. 47; GAUDICH.
Bot. Freyc. Voy. (1826) t. 103, non vidi; DECNE,
Herb. Tim. Descr. (1835) 120; Mig. Fl. Ind.
Bat. 1, 2 (1859) 677; BENTH. FI. Austr. 1 (1863)
376; F.-Vitt. Novy. App. (1880) 39; VIDAL,
Phan. Cuming. Philip. (1885) 101; Rev. Pl. Vasc.
Filip. (1886) 78; BatLey, Queensl. Fl. 1 (1899)
221; Back. Schoolfl. Java (1911) 194; Merr.
En. Philip. 2 (1923) 346; Back. Bekn. Fl. Java
(em. ed.) 6 (1948) fam. 146, p. 4.—Ebelingia
brownii SteuD. Nom. ed. 2 (1840) 535; O.K.
Rey. Gen. Pl. 1 (1891) 103.—Fig. 9f.

Leaves ternate; apical leaflet gradually narrowing
into a 0-1 cm long petiolule, 114—8(-13) by 14—-

towards the rachis, 1-S by 14-214 cm; petiole
14-3 cm. Cymes and thyrses up to 5(—7) cm long.
Bracts persistent, triangular, pubescent, c. 11 mm
long, once found like a small leaflet. Flowers
4(—S)-merous. Pedicels up to 6 mm. Calyx glabrous
or sparsely hairy, c. Y% mm high. Petals lanceo-
late to oblong, 3144-5 by 1144-214 mm. Anthers
114-2 by 1 mm; filaments c. 2mm; ligule 1-114 mm.
Disk + short-cylindrical, slightly 8- or 10-lobed
below the margin, 4-—'4 mm high. Ovary c.
14-1 mm high, rather deeply lobed; style 14-2'4
mm. Drupe c. 4-5 by 7-9 mm; exocarp thin,
fleshy; endocarp hard; each cell with an abaxial
suture.

208

FLORA MALESIANA

[sera 1, -volaaa2

Fig. 9. Harrisonia perforata (BLCO) MERR. a.

Flowering twig, * 24, b. flower, 2, c. gynae-

cium and disk, =< 4, d. stamens, 8, e. fruit,

nat. size-—H. brownii A. Juss. f. Fruit, x 2

(a MERRILL 433, b-d TEYSMANN & DE VRIESE
5.n., €@ KOORDERS 30017, f BACKER 19469).

Distr. North Australia (islands in the Gulf of
Carpentaria), S. Andaman, and Malaysia: S.
Philippines (Palawan, Mindanao, Cebu, Bohol,
Siquijor), E. Celebes (Banggai Pen., Muna I.),
E. Java (also Madura and Kangean), Lesser
Sunda Islands (Bali, Sumba, Sumbawa, Flores,
Timor, Wetar), S. Moluccas (Babar and Tanimbar
Is), and SE. New Guinea. Fig. 10.

Ecol. and Uses. See under the genus.

Vern. Philip.: kankasira, Tagb., malomanhak,
C. Bis.; kaju bilis, Md, tadaibana, tara kedauk,
Sumba, kai tudu, Timor.

Notes. In one sheet I found detached fruits
of which the cells had dehisced along the suture.

The species has erroneously been recorded from
Malaya by Kina (J. As. Soc. Beng. 62, ii, 1893,
227) and Rip.ey (FI. Mal. Pen. 1, 1922, 360) in
confusion with H. perforata.

The leaves of the S. Andaman specimen (KING
s.n. 7-3-1891, in SING) agree with those of H.
brownii, but the buds are immature and fruit is
lacking, defeating proper identification. KuRz also
with doubt referred material from the Andamans
to H. brownii (Rep. Veg. Andam. 1870, 33) but
his material was lost. Plant-geographically the
Andamans represent a marked and unexpected
extension of the range towards the west.

2. Harrisonia perforata (BLANCO) Merr. Philip.
J. Sc. 7 (1912) Bot. 236; Fl. Manila (1912) 272;
Sp. Blanc. (1918) 206; En. Philip. 2 (1923) 346;

Crais, Fl. Siam. En. 1 (1926) 243; GAGNEP.
Fl. Gén. I.-C. Suppl. 1 (1946) 659; Back. Bekn.
Fl. Java (em. ed.) 6 (1948) fam. 146, p. 4; FORMAN,
Kew Bull. 1957 (1958) 503.—Paliurus perforatus
Bianco, Fl. Filip. (1837) 174, ed. 2 (1845) 122,
ed. 3, 1 (1877) 220.—Paliurus dubius BLANCO,
Il. cc. 175, 123, and 221.—Lasiolepis paucijuga
BENN. Pl. Jav. Rar. (1844) 202, t. 42.—Lasiolepis
multijuga BENN. L.c. 204.—Lasiolepis bennettii
PLaNcH. in Hook. Lond. J. Bot. 5 (1846) 570,
nom. illeg., incl. var. a paucijuga (BENN.) PLANCH.
and var. B multiijga (BENN.) PLANCH.; Mia. FI.
Ind. Bat. 1, 2 (1859) 678.—Limonia pubescens
WALL. ex Hook. f. Fl. Br. Ind. 1 (1875) 507.—
H. paucijuga Oxtyv. Fl. Trop. Afr. 1 (1868) 312,
in obs.; BACK. Fl. Bat. 1 (1907) 257; Schoolfi.
Java (1911) 194; Heyne, Nutt. Pl. (1927) 871.—
H. bennettii BENN. in Fl. Br. Ind. 1 (1875) 519,
nom. illeg.; KURZ, For. Fl. Burma 1 (1877) 203;
F.-ViLL. Nov. App. (1880) 39, incl. var. paucijuga
and var. multijuga; VIDAL, Sinops. Atlas (1883)
19, t. 26 f. A; Phan. Cum. Philip. (1885) 101;
Rev. Pl. Vasc. Filip. (1886) 78; Lec. Fl. Gén.
I.-C. 1 (1911) 689; RipL. Fl. Mal. Pen. 1 (1922)
360.—Fagara piperita (non L.) NAvVEs in Blanco,
Fl. Filip. ed. 3, 1 (1877) t. 23, excl. syn. DC.,
cf. F.-ViLL. Novis. App. (1880) 39 and MeErr.
Sp. Blanc. (1918) 206.—Ebelingia paucijuga O.K.
Rev. Gen. Pl. 1 (1891) 103.—H. citrinaecarpa
Exim. Leafi. Philip. Bot. 8 (1915) 2828.—Fero-
niella pubescens TANAKA, Bull. Mus. Hist. Nat.
Paris II, 2 (1930) 161; ENGL. Pfl. Fam. ed. 2,
19a (1931) 354; SwINGLE in The Citrus Industry
1 (1943) 470; GAGNEP. FI. Gén. I.-C. Suppl. 1
(1946) 651.—Fig. 9a-e.

Leaves 1—15-jugate, up to c. 20 cm; rachis
narrowly winged, usually with a rib above, more
or less pubescent, especially above; leaflets 10-20
by 5-15 mm; petiole 14-3 cm. Branches of cymes
and thyrses usually for some length adnate to the
peduncle. Pedicels up to 2 mm. Calyx c. 14% mm
high, lobes c. 34 mm. Petals lanceolate, rarely
oblong, 6-9 by 2-4 mm. Anthers c. 144-44 mm;
filaments 7-10 mm; ligule densely woolly at the
margin, c. 2 mm. Disk cup-shaped, 1—2 mm high.
Ovary 4-1 mm high, slightly lobed; style pu-
bescent, 5-8 mm. Fruit 4-9 by 11-15 mm; exo-
carp coriaceous, at least 1 mm thick; endocarp
hard; no suture in the endocarp.

f. er

Fig. 10. Distribution of Harrisonia brownii A.
Juss.

Dec. 1962] SIMAROUBACEAE (Nooteboom) : 209

Distr. SE. Asia (Hainan, Cochinchina, Cam-
bodia, Siam, and Burma) and Malaysia: Malay x : - Ti SSS al
Peninsula (Perlis, ?Kedah, ?Perak), Philippines, ;
N. Borneo (Sandakan), Celebes (also Buton), Sas
Java (also Madura and Kangean), S. Sumatra | ~
(Lampongs), and Lesser Sunda Islands (Bali). )
Fig. 11.

Ecol. and Uses. See under the genus.

Vern. Philip.: asimau, laiya, mamigil, Tag.,
bakit, sapsapdng, \\k., Pang., dagiangas, Mbo.,
kamungi, Sul., muntani, Bis., sapléng, Sbl.; ri
kéng-kéng, J, Md., garut, sésépang, Lamp. i ;

Note. In the sterile state sometimes difficult to _ i
distinguish from certain spp. of Fagara (Zan- L— = J
thoxylum) which have pellucid glands in the leaf Fig. 11. Distribution of Harrisonia perforata
and large glands along the margin near the teeth. (Brco) Merr., from Asia following the drought
However, in H. perforata the latter may occa- corridor through Central Malaysia.
sionally also be observed.

5. BRUCEA

J. F. MILt. Icon. (1779) t. 25, nom. gen. cons.; L’ HERIT. Stirp. (1784) 19, t. 10;
Bruce, Travels 5, App. (1789) 69, tab.; Cartes & Fig. Voy. Nub. & Abess.
(1792) t. 21; ENGLER, Nat. Pfl. Fam. ed. II, 19a (1931) 386.—Lussa RUMPH.
[Herb. Amb. 7 (Auct.) (1755) 27, t. 15, nom. inval.| ex O.K. Rev. Gen. Pl. 1
(1891) 104.—Gonus Lour. Fl. Coch. (1790) 158.—Fig. 12.

Very bitter, monoecious or dioecious shrubs or small trees; at least the younger
parts pubescent or puberulous. Leaves exstipulate, imparipinnate, petiolar base
and rachis-joints shrunken in the herb.; leaflets 3-15, more or less oblique, ovate
to lanceolate, acuminate, entire or not, on the undersurface with scattered, flat,
spot-like glands along the margin, situated under the teeth if these are present.
Flowers uni- or bisexual, in axillary inflorescences (in Mal.) which consist of small
cymes, united into bracteate, mostly unbranched raceme-like thyrses. Sepals 4,
connate at the base, imbricate in bud, ovate-elongate or triangular, small. Petals
4, free, imbricate in bud, ovate-oblong, oblong or linear, small. Disk thick, with
4 lobes. Stamens 4, with short filaments, inserted between the lobes under the
outer margin of the disk; filaments attached in the middle, basal, between the
divergent latrorse cells of the cordate-ovate anthers. Stamens vestigial or absent
in the 2 flowers. Ovaries 4, free, ovate; ovule 1, anatropous, pendent, attached
above the middle at the adaxial side. Styles free or coherent at the base, absent
laterally, adaxially, short, subulate, widened in a thickened or club-shaped stigma,
bent outward over the top of the ovary. Fruit -- drupaceous, hardly fleshy. Mature
dried nuts ovoid, with 2 ribs; pericarp thin, endocarp wrinkled and hard. Seeds
ovoid, with a thin testa and a thin to very thin endosperm; embryo with a short
plumule and 2 planoconvex cotyledons.

Distr. Old World tropics, c. 4 spp. in tropical Africa and 2 in tropical Asia (to S. China & S. Formosa),
Malaysia and North Australia; B. javanica introduced in Fiji and Ponape.

Notes. Brucea was on the list of nomina generica conservanda because of the name Lussa RuMPH., but
as the RuMpHIUs names have no nomenclatural standing this conservation is unnecessary.

BACKER (FI. Bat. 1907, 260) mentioned the rare occurrence of 5-merous flowers in B. javanica; I have
not observed them.

MILLER’s Icones is inthe B. M. Library; although cited generally as [cones Animalium et Plantarum the
original title was Various Subjects in Natural History (STEARN in litt.).

210 FLORA MALESIANA [ser. I, vol. 6?

Fig. 12. Brucea javanica (L.) MERR. a. Twig in flower and fruit, < 24, 6. d flower, < 8, c. 2 flower, = 8,
d. fruits, * 2.—Brucea mollis WALL. ex Kurz. e. Leaf, x 24, f. fruit, x 2 (a after GRESHOFF, b ELBERT
2807, c CoLrs 174, d ENpERT 1971, ef Ramos 13612).

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

211

KEY TO THE SPECIES

1. Dried fruits 4-5 (-7) mm long, with pedicels of 2-6 mm. Endosperm 1/5—1/2 mm thick. Pedicels of the
2 flowers up to 214 mm long. Leaflets always bluntly serrate or crenate with 6-15 pairs of nearly par-

allel nerves which without anastomosing directly end ina marginal gland;

nerves 1-10 (—17) mm apart.

insertions of the lateral
1. B. JNO

1. Dried fruits 9-13 mm long, with pedicels of 4-10 mm. 1. Endosperm ‘membranous. Pedicels of the 9
flowers up to 6 mm long. Leaflets entire or a little toothed, sometimes bluntly serrate, crenate or un-
dulate, with 4—9 (-11) pairs of mostly not parallel nerves which usually anastomose before reaching

the margin; insertions of the lateral nerves 3-30 mm apart .

1. Brucea javanica (L.) Merr. J. Arn. Arb. 9
(1928) 3; Backer, Bekn. Fl. Java (em. ed.) 6
(1948) fam. 146, p. 5; A. C. Smitu, J. Arn. Arb.
(1955) 279; Nair & SUKAMARAN, Bot. Gaz. 121
(1960) 175-185 (floral morph. and embryol.).—
Rhus javanica LINNE, Sp. Pl. (1753) 265; ed. 2
(1762) 380 (T. in LINN).—Lussa radja RUMPH.
Herb. Amb. (Auct.) 7 (1755) 27, t. 15.—Gonus
amarissimus Lour. Fl. Coch. (1790) 658 (T. in
P).—Ailanthus gracilis SALIsB. Prod. (1796) 171.—
B. sumatrana Roxs. Hort. Beng. (1814) 12,
based on Lussa radja RUMPH.; SPRENG. Syst.
Veg. 1 (1825) 441; DC. Prod. 2 (1825) 88; BLUME,
Bijdr. 17 (1826) 1167; Roxs. FI. Ind. ed. Carey |
(1832) 449; BeNN. PI. Jav. Rar. (1844) 200; Mia.
Fl. Ind. Bat. 1, 2 (1859) 680; Bru. Fl. Austr. |
(1863) 373; BENN. in Hook. f. Fl. Br. Ind. 1 (1875)
§21; BAILEY, Queensl. Fl. 1 (1899) 218; BACKER,
Fl. Bat. (1907) 260; Schoolfl. Java (1911) 192;
Lecomte, FI. Gén. I.-C. (1911) 698; Ripv. FI.
Mal. Pen. 1 (1922) 361.—B. sumatrensis SPRENG.
Pl. Min. Cogn. 2 (1815) 90.—B. gracilis DC.
Prod. 2 (1825) 88.—B. glabrata Decne, Nouv.
Ann. Mus. Par. 3 (1834) 447, t. 20; Herb. Timor.
Descr. (1835) 119; Mig. Fl. Ind. Bat. 1, 2 (1859)
680.—B. amarissima Drsy. ex Gomes in Mem.
Acad. Sc. Lisb. n.s. 4, pars 1 (1872) 30; MeRR.
Philip. J. Sc. 10 (1915) Bot. 18; Int. Rumph.
(1917) 299; Backer, Trop. Natuur 11 (1922)
134; Merr. En. Philip. 2 (1923) 347; CRAIB,
FI. Siam. En. 1 (1926) 241; N. C. Nair, J. Bomb.
Nat. Hist. Soc. 57 (1960) 237-238, t—Fig. 12a—d.

Shrub or small tree, 0.3-10 m, up to 10 cm o.
Leaves 20-50 cm long; leaflets 3-15, ovate-
oblong to ovate-lanceolate, sparsely hairy above,
more or less pubescent beneath, sometimes com-
pletely glabrous, 314-11 by 114-5 cm, the younger
ones usually densely pubescent; petiole 5-10 cm,
lateral petiolules 2-9 mm, terminal one 3-40 mm.
Peduncle almost absent, rachis 7-60 cm. Bracts
deltoid, small. Flowers greenish-white to greenish-
red or purple.—<: Pedicels very slender, up to c. 3
mm long, sepals pubescent, sometimes toothed,
c. Y—-1 by 1/3—'4 mm; petals sparsely pubescent
to nearly glabrous, sometimes toothed, 1-2 by
14-1 mm; filaments subulate, c. 0.6 mm, anthers
c. 0.4mm long.—®%: Pedicels up to 244 mm, sepals
and petals as in 3; stamens 0 or vestigial. Mature
dried drupes 1-4 together, 4-5(-7) mm long,
pedicels 2-6 mm.

Distr. From Ceylon and the Deccan through

2. B. mollis

SE. Asia to S. China and S. Formosa, throughout
Malaysia to N. Australia (N. Territory and N.
Queensland); introduced in Fiji (A. C. Smitu,
1c.) and Micronesia: Ponape (KANEHIRA, En.
Micr. Pl. 1935, 343). Fig. 13. It is most remarkable
that the localities of this very tolerant plant are
very scattered in East Malaysia; there is no
material from the central Moluccas (Buru,

Ceram, Ambon) and only one recent sheet from
the Wassi Kussa area in New Guinea. RUMPHIUS
knew it only from the Lesser Sunda Islands.
From this distribution pattern it can be deduced
that man has probably imported it in several
places of its area, but this cannot be traced
or proved in detail. Fig. 13.

ek

Fig. 13. Distribution of Brucea javanica (L.) MERR.
(open dot after literature).

Ecol. A common, tolerant species, preferring
open sites and light secondary forest and thickets,
forest edges and ridges, even occurring in sunny
places in sandy dunes and on limestone rocks, under
both everwet and seasonal conditions (0-900 m).

In 1907 BAcKer stated that all specimens he had
seen had bisexual flowers; in 1911 he had received
material from Semarang with g and flowers.
According to BACKER (1922) in Java specimens
with bisexual flowers would be found only west of
the line Semarang-Djokjakarta, plants with uni-
sexual flowers east of it. In my opinion unisexual
flowers prevail.—F/. Jan.—Dec.

Uses. The very bitter roots and fruits are used
as a medicine against dysentery and other fevers,
and against diarrhoea. The leaves are applied
against spleenomegaly and internal pains, scurf,
ringworm, boils, and centipede-bites. The fruits
are well known under the name of Macassar
kernels, E, and Makassaarse pitjes, D (GREsS-

(1) The length of the pedicels in flower and fruit has been measured above the last bract or bracteole.

212

FLORA MALESIANA

[ser. I, vol. 6?

HOFF, Nutt. Ind. Pl. 1894, 71; Heyne, Nutt. Pl.
Ned. Ind. 1927, 871; BURKILL, Dict. Econ. Prod.
Mal. Penin. 1, 1935, 370).

Vern. Malaya: abélor, chérék jantan, émbalau,
é. bétina, &. padang, hebélur, hémpédu béruang,
kKusum (from the Chinese), lada barau, (mé)lada
pait, malau, sarai pusur, sérajat, sisek manek,
suntang hutan; Sumatra: dadih-dadih, Karo,
tambar-si-pogu, Toba Mal., malur, sikalur, tambar
bui, tambar sipago, Batak, bérul, Lamp.; Java:
kéndung peutjang, ki padésa, kuwalot, trawalot,
walot, S, kwalot, tambara maritja, J, morindja,
M; Philippines: balaniog, Chab., bogo-bogo, P.
Bis., C. Bis., magka payos, S. L. Bis., manongao-
bobi, C. Bis., selte, Yakan.

2. Brucea mollis WALL. [Cat. (1848) 8483] ex
Kurz, J. As. Soc. Beng. 42, ii (1873) 64; BENN.
Fl. Br. Ind. 1 (1875) 521; Merr. & ROLFE,
Philip. J. Sc. 3 (1908) Bot. 104; Lecomte, FI. Gén.
I.-C. 1 (1911) 698; Merr. En Philip. 2 (1923) 347;
Crais, Fl. Siam. En. 1 (1926) 241; MeRR. &
CuuN, Sunyatsenia 5 (1940) 89.—B. luzoniensis
VIDAL, Sinops. Atlas (1883) 19 t. 26, f. B; MERR.
Publ. Goy. Lab. Philip. no. 35 (1906) 26; Philip.
J. Sc. 1 (1906) Suppl. 70.—B. membranacea MERR.
Philip. J. Sc. 1 (1906) Suppl. 70.—B. macrobotrys
Merr. Philip. J. Sc. 10 (1915) Bot. 19; En.
Philip. 2 (1923) 347.—B. stenophylla MERR.
Philip. J. Sc. 12 (1917) Bot. 274.—B. acuminata
Lt, J. Arn. Arb. 24 (1943) 445, ex descr.—Fig.12e-f.

Shrub or small tree, 1-8 m, stem 3-10 cm o.
Leaves 20-60 cm long; leaflets 3—9(—13), ovate-
oblong to ovate-lanceolate or lanceolate, entire,
toothed, undulate, bluntly serrate, or crenate,
glabrous or pubescent, 5-16 by 1-8 cm; petiole
2-13 cm, lateral petiolules 2-10 mm, terminal one
3-40 mm. Inflorescence and flowers as in B.
javanica, but the pedicels of the 2 flowers up to
6 mm long. Flowers white, creamy, green or red.
Mature dried drupes 1—2(—3) together, 9-13 mm
long.

Fig. 14. Distribution of Brucea mollis WALL. ex
Kurz (delineated, dots) and Eurycoma apiculata
BENN. (dotted delineation, triangles).

Distr. From the East Himalayas, Burma, Siam,
Laos, Cambodia, and Hainan, to Malaysia:
throughout the Philippines. Fig. 14.

Ecol. Always in open forests, often in damp
places, usually on slopes and ridges, 0-1800 m.
Fl. Jan.—Aug.

Vern. Philippines: makamara, Mag., suga, Ig.

Notes. The form with narrow leaflets,
distinguished by MERRILL as B. stenophylla, occurs
only above 1500 m. The form with large leaflets,
distinguished by him as B. macrobotrys, occurs
only at a low altitude. In my opinion these varia-
tions are only due to the influence of the
altitude. As I saw a complete series of transitional
forms I reduced both to B. mollis. B. mollis is
more variable than B. javanica.

Excluded

Brucea quercifolia SEEM. Fl. Vit. (1865) 33, based
on SEEMANN 105, coll. a. 1860 (K), is according to
A. C. SmitH & W. L. STERN, Brittonia 14 (1962)
237-241, 16 fig. = Dysoxylum quercifolium (SEEM.)
A. C. SMITH (Meliaceae).

6. PICRASMA

BLumE, Bijdr. 5 (1825) 247.—Nima Ham. ex A. Juss. Mém. Mus. Hist. Nat.
Paris 12 (1825) 516.—Aeschrion VELLosO, Fl. Flum. (1825) 58, Ic. (1835) t. 152.—
Picraena LINDL. Fl. Med. (1838) 208.—Muenteria WALP. Rep. 5 (1846) 398.—
Triscaphis GAGNEP. Not. Syst. 13 (1948) 190; cf. Fl. Mal. I, 6 (1960) 49.—Fig. 15.

Monoecious or dioecious trees or shrubs. Branches pithy, almost glabrous.
Leaves imparipinnate; base of the petiole and usually the rachis nodes swollen,
++ shrunken when dry; leaflets opposite or subopposite. Stipules early caducous,
triangular, ovate or orbicular, sometimes seemingly absent, but present on inno-
vations. Inflorescences axillary, longish peduncled, compound-cymose, unisexual.
Bracts small, whether or not early caducous. Pedicels articulated in the lower half.
Flowers 4—5-merous, unisexual or functionally 9, the 2 ones usually twice as large
as the 3. Sepals small, free to united halfway up, persistent. Petals valvate or sub-
valvate in bud, the mucronate tips incurved, persistent in 9 flowers, much longer
than the sepals and sometimes accrescent. Stamens 4-5; anthers latrorse, emargi-

Dec. 1962] SIMAROUBACEAE (Nooteboom) 213

ele

Fig. 15. Picrasma javanica BL. a. Flowering twig, * 74, 6. fullgrown stipules, < 74, c. g flower, x 6
d. 2 flower, petals and sepals removed, » 6, e. fruits, nat. size (a, d DE VRIESE & TEYSMANN S. 11., 6 NGF
10115, ¢ JUNGHUHN s.n., e KOSTERMANS 431).

ze

FLORA MALESIANA

[ser. I, vol. 6?

nate at the top, split from the base to halfway up, basidorsifixed, versatile; filament
inserted at the narrowed base of the disk (torus). Disk rather thick, sometimes
accrescent in fruit. Carpels up to 7, free, vestigial or absent in ¢ flowers; styles
connate except at the base, sometimes | or 2 free; stigmas free, rather long, fili-
form. Oyule 1, basal. Fruits 1-4, drupaceous; exocarp thin, fleshy, wrinkled when
dry, endocarp hard. Seed with a broad hilum; placenta adaxial, basal; testa rather

thick and hard; no endosperm.

Distr. About 6 spp. in America (Mexico, West Indies, Venezuela, Brazil, Argentina), 2 spp. in Asia
(Korea, Japan, S. China, and SE. Asia), one of which through Malaysia to the Solomon Is.
Note. Triscaphis GAGN., described in Staphyleaceae, is reduced here at the instigation of Mr Airy SHAW.

1. Picrasma javanica Briume, Bijdr. 5 (1825)
248: BENN. Pl. Jav. Rar. (1844) 197, t. 41; Mia.
Fl. Ind. Bat. 1, 2 (1859) 679; BENN. in FI. Br.
Ind. 1 (1875) 520; Kurz, For. Fl. Burma 1
(1877) 201; Kina, J. As. Soc. Beng. 62, ii (1893)
227; K. & V. Bijdr. 4 (1896) 8; Merr. Bull. Bur.
For. Philip. 1 (1903) 27; Back. Schoolfl. Java
(1911) 192; Lecomte, FI. Gén. I.-C. 1 (1911) 699;
Koorp. Atlas 2 (1914) t. 319; Laut. Bot. Jahrb.
56 (1920) 344; Rip. Fl. Mal. Pen. 1 (1922) 361;
Merr. En. Philip. 2 (1923) 347; Cras, Fl. Siam.
En. 1 (1926) 240; GAGnep. FI. Gén. I.-C. Suppl.
1 (1946) 667; Back. Bekn. FI. Java (em. ed.) 6
(1948) fam. 146, p. 6.—Brucea dubia STEvuD.
Nomencl. 1 (1841) 230, nomen.—P. nepalensis
BENN. Pl. Jav. Rar. (1844) 201; in Fl. Br. Ind. 1
(1875) 520.—P. andamanica KuRz ex BENN. in
Fl. Br. Ind. 1 (1875) 520.—P. philippinensis ELM.
Leafl. Philip. Bot. 5 (1913) 1837.—Triscaphis kerrit
GAGN. Not. Syst. 13 (1948) 190; Fl. Gén. I.-C.
Suppl. 1 (1950) 999, f. 128 3-8.—Fig. 15.

Tree, up to 20 m, bole up to 15 m and 35cm 9.
Bark grey to brown, smooth, fissured. Leaves
2-3(—4)-jugate; petiole 2-6 cm, as the rachis terete,
sparsely puberulous to glabrous; petiolules 0-7
mm; leaflets entire, sometimes with a waved or
wrinkled margin, usually rather abruptly blunt-
acuminate, cuneate at the base, 4-20 by I-10 cm;
nerves 3-8 pairs; on upper surface midrib crested,
nerves narrowly sulcate, venation indistinct; under-
neath nerves and reticulate venation prominent.
Stipules foliaceous, flabellately veined, nearly
orbicular with rounded apex and acute base,
7-25 by 5-20 mm, usually early caducous, leaving
a rather large scar. Inflorescences up to 20 cm
long, planoconvex. Bracts obovate, rounded, very
early caducous. Flowers 4-merous, white to yellow
or green. Pedicels c. 10 mm in 9 and up to 7 mm
in 3 flowers. Sepals glabrous to puberulous,
triangular to ovate, acutish, c. 1 mm. Petals ovate-
oblong or oblong, often acute-acuminate to
mucronate, glabrous or sparsely hairy, with a
conspicuous midrib, in 3 flowers 2-5 by 1-2 mm;
in 2 flowers 3-7 by 3-5 mm, accrescent to 10-
15(-20) by c. 7(-10) mm. Disk hairy, 4-lobed,
14-1 mm high. Stamens usually longer than petals
in 3 flowers, shorter than petals in 2 flowers;
filaments gradually thinner towards the top, hairy
at the base, 14-2 mm in 9 and 1-5 mm in ¢ flowers;
anthers 1-2 by 34-114 mm in ¢ and up to | by
14 mm and barren in 2 flowers. Carpe/s up to 4,
glabrous or puberulous; styles 1-1! mm, stigmas

c. 2 mm. Fruits 1-4, green to red or blue, ovoid

to depressed-globose, c. 9-10 by 7-12 mm.
Distr. Tropical SE. Asia (from Sikkim, Assam,

Burma, and Tonkin southward), throughout

Malaysia to the Solomon Is. Fig. 16.

Fig. 16. Distribution of Picrasma javanica BL.

Ecol. Usually rather scarce, scattered in rain
forests from sea-level up to 1500 m. Fi. fr. Jan.—
Dec.

Uses. The bark contains quassiin, which gives
it a bitterness and causes it to be used in Burma
and Java in lieu of quinine, though there is no
alkaloid in it. In Java the leaves may be applied to
sores. The trunk is too small for timber and the
wood is not durable (HEYNE, Nutt. Pl. 1927,
872; Burk. Dict. 1935, 1723).

Vern. Philip.: nalis, Sul., palumpang, Bag.;
Celebes: tambara tédong, Bonthain; Sumatra:
tuba lalat, Karo, t. ulét, Palemb., émpédu kaju;
Java: ki bagara, k. brahma, k. pahit, k. tjadan,
k. tjanting, k. tjitan, S, pati laler, J; kaju chutu,
Minah.; New Guinea: snippa, Numfoor, annamur,
Biak.

Note. The second Asiatic species, P. quassioides
(D. Don) BENN., occurs from the Himalayas to
Japan and Korea; it is medicinal and is sometimes
cultivated in Europe for ornamental purpose-
It is easily distinguished by its crenate or serrate
leaflets, 5-merous flowers and smaller fruit (c.
5 mm).

Excluded

Picrasma denhamii Stem. Fl. Vit. (1865) 33 from
Aneityum, New Hebrides, is according to Mr J. E.
Danby (in litt.) = Evodia triphylla DC. Prod.
1 (1824) 724 (Rutaceae).

Dec. 1962] : SIMAROUBACEAE (Nooteboom) 215

7. AILANTHUS

Desr. Mém. Phys. Math. Ac. R. Sc. Paris (1786) 270 t. 8, nom. gen. cons.; DC.
Prod. 2 (1825) 88 (Ailantus); PIERRE, Fl. For. Coch. 4 (1892) t. 294-295; ENGLER
in E. & P. Nat. Pfl. Fam. 3, 4 (1896) 223; ed. 2, 19a (1931) 390; VAN TIEGHEM,
Ann. Sc. Nat. IX, 4 (1906) 272.—Pongelion ADANS. Fam. Pl. 2 (1763) 319;
VAN TIEGHEM, Ann. Sc. Nat. IX, 4 (1906) 272.—Albonia BuUCHoz, Herb. Color.
Am. (1783) t. 57, sine descr—Hebonga RADLK. Philip. J. Sc. 6 (1911) Bot. 365.
Fig. 7f-g, 17, 18.

Tall, fast-growing, sometimes deciduous, dioecious trees. Branches thick,
pithy; leaf-scars large. Leaves large, imparipinnate or paripinnate (with a pro-
longed rachis), sometimes on a single tree, more or less tufted at the ends of the
twigs, multijugate, base of petiole often shrunken when dry; leaflets opposite
or subopposite, oblique, usually acuminate, (in Mal.) entire, rachis and petiole
terete, pithy, nearly always with some glands on the undersurface, usually near
the base; midrib and lateral nerves prominent on the undersurface; petiole, rachis,
petiolules, and branchlets of the panicles usually striate when dry. Flowers in
axillary panicles, 5(—6)-merous. Calyx small, 5(—6)-lobed or closed in bud and
later irregularly dehiscing (often 2-lobed) to the base, rarely cupular. Petals 5(-6),
induplicate-valvate in bud, concave, oblong to narrowly oblong, still enlarging
during anthesis. Stamens 10, in 3 flowers inserted below the outer margin of the
disk, in Q flowers either of subnormal size (but without pollen) or vestigial or
absent; anthers oblong to broadly oblong, latrorse to extrorse, filaments dorsally
attached halfway, the 2 cells free in their lower half. Carpels 2-5, free, flat, in the
3 flowers vestigial or absent; styles 2-5, free or connate, inserted above the middle
on the adaxial side; ovule 1, epitropous, anatropous, adaxially attached in the
middle. Fruit a linear or oblong-lanceolate samara. Seed flat, orbicular or obovate
or somewhat triangular, exalbuminous; testa thin; cotyledons 2, flat, planoconvex,
radicle pointing upwards.

Distr. Five spp. in tropical and subtropical SE. Asia from Turkestan and India to China through
Malaysia to the Solomon Islands, Queensland, and northern New South Wales; in Malaysia 2 species,
not yet found in the Malay Peninsula.The seed of the Chinese A. a/tissima (MILL.) SWINGLE was received
in England in 1751. The species was soon cultivated in Europe and N. America where it became widely
naturalized and now occurs as a common weed on the outskirts of large cities; also naturalized in Australia.

According to the fossil record (fruits!) Ai/anthus occurred during the Tertiary in Europe and North
America.

Ecol. In Malaysia both in the rain-forest and in monsoon forest, below 1000 m, on the whole rather
uncommon and never gregarious, in valleys, along streams, and in open places.

In literature it has often been suggested that the flowers might also be bisexual (e.g. by ENGLER, 1931)
or the trees monoecious (ROXBURGH). I have, however, never observed bisexual flowers. The trees PRAIN
observed (Ind. For. 28, 1902, 131-134, 210-211, t. 1-3; Contr. Ind. Bot. 1906, 1-6, t. 1-3) were all
dioecious, as is the commonly cultivated A. a/tissima (MILL.) SwINGLeE. The stamens which are often
present in 2 flowers do not contain pollen.

During anthesis the 3 flowers emit a fetid, disgusting smell and for that reason ¢ trees are not desirable
to cultivate as an ornamental.

The rarity of the species is remarkable as trees produce an immense number of membranous sama-
ras fit for dispersal by wind. And if the species might be shade-intolerant one would expect them to settle
as a nomad tree in glades, clearings, and open secondary forests.

Uses. See under the species.

Taxon. Prerre, /.c., subdivided the genus (Pongelion) into two sections, Eupongelion RUMPH. ex
Pierre and Ail/anthus Pierre, according to the occurrence of 1-3 or 5 styles respectively, but he placed
P. moluccanum DC. erroneously in the first section. ENGLER, 1896, /.c., accepted the sectional names of
Pierre, changing Ailanthus into Euailanthus, but distinguished them on another character, viz Euponge-

lion with free styles and Euailanthus with connate styles, thereby redistributing the species in a different
way. No use was ever made of the structure of the calyx which seems to me a third character worthy of

216 FLORA MALESIANA [ser. I, vol. 6?

consideration. Among the five species which I admit under the genus, one, A. a/tissima, has free styles,
one, A. triphysa, has (2—) 3(-4) styles, and one, A. integrifolia, has a calyx closed in bud rupturing irreg-
ularly during anthesis. As these three characters are obviously not correlated, it appears that they are
of specific value and cannot serve for subdividing the genus. This was also the opinion of VAN TIEGHEM,
1.c. 272-280, who, in an elaborate anatomical study of the leaves proposed to distinguish two genera
based on vegetative characters, viz. Pongelion with entire leaflets without glands and Ailanthus with den-
tate leaflets with under each tooth a gland. However, in the species with entire leaves also glands occur,
either at the base or scattered, or in the forks of the lateral nerves near the margin. As the situation of the
glands varies with the species, I can only conclude to the specific value of this character.

Ricketr & STAFLEU (Taxon 8, 1959, 302) erroneously assumed that Pongelion ADANS. is a Leguminosa.

Notes. The name of the genus is derived from RuMmpuHius’s Amboinese vernacular name aylanto,
meaning tree of heaven, alluding to the lofty size of the Moluccan species.

Due to inadequacy of the material and the rarity of the species, specific distinction has been in great
confusion. I felt necessitated to extend my study to all species described and could examine the types of
many specific names. I have come to the conclusion that only 5 species can be distinguished; it is remar-
kable that p—E CANDOLLE had four of them. Although only two of them have as yet been found in Ma-
laysia, itis of interest to give a key to and synonymy of all of them. Though variable they can easily be
distinguished in flower, fruit, and vegetatively.

In the herbarium flowers and fruit are never present on one sheet; field collectors are therefore requested
to mark flowering trees to collect fruit later.

KEY TO THE SPECIES
based on flowering material

1. Leaflets entire.

2. Petals puberulous. 2 Flowers with 5 carpels. Stigmas long, stellately spreading. Leaflets ovate to
elliptic-oblong, rarely ovate-lanceolate, usually with a few a ee large, black glands on the under-
surface mostly near the base Bes é 1. A. integrifolia
. Petals glabrous or nearly so. Leaflets without large glands 1 near the base, ‘but some small glands may
occur over the surface, either scattered or in vein-forks.

3. Calyx lobes as long as the tube or longer. 2 Flowers with 3-4 glabrous carpels. Stigmas connate,
peltate, slightly 3-4-lobed. Leaflets ovate to oblong-lanceolate, rarely ovate to elliptic-oblong. In
the forks of the dichotomous-branching nerves usually a more or less hairy, often pitted gland,
sometimes obscured by a hair tuft. 4 . 2, A. triphysa

3. Calyx lobes shorter than the tube. 2 Flowers with ‘5 ‘puberulous carpels. ‘Stigmas free, stellately
spreading. Leaflets as in A. integrifolia but not with large black basal glands; a few small flat glands
scattered on the undersurface. (Hongkong.) ..............2.4. 5. A. ford

1. Leaflets toothed.

4. Leaflets coarsely toothed or lobed. Petiolules long, c. 2-4 cm. Petals glabrous or nearly so. Filaments
as long as the anthers or shorter. Carpels sparsely longish hairy or pubescent; styles attached near
the top, very short, free or connate, with long, outwards curling stigmas. (India and Ceylon.)

3. A. excelsa

4. Leafiets with a few short, rarely rather large teeth near the base. Petiolules short, 14(—1) cm. Petals
densely woolly hairy on the lower half of the margins and on the inner surface. Filaments twice as
long as the anthers or longer. Carpels glabrous; styles attached nearly halfway, connate, c. 144 mm
long, with a peltate, 5-lobed stigma c. 1 mm above the carpels. (China.) .. . 4. A. altissima

KEY TO THE SPECIES

based on fruit characters

1. Leaflets entire.
2. Main vascular bundle towards seed in the margin of the samara... .... 1. A. integrifolia
es Main vascular bundle towards seed intramarginal.
. Samaras 414-714 cm long. Scar of style at the same level as the middle of the seed.
2. A. triphysa
3. Samaras 3—5 cm. Scar of style at the same level as the apex of the seed. . . . . . 5. A. fordii
1. Leaflets toothed.
4. Scar of style nearly at the same level as the apex of the seed. Main vascular bundle towards seed
intramarginal. . . . . . 3. A. excelsa
4. Scar of style nearly at the s same level. as the middle of the seed. Main vascular bundle towards seed in
the margin of the samara Wehs Baotwh oat Sd aR ee ee 4. A. altissima

Dec. 1962] SIMAROUBACEAE (Nooteboom) ANG

ley a® te

Fig. 17. Leaflets of Ailanthus spp. a. A. integrifolia Fig. 18. Fruits of Ailanthus spp. a. A. integrifolia

Lamk, b. A. triphysa (DENNST.) ALSTON, c. ditto, LAMK, b. A. fordii NOooTEBOOM, c. A. triphysa

glandular domatium, d. A. excelsa Roxs., e. A. (DENNST.) ALSTON, d. A. excelsa ROxB., e. A.

altissima (MILL.) SWINGLE, all * 24 (a WaTER- — altissima (MILL.) SWINGLE, all * 74 (a WATER-

HOUSE 664, b-c VILLAMIL 20972, d DRUMMOND HOUSE 664, b. HANCE 1497, c CURRAN 3847, d
21597). ; / DRUMMOND 21597).

218

FLORA MALESIANA

[ser. I, vol. 6?

1. Ailanthus integrifolia LAMK, Dict. 3, 2 (1792)
417, based on Arbor coeli sive Caju langit, RUMPH.
Herb. Amb. 3 (1743) 205, t. 132; Merr. Int.
Rumph. (1917) 299.—A. pongelion GMEL. Syst.
Veg. 1 (1791) 726, nom. illeg., pro parte, tab.
Rumph.; Bico, Fl. Filip. (1837) 380; ed. 2 (1845)
268; ed. 3, 2 (1878) 134, excl. syn. malab.—A.
moluccana DC. Prod. 2 (1825) 89, nom. illeg.;
Mig. Fl. Ind. Bat. 1, 2 (1859) 679; Koorpb.
Minah. (1898) 374, incl. var. mollis (K. & V.)
Koorp.; VAL. Ic. Bog. 1 (1901) t. 82; BAcK.
Schoolfl. Java (1911) 191; Koorp. Atlas 2 (1914)
t. 321, I-K.—Dysoxylum dasyphyllum Mia.
Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 19, pro
sched. DE VR. & T., cf. Koorp. Minah. (1898)
374.—A. malabarica (non DC.) F.-VILL. Nov.
App. (1883) 349, non vidi; K. & V. Bijdr. 4
(1896) 3, incl. var. mollis K. & VY. pro parte.
—A. calycina Pierre, Fl. For. Coch. 4 (1892)
t. 294 A (plate); Lecomte, Fl. Gén. I.-C. 1 (1911)
696; Koorp. Exk. Fl. Java 2 (1912) 432; Atlas
2 (1914) t. 321 A-H; Koorp.-ScHum. Syst.
Verz. 1 § 1 fam. 138 (1913) 18; BAKn. f. in Back.
Bekn. FI. Java (em. ed.) 6 (1948) fam. 146, p. 7.—
Pongelion calycinum Pierre, Fl. For. Coch. 4
(1892) t. 294 A (text)—A. grandis PRAtn, Ind.
For. 28 (1902) 131, t. 1, 210, repr. Contr. Ind.
Bot. (1906) 1, 5.—Pongelion grandis VAN TIEGH.
Ann. Sc. Nat. IX, 4 (1906) 278.—A. blancoi
Merk. Sp. Blanc. (1918) 205; En. Philip. 2 (1923)
348.—A. peekelii MeLcu. Notizbl. Berl.-Dahl. 10
(1930) 893; C. T. Wuite, J. Arn. Arb. 31 (1950)
91, incl. var. glabrata.—Fig. 7f-g, 17a, 18a.

Tree, up to 60 m high, bole occupying c. 34
of the length, up to 75(-175) cm g; bark smooth,
light brown or grey; crown dense. Leaves 2—9—
jugate (apical leaflet mostly vestigial), c. 30-200
cm long; leaflets very oblique, usually falcate and
acuminate with an obtuse tip, glabrous above,
sometimes more or less pubescent beneath, espe-
cially on the midrib and the 6-13 pairs of nerves,
10-40 by 4-15 cm; usually a few black, flat, or-
bicular or oblong glands c. 1,-5 mm o mostly
near the base of the undersurface sometimes
seemingly separated from the parenchyma as a
loose membrane; petiole glabrous or puberulous,
5-20 cm; petiolules glabrous or puberulous, 4-114
cm. Panicles loose, up to c.40 cm or more, glabrous
or pubescent. Bracts small, triangular, very early
caducous. Pedicels up to c. 15 mm in anthesis.
Calyx more or less pubescent, closed in bud,
rupturing and toothed irregularly, rarely cupular,
1-4 mm high, rarely caducous. Petals puberulous,
acute or bluntish, up to c. 9 by 3 mm. Filaments
with many long spreading hairs to glabrous, usu-
ally thickened downwards, c. 4 mm in 9 to 4mm
long in 3; anthers c. 1 mm in @ to 214 mm long
in ¢g. Carpels 5, usually densely puberulous;
styles 5, connate at the base, including the long,
stellately spreading stigmas, up to c. 6 mm long.
Samaras (1—)3-5, with obtuse apex, more or less
prominently reticulate or lengthwise striate, c.
11-22 by 2'4-5 cm; main vascular bundle to the
seed in the adaxial margin; scar of the style beneath
the seed; pedicel 214-5 cm.

Note. Of A. integrifolia LAMK two replacing
races can be distinguished which differ in the size
of the flowers, but unfortunately cannot be
discriminated in sterile state or in fruit. I have
mapped the localities of flowering specimens
through which it appeared that the small-
flowered race is apparently restricted to seasonal
regions in SE. Asia and Central to East Java,
whereas the large-flowered form occurs only in the
everwet forest. The type of A. integrifolia LAMK,
obviously belongs to the large-flowered race.

ssp. integrifolia.—A. integrifolia LAMK.—A. pon-
gelion GMEL.—A. moluccana DC.—A. moluccana
var. mollis (non K. & YV.) Koorpb. 1898.—
Dysoxylum dasyphyllum Miq.—A. blancoi MERR.
—A. peekelii MELCH., incl. var. glabrata.

in F = : 73 as

Lu. = \e

Fig. 19. Distribution of Ailanthus integrifolia

LAMK ssp. integrifolia (delineation with continuous

line) and ssp. calycina (PIERRE) NOOTEBOOM
(delineation with dotted line).

Pedicels up to 15 mm. Calyx 2-3 mm high.
Petals 6-10 mm long, evenly hairy outside.

Distr. Melanesia (Bismarcks and Solomons),
in Malaysia: all islands, except Java and the
Lesser Sunda Islands. Fig. 19.

Ecol. In primary rain-forest, very rare, locally
rather common in North New Guinea, in New
Ireland said to be commoner than Pometia,
Alstonia, and Octomeles, both in the primary and
secondary forest, from the lowland up to 900 m.
Fl. Jan.—Dec., fr. March, April, Nov.

Uses. Made into planks for housing efc.
in New Guinea and the Bismarcks; the sapwood
is white, yellow, pale brown or creamish, and is
very soft and not durable; heartwood is absent
(Heyne, Nutt. Pl. Ned. Ind. 1927, 1499).

Vern. Philippines: balokas, Tagb., makaisa,
malaaduds, malasapsdp, Tag.; aylanto, kaju
langit, Ambon; New Guinea: won, Karoon lang.,
broes, Mooi lang., aisasa, Upper Waria, limoetiti,
Kebar lang., kokop, kun-kun, New Britain.

ssp. calycina (PIERRE) NooTEBOOM, comb. nov.—
A. malabarica var. mollis K. & V., pro parte.—
A. calycina PiERRE.—Pongelion calycinum PIERRE.
—A. grandis PRAIN. —Pongelion grandis VAN TIEGH.

Pedicels up to 5 mm. Calyx shorter than 2 mm.
Petals c. 4 mm long, the margins densely, the
outer surface laxly hairy.

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

Be

Distr. SE. Asia (Assam, Sikkim, Cochinchina),
in Malaysia: Central to East Java. Fig. 19.

Ecol. Mixed primary forest, obviously under
seasonal conditions.

Vern. Raden, tawa, J.

2. Ailanthus triphysa (DENNsT.) ALsTON, Handb.
Fl. Ceyl. VI, Suppl. (1931) 41.—A. integrifolia
var. B LaMK, Dict. 3, 2 (1792) 417, typ. rheed.—
Adenanthera triphysa DENNST. Schliiss. Hort.
Mal. (1818) 32, based on Pongelion RHEEDE,
Hort. Mal. 6 (1686) 27 t. 15.—A. malabarica DC.
Prod. 2 (1825) 89, nom. illeg.; W. & A. Prod.
(1834) 150; WiGuHT, Ic. (1850) t. 1604; Datz.
& Giss. Bomb. Fl. (1861) 46; BRANpIs, For.
FI. (1874) 58; BeNN. in Hook. f. Fl. Br. Ind. 1
(1875) 518; Pierre, Fl. For. Coch. 4 (1892) t.
294 B; TRIMEN, Handb. FI. Ceyl. 1 (1893) 230;
K. & V. Bijdr. 4 (1896) 4, incl. var. mollis K. &
V. pro parte; PRAIN, Ind. For. 28 (1902) 132,
t. 3 A, repr. Contr. Ind. Bot. (1906) 132; LECOMTE,
Fl. Gén. I.-C. 1 (1911) 692; Back. Schoolfl.
Java (1911) 191; Francis, Austr. Rain-For. Trees
(1929) 174, t. 110, 111; ed. 2 (1951) 196, t. 112,
113; Koorp. Atlas 2 (1914) t. 320; Baku. f. in
Back. Bekn. FI. Java (em. ed.) 6 (1948) fam. 146,
p. 6.—A. imberbiflora F. v. M. Fragm. 3 (1862)
42; BentH. FI. Austr. 1 (1863) 373; BAILEY,
Queensl. Fl. 1 (1899) 217; Francis, Austr. Rain-
For. Trees (1929) 174, t. 110, 111; ed. 2 (1951)
196, t. 112, 113.—A. fauveliana Pierre, Fl. For.
Coch. 4 (1892) t. 295 B (plate); Lecomte, FI.
Gén. I.-C. 1 (1911) 692; Crais, Fl. Siam. En. 1
(1926) 240.—Pongelion fauvelianum Pierre, FI.
For. Coch. 4 (1892) t. 295 B (text).—Pongelion
malabaricum PtERRE, /.c. t. 294.—Pongelion im-
berbiflora PieERRE, /.c. t. 294.—A. kurzii PRAIN,
Ind. For. 28 (1902) 133, t. 3 B, repr. Contr.
Ind. Bot. (1906) 3.—A. philippinensis MERR.
Publ. Goy. Lab. Philip. no 35 (1906) 25; Philip.
J. Sc. 1 (1906) Suppl. 70; ibid. 2 (1907) Bot. 431.—
Hebonga obliqua RaAvLK. Philip. J. Sc. 6 (1911)
Bot. 366; Boas, Beitr. Anat. Syst. Simar. (1912)
49; repr. in Fedde, Rep. 13 (1914) 290; Merr.
En. 2 (1923) 348.—Hebonga mollis RADLK. l.c.
367; Boas, /.c.; MeRR. /.c.—Hebonga siamensis
RADLK. ex CrRAis, Kew Bull. (1912) 264; Fl. Siam.
En. 1 (1926) 243; GAGnepP. Fl. Gén. I.-C. Suppl. 1
(1946) 668.—Fig. 17b—c, 18c.

Tree, up to 45 m high and 34(-114) m o.
Leaves 6-17(—30)-jugate, 20-70 cm long; leaflets
glabrous above, more or less pubescent beneath,
obliquely ovate-lanceolate to oblong-lanceolate,
rarely ovate or oblong, faicate, acuminate, (5—)
9-15(-26) by 214-5'%4 cm; nerves 8-20 pairs;
usually at the end of the midrib beneath a hairy
gland, making the apex obtuse or even emarginate,
apex rarely acute; on the undersurface in the forks
of the usually dichotomous-branching nerves
usually a circumvallate, more or less hairy, often
pitted gland, sometimes obscured by a hair tuft.
Panicles dense and many-flowered, more or less
pubescent, c. 20-60 cm long. Bracts small, ovate
to triangular, caducous. Pedicels up to c. 4 mm.
Calyx pubescent, less than 1 mm high, the trian-

gular, acute lobes as long as the tube or a little
longer. Petals glabrous or nearly so, c. 3-5 by
1-114 mm. Filaments tortuous-folded in bud,
filiform or sometimes attenuating from the base
to the top, usually with spreading hairs beneath,
c. 1-3 mm long in 2 and c. 3-6 mm long in 3
flowers; anthers c. 1,2 by 1 mm in ¢ flowers,
smaller in 2 flowers. Carpels (2—)3(-4), glabrous,
c. 2-244 by 1-114 mm; styles free or coherent at
the base, connate at the top, c. 1-114 mm; stigma
(2-)3(-4)-lobed, peltate, c. 2 mm o. Samaras
1—3(—4), obtuse at the apex, main vascular bundle
towards the seed intramarginal, scar of the style at
the same level as the seed, c. 414-8 by 114-214
cm; pedicels 8-20 mm.

Distr. SE. Asia: India (Concan, Malabar,
Canara), Ceylon, Burma, Siam, Vietnam, through
Malaysia (except the Malay Peninsula, Sumatra,
the Lesser Sunda Is, and New Guinea) to Queens-
land and the north of New South Wales. Fig. 20.

Fig. 20. Distribution of Ailanthus triphysa
(DENNsT.) ALSTON.

Ecol. In forests, usually very rare, under both
everwet and seasonal conditions, according to
BURKILL intolerant of shade, from the lowland
up to + 600 m. FI. Jan.—Dec.

Uses. The resin, obtained by making incisions
in the bark, is used as incense and medicinally in
India. In Indo-China the bark is burned as in-
cense. The bark and the leaves are in great repute
as a tonic, especially in debility after childbirth.
They also possess febrifuge properties and are
useful in dyspeptic complaints. The wood is used
for making wooden shoes in Luzon; in India for
fishing floats, catamarans, sword-handles, and
spear-sheaths, and in Ceylon for teaboxes (BUR-
KEE. Dict; Hex Prod, Male Pens 15.1985; 979):

Vern. Ki pahit, S, sélangké, Central J; Philip.:
empau, Tagb., kalauag, Bik., hebong, makaisa,
malaaduds, Tag.

3. Ailanthus excelsa Roxs. Pl. Corom. | (1795) 24,
t. 23; DC. Prod. 2 (1825) 89; Roxs. Fl. Ind. ed.
Carey 2 (1832) 450; W. & A. Prod. (1834) 150;
Grau. Cat. Bomb. Pl. (1839) 37; WiGur, I11.
Ind. Bot. 1 (1840) 170 t. 67; BENN. in Hook. f.
Fl. Br. Ind. 1 (1875) 518; Pierre, Fl. For. Coch.
4 (1892) t. 295 A; Pratn, Ind. For. 28 (1902)
t. 2 B; vAN TieGH. Ann. Sc. Nat. IX, 4 (1906)
277.—Pongelion excelsum Pierre, Fl. For. Coch.
4 (1892) t. 295 A.—Pongelion wighti VAN TIEGH.

220

FLORA MALESIANA

[ser. I, vol. 6?

Ann. Sc. Nat. IX, 4 (1906) 277.—Fig. 17d, 18d.

Distr. South, Central, and NW. India (in the
last probably introduced) and Ceylon.

Note. VAN TIEGHEM, /.c., rightly observed that
there is a discrepancy between the figures given
by RoxpurGH and by WiGuT, the latter being in
entire accordance with the material, that of
ROXBURGH deviating by very short petiolules and
very long filaments. As there is only one species
on the Coromandel Coast with dentate leaflets I
assume that ROxBURGH’s plate is inaccurate in
detail. Therefore I have reduced Pongelion wighti
TiEGH., based on WIGHT’s plate, to the synonymy.
Dr. SANTAPAU (in /itt.), after having examined the
specimens in the Blatter Herbarium, agreed with
this interpretation.

4. Ailanthus altissima (MILL.) SWINGLE, J. Wash.
Ac. Sc. 6 (1916) 495; REHDER & WILSON, J.
Arn. Arb. 9 (1928) 86; CroNnguistT, Brittonia 5
(1944) 146; E. ANDERSON, Bull. Mo. Bot. Gard.
49 (1961) 105-107, 2 fig.—Toxicodendron al-
tissimum MILL. Gard. Dict. ed. 8 (1768) n. 10.—
Rhus cacodendron Exru. Hann. Mag. (1783)
227, non vidi—Albonia peregrina BucHOoz, Herb.
Color. Am. (1783) t. 57.—A. glandulosa DesrF.
Mém. Math. Phys. Ac. R. Sc. Paris (1786) 265,
t. 8; Arron, Hort. Kew. 3 (1789) 443; DC. Prod.
2 (1825) 89; W. & A. Prod. (1834) 150; Mie.
Fl. Ind. Bat. 1, 2 (1859) 678; Kocu, Dendrol. 1
(1869) 569; BAtLEy, Queensl. Fl. 1 (1899) 218;
Prain, Ind. For. 28 (1902) t. 2 A.—A. cacoden-
dron WV HeEriv. Stirp. Nov. 6 (1790) 179; ScHINz
& THELL. Mém. Soc. Nat. Sc. Cherbourg IV, 38
(1911-12) 679.—A. pongelion GMEL. Syst. Veg. 1
(1791) 726, nom. illeg., pro parte, cit. tab. Desfont.
—A. procera SAuIisB. Prod. (1796) 171, ex Ind.
Kew.—A. rhodoptera F. vy. M. Fragm. 3 (1862)
43, ex descr.—A. erythrocarpa CARR. Rey. Hortic.
(1867) 419.—Pongelium glandulosum PIERRE, FI.
For. Coch. 4 (1892) t. 294.—A. macrophylla ex
Handl. Trees Kew pt 1 (Polypet.) (1894) 53, cult.,
ex Ind. Kew.—A. mascula l.c.—A. rubra l.c.—
A. vilmoriniana Dopg, Rey. Hortic. (1904) 444.—
Pongelium vilmorinianum VAN TigGH. Ann. Sc. Nat.
IX, 4 (1906) 278.—A. giraldii Dope, Bull. Soc.
Dendr. Fr. (1907) 191.—A. sutchuensis Dove,
1.c.—Pongelion cacodendron FARWELL, Am. Midl.
Nat. (1930) 67.—A. peregrina F. A. BARKLEY,
Ann. Mo. Bot. Gard. 24 (1937) 264.—Fig. 17e, 18e.
Distr. Native in China, cultivated in nearly all
countries of the world with a temperate or sub-
tropical climate, and often naturalized.
Note.The type specimen(from a cultivated tree)
of A. vilmoriniana Dove agrees in all respects save
that the branches and leaf rachis bear small spines
which are further unknown in the genus.

5. Ailanthus fordii NooTEBOOM, nov. sp.

Arbor parva, teste collectore palmae habitu
conspicua. Foliola 6-13-juga, integra, subtus
glandulis parvis applanatis sparse munita. Pedicelli
1-2 mm. Flores paniculati, ultimi 1-3 aggregati.

Calyx cupulatus c. Yz mm longus, breviter obtuse
5-lobatus. Petala 5, glabra, oblonga, 2-3 mm longa,
c. 1 mm lata. Stamina glabra, in floribus 3 in
alabastro plicata c. 3-5 mm longa, in floribus 2
c. 1-3 mm longa. Discus superne 5—lobatus, 4-1
mm altus. Carpella 5, dense puberula, stylis con-
natis Y.—1 mm longis terminata. Stigmata 5, libera,
basi excepta excurvata, 1-114 mm longa. Samara
3-5 cm longa, 1-134 cm lata; fasciculus vasorum
seminem versus intramarginalis; styli cicatrix
seminis apici opposita——Typus: CH. ForpD s.n.
(K; isotype BM), Hongkong, Cape Aquilar.—
Fig. 18b.

Small but conspicuous tree, sec. coll. ‘the bare
trunk surmounted with foliage like a palm’.
Leaves c. 40 cm long; leaflets 6-13 pairs, opposite
to subopposite, entire, glabrous or nearly so,
with few, small, flat glands scattered on the un-
dersurface; petiole 7-13 cm; rachis puberulous to
glabrous. Panicle large, dense-flowered, pyramidal,
20-40 cm long and c. 20 cm wide at the base;
branches of the first, and sometimes also of the
2nd and 3rd order with a conspicuous constricted
articulation at the base. Pedicels 1-2 mm. Bracts
small, triangular, not early caducous. Flowers
1-3 together, unisexual, the 2 ones with reduced
stamens. Calyx cupular, c. 4% mm high, with 5
short, obtuse lobes. Petals 5, glabrous, + oblong,
2-3 by c. 1 mm. Stamens glabrous, in ¢ flowers
sinuous in bud, 3—5 mm long, in 9 flowers c. 1-3
mm long; anthers c. 34 mm long, sterile in 2
flowers. Disk 5-lobed on top, 44-1 mm high.
Carpels 5, densely puberulous; styles connate,
14-1 mm high, more or less puberulous; stigmas
5, free, except at the very base, recurved-curled
outwards, 1-114 mm long. Samara 3-5 by 1-134
cm; main vessel to seed intramarginal; stylar scar
at the same level as the apex of the seed.

Distr. Hongkong, near Cape Aquilar quite
common (Forp; 1884-1886); emergent from
shrubberies along ravines.

Note. This species differs from A. triphysa in
having 5 pubescent carpels, long free stigmas, a
very short lobed cupular calyx, the stylar scar
being at the same level as the apex of the seed.
It differs from A. integrifolia in the leaflets never
having 2 large glands at the base, in the flowers
being much smaller and the petals being glabrous,
and in the main vascular bundle towards the seed
in the samara which is intramarginal.

Excluded

Ailanthus mairei GAGNEP. Not. Syst. 11 (1944)
164 from Yunnan = Toona sinensis (Juss.) ROEM.
(Meliaceae).

Ailanthus punctata F. v. M. Fragm. 3 (1862) 42,
cf. BENTH. Fl. Austr. 1 (1863) 373 = Pentaceras
australis Hoox. f. (Rutaceae).

Ailanthus scripta GAGNEP. Not. Syst. 11 (1944)
165 from Yunnan = Rhus vernicifera DC.
(Anacardiaceae).

Dec. 1962] SIMAROUBACEAE (Nooteboom)

8. SOULAMEA

LaMK, Dict. Enc. Méth. 1 (1783) 449; GuILLAUMIN, Bull. Soc. Bot. Fr. 85 (1938)
20.—Cardiocarpus REINW. Syll. Ratisb. 2 (1826) 14, 48, nom. illeg —Cardiophora
BENTH. in Hook. Lond. J. Bot. 2 (1843) 216.—Amaroria A. GRaAy in Wilkes,
U.S. Expl. Exp. | (1854) 356, t. 40.—Fig. 21.

Shrubs or small trees. Leaves simple (imparipinnate in some New Cal. ssp.),
sometimes with a few glands underneath. Flowers in axillary racemes or narrow
thyrses, 3(—4-5)-merous, bisexual or unisexual (in New Cal. and the Seychelles);
floral parts persistent. Bracts minute. Sepa/s more or less connate at the base,
slightly imbricate in bud. Peta/s not touching, longer than sepals. Stamens twice
as many as petals, in 2 distinct rows, inserted under the lower outer margin of the
disk; filaments attached adaxially, versatile; cells latrorse, diverging at both ends,
connective very short. Disk 3(—4—5)-lobed, each lobe forked. Carpels (1—)2(-3),
connate; ovules sessile, anatropous; style horizontally adnate to its carpel, ex-
cept for a short free patent tip; stigma small, rarely reniform. Fruit dry, (1—)2(-3)-
celled, indehiscent, flattened, distinctly winged, more or less emarginate, rarely
flattened, ovoid, acute. Seed attached adaxially nearly halfway down, with more

or less albumen; testa thin; cotyledons planoconvex.

Distr. One sp. endemic in the Seychelles (Mahé I.), one sp. widely distributed in Malaysia and
Polynesia, 6 spp. in New Caledonia, and one in Fiji.

Notes. The monotypic Fijian genus Amaroria can in my opinion not be upheld against Soulamea. It
differs merely by having one carpel, against 2-3 in Soulamea. The characteristic, emarginate fruit shape of
the 2-celled Soulameas cannot be expected to occur in the 1-celled Amaroria, as this shape is precisely
caused by the presence of 2 not entirely connate cells and absence of a terminal style; the difference
in fruit shape is thus a compulsory structural consequence. Geographically it fits well with the distribution
of the genus. Like the other inland species of the genus, S. soulameoides (A. GRAY) NOoTEBOoM, comb.
nov. (Amaroria soulameoides A. GRaAy, /.c.) has unisexual flowers.

The specimen I saw of S. terminalioides BAKER, from the Seychelles, is strongly suggestive of S. amara
LaMK. It differs in the leaves (rounded apex), fruit (3-winged), and unisexual flowers.

All New Caledonian species have also unisexual flowers and besides less emarginate fruits. They
differ inter se hardly in their generative parts, and are obviously very closely allied. The main differences
are in their leaf structure, varying from 1-11 leaflets, and the indument. The first character is, however,
not particularly important taxonomically as JADIN has demonstrated that simple and compound leaves
may occur on a single plant (Ann. Sc. Nat. VIII, 13, 1901, 283-285, t. 1).

Although the littoral S. amara has not been found in New Caledonia it seems that whereas the largest
(phenotypic and genetic) variability occurs in that island, and that inland, the origin of the genus and its
distribution must have taken place in Melanesia. It is not impossible that in its primitive state it was
unisexual — and in that case the Seychelles ‘species’ could be considered a marginal relic — and that it has
later become bisexual by selection on the littoral.

The ‘escape’ of a littoral species from an inland aggregate finds a marked parallel in Casuarina,
Spinifex, and some other genera.

Soulamea shows a striking resemblance in habit with Lunasia (Rutac.).

Amb. 2 (1743) 129, t. 41.—Cardiocarpus amarus
ReINW. Syll. Ratisb. 2 (1826) 14.—Cardiophora
hindsii BENTH. in Hook. Lond. J. Bot. 2 (1843)
216; Voy. Sulphur (1844) 181, t. 56, ic.—Fig.

1. Soulamea amara LaAmMk, Dict. Enc. Méth. 1
(1783) 449; BL. Bijdr. 2 (1825) 60; ENDL. Ann.
Mus. Wien | (1836) 188, t. 16; BENTH. Voy.
Sulphur (1844) 181, t. 56, textu; Mia. Fl. Ind. Bat.

1, 2 (1859) 129; Hemscey, Bot. Chall. 3 (1885) 235;
Wares. Bot. Jahrb. 13 (1891) 341; Back. Schoolfl.
Java (1911) 193; Merr. Int. Rumph. (1917)
300; Laut. Bot. Jahrb. 56 (1920) 344; HEYNE,
Nutt. Pl. (1927) 872; STEEN. Bull. Jard. Bot.
Btzg III, 12 (1932) 259, f. 13 map 3; KANEHIRA,
J. Dep. Kyushu Imp. Un. 4, 6 (1935) 343; K.
Sco. & Laut. FI. Schutzgeb. Siidsee (1901) 378
(Sulamea’); GUILLAUMIN, Ann. Mus. Col. Mars.
55/56 (1948) 28.—Rex amaroris RumpH. Herb.

21.

Shrub or small tree, up to 5(—15) m. Innovations
rusty tomentose. Branchlets 5-15 mm go, rather
abruptly narrowed at the apex, with a thick pith.
Leaves simple, crowded at the apex of the branch-
lets, leaving large scars, obovate-oblong and with
a blunt but never rounded apex which is sometimes
mucronate, cuneate at the base, hairy on midrib,
nerves and veins below, 10-35 by 4-12 cm; midrib
slightly immersed or inconspicuous above, strongly

222

FLORA MALESIANA

[ser. I, vol. 6?

Fig. 21. Soulamea amara LAMK. a. Fertile twig, X 24, b. flower, < 8, c. ditto, in section, pistil removed,
x 8, d. fruit, nat. size (a, d FosBeERG 33912, b, c FosBerG 26162).

prominent beneath; nerves straight, parallel,
ending in an intramarginal looped vein, sulcate,
slightly prominent or inconspicuous above, prom-
inent beneath; veins inconspicuous above,
finely dense-reticulate beneath; petiole pithy,
shrunken at the base when dry, sometimes also at
the apex, hairy, 3-8 cm. Racemes erect, shorter
than the leaves, 3-12 cm. Flowers bisexual,

3(—4-5)-merous, c. 2 mm long. Pedicels up to
5 mm. Sepals puberulous, erect, appressed, c.
14-1 mm long. Petals concave, spreading, finally
reflexed, sparsely hairy to glabrous, accrescent to
21% by 1 mm. Filaments glabrous, up to 1 mm long;
anther cells c. 34 mm long. Disk c. Y2 mm high.
Carpels 2(—3), never more than 2 fertile, connate,
except at the top, rather large, growing during

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

2

anthesis. Fruit obcordate, up to 2 by 244 cm,
strongly emarginate, wings often nearly touching
by the inward curved style-bases; pericarp hard-
corky.

Distr. From SW. Borneo eastwards to Mi-
cronesia (West and East Carolines and Marshalls)
and Melanesia (New Britain, Solomons, New
Hebrides), not in Australia and not in New Cale-
donia; in Malaysia: N. and SW. Borneo (only
Banguey, Sarawak, Karimata I.), Moluccas (Hal-
maheira, Sula, Batjan, Gebe, Buru, Ceram,
Ambon, Banda), and New Guinea (also Admiralty
Is). Fig. 22.

F ————— _— ——— —

(

Fig. 22. Distribution of Soulamea amara LAMk,
sheets seen (@), from literature (0), unlocalized

(+).

It was recorded from Java by MIQUEL, but this
rests On an error.

Ecol. A typical constituent of the Barringtonia
formation, but much rarer than most of the species
belonging to that formation, though locally com-
mon on the sandy beach and behind coral reefs,
below 3 m, associated with Messerschmidia

argentea, Scaevola taccada, and Ochrosia, some-
times (but not in Malaysia) dominant as a rather
small shrub along the shore, and of more scattered
occurrence as a treelet more inland. Under the
parent plant seedlings may be found in great
profusion. The majority of the localities are
situated on small islands or islets, and atolls, a
peculiarity which it shares with very many other
beach plants, such as Pisonia grandis, Suriana
maritima, etc. Though its distribution is less errat-
ic as compared with Suriana, it is less common
than could be expected; the reason of its preference
will probably rather liein a sort of exacting habitat
than in the chances offered by dispersal. Though
not expressly stated and justified by experiments
I agree with RIDLEY (Disp. p. 264) that the fruits
possess buoyancy power and seem adapted to
dispersal by seawater; HemsLey recorded that
seeds have been found in the crop of birds in the
Admiralty Is.

The reason why it is absent in West Malaysia,
the Philippines, Celebes, Java, and the Lesser
Sunda Islands, save for four localities in Borneo,
must remain a rather fascinating problem, similar
to that found in the erratic distribution of the
beach Triumfettas, Scaevola plumierii, Launaea
pinnatifida, Spilanthes urens, etc.

Uses. The roots and fruits of this very bitter
plant are often used against cholera, pleurisy, and
other fevers. Powdered and mixed with water the
beverage is taken against colic and cough. In
poisoning, e.g. by snake bites, the fruits are whole-
some by urging the patient to vomit (RUMPHIUS,
1.c., HEYNE, /.c.).

Vern. Bona atti, buwa hati, kaju sulamu, M,
pénawar pipis, sulamu pohon, Ternate, dschiri
pangpang, Tami.

9. IRVINGIA

Hook. f. Trans. Linn. Soc. 23 (1860) 167; BENTH. & Hook. f. Gen. Pl. 1 (1862)
314; Pierre, Fl. For. Coch. 4 (1892) t. 263; VAN TreGH. Ann. Sc. Nat. IX, 1 (1905)
247-320; ENGLER, in E. & P. Pfl. Fam. 19a (1931) 398.—Irvingella VAN TIEGH.
Ann. Sc. Nat. IX, 1 (1905) 276.—Fig. 23, 24.

Large trees. Branchlets with conspicuous annular scars of the stem-clasping,
very early caducous stipules forming a narrow-conical cap surrounding the ter-
minal bud. Leaves simple, glabrous, entire; midrib sulcate; petiole with a groove
above between the very narrow wings. Panicles terminal or axillary. Bracts small,
early caducous. Flowers (4—-)5-merous, bisexual. Sepals connate at the base,
imbricate in bud. Petals exceeding the sepals, imbricate in bud. Stamens twice as
many as petals, inserted beneath the disk; filaments slender, long, dorsally atta-
ched; anthers latrorse, emarginate at base and apex. Disk large, cushion-shaped,
intrastaminal. Ovary 2-celled, conical or somewhat flattened, sessile on the torus-
like disk; style 1, with an inconspicuous terminal stigma; ovules solitary, anatro-
pous, attached adaxially and apically. Drupe large, 1-(2—)-seeded, resembling a
mango. Albumen 0 or small (e.g. in I. malayana).

224 FLORA MALESIANA

[ser. I. vol. 6

Fig. 23. Irvingia malayana OLty. ex BENN. in cleared area with secondary forest behind, grassland and

Musa in foreground, near Gemas, Malay Peninsula (Photogr. CORNER).

Dec. 1962]

SIMAROUBACEAE (Nooteboom)

225

Distr. About 3 spp. in tropical Africa and | sp. in tropical SE. Asia and W. Malaysia.

Ecol. The African species are sometimes a conspicuous constituent of the tropical rain-forest. The
Asiatic species is not seldom frequent in the lowland forest. Because of its hard wood and big buttresses
it is often left in forest clearings and thus by its great size stands out on the landscape (CORNER).

Uses. The fruit of all species is edible, but usually only the seeds are eaten. The cotyledons (as the
bark and pith of branchlets and petioles) contain large lysigenous arabin-containing mucilaginous cells,
surrounded by cells containing fat. The seeds are used for the manufacturing of wax, cacao, and soap.

1. Irvingia malayana OLiv. ex BENN. in Fl. Br.
Ind. 1 (1875) 522; Oxtv. in Hook. Ic. Pl. (1877)
t. 1247; Pierre, Fl. For. Coch. 4 (1892) t. 263 A;
Kina, J. As. Soc. Beng. 62, ii (1893) 230; BACK.
Schoolfl. Java (1911) 194; Crats, Fl. Siam. En. 1
(1926) 243; Ripey, FI. Mal. Pen. 1 (1922) 364;
Heyne, Nutt. Pl. (1927) 872; Burk. Dict. 2 (1935)
1251; CorNeER, Ways. Trees (1940) 604; GAGNeEP.
Fl. Gén. I.-C. Suppl. 1 (1946) 669.—/. oliveri
Pierre, Fl. For. Coch. 4 (1892) t. 263 B; LEComMTE,

=< JOSSEN

Fig. 24. Irvingia malayana OLIv. ex BENN. a. Flowering twig,

* %, c. flower, * 4, d. ditto, petals and stamens removed, *
carp and remains of exocarp, f. ditto, in section, »

Fl. Gén. I.-C. 1 (1911) 701; Crats, Fl. Siam. En.
1 (1926) 243; GAGNeEP. FI. Gén. I.-C. Suppl. 1
(1946) 670.—Irvingella malayana VAN TiEGH. Ann.
Sc. Nat. IX, 1 (1905) 276.—/rvingella oliveri VAN
TieGH. /.c.—Irvingella harmandiana VAN TiEGH.
1c. 279.—I. harmandiana Pierre [in De Laness.
Pl. Ut. Col. Fr. (1886) 306, nomen] ex Lecomte,
Fl. Gén. I.-C. 1 (1911) 701.—I. longipedicellata
GaGneP. ibid. Suppl. 1 (1946) 670.—Fig. 23, 24,

2/,, b. stipules clasping terminal bud,
4, e. withered fruit showing fibrous meso-
2/4, (a THORENAAR 29-E-1P-362, b bb 8375, c—d

29-E-1P-424, e-f 29-E-1P-484).

More or less deciduous large tree, up to 60 m
and 114 m 9; bole straight with very prominent,
thin, steep buttresses; bark grey, sometimes peeling
off in large pieces. Leaves elliptic-oblong to lan-
ceolate, slightly acuminate, broad-cuneate to
roundish, or even subcordate at the base, dis-
tinctly prominent-reticulate-veined at either side,
8-20 by 214-9 cm; petiole 1-2 cm. Stipules sur-
rounding the buds as a sharp, narrow-conical cap
up to 3(-4) cm long. Panicles 5-15 cm long.

Bracts ovate, acute, c. 11 mm. Pedicels up to
3(—5) mm, articulate at the base. Flowers glabrous,
greenish-white or yellowish. Calyx c. 114 mm long,
the lobes rounded, with membranous margin, c. 1
by34-1 mm. Petals elliptic-oblong, rounded, finely
reticulate, c. 3-4 by 2 mm, spreading finally or
reflexed and with often involute margin in the
upper half. Disk plicate outside and with sinuous
margin, apex excavated in the middle. Filaments
sinuous in bud, 3—6 mm, their base situated in the

226

FLORA MALESIANA

[ser. I, vol 6?

concavities of the disk; anthers c. 44 by 44 mm.
Ovary c. 1 mm high, style 1-2 mm, sinuous in
bud. Drupe with thick, fleshy, very fibrous, orange
exocarp and hard endocarp, somewhat flattened-
ellipsoid, up to 6 by 4cm when dry. Seed with small
albumen, only at the back of the cotyledons.
Distr. Siam, Indo-China (Laos, Cambodia,
Cochinchina); in Malaysia: Sumatra, Malay
Peninsula, Borneo, and Bawean. Fig. 25.

Ecol. Scarce or rather common, scattered in
dryland primary rain-forest, below 250 m. Flow-
ers before or with the new leaves. Fi. fr. Jan.—Dec.

Uses. The yellow-coloured wood is too hard to
manufacture, and, besides, not very durable.
MAINGAY recorded its use for making kris-handles
and Foxwortuy for handles of tapping knives
(BURKILL).

The seeds contain a white or sometimes yellow

fat with an agreeable smell and taste, which is
known as ‘dika’ fat in Europe and is used for
making soap, wax, and candles. The seeds are
also eaten (HEYNE, /.c.; FoxwortHy, Mal. For.
Rec. 8, 1930, 26; BuRKILL, /.c.).

Vern. Mal. Pen.: bunga paukijang, kébayang,
mérélang, mirlang, pauh kidjang, pauh kijang,
M, perseh, Sakai; Sumatra: émplas batu, kulut,
Lampong, pauh bayan, p. kijang, p. rusu, sépah
wv bongin, s. bungin, Palembang, kalék karsik,
sy i Minangk.; kaju bongin, sépah, Kubu, pauh ménté;
Borneo: kaju batu, k. tulang, k. tulung, kérangi,
kéranji, pauh kijang, M, téngilan, Kedayan,
malenna gunung.

Note. The vegetative parts show some resem-
Fig. 25. Distribution of Irvingia malayana Ou1v. blance to those of Samadera and Inocarpus
ex BENN., examined sheets (@), from literature (Legum.) but are easily distinguished by the con-

(o). spicuous stipules leaving annular scars.

Ib Z

Excluded

Ailantopsis poilanei GAGNEP. Not. Syst. 11 (1944) 163, a monotypic genus from Indo-China, was
reduced to Heynea by GAGNEPAIN at the instigation of PELLEGRIN, Not. Syst. 13 (1947) 63; Fl. Gén.
I.-C.Suppl. 1 (1948) 727 is according to BENTVELZEN, Act. Bot. Neerl. 11 (1962) 14 = Trichilia connaroides
(W. & A.) BENTVELZEN f. connaroides (Meliaceae).

Philagonia BL. referred by Miquel, FI. Ind. Bat. 1, 2 (1859) 679 to the Simaroubaceae = Evodia (Ru-
taceae).

Picroderma laotica THOREL ex GAGNEP. Not. Syst. 11 (1944) 165 is according to BENTVELZEN, Act. Bot.
Neerl. 11 (1962) 17 = Trichilia connaroides (W. & A.) BENTVELZEN f. glabra BENTVELZEN (Meliaceae).

Quassia simaruba (non L.)BLco, FI.Filip. ed. 2 (1845) 247, ed. 3, 2 (1878) 94 is according to MERRILL, Sp.
Blane. (1918) 241 = Guioa koelreuteria (BLCO)MERR.; RADLKOFER, Pfl. R. Heft 98 (1933) 1273 reduced it
with doubt to Aryrera litoralis BL. Anyway it belongs to Sapindaceae.

Quassia tricarpa Buco, FI. Filip. (1837) 351; ed. 2 (1845) 206, ed. 3, 2 (1878) 94, t. 388, p.p. is according to
MERRILL, Sp. Blanc. (1918) 238 and RapLKorer, Pfl.R. Heft 98 (1932) 642 = Sapindus saponaria L.
(Sapindaceae).

Tetramyxis GAGNEP. Not. Syst. 11 (1944) 166, of which 3 species were described, is, according to FORMAN,
Kew Bull. 16 (1962) 158 and TarpiEu-BLot, Fl. Cambodge efc. n. 2 (1962) 130—131 = Allospondias
lakonensis (PIERRE) STAPF (Anacardiaceae).

CELASTRACEAE—I (Ding Hou, Leyden)

Trees, erect or scandent shrubs; stems sometimes producing rootlets (Euonymus
spp.), rarely buttressed at the base (e.g. Bhesa) or with aerophores (Lophopetalum
multinervium), sometimes thorny (Maytenus spp.); sometimes with elastic or resin-
ous threads in the leaves, inflorescences, floral parts, fruits, or branchlets, show-
ing on fractures. Leaves simple, alternate, spiral, decussate or opposite, some-
times fascicled on short branchlets, penninerved, sometimes black-dotted be-
neath, rarely so on both surfaces, often crenate, more rarely entire. Stipules
small, simple or laciniate, caducous, or none. Inflorescences axillary and/or ter-
minal, sometimes extra-axillary, or ramiferous, cymose, thyrsoid, paniculate,
racemose, fasciculate, sometimes 1-flowered, usually bracteate. Flowers generally
small, actinomorphic, bisexual or unisexual, in the latter case the plants usually
dioecious or sometimes polygamous. Calyx 4- or 5-lobed, lobes imbricate,
rarely valvate, usually persistent. Petals 4 or 5, imbricate, contorted, rarely val-
vate, caducous, sometimes persistent, rarely slightly connate at the base and
sometimes also united with the staminal ring below the connate filament bases
(i.e. the so-called ‘disk’ in Microtropis), upper surface usually smooth, sometimes
partly covered with cristate, lamellate, fimbriate, or fleshy papilla-like appendages
(e.g. Lophopetalum). Stamens (2-) 3, 4, or 5, rarely 8-10 (extra-Mal. gen. Forsellesia),
alternate with the petals (except in Forsellesia), filaments inserted on or within
the disk, on its margin or slightly below it, or on a basal ring (Microtropis), ca-
ducous or persistent; anthers mostly 2-celled, very rarely 1-celled (extra-Mal.
spp.), usually ovoid, ellipsoid, or subglobose, rarely reniform, sometimes diver-
gent, longitudinally, laterally, or very rarely apically (extra-Mal. spp.) dehiscent,
introrse or extrorse, basifixed, dorsifixed, or dorso-basifixed. Disk various, often
present and conspicuous, fleshy or membranous, patelliform or cupular, or flat,
entire, dentate, angular, or lobed; extrastaminal to intrastaminal, sometimes
adnate to the torus or partially free at the margin, usually annular and continuous,
rarely discontinuous and lobed, or even forming staminiferous pockets (extra-
Mal. genera Cheiloclinium and Apodostigma), rarely obscure (Microtropis),
usually smooth, rarely covered with papilla-like or fleshy subulate processes).
Ovary partly or entirely immersed in the disk, sometimes concealed within it or
adnate to it, or free from it, usually glabrous, sometimes with a tuft of hairs at
the top (Bhesa), rarely puberulous (extra-Mal. spp.), or covered with papilla-
like or fleshy subulate processes at the base (Evonymus spp.), (1—)2—5-celled or
rarely many-celled (Siphonodon), mostly completely, very rarely incompletely
celled; usually ending in a style, or very rarely hollow at the top (Siphonodon);
style distinct, short, or obscure, or lacking (Brassiantha, Siphonodon and extra-
Mal. genus), simple, rarely almost divided to the base (Bhesa), terminal, rarely
lateral in fruit (Pleurostylia); stigma(s) simple, or lobed. Ovules mostly 2 in each
cell, sometimes 1, or 3-18, anatropous, inserted at the inner angle, erect and
inserted at the base or slightly higher, or pendulous, collateral, superposed or in
2 series. Fruit capsular, loculicidal or with 3 divergent separate or laterally con-
nate ‘follicles’, or drupaceous, dehiscent, and sometimes leaving a columella,
or indehiscent, smooth, sometimes echinate. Seeds erect or pendulous, sometimes
winged; aril present or none, when present usually partly or entirely enveloping

(227)

228 FLORA MALESIANA [ser: Ty voliiG2

the seed or cushion-like situated at the base of it; usually orange or orange-red,
rarely white; albumen present or 0; embryo erect; cotyledons flat, foliaceous.

Distribution. The family Celastraceae (including Hippocrateaceae) comprises c. 90 genera and over
1000 spp., distributed in both hemispheres except the arctic regions, predominantly occurring in the
tropics and subtropics.

Ecology. The Malaysian spp. occur mostly in primary, occasionally in secondary rain-forests, some
in peat or freshwater swamp forests. They are mostly shrubs or small trees of the substage, some species
are climbers, but some may grow to large-sized trees, e.g. spp. of Bhesa, Kokoona, Lophopetalum, and
Siphonodon. They commonly occur at low and medium altitudes, but some species grow above 1000 m up
to 3200 m, e.g. species of Euonymus, Microtropis, and Perrottetia.

Almost all Malaysian species are adapted to everwet climatic conditions but a few, notably Mayrenus
sp. and Cassine glauca (Rotrs.) O.K. prefer or are characteristic for seasonally dry climatic conditions
and are consequently found in Central and East Java and the Lesser Sunda Islands.

Cassine viburnifolia (JUss.) DING Hou is a typical mangrove plant.

Some species occur in peat swamps, notably Lophopetalum multinervium RIDL., L. sessilifolium R1DL.,
L. rigidum RiDL., L. javanicum (ZOLL.) Turcz. (occasionally), and Kokoona ovatolanceolata RIDL.
Under such conditions L. multinervium RIDL. possesses prominent buttresses and cylindrical aero-
phores (fig. 13).

The only spiny plants occur in the genus Maytenus.

Pollination. The flowers of most of the members of this family have nectariferous disks which sometimes
show a marked contrast with the colour of the ovary or some other floral parts and are visited by various
kinds of insects, e.g. ants, bees, beetles, and flies. Some species are dioecious, or polygamous and require
presumably insect-pollination. Unfortunately we have no pertinent data on pollination of tropical
Celastraceae.

Dispersal. The seeds of most of the members of this family are generally provided with a bright-col-
oured aril which contrasts to the colour of the pericarp and testa, if the latter is only partially covered
by the aril. These arillate seeds are a great attraction to birds.

In Celastrus the ripe fruits are usually yellowish with seeds enclosed in a showy red or golden-yel-
lowish aril; in the widely opening fruits the bright-coloured arillate seeds display a showy contrast. In
Glyptopetalum and some Euonymus spp. the arillate seeds are hanging out from the capsules. The cap-
sules of Microtropis and some species of Bhesa are splitting lengthwise on one side exposing the seeds
covered with bright-coloured arils lying inside.

The winged seeds of Lophopetalum and Kokoona are wind-dispersed.

Some of the drupaceous fruits, e.g. in Cassine (syn. Elaeodendron), are also bird-dispersed. The fruits
of some African Cassine spp. have been reported to be dispersed by elephants, which browse on the fo-
liage swallowing the fruit with it; the seed germinates well after having passed the digestive tract.

In Malaysia Elaeodendron subrotundum = Cassine viburnifolia (Juss.) DING Hou occurs along tidal
rivers and seashores. Its fruit has a thin exocarp and a not evenly thickened, corky mesocarp; according
to RIDLEY it is well adapted for sea-dispersal (Disp. 1930, 120, 267, 357, 426).

It is remarkable that in some species of Microtropis full-grown fruits do not contain a single mature
seed. The fruits I saw of Pleurostylia contain either no seed or this was damaged by insects. It could be
that this deficiency of setting seed is correlated with the very restricted distribution or rarety of species
of these genera.

Morphology. Elastic threads. In breaking young leaves, branchlets, inflorescences, floral parts, fruits,
or seeds of some species, the two parts remain sometimes connected by a number of fine, elastic threads,
e.g. in the leaves of the African Maytenus acuminatus and spp. of the American genus Wimmeria, They
can be dissolved in benzene and are classified as gutta-percha.

The funicle of the S. American Maytenus magnifolia is also composed of fine threads but these are of
an entirely different nature, namely spiral thickenings of vessels such as found under the seeds of
Talauma, Magnolia, and other Magnoliaceae. The loosening of spiral thickenings from vessels is not un-
common in Monocotyledons, but according to Dr. METCALFE it is probably more rare in Dicotyledons.

Floral tube and disk. \n the celastraceous flower there is generally a conspicuous cupular or plate-like
structure which is commonly called the disk.

BERKELEY (J. Elisha Mitchell Sc. Soc. 69, 1953, 185-206, t. 3-4) has studied the floral morphology and
anatomy of some representatives of Evonymus, Celastrus (2 spp.), and Pachistima. He interpreted the
cup-shaped structure in Celastrus (i.e. C. scandens and C. orbiculatus), which surrounds the ovary but
is free from it, as a ‘floral tube’ consisting of the fused basal portions of the floral whorls. The tissue be-
tween the stamens he called the ‘disk’ and considered it to represent the vestiges of reduced stamens. The
broad flat structure surrounding the ovary, e.g. in Euonymus, to the edge of which are attached the floral
whorls he also interpreted as being a ‘floral tube’ comparable to that of Ce/astrus; the ‘disk’ is considered
to occupy the same position as in Ce/astrus and has been given the same interpretation.

The term disk used in this treatment, as I have done in my revision of Celastrus (cf. Ann. Mo. Bot.
Gard. 42, 1955, 220) is equivalent to the ‘floral tube’ of BERKELEY while what I call disk lobes is equivalent
to his ‘disk’.

Dec. 1962] CELASTRACEAE (Ding Hou) 229

Arils and arillodes. According to PLANCHON (cf. Ann. Nat. Sc. III, 3, 1845, 281, 308, t. 11, f. 3-5,
t. 12, f. 9) the so-called arils of the seeds of Euonymus and Celastrus are actually false arils or arillodes.
They are formed by the dilatation and expansion of the edge of the ‘“exostome’, the foramen of the outer
integument which is often reflexed around the micropyle. He stated that one would find the same kind of
arillodes in other genera of Celastraceae. A true aril is formed, after fertilization, as an expansion of the
funicle, which is initially only adherent with the seed around the hilum, but it may eventually cover al-
most the entire surface of the seed, though often not covering the micropylar part. In the herbarium the
origin of the arillate structure is difficult to trace and in this revision I have used the term ‘aril’ for the
coloured body which is partly or completely enveloping the seed, sometimes cushion-like situated at the
base of it, regardless of its origin.

Embryology. The embryology in Microtropis is not clear; see p. 273.

Indument. Hairs occur infrequently in this family. When present they are usually unicellular, e.g. in
Perrottetia spp., sometimes multicellular-uniseriate, and occasionally associated with some tufted ones,
e.g. in Hexaspora pubescens C.T. Wuite, or papilliform, e.g. in Lophopetalum floribundum WIGHT
(cf. also METCALFE & CHALK, Anat. Dic. 1, 1950, 389).

Briquet (C.R. Soc. Phys. Hist. Nat. Genéve 33, 1916, 65) reported that the margins of sepals of
Maytenus spinosa (Gris.) Lourt. & O’DoNELL (syn. Moya spinosa Gris.) are fringed by multiseriate
trichomes which function as colleters exuding blastocolla when young and persisting as ciliae in the adult
stage. He found this kind of trichomes in Maytenus also on the margins of sepals, very rarely on petals.
e.g. in M. ilicifolia MART.

I have examined the sepals of M. ilicifolia, which are short-fimbriate or -ciliate on the margin in agree-
ment with BRIQUET’S description. According to BriqueT the margin of sepals or petals is in most of the
cases reduced to two layers of epidermis without mesophyll between, so that these fringes or cilia are
of epidermal origin and should be called trichomes. In my opinion, however, these fringes or cilia are
marginal outgrowths of the sepals or petals, a sort of teeth, but not trichomes proper.

Galls. Leaf-galls caused by gall-midges have been found on the leaves of Maytenus emarginata,
Bhesa paniculata, and Siphonodon celastrineus; the galls are+ globose or ovoid, c. 4-5 mm long.

Flower galls occur in Perrottetia alpestris; they are caused by gall-midges. The ovary is swollen and
resembles a fruit, sometimes hairy outside. Sometimes small lenticular or oval swellings, or small ca-
vities situated close to the midrib or between the nerves and veins, have been frequently found in Per-
rottetia spp. (cf. DoCTERS VAN LEEUWEN. Zoocecidia N.I. 1926, 328-329).

Phytochemistry. Many highly characteristic compounds are known from this family, but most
of them have thus far been found in a few species only.

Two exceptions may be made, however, with regard to this statement. All Ce/astraceae seem to accu-
mulate the hexitol dulcitol in leaves and bark (V. PLouvier, C. R. Paris 227, 1948, 85; ibid. 228, 1949,
1886). This chemical compound may be looked upon as a highly characteristic biochemical feature
of the family. Dulcitol has been isolated from many species of Euonymus and Celastrus and also from
species of the genera Catha, Gymnosporia, Lophopetalum, Maytenus, Siphonodon, and Tripterygium. The
second compound which seems to occur rather universally in the family is gutta. Gutta like rubber is a
polymer of isoprene with the empirical formula (C5H,)n; gutta has, however, trans-configuration while
rubber has cis-configuration. Rubber is accumulated much more frequently by plants than gutta, which
is known only from Sapotaceae, Eucommiaceae, Hippocrateaceae, and Celastraceae. Gutta was detected
in Celastraceae in the bark of several species of Euonymus by Cot in 1901 (C. R. Paris 132, 1901, 1354);
he also noticed the anisotropy of the contents of the laticiferous tubes. Birefringency differentiates
gutta deposits in plant tissues clearly from rubber deposits. According to Metz (Bot. Centr. Beih. 15,
1903, 309) rubber (in fact gutta!) occurs in many species of Ce/astraceae; in the leaves of species of Wim-
meria and Mystroxylon it is deposited in special idioblasts but in many Ce/astraceae gutta accumulates in
ordinary cells of the mesophyll. In the bark latex tubes are often present according to CoL. A number
of species of Euonymus was investigated, especially in Russia, for the possibility to produce gutta commer-
cially in a temperate climate.

The seeds of Ce/astraceae contain a large amount of fatty oils. These oils are unusual in as far as they
have a high saponification number and give rise to volatile acids on saponification. Among the latter
formic, acetic and benzoic acids seem to be always present. It is probable, however, that these volatile
acids are not esterified with glycerol but with polyhydric alcohols. The corresponding polyesters are sol-
uble in the fatty oils and are therefore always extracted together with the seed fats. The fatty oil proper
has a normal composition being composed predominantly of glycerides of palmitic, oleic, linoleic and
linolenic acids. Therefore, the characteristic feature is not due to the composition of the fatty oils, but to
the fact that the oils always contain appreciable amounts of polyesters of polyhydric alcohols in solution.
This state of affairs has been proved in the case of the fatty oil of the seeds of Celastrus paniculatus WILLD.
(B.G. GuNDL & T. P. Hitpircnu, J. Chem. Soc. 1938, 1980).

It is interesting to note that the insecticidal alkaloids of the Chinese Tripterygium wilfordii HOOK.
f. are also polyesters yielding acetic acid, benzoic acid, furan-f-carboxylic acid and a substituted nicotin-
ic acid on saponification (M. BErozA, J. Am. Chem. Soc. 75, 1953, 44). Insecticidal properties are also
ascribed to other Ce/astraceae, e.g. Euonymus europaeus L. and the very rare furan-f-carboxylic acid
was first isolated from the bark of Euonymus atropurpureus JACQ. Furthermore the presence of alkaloids

230 FLORA MALESIANA [ser. I, vol. 6?

has been noticed in seeds of different species of Euonymus and Celastrus and in the bark of Lophopetalum
toxicum LOoHER. Many features described in chemical literature indicate that in barks and seeds of Ce-
Jastraceae polyesters of polyhydric alcohols are accumulated. If, as in the case of the insecticidal princi-
ples of Tripterygium wilfordii, derivatives of nicotinic acid are involved in esterification, the resulting poly-
esters contain heterocyclic nitrogen and are classified as alkaloids.

Bitter, cardiotoxic glycosides have been isolated from the seeds of Euonymus europaeus L. (glycosides
of digitoxigenin; CH. TAMM & J. P. RossELeT, Helv. Chim. Acta 36, 1953, 1309), the rootbark of Euony-
mus atropurpureus JACQ. (glycosides of digitoxigenin; TsCHESCHE R. et al. Chem. Ber. 88, 1955, 1619;
C. A. Buiss & E. RAMsTAD, J. Am. Pharm. Ass. 46, 1957, 423). Digitaloid glycosides have further been
detected in the bark of Lophopetalum toxicum LOHER (used as an arrow poison in the Philippines) and
in the seeds of different species of Euonymus.

Triterpenoids are probably accumulated in large amounts in leaves and bark by many Celastraceae.
Lupeol and betulin represent approximately 3°% of the dry weight of the bark of Lophopetalum toxicum
LoHER (H. DIETERLE c.s. Arch. Pharm. 271, 1933, 264). The leaves of Celastrus scandens L. contain lu-
peol and f-amyrin (E. J. Corey c.s., J. Am. Pharm. Ass. 46, 1957, 183). From the leaves of Euonymus
alatus Stes. triterpenes of the friedelane series were isolated (T. MAzAK1 & M. Aritomt, J. Pharm. Soc.
Japan 77, 1957, 1353) and the bark of Siphonodon australe BENTH. contains at least 12 triterpenes of the
friedelane series (J. L. CoURTNEY c.s., J. Chem. Soc. 1956, 2115, 2119).

Taxonomically more important, however, are the red bark pigments celastrol (= tripterin) and pristi-
merin (= monomethylcelastrol), which according to GRANT et al. (J. Chem. Soc. 1960, 549) seem to be
modified triterpenes. Celastrol was isolated from the rootbark of Celastrus scandens L. and Tripterygium
wilfordii Hoox. f. and pristimerin from the rootbark of Celastrus (= Maytenus) dispermus and Denhamia
pittosporoides. The isolation of these pigments from species of different genera and different continents
indicates that they are probably of frequent occurrence in the family.

Finally two species have to be mentioned. The aril of Maytenus species is used in Brazil as a diuretic;
according to FRreIsE (Pharm. Zentralhalle 76, 1935, 704) the drug contains 0,85°% of coffeine. Catha edulis
Forsk. yields the so-called khat or abyssinian tea; this drug contains an appreciable amount of the
alkaloid norpseudoephedrine (= cathine), a stimulantof the central nervous system (compare R. PARIS
& Mme H. Moysg, Le thé des Abyssins, Bull. Stupé fiants 10, 7. 2, 1958, 10 and K. WINTERFELD & G.
BERNAUER, Arch. Pharm. 293, 1960, 991).

Comparing the thus far known chemical characters of Celastraceae with those of other families we
come to the conclusion that there are close biochemical affinities to one family only, Hippocrateaceae.
These affinities are so striking (dulcitol, gutta, pristimerin) that from a phytochemical point of view the
separation of these two families seems hardly justified. —R. HEGNAUER.

W ood-anatomy. DEN BERGER, Determinatietabel houtsoorten van Malesié, Veenman, Wageningen
(1949) 30, 33, 34, 43, 46, 67; Descu, Mal. For. Rec. 15! (1941) 77; ibid. 152 (1954) 430, 525 (hand lens);
CHALK & CHATTAWAY, Proc. Roy Soc. B (1933) 82; Hemscn, Lilloa 8 (1942) 182, 189; METCALFE &
CHALK, Anat. Dic. 1 (1950) 393; MoLL & JANSSONIUS 2 (1908) 254.

The family is interesting because of the distribution of derived and primitive features among the genera:
libriform fibres or septate libriform fibres and fibre tracheids; simple and scalariform perforation plates
(with few or many bars); radial multiples and solitary pores; wood parenchyma absent or sparse, and
multiseriate bands; multiseriate rays with short and such with very long wings. HErmscH, /. c., suggested
affinity of Staphyleaceae to Celastraceae, cf., however, Fl. Mal. 6 (1960) 53—C.A.R.-G.

Taxonomy. In Prerre’s opinion (FI. For. Coch. 4, 1893, sub t. 296B) Bhesa (syn. Kurrimia), though
closely related to Celastraceae, differs by its small embryo, distinct styles, and additional anatomical features
and represents the type of a family of its own,*Kurrimiacées’ (Kurrimiaceae). This group, which is char-
acterized by: spirally arranged leaves with crossbar veins, three vascular bundles in the petiole, two or
three vascular bundles in the pith of the petiole, free styles, and an embryo which is much shorter than
the albumen.

METCALFE & CHALK (Anat. Dic. 1, 1950, 395) have stated that from the wood-anatomical strandpoint
Perrottetia (syn. Caryospermum), Bhesa (syn. Kurrimia), and Siphonodon represent aberrant genera by
the presence of paratracheal parenchyma and the absence of fibre tracheids. They quoted and followed
the opinion of SpRAGUE, who said that Tripterygium. Perrottetia, and Kurrimia are not sufficiently dis-
tinct to exclude them from the Celastraceae.

METCALEE (in /itt.) stated that the wood structure of Siphonodon differs to a certain extent from that of
most of the Celastraceae represented in Jodrell Laboratory collection, ‘but that the wood structure of the
family as a whole is rather heterogeneous, and, if Siphonodon is to be excluded certain other genera such
as Microtropis, Bhesa, and Tripterygium could with equal justice be excluded as well’. Mr H. K. Airy
SHAW (in litt.) feels that ‘from evidence from external morphology Siphonodon should not be excluded
from the family, although agreeing that the floral characters are somewhat aberrant.’

From external morphology, anatomical characters, pollen morphology and chemotaxonomical char-
acters, Hippocrateaceae and Celastraceae are very closely related to one another. A.C. SMITH & BAILEY
(J. Arn. Arb. 22, 1941, 389-394, t. 1) have, in a detailed discussion of the characters and relationships of
the new Malaysian genus Brassiantha, pointed out that these two families ‘exhibit parallel series of varia-
tions in characters pertaining to wood-anatomy and pollen-structure’ and in their opinion their separa-

Dec. 1962] CELASTRACEAE (Ding Hou) 231

tion is quite artificial (cf. /.c. 394). On the basis of chemical characters HEGNAUER (see p. 230) has concluded
that ‘from a phytochemical point of view the separation of these two families seems hardly justified’. It is
also my contention that Celastraceae and Hippocrateaceae cannot be upheld as two separate families
and they will be treated in this revision as one.

Generic delimitation. In this revision the following new generic reductions have been made:
Quadripterygium TarviEU has been reduced to Exonymus L., Phanrangia TARDIEU to Mangifera L.,
Solenospermum ZOLL. to Lophopetalum Wicut, and Monocelastrus WANG & TANG to Celastrus L.

Although not occurring in Malaysia, there is another pair of genera which deserve attention, viz
Tripterygium Hoox. f. in B. & H. Gen. Pl. 1 (1862) 368 from East Asia and Wimmeria SCHLECHT.
Linnaea 6 (1831) 427 from Central America.

In the key by LoESENER (in E. & P. Pfl. Fam. ed. 2, 20b, 1942, 110) they are merely contrasted by the
number of ovules, 2 basal and collateral in Tripterygium, 6-8 (in his description 4-8) axile in two series in
Wimmeria. This single character is feeble, as for example in Evonymus the number of ovules per cell
varies from 2-12. If this would be the only differential character I would have reduced Tripterygium to
Wimmeria. There are, however, a number of other macroscopical differential characters observed on the
available material, and besides anatomical differences, which I owe to Prof. HEGNAUER, worthwhile to
put on record here:

Tripterygium: Climbers. Bark of twigs lenticellate, often warted. Leaves herbaceous, broad-ovate;
base of blade rounded; petiole and base of midrib sulcate on the upper surface. Inflorescences large, ter-
minal thyrses leafy at the base. Ovules 2 per cell. In breaking the leaves no gutta-percha threads. Veins
and nerves with well-developed tracheid sheath; Ca-oxalate crystals in the leaf parenchyma restricted to
rows of cells of normal size adjacent to the tracheids of these sheaths.

Wimmeria: Shrubs or trees. Bark of the twigs not lenticellate or warted. Leaves chartaceous to coria-
ceous, narrower; base of blade cuneately attenuated into the petiole; base of midrib and petiole flat above,
with prominent midrib. Inflorescences smaller, axillary, not forming large terminal thyrses. Ovules 4-8
per cell. Leaves with gutta-percha threads showing on the fractures. Veins and nerves without tra-
cheid sheath; leaf parenchyma with large Ca-oxalate crystals in crystal-idioblasts.

Some of the leaf characters may be correlated with the more sclerophyllous texture of the leaves of
Wimmeria. In each case these two genera are closely allied and one may quibble to unite them and give
them subgeneric rank, but one cannot escape the definite impression that they are separate taxa. Their
distribution coincides exactly with the North Pacific Asa Gray disjunction.

Uses. Some Celastraceae are used for timber, notably Lophopetalum wightianum ARN. and L. mul-
tinervium RipL., Kokoona littoralis Laws. and K. reflexa (LAws.) DING Hou, Bhesa paniculata ARN.,
and Siphonodon celastrineus GrirF. Timber of Euonymus javanicus BL. is said to be very resistant against
weathering and insects, but little is known of it because timber of good size is extremely rare.

Bark of Kokoona and Lophopetalum species contains oilin a thin outer layer and is easily inflam-
mable, even in a wet state; it is used as fire-lighter in the forest.

The fruits of Salacia and Bhesa have seeds with a rather large aril which is edible.

Leaves of Celastrus paniculata WILLD., Cassine spp., Salacia spp., and Hippocratea spp. are used for
medicinal purpose. BURKILL discussed the use of a decoction of the bark of Lophopetalum pallidum
Laws. for dart poison, together with many other ingredients; it is not clear which role is played by Lo-
phopetalum.

For further details see HEYNE, Nutt. Pl. (1927) 983-985 and BurRKILL, Dict. (1935) 505, 904, 1288,
1365.

Note. For practical purposes the revision of this family will be published in two parts; the second part
will contain the treatment of the genus Siphonodon and the genera formerly assigned to the Hippocratea-
ceae. In the second part I will discuss more fully the subdivision of the family, frame my final generic
delimitation, and provide an emended key.

KEY TO THE GENERA
(based on flowering material)

1. Leaves spirally arranged or alternate.
2. Leaves with crossbar veins between the nerves; petiole thickened at the apex beneath. Petals contort-
ed. Styles 2, free or slightly united at the base . . 2) See lee bhesa
2. Venation reticulate. Petals usually imbricate, rarely valvate (Perrotiennys Style simple.
3. Petals always larger than the calyx lobes and usually different in shape, imbricate. Ovary 3—4-cel-
led (2-celled in Maytenus diversifolia).
4. Ovary (2—)3-celled, each cell with two collateral ovules.
5. Ovary free from the disk. Ovules with a cup-shaped aril at the base. Scandent shrubs, always

unarmed : ee 2. Celastrus
5. Ovary usually partly immersed in the disk. Ovules without arillar cup ‘at the base. Erect (some-
times scandent?) shrubs or small trees, sometimes spinous ....... ... . 3. Maytenus

4. Ovary 4-celled, each cell with c. 10 ovules arranged intwo series . . . .. . . . 4. Xylonymus

232 FLORA MALESIANA [ser. I, vol. 6?

3. Petals usually similar to calyx lobes both in size and shape, usually valvate. Ovary 2-celled.
5. Perrottetia

1. Leaves decussate or opposite.

6. Petals slightly united at the base (very rarely free, in Microtropis filiformis). Disk proper absent,
filaments united at the base in a ring or short tube, by some interpreted to represent a disk, usually
united with the petals... . sec peel! Microtropis

6. Petals free. Disk conspicuous, fleshy, ‘cupular ¢ or ‘flat (Covering the. ovaey and coherent with it in
Glyptopetalum sp.), free from the petals.

7. Ovary 3-, or 4~S-celled. Ovules 1—18 in each cell.
8. Ovary 4-5-celled.

Or tOvulesi2vinieachscellieien 2) < Soo. Seth se eal Been oo ae ee OOD SITIES:
9. Ovule lineachcell .. . Sele ijel op Sr te Seeleda tet ve GlyptOpetalann
8. Ovary 3-celled. Ovules (4-6)8- 18. in 1 each cell.

10. Petals contorted, without appendages . . . . . 8. Kokoona

10. Petals imbricate, usually with appendages on the i inner side, very rarely naked . 9) Lophopetalum
ie nee 2-celled, or 1-celled by abortion (Pleurostylia). Ovules 2 in each cell.
. Disk more or less flat. Anthers subglobose and rounded at the apex, connective invisible on the

dorsalside a -wn i ake 11. Cassine
11. Disk cupular. Anthers ovoid “and ‘short- apiculate, connective distinct and broad on the dorsal
Stal ea eee te RSG enon bo. a neti ace kane) eA COSEES Sake, Seek eer iAM ie iL, [Aims eine

KEY TO THE GENERA

(based on fruiting material)

1. Leaves spirally arranged or alternate.
2. Fruit a loculicidally dehiscent capsule, sometimes splitting on one side only (Bhesa). Seeds smooth or
sometimes areolate. Petals usually caducous after anthesis.
3. Fruit 2-3-celled, or 1-celled by abortion, 2—3-valved, or splitting on one side; valves usually leath-
ery, less than 2 mm thick; each cell I— or 2-seeded, in the latter case seeds collateral.
4. Leaves with crossbar veins between the nerves; petiole thickened at the apex beneath. Fruit usually
not subglobose, sometimes 2-lobed, 2-valved or splitting on oneside. . . | nao esa
4. Venation reticulate; petiole not thickened at the apex beneath. Fruit usually subglobose, 3-valved
oe valved in Maytenus diversifolia).

. Seeds completely enveloped by the aril. Scandent shrubs . . . . 2. Celastrus

2 Seeds incompletely (at the lower half or at the base) enveloped by the aril. Erect (sometimes
scandent?) shrubs or small trees . . . . . 3. Maytenus

3. Fruit large, 4-celled, 4-valved, valves woody, qe 9 mm : thick, "each cell Cs “10- seeded: seeds arranged
in two rows. . . . . . 4. Xylonymus

2. Berry indehiscent. Seeds muricate- foveolate or r tuberculate. Petals ‘usually persistent.
5. Perrottetia
1. Leaves decussate or opposite.
6. Fruit a dehiscent capsule.
7. Fruits 3—S-celled, loculicidally dehiscent, usually 3—S-valved, 3—co-seeded.
8. Fruits usually 4-S-angular or -lobed, 4—S-celled, Bieter 1-3-celled by abortion, each cell
1— or 2-seeded. Seeds not winged, completely or incompletely enveloped by the aril.
9. Axis of the fruit splitting completely together with the valves when the fruit dehisces. Seeds
usually 2 in each cell, attached to the top or base at the inner angle, raphe not branched.
6. Euonymus
9. Axis splitting or not, free from the valves when the fruit dehisces (columella). Seeds only one in
each cell, hanging from the top of the persistent axis, raphe branched usually at the morpholo-
gical base of the seed, the bands ascending on the other side towards the hilum.7. Glyptopetalum
8. Fruits 3-angular, -lobed, or + winged. Seeds winged. No aril.
10. Seeds attached at their base, wing at the apicalend . . . . . . . 8. Kokoona
10. Seeds attached at their + centre on one side, the wing surrounding the entire seed.
9. Lophopetalum

7. Fruits usually 1-celled, splitting on one side, usually 1-seeded.. . . . . . . . 10. Microtropis
6. Fruit drupaceous, indehiscent.
11. Fruit with terminal persistent style oritsscar ...............- . M1. Cassine

1., Fruit) with Jaterali persistent style. = -, 0 6 5 ss) se eee sen eeiblennosnylia

Dec. 1962] CELASTRACEAE (Ding Hou) a — = 233

1. CELASTRUS

Linng£, Gen. Pl. ed. 5 (1754) 91; Sp. Pl. (1753) 196; Logs. in E. & P. Pfl. Fam. ed.
2, 20b (1942) 131; Dinc Hou, Ann. Mo. Bot. Gard. 42 (1955) 227.—Celastrus
§ Eucelastrus W. & A. Prod. (1834) 158.—Monocelastrus WANG & TANG, Acta
Phytotax. Sin. 1 (1951) 36.—Fig. 1.

Fig. 1. Celastrus novoguineensis MERR. & PERRY. a. Habit, = 24, b. flower, 6, c. section of flower,
pistil removed, * 6, d. pistil, * 12, e. petal, <6, f. dehisced fruit, nat. size, g. seed, aril largely removed,
* 2.—C. stylosus WALL. h. Branch with fruit, * 24, i. stamen, x 6, 7. dehisced fruit, seeds removed, nat.
size, k. seeds, = 2, /. remains of central axis and septa after valves and seeds have dropped (a-e BRASS
32222, f-g WomeRSLEY 6051, h, j. k (right), / BACKER 22483, i BAKHUIZEN 2534, j BACKER 22483, k (left)
FORBES 3829).

Scandent deciduous, rarely evergreen shrubs, sometimes very large. Branchlets
(in Mal.) terete, glabrous rarely pubescent, solid very rarely hollow, almost always
lenticellate except sometimes on the flush. Leaves spirally arranged, petioled,
elliptic to orbicular, dentate, crenate or subentire, rarely finely ciliate-serrulate
(extra-Mal. sp.). Stipules small, usually laciniate and caducous. Inflorescences
pyramidal or narrow raceme-like thyrses with cymosely arranged flowers, some-

234 _FLorA MALESIANA [ser. I, vol. 6?

times branched almost from the base, solitary, axillary, and/or terminal, peduncled
or sessile, few- to many-flowered. Flowers 5-merous, small, usually light greenish,
unisexual and the plants usually dioecious, or bisexual (extra-Mal. spp.), pedicelled
and articulated. Calyx campanulate, persistent; lobes imbricate or valvate (extra-
Mal. sp.). Disk usually membranous and cupular, or fleshy and flat, entire or
5-lobed, lobes alternate with the stamens. Fertile stamens inserted on the margin
of the disk or immediately under the outer margin; filaments glabrous or papillose;
anthers ovoid or oblong-ellipsoid, obtuse or apiculate, latrorse or introrse (not
extrorse as stated by me, /.c. 228), dorsifixed, versatile, cells separated at the
lower half or third; sterile stamens much smaller in size, their anthers usually
ovate-oblong, acute or acuminate. Ovary free from the disk or its base slightly
confluent with it, completely or incompletely 3-celled; style usually columnar,
stigma usually 3-lobed, lobes obscure, or rarely each 2-fid; sterile pistil in 3 fl.
much smaller than the fertile one, sometimes columnar, its stigma usually not
lobed. Ovules 2 or | (extra-Mal. spp.) in each cell, attached at the inner angle at
the base, anatropous, sessile or on a short funicle; with a cupular aril at its base.
Capsule usually subglobose, rarely subcylindric, tipped by the persistent style,
loculicidally 3-valved, the central axis splitting close to the insertion of the seeds,
both valves and axis ridged by the remains of the dissepiments. Seeds 1-6, en-
veloped by a fleshy crimson aril, sticking together, the valves spreading; testa with
obscure or distinct areolae; albumen copious; embryo erect, cotyledons thin and

broadly spathulate.

Distr. About 31 spp. largely in the tropical and subtropical zones, widely distributed chiefly in eastern
Asia, Latin America (7 spp.), North America (1 sp.), New Caledonia and Australia (1 sp.), Fiji (1 sp.),
and Madagascar (1 sp.).

Ecol. Chiefly in forests or thickets, from the lowland up to 3000 m.

Taxon. I have divided Celastrus into two subgenera, /.c. 216-217, pl. 31. All the Malaysian species
belong to the subg. Celastrus which is characterized by: plants usually dioecious, flowers mostly unisexual,
ovary 3-celled with 2 ovules in each cell.

This subgenus has been subdivided into two series, viz ser. Paniculati REHD. & WILs. characterized by
species bearing terminal inflorescences only to which in Malaysia belong C. paniculatus and C. novoguineen-
sis, and ser. Axillares REHD. & WILS. characterized by species with both terminal and axillary inflore-
scences, or axillary ones only to which in Malaysia belong C. monospermoides, C. hindsii, and C. stylosus.

For the distinction of the genera Celastrus, Maytenus, and Gymnosporia see under Maytenus.

Notes. WANG & TANG (/.c.) have described a new genus Monoce/astrus from Yunnan (China), com-
prising M. virens WANG & TANG (Type: C. W. WANG 79015) and M. monosperma (ROxB.) WANG &
TANG (based on Celastrus monospermus ROxB.). They stated that the new genus has the scandent habit of
Celastrus but differs from it by the stamens inserted under the disk and by the ovary which is not immers-
ed in the disk but only confluent with it at its base. As the latter character holds, however, for the entire
genus Ce/astrus and the insertion of the stamens is variable, sometimes even in one species for the two
sexes, there is no reason for segregation.

Except one sheet (Tar 52465) I have seen all specimens cited to belong to ‘Monocelastrus’ and in my
opinion they all belong to one species, C. monospermus ROxB., except one (C.W. WANG 76880) which is a
doubtful specimen in fruit which I believe belongs to C. hindsii BENTH. (cf. Ann. Mo. Bot. Gard. 42,
1955, 245, 255, 297-298).

On a few field labels it was noted that the plant would be a tree; I believe this to be due to erroneous
observation.

The synonyms cited under the species are those which are based on Malaysian collections, extra- Ma-
laysian ones which are important for nomenclature or discussion, and a few which I missed in my revi-
sion of the genus.

KEY TO THE SPECIES

1. Inflorescences terminal only.

2. Branches of the inflorescence + obliquely erect. Pedicel distinct usually 1'/.-3'4 mm (3-6 mm on
fruits). Stamens in the 3 flowers distinctly filamentous; filaments sometimes longer than the anthers
in an open flower; anthers usually truncate or obtuse at the apex . . . . . . . 1. C. paniculatus

Dec. 1962]

CELASTRACEAE (Ding Hou)

235

2. Branches of the inflorescence usually at + right angles with the rachis. Pedicel obscure or very short,
c. Y% mm (c. 1 mm on fruits). Stamens in the 3 flowers usually subsessile; anthers apiculate.

2. C. novoguineensis.

1. Inflorescences both terminal and axillary, or axillary or laterally only.
3. Inflorescences usually without accompanying bud. Stamens with glabrous filaments. Fruits usually
i-seeded. Seeds broad-ovoid or -ellipsoid, 5-12 by 5-10 mm, obtuse at both ends.
4. Inflorescences few-flowered, in simple lax cymes or 1-flowered; disk membranous, cupular, 5-lobed;
stamens inserted usually between the lobes of the disk. Leaves densely reticulate. Young branchlet

(in the herbarium) light green, smooth and lacking lenticels.

3. C. hindsii

4. Inflorescences usually many-flowered, in dense umbelliform cymes; disk. + ’ fleshy, flat, rounded;
stamens inserted just beneath the margin of disk. Leaves loosely reticulate. Young branchlets (in
the herbarium) usually light brown, densely covered with slightly elevated lenticels

4. C. monospermoides

3. Inflorescences usually each accompanying a bud. Stamens with papillose filaments. Fruits 3-6 seed-

2d. Seeds -- plano-convex to slightly lunar, 4-6 by 1-2 mm, attenuate at both ends.

1. Celastrus paniculatus WILLD. Sp. Pl. 1 (1797)
1125; Roxs. Fl. Ind. ed. Wall. 2 (1824) 388;
DC. Prod. (1825) 6; Roxs. Fl. Ind. ed. Carey 1
(1832) 621; WiGurT, Ill. (1840) 176, t. 72; Ic. 1
(1840) t. 158; THwarrtes, En. Pl. Zeyl. (1864) 72;
Laws. in Hook. f. Fl. Br. Ind. 1 (1875) 617;
Trim. FI. Ceyl. 1 (1893) 272; BAKER in Andrews,
Monogr. Christmas Isl. (1900) 175; Logs. Bot.
Jahrb. 30 (1902) 470; Pratn, J. As. Soc. Beng.
73, ii (1904) 195; Logs. Bot. Jahrb. 39 (1906)
160, pro var. balansae Lors.; MERR. Philip. J. Sc.
1 (1906) Suppl. 85; BAcKeR, Schoolfl. (1911) 234;
PirarD, FI. Gén. I.-C. 1 (1912) 890; Koorp.
Exk. Fl. Java 2 (1912) 524; Ruep. & WILS. in

Sargent, Pl. Wilson. 2 (1915) 355; GAMBLE,
Fl. Madras (1918) 208; Rrpv. Fl. Mal. Pen. 1
(1922) 451; Crats, Fl. Siam. En. 1 (1926) 284;

Merr. Lingn. Sc. J. 5 (1927) 116; KANuILAL &
Das, Fl. Assam 1, 2 (1937) 268, pro var. venulo-
soides KANS. & Das; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 132, f. 31, K & L; TARDIEU,
Suppl. Fl. Gén. I.-C. (1948) 803, f. 98, 4-6,
incl. var. poilanei TARD.; Not. Syst. 14 (1950)
45; Merr. J. Arn. Arb. 35 (1954) 141; Dinc Hou,
Ann. Mo. Bot. Gard. 42 (1955) 229, t. 32 & 34,
f. 3, map 1, incl. ssp. paniculatus, serratus et
multiflorus—C. multiflorus Roxs. [Hort. Beng.
(1814) 18, nomen] Fl. Ind. 2 (1824) 389, non
LAMK, 1785; ed. Carey 1 (1832) 622; PRAIN,
J. As. Soc. Beng. 73, ii (1904) 196.—C. nutans
Roxs. [Hort. Beng. (1814) 18, nomen] Fl. Ind.
ed. Carey 1 (1832) 623.—Diosma serrata BLANCO,
Fl. Filip. (1837) 168; ed. 2 (1845) 119; ed. 3,
1 (1877) 213.—C. subspicatus Hoox. Ic. Pl. 5
(1842) t. 482; Dinc Hou, Ann. Mo. Bot. Gard.
42 (1955) 236, map 1.—Alsodeia glabra BurR-
GERSD. in Mig. Pl. Jungh. (1852) 122; Mia. Fl.
Ind. Bat. 1, 2 (1859) 116; OupeM. Arch. Neéerl.
Sc. Exact. & Nat. 2 (1867) 199, t. 9.—C. australis
Hary. & F. v. M. Trans. Phil. Soc. Vict. 1 (1855)
41; Bentu. FI. Austr. 1 (1863) 398; Dinc Hou,
Ann. Mo. Bot. Gard. 42 (1955) 236, t. 32.—
C. polybotrys Turcz. Bull. Soc. Nat. Mose. 31,1
(1858) 449.—C. racemosa Turcz. ibid. 36, 1
(1863) 599, ex descr., photogr. of type seen.—
Rinorea glabra O.K. Rey. Gen. Pl. 1 (1891) 42.—
C. papuana Wars. Bot. Jahrb. 13 (1891) 366.

Up to 10 m. Innovations usually pubescent.
Leaves glabrous, sometimes pubescent beneath

. 5. C. stylosus

especially on the venation, elliptic to elliptic-
oblong, broad-obovate, or suborbicular, ovate or
ovate-oblong, 5-15 by 214-6 cm; base cuneate,
obtuse or rounded; apex acute, acuminate, obtuse,
rarely emarginate; midrib elevated on both sur-
faces; nerves 5-8 pairs; petiole '4-114 cm.
Panicles usually thrice to multi-compound, rarely
once compound, spreading, (2-)5-10 cm long,
sometimes up to 20 cm long, usually puberulous
when young; peduncle 6-10 mm. Pedicels 114-314
mm (3-6 mm on fruits), the articulation at the
base.—¢: calyx lobes semi-orbicular, short-
ciliate, 24-1 by 114 mm. Petals oblong or obovate-
oblong, obtuse, entire, 214-3 by 1-114 mm. Disk
cupular, the lobes obscure or slightly triangular.
Stamens c. 3 mm long; filaments subulate; anthers
ovoid, obtuse. Sterile pistil columnar, c. 11/, mm
long.—®: calyx lobes, petals, and disk as in the
male. Sterile stamens 11/, mm long. Pistil 2-244
mm long; ovary globose; style columnar; stigmas
3-lobed and each sometimes bifid, slender. Fruits
subglobose; valves broad-elliptic, 5—10 bys 5—8 mm,
3-6-seeded. Seeds ellipsoid, 344-5 by 2-3 mm,
yellowish to reddish brown, smooth, or with
obscure areoles.

Distr. Widely distributed in India, Burma,
Siam, Indo-China, southern China, throughout
Malaysia (except Borneo), to Australia and New
Caledonia.

Ecol. Chiefly in thickets, 200 to 1800 m.

Notes. The type specimen of C. polybotrys
Turcz. from the Philippines is CUMING 1324, not
1321 as cited in the original description.

In my former revision I have distinguished
three subspecies within C. paniculatus but the
distinctions given, mainly based on dimensions of
leaves and geographical distribution, have broken
down through the study of abundant material
from Bogor. There are local forms but all are
connected by intermediate forms and cannot be
keyed out; consequently I have withdrawn them.

C. paniculatus is very closely allied to C. novo-
guineensis.

2. Celastrus noyoguineensis Merr. & Perry, J.
Arn. Arb. 22 (1941) 260; DinGc Hou, Ann. Mo.
Bot. Gard. 42 (1955) 234, t. 32 in part, map 1.—
Fig. la-g.

Shrub up to 10 m tall. Innovations glabrous.

236

FLORA MALESIANA

[sera ie volute?

Leaves oblong, elliptic-oblong, or broad-ovate;
base obtuse or rounded, sometimes acute; apex
acute or obtuse; nerves 5—7 pairs; petiole 1—21/,
cm. Panicles terminal, up to 20 cm, divaricately
branching, usually thrice compound. Pedicels c.
14 mm (c. 1 mm on fruits), distinctly articulated.
—¢: calyx lobes subreniform, slightly erose, c.
1 mm long. Petals oblong, obtuse, subentire,
scarious-marginate, 244 by 114 mm, subcarnose,
sometimes brownish-punctate. Disk cup-shaped,
membranous, lobes obscure, truncate or mucrona-
te. Stamens subsessile or with very short flat
filaments; anthers narrow-ovoid, obtuse, apicu-
late, brownish-dotted; sterile pistil 1 mm long.—
2: calyx lobes, petals, and disk as in the male.
Sterile stamens 1 mm long. Pistil c. 114 mm long;
ovary subglobose; style obsolete; stigmas 3-lobed,
each bifid. Fruits subglobose, the valves broadly
ovate, 9-12 by 7-10 mm, 3-6-seeded. Seeds
ellipsoid, 5-8 mm by 2!4-314 mm, reddish-brown,
smooth.

Distr. Malaysia: New Guinea, throughout the
island.

Ecol. In thickets or forests, usually at 1220-
1830 m, sometimes found in the forest at 2400—
2900 m, rarely occurring along river-banks at
100 m.

Vern. Raw wijh, remohnremohn, Enga language.

3. Celastrus hindsii BENTH. in Hook. J. Bot. Kew
Misc. 3 (1851) 334; Maxim. Bull. Ac. Imp. Sc.
Pétersb. 27 (1881) 455; Fors. & Hems-. J. Linn.
Soc. Bot. 23 (1886) 455; HALL. f. Med. Rijksherb.
1 (1910) 8; Pirarb, FI. Gén. I.-C. 1 (1912) 892;
DUNN & TUTCHER, Kew Bull. add. ser. 10 (1912)
61; ReHD. & WILS. in Sargent, Pl. Wilson. 2
(1915) 357; Merr. Lingn. Sc. J. 5 (1927) 116;
KANEHIRA, Formos. Trees. ed 2 (1936) 384,
t. 341; WANG, Chin. J. Bot. 1 (1936) 63; Logs. in
E. & P. Pfl. Fam. ed. 2, 20b (1942) 134; TARDIEU,
Suppl. Fl. Gén. I.-C. (1948) 802; DinGc Hou,
Taiwania | (1950) 175; Ann. Mo. Bot. Gard. 42
(1955) 249, f. 7, map 2, 10.—C. racemulosus Hassk.
Hort. Bogor. Descr. 1 (1858) 155, non FRANCH.
1886; Mia. Fl. Ind. Bat. 1, 2 (1859) 590; BACKER,
Schoolfl. (1911) 234; Koorp. Exk. Fl. Java 2
(1912) 524; FI Tyjib. (1923) 145.—Fliieggea
serrata Mig. Fl. Ind. Bat. 1, 2 (1859) 356, cf.
HAvuier f. Med. Rijksherb. 1 (1910) 7.—C.
racemulosa FRANCH. Bull. Soc. Bot. Fr. 33 (1886)
455, non HAssk. 1858; Pl. Delav. pt 2 (1889) 132.
—C. franchetiana Lots. Bot. Jahrb. 30 (1902)
470, new name for C. racemulosa FRANCH.;
in E. & P. Pfl. Fam. ed. 2, 20b (1942) 133.—C.
tonkinensis Pirarb, Fl. Gén. I.-C. 1 (1912) 892,
f. 112, 1-2.—C. marianensis Koipz. Bot. Mag.
Tokyo 30 (1916) 400, ex descr.—C. axillaris R1IDL.
J. Mal. Br. R. As. Soc. 1 (1923) 56.

Evergreen, up to 30 m. Young branchlets usually
without lenticels. Leaves usually coriaceous,
glabrous, elliptic- or obovate-oblong, sometimes
elliptic or broadly elliptic, 4-1444 by 114-6 cm;
base cuneate, obtuse or rounded; apex abruptly
acute or acute; nerves 6—9 pairs; veins and veinlets
almost always distinctly elevated and densely

reticulate on both surfaces; petiole 4-1 cm.
Flowers usually in axillary as well as terminal
3—5-flowered cymes, these not unfrequently com-
bined in axillary narrow racemes of cymes which
may exceed the subtending leaf; peduncle very
short sometimes up to 214 cm. Pedicels obsolete,
or very short (14-114 mm), rarely up to 4 mm.—
3: calyx lobes semi-orbicular, imbricate, obtuse,
short-ciliate, sometimes erose, c. 114 mm long.
Petals oblong or obovate, obtuse, glandular-
ciliate, 2-5 by 11/,-244 mm. Disk cup-shaped,
lobes usually deltoid, rarely slightly oblong,
shortly acute, rounded, or slightly dentate. Sta-
mens c. 2'/,; mm; filaments subulate, glabrous;
anthers ovoid, obtuse, cordate. Sterile pistil
ovoid, c. 14% mm long.—: calyx lobes, petals,
and disk as in the males, but the petals smaller.
Sterile stamens c. 114 mm. Pistil flask-like, c. 2
mm long; ovary subglobose; style columnar;
stigma 3-lobed. Fruits ovoid to subglobose; valves
broadly-ovate or -elliptic, or suborbicular, 8-11
by 7-9 mm, usually l-seeded. Seeds broad-
ellipsoid, 5-8 by 5 mm, reddish-brown, smooth.

Distr. India, Burma, Siam, Indo-China, Central
to S. China (also Hainan), Bonin Is., Marianas,
and Malaysia: Sumatra (East Coast Res.),
Borneo (Kinabalu), and Java (in West, once in
Central Java).

Ecol. Chiefly in thickets, rarely in mossy forest,
1000-1800 m, only once at 550 m.

Vern. Areuj ki sorot, reunghas areuj, S.

4. Celastrus monospermoides Lors. Nova Guinea
8 (1910) 280; in E. & P. Pfl. Fam. ed. 2, 20b (1942)
134; Dinc Hou, Ann. Mo. Bot. Gard. 42 (1955)
247, f. 6, map 2, 9.—C. championii (non BENTH.)
Kina, J. As. Soc. Beng. 65, ii (1896) 639; RIDL.
J. Fed. Mal. St. Mus. 4 (1901) 11.—C. malayensis
Ripe. J. ‘Str. Br: R. As. Soc. 75 75a (Siijmes
Fl. Mal. Pen. 1 (1922) 451; Crais, Fl. Siam.
En. 1 (1926) 283.—C. apoensis Emer, Leafl.
Philip. Bot. 7 (1915) 2579; Merr. En. Philip.
2 (1923) 482.

Up to 17 m tall. Leaves elliptic to elliptic-
oblong, ovate, rarely broad-ovate or subrotundate,
5-16 by 21!4-7!4 cm; base cuneate, obtuse or
rounded; apex acute to acuminate; nerves 4-7
pairs, obliquely spreading and curving upwards;
veins distinct and slightly raised below, obsolete
above; petiole 6-13 mm. Jnflorescences axillary
or also terminal, 1—3 together, frequently forming
narrow raceme-like thyrses up to 10 cm long,
once to thrice compound. Bracts deltoid, irregu-
larly erose. Peduncle up to 114—3'4 cm, sometimes
very short or obscure. Pedicels 314-5 mm, the
articulation at the base.—¢: calyx lobes sub-
orbicular, c. 24 mm long, entire, short-ciliate,
sometimes erose. Petals oblong or oblong-elliptic,
(1—)114-2!4 mm long, obtuse or rounded, entire,
usually pink-punctate. Disk fleshy, flat, obscurely
lobed, the lobes transverse-oblong. Stamens
114-214 mm, attached slightly under the margin
of the disk; filaments filiform; anthers ovoid,
obtuse; sterile pistil conoid, c. 4% mm long.—
2: calyx lobes, petals, and disk as in the males.

Dec. 1962]

CELASTRACEAE (Ding Hou)

237

Sterile stamens c. 24 mm long. Pistil flask-
shaped, c. 11/, mm long. Ovary globose; style
slender, distinct; stigma discoid or slightly 3-
lobed. Fruits angular-ovoid, obovoid, or some-
times subglobose; valves ovate or obovate, or
broadly elliptic, 12-17 by 7-11 mm, once noted
to be rich-yellow. Seeds broad-ovoid, 7-12 by
5-10 mm, blackish brown, smooth; aril bright red.

Distr. Malaysia: Sumatra (N. half), Malay
Peninsula (also Penang), Borneo (N._ half),
Philippines (Mindanao), Moluccas (Ceram, Am-
bon), and New Guinea.

Ecol. Forests, from the hills up to 3000 m.

Vern. Mangauat, Bag., djirak, Sum. W.C.

5. Celastrus stylosus WALL. in Roxb. FI. Ind. ed.
Wall. 2 (1824) 401; Laws. in Hook. f. Fl. Br.
Ind. 1 (1875) 618, pro parte; PRAtNn, J. As. Soc.
Beng. 73, ii (1904) 196; AmsHorr, Blumea 5
(1945) 517.—Fig. 1h-1.

ssp. Stylosus.—Di1nG Hou, Ann. Mo. Bot. Gard.
42 (1955) 272, f. 12, map 3, 22A.—Gymnosporia
neglecta WALL. ex Laws. in Hook. f. Fl. Br.
Ind. 1 (1875) 619; PRaINn, J. As. Soc. Beng. 73,
ii (1904) 198; Nov. Ind. (1905) 419.

Shrub 3-4 m tall. Innovations puberulous or
pubescent. Leaves glabrous, or pubescent on the
nerves and veins beneath, glabrescent, elliptic-
or obovate-oblong, rarely subrotundate, 5-8 by
3-5 cm; base acute, or obtuse; apex acute; nerves
5-7 pairs, obliquely spreading and slightly curving
upward, as the veins raised beneath, not so above;
veins loosely reticulate; petiole 1-2 cm, pubescent,
glabrescent. Peduncle 5-11 mm, puberulous.
Pedicels 2-5 mm, puberulous, articulated usually
at the middle. Inflorescences axillary, as well as
cauline at the basal part of the new shoot, rarely
also terminal, cymose, short.—¢: calyx lobes
ovate, or oblong, 114 mm long, obtuse, slightly
erose to entire. Petals obovate, obtuse, slightly
erose, 2-4 by 1-114 mm, usually papillose on both
surfaces, white. Disk membranous, cup-shaped;
lobes distinctly arcuate or depressed-quadrate.
Stamens inserted between the disk lobes, 244 mm
long; filaments filiform, fleshy, usually densely
rarely sparsely papillose; anthers ovoid, obtuse.
Sterile pistil c. 114 mm.—: calyx lobes, petals
and disk same as in the male. Sterile stamens c.
1 mm. Pistil flask-shaped, c. 3 mm; ovary sub-
globose; style distinctly columnar; stigmas 3-
lobed, each lobe bifid, flat, reflexed. Fruits sub-
globose, 7-12 by 5-10 mm, 3- to 6-seeded, valves
broadly elliptic. Seeds more or less plano-convex
to slightly lunar, attenuate at both ends, reddish-
to blackish-brown, 4-6 by 1-2 mm, with distinct
areoles.

Distr. India (Assam, Bengal, and Sikkim) and
Malaysia: W. Java (Preanger) and Lesser Sunda
Is. (Lombok and Timor).

Ecol. In thickets, or forests, 1000-2750 m.

Note. The specimens of Malaysia match very
well those of India.

A slightly different ssp. glaber DING Hou with
usually glabrous filaments and elliptic-oblong

leaves occurs commonly in S. China and N. Indo-
China.

Excluded or doubtful

Celastrus lucida WALL. in Roxb. Fl. Ind. ed. Wall.
2 (1824) 400, non C. lucidus L. Mantissa 1 (1767)
49.—C. wallichii G. Don, Gen. Syst. 2 (1832) 8,
nom. novy.—C. jackianus StEuD. Nomencl. (1841)
314, nom. noy., illegit.

Based on a collection by JAck, in Penang.

According to Mr Arry SHAW (in /itt.) there is an
empty sheet in the Wallich Herbarium which
evidently never bore a specimen. The description
is very brief; I agree with Airy SHAW that if the
plant is celastraceous, there is every likelihood
of the plant being C. monospermoides which is the
only Celastrus species occurring in Penang.

Celastrus micrantha Roxs. [Hort. Beng. (1814)
86, nomen] Fl. Ind. ed. Wall. 2 (1824) 393;
ibid. ed. Carey 2 (1832) 625.

Based on a specimen from the Moluccas.

According to the isotype in the Martius Her-
barium at Brussels, kindly sent on loan by Prof.
RosuNs, it is an Ag/aia. Although the isotype
bears no flowers, distinctly conspecific fertile
material is collected in the Moluccas and New
Guinea; it appears to be distributed from the
Moluccas to Queensland. It is reduced here to
Aglaia sapindina (F. v. M.) Harms in E. & P.
Pfil. Fam. 3, 4 (1896) 298.—Celastrus micrantha
Roxs. /.c. non A. micrantha MERR. 1905.—Aglaiop-
sis glaucescens Mig. Ann. Mus. Bot. Lugd. Bat. 4
(1868) 58, non A. glaucescens KING, 1895.—A.
miquelii MERR. Philip. J. Sc. 11 (1916) Bot. 280
(Meliaceae).—Ed.

Celastrus pauciflora WALL. in Roxb. Fl. Ind. ed.
Wall. 2 (1824) 400.

Based on a specimen from JACK, in Penang.
WALLICH noted that he had not seen a specimen,
but had received the description from JACk.

By the 1-celled ovary and hairy pistil impossible
to refer to any known Malaysian Celastracea.
Mr. Airy SHAW (in Jitt.) found the description
clearly indicating a Rinorea, and very probably
R. lanceolata (ROXxB. 1832) O.K., which is common
in Penang. As the epithet pauciflora is already
occupied no name change is necessary (Violaceae).

Celastrus repandus Bu. Bijdr. (1827) 1145 is
according to HALuier f. Med. Rijksherb. 1 (1910)
8 = Maesa membranifolia Merz, but according
to Mez’s and BACKER’s identifications of the type
in the Rijksherbarium = Maesa ramentacea
(Roxs. 1824) A. DC. (Myrsinaceae).

Celastrus trigyna Roxs. Fl. Ind. ed. Wall. 2 (1824)
391; ibid. ed. Carey 1 (1832) 624, non LAMK.

Provenance: Moluccas.

WALLICH noted already the homonymy and
wrote in a footnote that this species should be
renamed; he made a cross-reference to C. ob-
tusifolia ROTH on page 394 of FI. Ind.; it is not
clear why, because the latter is from Mauritius.

Baker, in Hook. f. Fl. Br. Ind. 1, p. 618,
erroneously equalized RoxBuURGH’s and La-

238 FLORA MALESIANA [ser. I, vol. 6?

MARCK’S C. trigyna, treated by BAKER (FI. Maur. RoOxBURGH’s plant could be celastraceous. Mr
Seych.) as Gymnosporia trigyna. Airy SHAW wonders whether it could not be a
According to the very inadequate description Dichapetalum (Dichapetalaceae).

2. MAYTENUS

MOLINA, Saggio Chile (1782) 177, em. Bosc, Nouv. Dict. Hist. Nat. 14 (1803) 211;
cf. BLAKELOCK, Taxon 3 (1954) 196; ibid. 9 (1960) 15; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 134; LourTEIG & O’DONELL, De Natura | (1955) 184.—Ce-
lastrus § Gymnosporia W. & A. Prod. 1 (1834) 159.—Gymnosporia Hook. f.
in Benth. & Hook. f. Gen. Pl. (1862) 356; Logs. in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 147.—Fig. 2.

Shrubs or small trees; rarely scandent? Branchlets glabrous; young parts some-
times pubescent. Stipules small, lanceolate, sparsely laciniate, caducous, or ex-
stipulate (in some extra-Mal. spp.). Spines terminating a short-shoot, and/or in
a leaf axil, or none. Leaves spiral, alternate, rarely opposite (extra-Mal.), or in
fascicles especially on short-shoots, variable both in shape and texture even in
one species. Cymes axillary, one to several in a leaf axil, sometimes crowded at
the uppermost part of a short-shoot, or flowers sometimes in axillary fascicles.
Pedicels articulated. Flowers bisexual, sometimes unisexual. Calyx 5(—4)-lobed.
Petals 5(-4), patent sometimes reflexed after anthesis. Disk fleshy, flat or rarely
cupular, rounded, or slightly angular. Stamens inserted on the margin of the disk
or slightly just beneath its outer margin, in Q fl. abortive; anthers -+ introrse.
Ovary partly or rarely entirely immersed in the disk, the emerging part subglobose
or slightly 3-angular; completely or incompletely 3- (or 2-)celled, each cell with
2 ovules; stigmas 3 (or 2), slender, lobed or obscure. Ovu/es attached on the inner
side of the septum near the base. Capsule subglobose, sometimes slightly 3(—2)-
angular, loculicidal, 2-6-seeded. Seeds ellipsoid, at least at the base enveloped by
the aril, after dehiscence remaining and exposed in the spreading valves, or erect
and sticking together (in extra-Mal. spp.).

Distr. The genus is distributed in the tropics and subtropics of both the Old and New Worlds. It is
very difficult to estimate the total number of species; 5 occur in Malaysia.

Ecol. Lowland rain-forest, dry thickets at low altitudes, and on the beaches or at the inner mangrove.
As far as noted on labels the impression is gained that the seed is red and the aril white.

Taxon. In my revision of the genus Ce/astrus (Ann. Mo. Bot. Gard. 42, 1955, 216, p. 31) I have com-
pared the characters of the genera Celastrus, Gymnosporia, and Maytenus in tabular form. Keeping to
these provisional definitions there are, in America, especially Latin America, few species of Celaszrus
and many of Maytenus. Three of the latter have spiny branchlets similar to those in Gymnosporia; for
that reason they have been transferred to Gymnosporia, and later to the genus Moya (cf. LOESENER in
E. & P. Pfl. Fam. ed. 2, 20b, 1942, 109, 146-147). The genus Moya has recently been reduced to Maytenus
by LourTEIG & O’DoNELL (De Natura 1, 1955, 188).

In Asia the generic name Maytenus has never been used; there are two distinct groups of species,
referred to Celastrus and Gymnosporia.

In Africa the situation is again different: the species formerly described under Ce/astrus have been
transferred to either Gymnosporia or Maytenus, or first to Gymnosporia and then to Maytenus. EXELL
has pointed out (Bol. Soc. Brot. II, 26, 1952, 222 and Kew Bull. 1953, 103) that the separation of Gym-
nosporia and Maytenus as defined by LogsENerR (/.c. 109) appears artificial and there seems little point in
keeping the two genera distinct. His view is generally accepted and followed by botanists in treating that
group of plants in the African floras or revisions, e.g. EXELL & MENDONGA (Consp. FI. Angol. 2, 1954,
1-10), HUTCHINSON & DawzieL (Fl. W. Trop. Afr. ed. 2, 1 (2), 1958, 623-624), C. WiLczek (FI. Cong.
Belg. 9, 1960, 114-125), and W. Marais (Bothalia 7, 1960, 381-386). It seems also agreed that there is
no Celastrus in the Africa proper.

The differences between Gymnosporia and Maytenus listed in the table of my revision of Celastrus as
mentioned above can not be maintained, specially not in the African flora as expressed by MARAIS (/.c.).

After a discussion with Mr ExeLL, Mr BLAKELOCK, and Mr Marais I agree with the latter (/.c.) that

Dec. 1962] _ 28

Fig. 2. Maytenus curtisii (KING) DiNG Hou. a. Habit, with fruits, * 24, b. flower in section, 8, c. one
fruit valve with 2 seeds, nat. size, d. seed, with raphe and aril, * 2.—M. emarginata (WILLD.) DING Hou.
e. Seed, with raphe and aril, * 2.—M. cupularis DinG Hou. f. Flower in section, * 8, g. sections of ovary,
*12 (a SF. 15137, b. SF. 7529, c-d SF. 21368, e C.H.B. xv. J-B-ili-7a, f-g BRAss 22124).

240 ; FLORA MALESIANA [ser: I> voliG?

‘there seems no character or combination of characters constant enough to justify the retention of
Gymnosporia .

Pending the decision by a competent world monographer of the whole assemblage I retain Celastrus
and Maytenus which can be distinguished by a combination of three characters: habit, degree of adnation
of ovary and disk, and fruit structure.

So far known Celastrus spp. are scandent, Maytenus spp. are erect shrubs or trees. In a few sheets the
data of field labels were probably erroneous.

The ovary in Cel/astrus is free from the disk and only confluent with it at the base; in Mayfenus it is
partly or, rarely, entirely immersed in the disk. There are two exceptions in the latter genus, however,
e.g. M. senegalensis (LAMK) EXELL (cf. HutcH. & DAwz. Fl. W. Trop. Afr. ed. 2, 1 (2), 1958, 623-624,
f. 117 B & C) and M. cupularis DinG Hou.

After dehiscence of the fruit in Ce/astrus the central axis splits close to the insertion of the seeds and
after the seeds and valves have dropped the thickened placenta can still be observed on the pedicel. In
Maytenus, however, the central axis splits to the very base and after the seeds and valves have fallen al-
most nothing can be discerned on the top of the pedicel.

Whether this combination of differences will hold for the numerous species described in Maytenus and
Gymnosporia is not yet established.

The Australian genus Denhamia MEIsn. 1837 is also probably not distinct from Maytenus, the differ-
ential characters being normally a 1-celled ovary with 3 parietal placentas with 6—8 ovules per placenta
in Denhamia, against a completely or incompletely 2—3-celled ovary with 2 ovules at or near the
base of each cell in Maytenus. Of the four species of Denhamia, however, two (D. parvifolia L. S. SMITH
and D. pittosporoides F. vy. M.) possess normally 2 ovules per placenta or cell, with occasionally a
third or fourth ovule. Besides, in D. pittosporoides F. y. M. the capsule is 3-celled before dehiscence, the
dissepiments touching axially, and the seeds are attached at or near their base. From this it appears that
the distinction between Mayrenus and Denhamia is very meagre indeed, essentially remaining limited to
the occurrence of 3-4 seeds per placenta which in two Denhamias is only occasional. L. S. SmirH, in his
revision (Proc. R. Soc. Queensl. 67, 1956, 30-31), pointed to the anomalous characters in D. pitto-
sporoides, but he restricted his observations in Maytenus to the few Australian representatives. Really,
D. pittosporoides and D. parvifolia form a transition between Maytenus and Denhamia, in that order.

In North Queensland there is another genus, Hexaspora C. T. WuitTe (Contr. Arn. Arb. 4, 1933, 58,
t. 6). In habit, especially the 2-ranked leaves with zigzag twigs, it reminds of Perrottetia, but it has a
distinct differentiation of petals and sepals, apically dehiscing depressed-oblong anthers, and a 3-celled
ovary with 2-4 ovules per cell. Besides, it has a well-developed indument consisting of multicellular uni-
seriate hairs. C. T. Wuire recorded the ovules to be pendulous, but I found them attached clearly hori-
zontally or slightly obliquely at the inner angle at the base or sometimes slightly above it. The fruit of
Hexaspora is unfortunately unknown.

The delimitation and naming of the species of Maytenus given here is rather tentative. A thorough treat-
ment can only result from a large study including the African and continental Asian material; the impres-
sion is that there are in Asia few but variable species.

Note. The type species of the genus is M. boaria MOLINA, hence in accordance with the 1956 Code Rec.
75A the generic name Mayrenus should be treated as feminine.

KEY TO THE SPECIES

1. Ovary or fruits 3-celled, occasionally some 2-celled. Disk flat. Fruits subglobose, obovoid, or ellipsoid,
10-15 mm long, not compressed.
2. Fruits subglobose or slightly depressed-globose. Pericarp thin (c. !/, mm).
3. Fruits c. 15 mm long. Seeds (excluding the aril) c. 7 mm long; aril flat or shallow disk-like, laterally
attached at the base of the seed. Leaves pene ia ae sometimes broad-elliptic; apex usually

acute to short-acuminate .. . . 1. M. curtisii
3. Fruits 9-12 mm long. Seeds (excluding the aril) 2 y— 31/4 mm ‘long: aril fleshy, attached at the base
of the seed. Leaves usually obovate; apex obtuse or rounded . .... . 2. M. emarginata
2. Fruits obovoid or ellipsoid. Pericarp rather thick (c. 1144 mm). Leaves usually elliptic or elliptic-
oblonga: = 6 - os (Sa UMegerassa’

. Ovary or fruits 2-celled, ‘occasionally s some - flowers with ae celled ovary. Disk ‘cupular. Fruits broad-
obcordate, smaller, 5-6 mm long, + compressed (not known in M. cupularis).

4, Plant with axillary and/or terminal spines. Leaves obovate-oblong, oblanceolate, or broad-obovate,

114-414 by 1-2 cm; apex obtuse, sometimes slightly emarginate. . . . . . 4. M. diversifolia

4. Plant unarmed. Leaves elliptic, 7/4-11 by 4-5 cm; apex acuminate . . . . . 5. M. cupularis

1. Maytenus curtisii (KING) DinG Hou, comb. 2a-d.

noy.—Gymnosporia curtisii KiNG, J. As. Soc. An erect (or scandent?) shrub, or small tree.
Beng. 65, ii (1896) 353; PRAIN, J. As. Soc. Beng. Short-shoots very rarely terminating into a spine.
73, 11 (1904) 198; Rip. Fl. Mal. Pen. 1 (1922) Leaves chartaceous to subcoriaceous, elliptic,
451; Crats, Fl. Siam. En. 1 (1926) 284.—Fig. sometimes broadly elliptic, suborbicular, rarely

Dec. 1962]

CELASTRACEAE (Ding Hou)

241

obovate or ovate, 7-16 by 4-9 cm; base cuneate,
narrowed to the petiole; apex acute or short-
acuminate, rarely obtuse; margin shallow-crenate;
nerves 7—9(—15), obliquely spreading towards the
margin and then curved upwards; veins elevated
and reticulated beneath, visible above; petiole
ec. 10 mm. Cymes usually crowded towards the
apex of the short-shoots, axillary, sometimes
puberulous when young, c. 144 cm. Peduncle
5-10 mm. Bracts lanceolate, short-fimbriate.
Pedicels 2-5 mm. Calyx lobes deltoid or semi-
orbicular, 1-34 mm go, the margins sparsely
short-ciliate. Petals ovate, ovate- or obovate-
oblong, 2144-3 by 144 mm, obtuse, + entire.
Stamens inserted just beneath the outer margin
of the disk, usually c. 2!4 mm, in the 2 or function-
ally 2 sometimes very short (c. 24 mm) or abortive;
anthers broad-ovoid, c. 14 mm long, slightly
apiculate. Disk fleshy, rounded, 114-2 mm go.
Ovary semi-inferior, narrowed towards the apex
into a short style; stigma obscure sometimes 3-
lobed. Fruits depressed-globose, + flat or slightly
concave at the tip, 3-furrowed, c. 114 by 1144 cm,
3-celled, each cell 1- or 2-seeded. Seeds ellipsoid,
slightly, irregularly rugose, 744-8 by 4-5 mm;
aril + flat or shallow disk-like and attached
laterally at the base.

Distr. Siam (Surat, Lower Siam, and Bangta-
phan) and Malaysia: Malay Peninsula (Langkawi
Is., once in Perlis).

Ecol. On limestone at sea-level, or in lowland
forests.

Vern. Simah bater, M.

Note. Kina described the type as ‘scandent’
but no habit is recorded on the label. Two other
specimens from Langkawi Is. collected by Ha-
NIFF & Nur (SF 7085, 7529) are marked to be
scandent, but all others are defined as shrubs. The
first-mentioned may have been ‘sprawling’ shrubs.

2. Maytenus emarginata (WILLD.) DING Hou,
comb. noy.—Celastrus emarginatus WILLD. Sp.
Pl. 1, 2 (1798) 1128, ex descr., non R. & P. 1802;
R. & S. Syst. Veg. 5 (1819) 424; RotH, Nov.
Pl. Sp. (1821) 155; Roxs. Fl. Ind. ed. Wall. 2
(1824) 387; ed. Carey 1 (1832) 620.—Celastrus
montanus RoTH in R. & S. Syst. Veg. 5 (1819)
427; Nov. Pl. Sp. (1821) 154; Roxs. Fl. Ind. ed.
Wall. 2 (1824) 387 (montana); ed. Carey 1 (1832)
620; W. & A. Prod. (1834) 159.—Catha montana
G. Don, Gen. Syst. 2 (1832) 9; Hassk. Tijd. Nat.
Gesch. Phys. 10 (1843) 140; Mia. Fl. Ind. Bat.
1, 2 (1859) 589.—Cupania spinosa BLANCO, FI.
Filip. (1837) 184; ed. 2 (1845) 204; ed. 3, 2 (1878)
17.—Celastrus semiarillata Turcz. Bull. Soc.
Nat. Mosc. 36,1 (1863) 599.—Elaeodendron horizon-
tale Turcz. Ic. 603, ex descr.—Gymnosporia
montana BENTH. Fl. Austr. 1 (1863) 400, pro
comb., pro specim.?; VIDAL, Sinopsis Atl. (1883)
20, t. 31, f. A; Rev. Pl. Vasc. Filip. (1886) 88;
Merr. Bull. Bur. For. Philip. 1 (1903) 34; BACKER,
Voorl. Fl. Java (1908) 55, pro var. littoralis BAc-
KER; Schoolfl. (1911) 235; Koorp. Exk. Fl. Java
2 (1912) 524.—Gymnosporia emarginata THw.
En. Pl. Zeyl. (1864) 409; Trim. Fl. Ceyl. 1 (1893)

273; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
150.—Gymnosporia ambigua VipAL, Sinopsis
Atl. (1883) 20, t. 31, f. B——Gymnosporia spinosa
(BLANCO) Merr. & ROLFE, Philip. J. Sc. 3 (1908)
Bot. 109, incl. var. parva MERR. & ROLFE, non
(Forsk.) C. CHRISTENS. 1922; Merr. Fl. Manila
(1912) 302; Sp. Bl. (1918) 235; En. Philip. 2
(1923) 483; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 151.—Gymnosporia inermis MERR. &
Perry, J. Arn. Arb. 20 (1939) 335.—Fig. 2e.

Shrub 2-4 m. Innovations sometimes puberulous.
Spines terminating short-shoots, and/or axillary,
or unarmed. Leaves chartaceous to coriaceous,
usually obovate to subspathulate, sometimes
elliptic to elliptic-oblong, very rarely subrotun-
date, (214-)3144-14144 by (114-)2-9 cm; base
attenuate; apex obtuse or rounded, sometimes
emarginate; margin distinctly shallowly crenate
to entire; nerves (4—-)5—7 pairs; petiole 2-15 mm.
Cymes axillary, simple or fascicled at the leaf axil,
sometimes crowded at the apex of the brachyblast,
very short, rarely up to 314 cm. Peduncle very
short sometimes up to 2 cm. Bracts deltoid, short-
fimbriate. Pedicels 314-10 mm. Flowers white.
Calyx lobes deltoid, acute rarely obtuse, slightly
erose or sparsely short-laciniate, c. 1 mm long.
Petals obovate-oblong or oblong, sometimes ovate-
oblong, 2-314 by 1-114 mm, obtuse, entire. Disk
fleshy, rounded. Stamens inserted slightly beneath
the outer disk margin, 2-3 mm, sometimes small
or even abortive; anthers broadly ovoid, /4—24 mm
long, in functionally 2 fl. smaller or abortive,
obtuse or slightly apiculate. Pistil 1-114 mm above
the disk; ovary semi-immersed, 3-celled, narrowed
into a distinct or very short cylindric style;
stigmas 3, distinct, slender, reflexed, lobed, or
obscure. Fruits broadly obovoid or subglobose,
10-12 by 8-9 mm. Seeds ellipsoid, red, 3-3! by
2-234 mm, aril fleshy, white, attached at the base,
2-214 mm long.

Distr. Ceylon and SE. Asia to N. Queensland
(Cape York); in Malaysia: Malay Peninsula
(Johore), Java (throughout), Philippines (Luzon,
Lubang I., and Mindanao), Celebes(SE. Peninsula:
Kendari; Bonerate I., and Lembeh Strait), Mo-
luccas (Sulu Sanana, Taliabu, Kai Is., and Te-
nimber Is.), and New Guinea (Western Div.,
Hisiu, Daru I., Pt Moresby, and Merauke).

Ecol. In dry thickets at low altitudes, behind
the beach or at the inner mangrove.

Notes. In his key to the species of Gymnosporia,
LOESENER (1942, p. 147) divided the species into
two groups, one with a normally 3-celled ovary
and fruit and one with a 2-celled ovary and fruit.
In the latter group he recorded the African G.
senegalensis (LAMK) Logs. (= Maytenus sene-
galensis (LAMK) EXELL) as distributed from
tropical Africa through India and Malaysia to
Australia. He may have confused this African
species with the Asian-Malaysian G. diversifolia
Maxim. (= Maytenus diversifolia (MAxIM.) DING
Hou) which has also 2-celled ovaries, but which
differs by few-flowered cymes, smaller leaves,
flowers, and fruit, and a short aril situated at
the base of the seed. This error may have been

242

FLORA MALESIANA

[ser. I, vol. 6?

induced by BENTHAM’s remark (FI. Austr. 1,
1863, 400) that Celastrus montanus Roxs. ‘is
apparently the same as the tropical African
Celastrus senegalensis Lam.’ However, the latter
species has always 2-celled ovaries. As a matter of
fact the present species occurs in Australia; I
have seen a specimen from Cape York Peninsula
which exactly matches Malaysian material.

Technically Celastrus montanus RotTH and
Celastrus montanus Roxs. are different names, the
latter being a later homonym; however, WALLICH
already correlated ROXBURGH’s name in the
original publication with that of RoTH; later
authors (W. & A., BENTH., efc.) have not always
realized that RoTH’s publication preceded that of
ROXBURGH.

ROXBURGH distinguished C. montanus from C.
emarginatus by minor characters, viz larger, not
emarginate leaves which were not fascicled and
had a slightly serrate margin, its flowers in more
lush, dichotomous panicles, flowers white (instead
of yellow), three styles (instead of a style halfway
split into three branches), and a not inflated
capsule. The species is, however, very variable in
vegetative characters and the other characters
vary in degree, and I cannot attach much value to
them. The variability seems, at least in part, due
to variable environmental conditions.

In the present wide circumscription of this
common plant more names in use for Asiatic spec-
imens may fall into its synonymy, but this re-
quires an extensive study and falls outside the
scope of the present revision.

The species is distinctly variable but cannot be
subdivided. Specimens from the beach in Java and
Johore have usually both prominent short-
shoots terminating into a stout spine and axillary
spines, while those from other areas are bearing
either only distinct sometimes small axillary
spines, or are unarmed. Most specimens from the
Philippines, and some from Celebes and the
Moluccas have flowers with long stamens, a
normal ovary with a short, 3-lobed but not spread-
ing style, and obscure stigmas, while those from
New Guinea, Java, and a few from Celebes and
Philippines, have flowers which have small or
even abortive stamens, a normal ovary with a
distinct style, and 3 slender, reflexed stigmas.
ELMER 12573 collected in Luzon has both forms of
flowers mentioned aboye, i.e. the flowers on the
duplicates at Bogor and Leyden are the same as
those commonly on the specimens from the
Philippines, while those on the duplicates at
Geneva and Florence are similar to those on the
specimens from New Guinea or Java; the du-
plicates are very homogeneous, but I do not know
whether they stem from the same plant.

As to the leaf margin, there is a tendency to be-
come entire, which is distinctly so in the type of
Maytenus emarginata.

The closely allied Polynesian Maytenus vi-
tiensis (A. Gray) DING Hou, comb. nov. (Catha
vitiensis A. GRAY, Bot. Wilkes Exp. Phan. (1854)
287, t. 23) differs by ciliate calyx lobes.

According to the detailed original description,

M. rapakir Loes. (Notizbl. Berl.-Dahl. 13, 1936,
217) from the Bismarcks would be distinct by the
broadly ovate-oblong to ovate leaves and elliptic
petals.

For the reduction of Cupania spinosa BLANCO
and Gymnosporia ambigua ViDAL I have followed
MERRILL.

3. Maytenus crassa DING Hou, nom. nov.—
Gymnosporia nitida MeRR. Philip. J. Sc. 9 (1914)
Bot. 311; En. Philip. 2 (1923) 483, non Maytenus
nitida MART. 1841.

Shrub or small tree, glabrous, unarmed or with
a small, short axillary spine. Leaves coriaceous,
strongly shining above, rather dull beneath,
ovate-oblong to obovate-oblong, 4-7 by 2-4 cm;
base acute or cuneate; apex obtuse or rounded;
margin crenulate; nerves c. 8 pairs, slender,
obliquely ascending towards near the margin
and anastomosing; veins and veinlets reticulate;
petiole 5-10 mm. Flowers unknown. Infructe-
scences up to 3 cm long (ex descr.). Capsule
obovoid or ellipsoid, c. 15 mm by 8-10 mm;
rounded at apex; pericarp thick (c. 1144 mm).

Distr. Malaysia: Philippines (Luzon: Proy.
Pangasinan), once found.

Ecol. Forests at low and medium altitude.

Note. The isotype in US consists of a sterile
branch with some detached, dehisced, empty
fruits. The fruits are obovoid or ellipsoid and the
pericarp is much thicker than in any of the other
species I know. The species was described as
unarmed, but there is a very small spine in the
leaf axil.

4. Maytenus diversifolia (Maxim.) DinG Hou,
comb. nov.—Gymnosporia diversifolia MAXIM. Bull.
Ac. Imp. Sc. St. Pétersb. 27 (1882) 459; MerR.
En. Philip. 2 (1923) 483; Lingn. Sc. J. 5 (1927)
116; KANEHIRA, Formos. Trees, rev. ed. (1936)
393, f. 350; Logs. in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 149; TarpieEv, Suppl. Fl. Gén. I.—C. (1948)
799; Not. Syst. 14 (1950) 44; DinG Hou, Taiwania
1 (1950) 177; Hara, En. Sperm. Jap. 3 (1954) 93.
— Celastrus diversifolius HEMSL. in Forb. & Hemsl.
J. Linn. Soc. Bot. 23 (1886) 123; HayatTa, Ic.
Formos. Pl. 1 (1911) 139.— Gymnosporia mon-
tana var. parvifolia PirarD, Fl. Gén. I.-C. 1
(1912) 884.

Erect, sometimes scandent (?) shrub. Spines
terminating a short-shoot, and/or a small axillary
spine. Leaves oboyate-oblong, oblanceolate, or
broad-obovate, 1!14-414 by 14-2 cm; base attenu-
ate narrowed into the petiole; apex obtuse, some-
times slightly emarginate; margin sparsely den-
ticulate or crenulate, sometimes subentire; nerves
3-4; petiole very short or obscure. Cymes fas-
ciculate or solitary, sometimes 1-flowered, axillary
or crowded at the upper part of a short-shoot.
Peduncle 2-6 mm. Bracts elliptic, short-fimbriate,
c. 274 mm long. Pedicels 1-4 mm. Calyx lobes
deltoid or suborbicular, 14-24 mm g, short-
fimbriate. Petals oblong or elliptic, obtuse, 1-2
mm long. Disk cupular. Stamens c. 1% mm long,
sometimes small or abortive. Pistil flask-like, c.

Dec. 1962]

CELASTRACEAE (Ding Hou)

243

1 mm long. Ovary subglobose, narrowed into a
distinct style, 2-celled; stigmas 2, each 2-lobed;
sometimes the style very short and stigmas obscure.
Fruits compressed-obcordate, 5-6 by 5-8 mm.
Seeds ellipsoid, c. 3 by 144 mm (excluding the
aril), smooth and shining; aril at the base of the
seed, 1-2 mm long.

Distr. Siam (Soematjaj), Indo-China (Tonkin
and Annam), Ryu-Kyu, China (Fukien, Hainan,
and Formosa), and Malaysia: Philippines (Luzon)
and Lesser Sunda Is. (Bali).

Ecol. In dry thickets at low altitude.

Vern. Kum, Bali.

5. Maytenus cupularis DING Hou, nov. sp.

Arbor 12-15 m alta. Folia chartacea, elliptica,
7-11 cm longa, 4-5 cm lata, basi cuneata, apice
acuminata, margine serrulata, nervis utrinque
7-10. Inflorescentiae racemiformes, vel paniculatae,
axillares, 1¥%-2\%4 cm longae. Pedicellus c. 1% mm
longus. Flores albi. Calyx 5(-4)—lobatis, lobis late
ovatis, c. 1 mm longis. Petala 5(-4), ovata, 2. mm
longa, 14 mm lata, acuminata. Discus crassus,
cupularis. Stamina 5(-4), ad disci marginem
inserta. Ovarium basi discum late adnatum, im-
perfecte 2(-3)-loculare. Ovula in loculis 2, basalia.
Typus Brass 22124, K.—Fig. 2 f-g.

Tree 12-15 m. Branchlets terete, or slightly
angular. Leaves chartaceous, pale brown above,
greyish beneath, elliptic, 744-11 by 4-5 cm; base
cuneate; apex acuminate; margin crenulate;
midrib elevated on both surfaces; nerves 7—10
pairs, slightly elevated on both surfaces, + oblique-
ly ascending and anastomosing towards the
margins; veins and veinlets fine, loosely reticulate
on both surfaces; petiole 8-10 mm. Stipules
very small, filiform 14~—%4 mm. Inflorescences race-
mose or with a few short branches at the base,
axillary, solitary or 3 together, 1 44—21!4 cm long,

few-flowered. Peduncles very short or none.
Bracts triangular, c. 34 mm. Pedicels c. 14 mm.
Flowers white. Calyx lobes 5(—4)-lobed, lobes
broad-ovate, c. 1 mm long, slightly erose or den-
ticulate. Petals 5(-4), ovate, 244 by 14% mm,
acuminate. Disk rather fleshy, cupular. Stamens
5(-4), inserted + on the margin of the disk;
filaments filiform; anthers ovoid, c. 144 mm long,
introrse. Pistil flask-shaped, c. 1 mm above the
disk, the base confluent with the thick, cupular
disk. Ovary incompletely 2(—3)-celled; style short;
stigma slightly 2-lobed. Ovules 2 in each cell,
attached at the base, erect. Fruits unknown.

Distr. Malaysia: New Guinea (Milne Bay
District: BRAss 22124, type, K), once collected.

Ecol. Rain-forest, 30 m.

Note. The type number has been distributed as
Celastrus monospermoides Loes.; | have examined
the duplicate in the Kew Herbarium. It is an
interesting species of which the inflorescence
resembles that of certain Denhamias, but it is
also allied to some of the Australian species of
Maytenus. From Denhamia it differs by the in-
completely, usually 2-celled ovary with 2 basally
attached ovules in each cell. From M. bilocularis
(F. v. M.) Logs. it can be separated by the sparsely
crenulate leaf margin, the short pedicel (c. 144 mm,
not 3-5 mm), by the cupular (not flat) disk, and
by the ovary of which the base is confluent with
the disk but not immersed in it. Though a cupular
disk is rare in Mayrenus, its other floral characters
agree with that genus.

Excluded

Catha fasciculata TuL. Ann. Sc. Nat. IV, 8 (1857)
98 has by Ind. Kew. erroneously been credited
to hail from Malaya; it was described from
Madagascar.

3. XYLONYMUS

KALKMAN, nov. gen.—Fig. 3.

Glabrous shrub. Leaves alternate (distichous). Inflorescences cymose, solitary,
axillary, few-flowered. Flowers 4-merous. Disk flat, fleshy. Stamens 4, inserted on
the disk towards the margin. Ovary 4-celled, partly immersed in the disk. Ovules
c. 10 in each cell, arranged in 2 axial rows. Fruit a 4-angular loculicidally dehiscent
capsule, leaving a columella in the lower half; valves 4, thick and woody, com-
posed of strong radial bundles of fibres. Seeds distinctly arillate; albuminous;

embryo rather large, axile, cotyledons foliaceous.

Distr. Monotypic, in Malaysia: New Guinea.

Ecol. Primary rain-forest, at low altitude.

Notes. The generic name is a contamination of ‘evonymus’ and ‘xylos’ referring to its close affinity
with the genus Euonymus as well as indicating its characteristic woody fruit.

The floral characters of this new genus agree entirely with those of Euonymus, although the multi-
ovulate ovary-cells are not common in that genus. However, the strictly distichous leaves and the char-
acteristic woody fruit-valves have induced us to keep it as a genus separate from but closely allied to
Euonymus.

In Euonymus some species are credited with alternate leaves; in E. nana M.B. (E. Europe to China)
the leaves are very variable in phyllotaxis, on one specimen they may be strictly opposite, verticillate, or
spiral probably in 4 rows but not strictly alternate. This is probably similar in E. platyclinis OHw1 from

244 FLORA MALESIANA (ser, 1 voleez

Fig. 3. Xylonymus versteeghii KALKMAN in DinG Hou. a. Twig with fruit, 7, 6. bud in section, * 8,
c. pistil and disk, = 8, d. fruit in section, < 74, e. seed enveloped by aril, f. ditto, in section, both nat. size
(VERSTEEGH BW 4686).

Dec. 1962] CELASTRACEAE (Ding Hou) _ 245

Formosa, EF. yunnanensis FRANCH., E. decora W. W. SM., and E. linearifolia FRANCH. from China (the
latter closely allied to E. lichiangensis W. W. SM. with opposite leaves). Another Chinese species from
Szechuan, E. omeiensis FANG, has tomentose leaves which are said to be alternate but would appear

opposite from the photograph; this may be not Exonymus.
In Xylonymus, however, the phyllotaxis is strictly alternate, i.e. distichous.
in a European species of Euvonymus there is also a columella left after dehiscence, viz in E. latifolia

(L.) MILL.

1. Xylonymus versteeghii K ALKMAN, nov. sp.—Fig. 3.

Frutex glaber; folia alterna; cymae axillares
pauciflorae pedunculatae; flores hermaphroditi,
tetrameri; ovarium disco semimmersum, ovulis in
loculis circiter 10, biseriatis. Capsula 4-loculicida,
dehiscens, valvis crassis lignosis, columellata;
semina albuminosa, arillata. Typus \VERSTEEGH
4686, L, isotypes K, MAN.

Shrub up to 7 m tall. Branchlets angular, +
laterally flattened under the nodes. Stipules
present, caducous (only scars seen). Leaves
chartaceous, greyish green above, pale brownish
beneath, elliptic-lanceolate, 20-23 by 6-714 cm;
base cuneate; apex acuminate; margin entire;
midrib elevated on both surfaces; nerves 10—14
pairs, elevated on both surfaces, divaricate, or
slightly obliquely spreading towards near the
margin, archingly connected, to c. 3-5 mm from
the edge; veins loosely reticulate, slightly elevated
on both surfaces. Cymes (very young) probably
few-flowered; peduncle 10-12 mm. Rather young
flower. Sepals triangular, pale green. Petals broad-
ovate or subrotundate, 214-3 by 2!4 mm, yellow, in
the dry state dark-brown or with dark-brown pig-
ments especially at the upper 24. Disk fleshy, cush-
ion-shaped, rounded-quadrangular, c. 244 mm
across, theangles alternate withthe petals. Stamens

c.1mm long, inserted on the disk towards the margin
at the angles; filaments filiform; anthers suborbi-
cular, c. ?/; mm long, cells spreading at the lower
part but not free from the consequently triangular
connective, latrorse. Pistil partly immersed, the
free part above the disk resembling a + 4-pointed
pyramidal star; style and stigma obscure. Ovules
c. 10 in each cell, arranged in 2 rows, ascending,
apotropous-anatropous. Fruits dark red when
ripe, oblong, 614 by 3 cm, tetragonal on cross-
section, opening with 4 woody valves 7-9 mm
thick, the septa rupturing in dehiscence, leaving
a free central column; pedicel in fruit c. 1 cm.
Seeds ellipsoid, 11 by 6 mm, enveloped by a
fleshy aril except the uppermost part on one side;
testa horny; endosperm copious.

Distr. Malaysia: New Guinea (W. extremity
of the Vogelkop Peninsula, near Kalagilik between
Sorong and Klamono).

Ecol. Primary rain-forest, flat country, clay
soil, temporarily inundated, scarce, 10 m.

Note. The epithet of this new species is chosen
in honour of the collector, Mr CHR. VERSTEEGH,
at present assistant-botanist at Manokwari, who
in the course of his years in New Guinea has
collected so many new or otherwise interesting
plants.

4. EUONYMUS

TOURNEF. ex LINNE, Gen. Pl. ed. 5 (1754) 91; Sp. Pl. (1753) 197; Logs. in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 115; BLAKELOCK, Kew Bull. (1951) 232.— Evonymus’
Auct.—Pragmotessara PIERRE, Fl. For. Coch. (1894) sub t. 309.—Pragmatropa
Pierre, /.c.—Sphaerodiscus NAKAI, J. Jap. Bot. 17 (1941) 686.—Quadripterygium
TARDIEU, Bull. Soc. Bot. Fr. 95 (1948) 179; Fl. Gén. I.-C. Suppl. (1948) 809.—
Fig. 4.

Usually shrubs or small trees, rarely of moderate size (25 m), erect,
rarely scandent, glabrous, evergreen or deciduous. Leaves opposite, usually
decussate, very rarely spiral, alternate or verticillate (extra-Mal. spp.), sometimes
with dark dots underneath. Stipules lanceolate, caducous. Inflorescences axillary,
cymose, rarely flowers in fascicles (E. javanicus); pedicels articulated. Flowers
bisexual, 5- or 4-merous. Calyx deeply lobed, imbricate, entire, erose, or minute-
denticulate. Petals imbricate, spreading or reflexed, entire, erose, short-ciliate,
or long-fimbriate, smooth or finely areolate (strong lens!) on the inner surface.
Disk distinct, fleshy or thin, flat, 5- or 4-angular, or 5- or 4-lobed, or rounded,
very rarely membranous cupular (in the African E. congolensis WILCZEK), smooth,
or covered with fleshy papilla-like or subulate processes. Stamens inserted on the
disk, at the margin or halfway; anthers deltoid, -+- depressed-oblong, or broad-
ovoid, obtuse or short-apiculate, slightly free at the base, dehiscent on the top,

246 FLORA MALESIANA [ser. I} voliG2

Fig. 4. Euonymus castaneifolius RiDL. a. Twig with flowers, = 24, b. flower, petals and stamens removed,

<6, c. petal, < 6, d. young stamen, * 12, e. fruit, f. one valve, both nat. size. — E. japonicus THUNB. g.

Flower, in section, 6, h-i. petals, = 6, j. fruit, in section, seeds removed, <2 (a-d KOsTERMANS 7476,
e-f ENDERT 3159, g-i VAN STEENIS 18264, j ditto 4954).

lateral, or introrse, either 2-celled or 1-celled (extra-Mal.); filaments obscure or
distinct. Free part of the pistil terete, or slightly 4-5-angular towards the base,
gradually narrowed to the apex; the basal part sometimes covered with papillose
or subulate processes. Stigma obscure, obtuse, or discoid; ovary partly or +
wholly immersed in the disk; if the disk is angular the angles are episepalous.
Ovules mostly 2 in each cell (rarely 3-12 in each cell in a few extra-Mal. spp.),
attached to the inner angle near the base, or pendulous. Fruit a loculicidal capsule,
often coloured, when dehiscing the valves splitting and leaving no central axis
(except in the extra-Mal. sp. E. latifolia (L.) MILL.), usually (3—)4—S-angular or
-lobed, sometimes globose, rarely fusiform, smooth or echinate; apex obtuse,

Dec. 1962] CELASTRACEAE (Ding Hou) 247

acute, + truncate, or concave. Seeds (usually black) with (mostly orange) aril at
the base, or enveloped by it.

Distr. In the latest synopsis 176 specific names have been enumerated by BLAKELOCK (Kew Bull.
1951, 210-290). The genus is distributed chiefly in tropical and suo Opie Asia (especially in the Hi-
malaya, China, and Japan), 4 spp. in Europe, | sp. in Africa (Congo), 2 spp. in Madagascar, | sp. in
NE. Queensland, and 9 Spp- in North and Central America, but none in the Pacific; in Malaysia 12 spp.

Ecol. In Malaysia in primary and secondary forests from the lowland up to 3200 m.

Taxon. BLAKELOCK (/.c. 211-219, 232-238) followed Beck (FI. Nied.-Oesterr. 2, Abt. 1, 1892, 588)
in distinguishing two subgenera, viz subg. Kalonymus Beck characterized by 1-celled anthers dehiscing
with one continuous slit, winter buds usually conic, very acute, large, and capsules oblate-globose,
winged, and subg. Euonymus characterized by anthers with two cells each dehiscing with one slit, winter
buds usually ovoid, acute, and small, and capsules of various shape but if oblate-globose then not winged.
All Malaysian spp. belong to the subg. Euonymus.

This has been further subdivided into six sections comprising twelve series. I have not gone very deeply
in scrutinizing the characters on which they are based (fruit-shape, evergreen or deciduous, erc.) but the
differences between them are found ‘difficult’.

I have reduced the monotypic Indo-Chinese genus Quadripterygium poilanei TARDIEU to Euonymr
after a study of the description, plate, and an isotype specimen. It was proposed because it was assumed
that though fruits were not available they would be ‘very probably winged and indehiscent’. How-
ever, the free part of the ovary is slightly 4-angled as occurs in several species of Euonymus. Quadriptery-
gium poilane TARDiEU is closely related to E. tonkinensis Loes. and may be conspecific with it.

Note. Sterile material of this genus cannot be identified.

KEY TO THE SPECIES

1. Flowers (and fruit) 5-merous.

2. Petals usually long-fimbriate (short-fimbriate or denticulate in some specimens of E. cochinchinensis
from Celebes). Bud-scales or bracts crowded at the base of inflorescence and with reddish-brown,
long-fimbriate margins.

3. Inflorescences distinctly dichotomously (up to 5 times) lax-branched. Peduncles always distinct,
114-8 cm. Pedicels 14-4 cm. Fruits broadly-obovoid, rather small, c. 1 cm long, apex concave.

1. E. cochinchinensis

3. Flowers in almost sessile or short-peduncled, simple, axillary, 1-3-flowered cymes, sometimes
crowded or fascicled on minute knoblike brachyblasts. Peduncles usually obscure or very
short, very rarely up to 2 cm; pedicels (14—)1—2 cm. Fruits larger, usually clavate, or broad-obovoid,
rarely globose, 114-23/, cm long, the upper end conical, obtuse, or truncate, sometimes concave.

2. E. javanicus

2. Petals entire, erose, or obscurely denticulate. Bud-scales or bracts at the base of inflorescence lax,
usually entire, sometimes short-ciliate or fimbriate.

4. Leaves chartaceous to subcoriaceous; apex short-acuminate, or acuminate. Stigma obtuse or rarely

truncate.

5. Petals (when boiled) transparent with distinct, longitudinal nerves. Disk rather thin, obscurely

5-angular or suborbicular. Filaments obscure or very short. Pistil 5-angular towards the base.

6. Branchlets 4-angular. Nerves usually impressed above. Flowers 12-15 mm @ at anthesis. Fruits

(immature) obovoid, slightly lobed, + truncate at the top . . . .. 3. E. impressus

6. Branchlets terete. Nerves slightly elevated above. Flowers smaller, 6-71 5 mm @ at anthesis.
Fruits broad-obovoid, or slightly depressed-globose, deeply lobed, concave at the top.

7. Flowers rose or dull-red. Leaf margins usually entire, sometimes obscurely, remotely serrulate.

Pedicels (S—)8—14 mm. Fruit longer than wide, 18-25 by 13-17 mm... . 4. E. wrayi
7. Flowers cream-coloured. Leaf margins serrulate usually towards the upper half. Pedicels 2-4
mm. Fruit wider than long, 8-12 by 12-17 mm... > = » 5. EB. glaber

5. Longitudinal nerves on petals invisible. Calyx lobes unequal, reflexed i in anthesis. Disk orbicular,
fleshy. Filaments distinctly as long as or longer than the anthers. Pistil terete, slightly enlarged
towards the base, smooth. Fruits suboblanceolate or subfusiform .. . . 6. E. recurvans

4. Leaves usually membranous, sometimes chartaceous especially those from high altitudes; margin
dentate or crenate; apex usually long-acuminate. Stigma discoid, S-angular. 7. E. acuminifolius

1. Flowers (and fruit) 4-merous (very rarely and only occasionally a few flowers 5-merous).
8. Fruits broad-obovoid or subglobose, 4-angular, 4-lobed, smooth, or echinate, the apex round,
truncate, or concave.

9. Disk rather thin, c. 114 mmo, smooth. Stamens inserted between the pistil and the edge of the disk,
c. 4 mm long; anthers depressed oblong or deltoid, dehiscent on the top. Ovules basally attached.
Fruits broad-obovoid. Seeds with a short cup-shaped aril at the base.

10. Leaves slightly bullate by the distinctly sulcate nerves and major veins; nerves distinct. Fruits 4-

angular, slightly irregularly wrinkled outside, + truncate at the top. 8. E. castaneifolius

248

FLORA MALESIANA

[sera > volaiG=

10. Leaves not bullate by the distinctly sulcate nerves and major veins; nerves obscure above.

Fruits 4-lobed, smooth outside, concave at the top

9. E. glandulosus

9. Disk fleshy, 2-214 mm g, covered with sparse papillae, sometimes fleshy processes, or smooth.
Stamens inserted on the margin or each in an obscure marginal notch of the disk, 2-3 mm long;
anthers broad-ovoid, introrse. Ovules pendulous. Fruits + globose. Seeds completely covered by

the aril.

11. Pedicels 3-5 mm. Calyx lobes, at least the inner two, usually entire and the margin light yellowish
brown (in the herbarium); the base of the free part of pistil usually glabrous. Fruits smooth.

Erect or climbing with rootlets

10. E. japonicus

11. Pedicels 7-9 mm. Calyx lobes slightly denticulate and the margin always reddish brown (in the
herbarium); the base of the free part of pistil covered with fleshy, subulate processes. Fruits

echinate. Obviously decumbent or climbing shrub .

11. E. benguetensis

8. Fruits clavate, slightly 4-ridged and short-apiculate, smooth. Infructescence a simple cyme. Leaves

elliptic to elliptic-oblong, usually ash-coloured on both surfaces .... 12.

1. Evonymus cochinchinensis Pierre, Fl. For.
Coch. 4 (1894) t. 309A; PirarD, FI. Gén. I.-C. 1
(1/12) 873, f. 108, 6; MerR. Philip. J. Sc. 16 (1920)
450; En. Philip. 2 (1923) 480; Logs. in E. & P.
Pfil. Fam. ed. 2, 20b (1942) 119; TARDiEU, Suppl.
Fl. Gén. I.-C. (1948) 781; BLAKELOCK, Kew Bull.
(1951) 255.—E. timorensis (non Zipp. ex SPAN.)
Turcz. Bull. Soc. (Imp.) Nat. Mosc. 31, i (1858)
447; F.-Vitt. Nov. App. (1880) 46; VIDAL,
Phan. Cuming. (1885) 103.—Glyptopetalum scor-
techinii KiNG, J. As. Soc. Beng. 65, ii (1896)
345; Ripv. Fl. Mal. Pen. 1 (1922) 447.—E. philip-
pinensis MerR. Philip. J. Sc. 3 (1908) Bot. 238.—
E. viburnifolius (non Aegiphila viburnifolia Juss.)
Merr. ibid. 9 (1941) Bot. 312, pro specim.—E.
oliganthus Merr. ibid. 10 (1915) Bot. 320; En.
Philip. 2 (1923) 480.—E. pahangensis RIDL. FI.
Mal. Pen. 5 (1925) 299.—Sphaerodiscus co-
chinchinensis NAKAI J. Jap. Bot. 17 (1941) 686.
Small tree, up to 12 m by 14 cm o. Branchlets
terete, rarely angular, greenish, or reddish brown.
Leaves chartaceous to subcoriaceous, elliptic to
elliptic-oblong, sometimes obovate-oblong, 41-16
(-25) by 2!4-7(-10) cm; base cuneate, rarely ob-
tuse or rounded; apex acute, acuminate; margin
entire, sometimes remotely serrate; nerves and
veins obscure or slightly elevated on both surfaces;
petiole 3-8(—15) mm. Jnflorescences axillary, extra-
axillary, and sometimes on internodes, usually
at the basal part of the new twig, 3—10!4 cm long,
loosely, dichotomously (up to 5 times) branched;
peduncle 114-8 cm; bracts lanceolate, c. 2 mm
long, arenes Pedicel 3-5 mm. Flowers light
yellowish or greenish yellow. Calyx lobes subre-
niform, 114-244 by 2-4 mm, short-fimbriate.
Petals broad-obovate, 444 by 4 mm, fimbriate,
or rarely denticulate, areolate on the inner surface.
Disk fleshy, suborbicular or obscurely 5-angular,
c. 3 mm go. Stamens inserted between the pistil
and disk margin; filaments c. 2 mm, flat, subulate;
anthers + deltoid, c. 4/5 mm long, obtuse or
slightly apiculate. Pistil c. 2 mm above the disk,
gradually narrowed upwards, Stigma obscure.
Fruits broad-obovoid or subglobose, c. 1 cm long,
concave at the apex, deeply 5-lobed. Seeds ellip-
soid, obtuse at both ends, 5—6 by 3-4 mm.
Distr. Siam (Kaw-koh-suwan, Koh-si-kah),
Indo-China (Cambodia), China (Hainan), and
Malaysia: N. Sumatra (Atjeh), Malay Peninsula,
N. Borneo, Philippines (Batan Is., Luzon, Min-

E. moluccensis

doro, Polillo, Sulu Is., Palawan, Masbate, Min-
danao), SW. Celebes, Lesser Sunda Is. (Sumbawa),
Moluccas (Sula Is. and Kai Is.), and W. New
Guinea (also Waigeo and Aru Is.).

Ecol. Forests, from the lowland up to 1300 m.

Vern. Ampah loloh, bunga sanggara, Celebes;
Philip.: alumangug, P. Bis., baras-baras, Ilk.,
burubatuan, Tag., kéyum-bakiu, Luzon.

Notes. The fruits of MERRILL 9644 (type of
E. oligantha) is broad-obovoid and not sulcate,
which is obviously due to insects. Another
Philippine specimen (BS 76829, SING), in which
the rather young fruits are also not sulcate, is also
damaged by insects. Normally young fruits are
always sulcate or lobed.

2. Euonymus javanicus BL. Bijdr. (1827) 1146;
Benn. & R. Br. Pl. Jav. Rar. (1840) 130, t. 28;
Hassk. Tijd. Nat. Gesch. & Phys. 10 (1843) 140;
Cat. Hort. Bog. (1844) 229; Pl. Jav. Rar. (1848)
230, incl. var. sphaerocarpus Hassk.; Miq. FI.
Ind. Bat. 1, 2 (1859) 588, incl. var. sphaerocarpus;
SCHEFFER, Nat. Tijd. Ned. Ind. 34 (1870) 98;
Kurz, J. As. Soc. Beng. 39, ii (1870) 73; Laws.
in Hook. f. Fl. Br. Ind. 1 (1875) 607; Kurz,
J. As. Soc. Beng. 45, ii (1876) 123; For. FI.
Burma 1 (1877) 249; F.-ViLL. Nov. App. (1880)
46; VIDAL, Sinopsis (1883) 20, t. 31, f. C; Rev.
Pl. Vasc. Filip. (1886) 87; Pierre, Fl. For. Coch.
4 (1894) t. 308C, pro var. talungensis PIERRE;
Kina, J. As. Soc. Beng. 65, ii (1896) 339; Koorp.
Minah. (1898) 396; K. & V. Bijdr. 7 (1900)
89-90, incl. var. genuina et sphaerocarpus; MERR.
Philip. J. Sc. 2 (1907) Bot. 278; BAcK. Schoolfl.
(1911) 233, incl. var. sphaerocarpus et horsfieldii,
et f. genuinus; Koorp. Exk. Fl. Java 2 (1912)
523; Koorp.-ScHuM. Syst. Verz. 1, Fam. 158
(1912) 1; Pirarp, Fl. Gén. I.-C. 1 (1912) 869;
K. & V. Atlas 1 (1913) t. 139; Gipps, Arfak
(1917) 214; Ript. Fl. Mal. Pen. 1 (1922) 445;
Merr. En. Philip. 2 (1923) 480; Crais, Fl. Siam.
En. 1 (1926) 279, incl. var. talungensis; TARDIEU,
Suppl. Fl. Gén. I.-C. (1948) 788, f. 95, 4-5,
pro var. talungensis; BLAKELOCK, Kew Bull.
(1951) 257, incl. var. genuinus, sphaerocarpus,
timorensis, elmeri et coriaceus.—E. timorensis
Zipp. ex SPANOGHE, Linnaea 15 (1841) 186;
Koorp. Minah. (1898) 396. —E.sumatranus
Mia. FI. Ind. Bat. 1, 2 (1859) 589; Sum. (1861)
512; BLAKELOCK, Kew Bull. (1951) 257.—E.

Dec. 1962]

CELASTRACEAE (Ding Hou)

249

bancanus Mig. Sum. (1861) 513; Kurz, Nat.
Tijd. Ned. Ind. 27 (1864) 194.—E. horsfieldii
Turcz. Bull. Soc. (Imp.) Nat. Mosc. 36, 1 (1863)
598, ex descr., photogr. of type seen!; K. & V.
Bijdr. 7 (1900) 90.—E. alatus EvM. Leafl. Philip.
Bot. 4 (1912) 1484, non (THUNB.) SIEBOLD 1830.
—E. elmeri MeRR. Philip. J. Sc. 12 (1917) 281,
new name for E. alatus ELm.; En. Philip. 2 (1923)
480.—E. coriaceus Riv. Fl. Mal. Pen. 5 (1925)
299.—E. micropetalus Ripv. /.c.; BLAKELOCK,
Kew Bull. (1951) 258.

Tree up to 23 m by 53 cm @, sometimes shrub.
Branchlets terete. Leaves chartaceous to sub-
coriaceous, elliptic to elliptic-lanceolate, some-
times broad-elliptic, obovate, or obovate-oblong,
5-20 by 2-9 cm; apex acute to acuminate; base
cuneate, rounded, or attenuate; margins entire
rarely crenate in the upper half; nerves 4~7 pairs,
obliquely arcuate, ascending and loosely anasto-
mosing near the margin; petiole 5—8 mm, furrowed
above. Flowers 1-co on a condensed, short, tuber-
cular axillary and extra-axillary glomerule, very
rarely in a simple cyme with a peduncle (74-2 cm).
Bracts lanceolate, fimbriate, c. 1 mm _ long.
Pedicels (14—)1—2 cm, articulated at the base.
Flowers 5-merous. Calyx brown to reddish, lobes
unequal in size, suborbicular, reniform, or ovate,
1-314(-5) by 114-4(-6) mm, slightly concave,
minutely denticulate. Pefals light green or yellow-
ish, broadly obovate or suborbicular, 414-5 by
314-614 mm, areolate, fimbriate. Disk fleshy,
214-3 mm @g, 5-angular, flat or slightly concave,
sometimes slightly swollen at the base of the
filaments. Stamens inserted on the margin of the
disk or quite near to it, c. 244 mm; anthers
triangular, divaricate, 4/5-1 mm long and wide,
obtuse or short-apiculate, the base reddish to
darkish brown (in herbarium). Pistil conical,
emerging 114-2 mm from the disk, smooth or
obscurely 5-angular towards the base, gradually
narrowed into a short style; stigma obtuse. Ovules
inserted near the base. Fruits red, usually clavate
or broadly obovoid, rarely globose, the upper end
conical, obtuse, truncate, or concave, bearing an
apiculate apex, 5(—4)-celled, one or two seeds in
each cell.

Distr. India (Andamans; Nicobars, _ sec.
Kurz, 1876), Burma (Tenasserim), Siam (Tassan
Champhon and Huey Mut), Indo-China (Co-
chinchina), and Malaysia: Sumatra (also Simalur,
Batu, and Banka Is.), Malay Peninsula (also
Penang I.), SE. Borneo (Téwe R.) and Natuna
Is., Java (common), Lesser Sunda Is. (Bali,
Sumba, Timor, and Wetar), Celebes (also
Salajar I.), Philippines, Moluccas (Talaud and
Halmaheira), and New Guinea (Manokwari; also
in Misool, Sorong, and Radjah Ampat Is.).
Fig. 5.

Ecol. Primary dryland rain-forests, from the
lowland up to 1500 m.

Vern. Sumatra: awa kudang kudang, bientol
pajo, bintol eiafai, kudang kudang balah, lamélul,
simani dotan, tutun bintol lantja, Simalur, kaju
kumbang kétjil, kumbang, Palembang, kienjiens,
Banka; Mal. Pen.: bélimbing hutan, kémuning

a = ed

Fig. 5. Distribution of Euonymus javanicus BL.

ayer, k. gading, logan, Temuan, bélungkas, Perak,
gading, Pahang; Java: atiati, E. Java, djambon,
djirak, pajangan, ragen, sapén, Central Java,
képiting, M, kakatjangan, ki keujeup, kihapit
lalaki, ki-katjang, S; Lesser Sunda Is.: aadakurang,
aitada kuru, Sumba; Natuna Is.: gunu; Philip.:
kalimétas, malasangki, surag, tubatubhan, Tag.,
tabdan, Mbo., talangutifigon, C. Bis.; Celebes:
sandu molaba, tadugheho, Tobela; Moluccas:
ligisata’a, Talaud I., radja, Weda.

Notes. The fruits of this species are rather
variable in shape and size. They are commonly
clavate or broad-obovoid, sometimes being
globose; intermediate forms also occur. Several
varieties have been described based chiefly on the
shapes and sizes of fruits. From the many fruiting
specimens examined it is very difficult to distin-
guish well defined varieties.

Under E. javanicus, HAssKARL (Tijd. Nat.
Gesch. Phys. 10, 1843, 140; Cat. Hort. Bog.
1844, 229) named two varieties, viz var. conocarpus
and var. sphaerocarpus, but he provided no de-
scriptions. In 1848 (Pl. Jav. Rar. 229-230) he gave
a rather detailed description of E. javanicus and
of its var. sphaerocarpus, but did not mention
var. conocarpus; evidently he treated the latter as
the type variety. The fruit of E. javanicus he defined
as ‘ob-pyramidal, prominently angular, with co-
nical, acute apex’. In var. sphaerocarpus the fruit
is ‘obconical-subglobose when young and pro-
minently S-angular at the base and rounded or
mucronate-acute at the apex at maturity’. Un-
fortunately I could not locate the type specimens
of these varieties.

Koorpers & VALETON (1900) interpreted var.
conocarpus HAssk. in the sense of the type variety
and named it var. genuinus. They also accepted
var. sphaerocarpus Hassk. and amplified its de-
scription by including specimens with obcordate
fruits, which was also accepted by BACKER (1911)
and BLAKELOCK (1951).

Lawson (/.c. 610) identified two specimens,
collected by HELFER in Tenasserim or the An-
daman Is. and by Kurz in Pegu respectively, as
*E. timorensis Zirp.’. In the description he stated:

250

FLORA MALESIANA

[ser. I, vol. 6?

‘cymes as long or nearly as long as the leaves’;
these two specimens evidently do not belong to the
present species.

3. Euonymus impressus BLAKELOCK, Kew Bull.
(1951) 256, f. 5, left.

Small tree up to 6 m. Branchlets 4-angular.
Leaves chartaceous to coriaceous, usually bullate
(i.e. nerves and some veins depressed above),
elliptic to narrow-elliptic, sometimes ovate,
7-12 by 214-4 cm; base cuneate; apex acuminate;
margin remotely crenate-serrate usually at the
upper half; nerves 5—6 pairs, arching c. 4% cm
near the leaf margin; veins obscure on both
surfaces; petiole 3-5 mm. Cymes up to 44% cm
long, very slender and lax, few-flowered, axillary
or on a very short bracteate brachyblast, or extra-
axillary. Peduncle very fine, 2-21 cm, sometimes
very short. Bracts lanceolate, c. 1 mm _ long,
sparsely short ciliate. Pedicels 8-12 mm, very
fine. Flowers 5-merous. Calyx lobes semi-orbicular
or reniform, 34-1 by 114-3 mm, slightly concave,
transparent, irregularly denticulate. Peta/s sub-
orbicular or broad-obovate, unequal in size,
3-6 mm 49, slightly contracted at the base, ir-
regularly denticulate, distinctly veined, areolate
on the inner surface. Disk suborbicular, rather
thin, c. 3 mm @. Stamens inserted + halfway on
the disk; filaments very short; anthers c. 4. mm
long. Pistil conical, c. 1 mm above the disk,
slightly 5-angular; stigma obscure. Fruits (im-
mature) obovoid, slightly lobed, + truncate at
the top.

Distr. Malaysia:
once collected.

Ecol. Rain-forest, along stream, 1100 m.

Celebes (SW. Peninsula),

4. Euvonymus wrayi KiNG, J. As. Soc. Beng. 65,
ii (1896) 344, excl. RipLEY 2652; PRAIN, ibid.
73, 11 (1904) 194; RipL. J. Fed. Mal. St. Mus. 4
(1909) 11; Fl. Mal. Pen. 1 (1922) 446; BLAKE-
Lock, Kew Bull. (1951) 254.—E. rufulus R1DL.
J. Str. Br. R. As. Soc. m. 75 (1917) 19; Fl. Mal.
Pen. 1 (1922) 446; BLAKELOCK, Kew Bull. (1951)
255.

Shrub or small tree, up to 5 m. Branchlets
terete. Leaves chartaceous to subcoriaceous, rarely
coriaceous, oOvate-oblong, elliptic to elliptic-
oblong, sometimes obovate to obovate-oblong,
744-15 by 214-7 cm; base cuneate to narrow-
cuneate; apex acuminate; margins entire, some-
times obscurely, remotely serrulate; nerves 7—9
pairs, towards the margin loosely arching-
anastomosing; petiole 5-8 mm. Inflorescences
cymose, axillary, at the base of the new twigs, or
crowded on the undeveloped shoot in the leaf-
axil, 1-714 cm long, sometimes very short.
Peduncle very short, sometimes up to c. 5 cm.
Bracts lanceolate, 4-214 mm long, fimbriate.
Pedicels (S—)8—-14 mm. Flowers p'nk or dull red,
5-merous. Ca/yx lobes unequal in size, subreniform
or suborbicular, 1-2 by 1-214 mm, slightly erose.
Petals broad-obovate, broad-elliptic or suborbicu-
lar, 3-4 by 214-3 mm, irregularly minutely den-
ticulate, areolate. Disk 114-2 mm g, + round, or

obscurely 5-lobed. Stamens inserted on the disk
between its margin and pistil; anthers + deltoid;
filament short. Pistil very short, sometimes emerg-
ing c. 14 mm from the disk, S-angular at base.
Fruits broadly obovoid, 17-25 by 13-17 mm,
concave at the apex, distinctly 5-lobed, gradually
narrowed towards the base. Seeds ellipsoid, 13
by 6 mm, reddish- or dark-brown.

Distr. Malaysia: Sumatra (Atjeh and Gajo-
lands) and Malay Peninsula (Perak, Kedah,
Trengganu, Pahang, and Selangor).

Ecol. In forests, usually between 1300-2500 m,
rarely in the lowland.

Vern. Ségading bukit, M.

Note. Very close to E. javanicus from which
it can only be distinguished by laxer, more branch-
ed and longer peduncled cymes and minutely,
irregularly denticulate petals. Leaves and fruits
are exactly matching those of E. javanicus. Bracts
of axillary buds are usually rusty fimbriate.

5. Euonymus glaber Roxs. [Hort. Beng. (1814)
86, nomen] FI. Ind. ed. Wall. 2 (1824) 403;
ibid. ed. Carey 1 (1832) 628; LAws. in Hook. f.
Fl. Br. Ind. 1 (1875) 609; BLAKELOCK, Kew Bull.
(1951) 254.—Lophopetalum pedunculatum RIDL.
J. Str. Br. R. As. Soc. n. 59 (1911) 85; Fl. Mal.
Pen. | (1922) 449.

Small tree up to 5 m. Branchlets terete. Leaves
chartaceous, elliptic, sometimes elliptic-oblong,
7-8 by 3-41, cm; base narrow-cuneate; apex
acute to short-acuminate; margins usually dentate-
crenate in the upper half; nerves S—6 pairs, rather
fine; petiole 5-7 mm. Cymes axillary or slightly
extra-axillary, usually at the base of a new shoot,
up to 4144 cm long, sometimes on a leafy axillary
brachyblast. Peduncles up to 2! cm. Bracts ovate,
short-ciliate. Pedicels 2-314 mm. Flowers cream-
coloured, S-merous, some flowers occasionally
4-merous. Ca/yx lobes unequal, usually the outer
two smaller, suborbicular or broadly-obovate,
rarely subreniform, slightly erose, sometimes short-
ciliate. Perfals suborbicular, sometimes broad-
obovate, 2-214 by 2-214 mm, sometimes slightly
contracted at the base, the margin usually min-
utely denticulate. Disk ++ orbicular, c. 144 mmo.
Stamens very short, inserted near the margin of
the disk; anthers + triangular. Pistil c. 74 mm
emerging from the disk, 5(-4)-angular at the base,
narrowed into a short, cylindric style. Fruits
slightly depressed-globose, 8-12 by 12-17 mm,
deeply 5(—4)-lobed, usually concave at the apex,
narrowed towards the base. Seeds usually only 1
in each cell.

Distr. E. Pakistan (Chittagong), Siam, and
Malaysia: Malay Peninsula (Perlis), twice col-
lected.

Ecol. On hill slope.

6. Euonymus recurvans Mig. Sum. (1861) 513;
BLAKELOCK, Kew Bull. (1951) 262.

Branchlets terete. Leaves chartaceous, light
brown to brown beneath, elliptic, 7-8 by 314-4
cm; base cuneate rarely obtuse; apex short-
acuminate; margin usually crenate at the upper

Dec. 1962]

CELASTRACEAE (Ding Hou)

251

half, slightly recurved; nerves 4~7 pairs, obliquely
ascending towards near the margin and curving
upwards; petiole 5-8 mm. Cymes axillary or
extra-axillary near the base of a new twig, simple
or once forked. Peduncle very short, sometimes
up to 18 mm. Bracts deltoid, fimbriate. Pedicels
4-5 mm. Flowers 5-merous. Calyx lobes unequal,
the inner two much larger, semi-orbicular to
reniform, %—2 by 1-2!4 mm, slightly erose,
reflexed at anthesis. Petals (from floral bud)
suborbicular, 2 mm 9g, short-ciliate. Disk fleshy,
orbicular, c. 2/4 mm 9g. Stamens inserted on the
disk about halfway the edge and pistil; anthers
-+ triangular, free at lower half, slightly apiculate;
filaments c. 34 mm. Pistil c. 2 mm emerging from
the disk. Ovules attached at the base. Fruits
(young) suboblanceolate or subfusiform, 11 by
Vy cm.

Distr. Malaysia:
Singalang; Priaman).

Sumatra (Padang: Mt

7. Euonymus acuminifolius BLAKELOCK, Kew Bull.
(1951) 253, f. 5, right, incl. var. borneensis BLAKE-
LOCK.

Shrub up to c. 314 m. Branchlets 4-angular or
subterete. Leaves membranous, sometimes char-
taceous (especially at high altitude), ovate-oblong
to lanceolate, or elliptic to elliptic-oblong,
7144-1014 by 2-4 cm; base cuneate sometimes
obtuse; apex usually long-acuminate (acumen up
to 2 cm); margin crenate-serrulate or serrulate;
nerves 4-6 pairs, spreading towards 3-5 mm from
the edge and then arching; petiole 2-6 mm.
Cymes 1-3 times branched, axillary or extra-
axillary, or crowded on axillary brachyblasts,
very short to sometimes up to 8 cm long, few-
flowered. Peduncle short, sometimes up to 4 cm.
Bracts triangular, 1-2 mm long, entire or some-
times short-ciliate. Flowers purplish-red, 5-merous.
Calyx lobes semi-orbicular or subreniform, 1-11
mm long, the inner two usually larger, irregularly
minute-denticulate. Peta/s suborbicular, 21-3
by 2!4 mm, minute-denticulate, slightly contracted
at the base. Disk obscurely S-angular, c. 2 mm @.
Stamens small; filaments very short; anthers c.
14 mm long. Pistil short, emerging c. 3 mm from
the disk, slightly 5-angular; stigma discoid, ob-
scurely 5-angular. Fruits obcordate, distinctly
5-lobed, 11-2 by 114-134 cm. Seeds (very young)
with aril at the base.

Distr. Malaysia: Central West Sumatra
(Ophir Distr.), N. Borneo (Mt Kinabalu) and
SW. Celebes (Enrekang).

Ecol. Forests, from 700-3200 m.

8. Euonymus castaneifolius RIDL. Kew Bull. (1931)
36; BLAKELOCK, ibid. (1951) 255.—E. moultoni
RipL. jhid. (1931) 36; BLAKELOCK, ibid. (1951)
256.—Fig. 4a-f.

Tree, 5 up to 15 m by 20 cm g. Branchlets
terete. Leaves chartaceous to subcoriaceous,
elliptic to elliptic-oblong, sometimes ovate-oblong,
rarely broad-elliptic, 514-1314 by 214-714 cm,
sometimes with scattered, black dots beneath; base
acute to cuneate, sometimes attenuate; apex

short-acuminate to acuminate, rarely obtuse;
margin slightly cartilaginous, usually remotely
serrulate at the upper half, slightly recurved;
nerves 5-9 pairs, strongly prominent beneath,
depressed above, sometimes flat above (especially
specimens from higher altitude), obliquely as-
cending near the margin and then curving upward;
veins obscure sometimes visible on both surfaces;
petiole 12-15 mm. Cymes 1-4 times branched,
axillary, extra-axillary, solitary or sometimes
fasciculately crowded on an axillary brachyblast,
1-3(-7) cm long, usually with several small,
fimbriate bracts at the base, sometimes with
subulate ones up to 6 mm. Peduncle very short,
sometimes 114-5 cm. Pedicels 3-6 mm, articulated
at the base. Flowers white, 4-merous. Calyx lobes
suborbicular, 1-114 mm 9, the inner pair largest.
Petals elliptic, or broad-elliptic, 2-24%4 by 14-2
mm, obtuse and erose, with a few longitudinal
veins. Disk slightly 4-angular. Stamens inserted
on the disk between its edge and the pistil; anthers
slightly depressed-oblong and contracted at the
base, sometimes deltoid, dehiscing at the top.
Pistil emerging c. 34 mm from the disk, 4-angular
towards the base. Fruits subobovoid, obscurely
4-angular, red when mature, 10-18 by 7-11 mm,
flat or slightly concave sometimes slightly obtuse
at the top, slightly irregularly wrinkled when dry.
Seeds ellipsoid, 12 by 10 mm, irregularly wrinkled,
with a cup-shaped aril at the base.

Distr. Malaysia: Central Sumatra (Pajakum-
buh) and Borneo (Mt Kinabalu, Balikpapan,
Kutai and Sarawak).

Ecol. Primary forests, sandy ridges, mossy
forest, from low ridges up to 1500 m.

Note. The dark dots on the underside of the
leaves are not caused by a fungus; they appear to
be resinous (?), but the coloured substance is not
soluble in alcohol. The only other species in which
they also occur is the following one.

9. Euonymus glandulosus (Merr.) DING Hou,
noy. comb.—Glyptopetalum glandulosum MEeERR.
Philip. J. Sc. 12 (1917) Bot. 279; En. Philip. 2 (1923)
481.

Shrub or small tree 3 to 5 m. Branchlets terete,
sometimes sulcate or slightly 4-angular. Leaves
chartaceous to subcoriaceous, always with scatter-
ed black dots beneath, elliptic to elliptic-lanceolate,
rarely ovate, or obovate-oblong, 414-1214(-1414)
by 2-414(-5!4) cm; base cuneate; apex acuminate;
margin slightly recurved, serrulate at the upper
half; nerves 4-5 pairs, slightly elevated below,
visible or obscure, sometimes depressed above,
sometimes obscure on both surfaces, spreading
towards near the margin, arching; veins usually
obscure on both surfaces; petiole 5-8 mm.
Cymes 2-4 times branched, sometimes a triad or
even a solitary flower, axillary, or towards the
base of the new twigs, short, sometimes up to
314 cm, few-flowered. Peduncle short, sometimes
up to 244 cm. Pedicels 4-6(-8) mm. Flowers
dark purple, 4-merous. Ca/yx lobes subreniform
or suborbicular, 34-1 by I-14 mm, slightly
erose, inner lobes slightly larger, thinner, and

252

FLORA MALESIANA

[ser. I, vol. 6?

transparent. Petals suborbicular or broad-obovate,
24%,-3 by 2-3 mm, slightly erose, with a few,
longitudinal veins. Disk obscurely 4-angular, c.
14%, mm g. Stamens small; anthers slightly de-
pressed-oblong, 14 by 4/5 mm, dehiscing at the
top; filaments very short, inserted between the
pistil and the edge of the disk. Pistil c. 4% mm
emerging from the disk, slightly 4-angular towards
the base. Fruits broad-obovoid, 18 by 15 mm,
4-lobed, concave at the apex, cuneate at the base,
smooth, ochre-yellow suffused with rose-red, finally
red. Seeds ellipsoid, smooth, 8 by 5 mm, aril at
the base disk-like.

Distr. Malaysia: Borneo (Mt Kinabalu) and
Philippines (Mindanao and Palawan).

Ecol. In forests, 900-2700 m.

Note. This species has been described under
Glyptopetalum because MERRILL found only one
ovule in each cell of the ovary; in the duplicates of
the type (MERRILL 9547, Bo, K, L), however,
the ovary distinctly bears two ovules in each cell.
The black dots on the leaves are similar to those
in the foregoing species.

10. Euonymus japonicus THuNB. Nov. Act. R.
Soc. Sc. Upsal. 3 (1780) 198 & 208; Fl. Jap.
(1784) 100; BANkKs, Ic. Kaempf. (1791) t. 8;
Biume, Bijdr. (1827) 1147; LinpL. Bot. Reg.
n.s. 17 (1844) t. 6; Mie. Ann. Mus. Bot. Lugd.
Bat. 2 (1865) 86, incl. var. radicans M1Q.; FRANCH.
& SAVAT. En. Pl. Jap. 1 (1875) 79; Maxim. Bull.
Ac. Imp. Sc. St. Pétersb. 27 (1882) 441; Fors. &
Hems-. J. Linn. Soc. Bot. 23 (1886) 120; Koorp.
Minah. (1898) 395; Logs. Bot. Jahrb. 29 (1900)
441; K. & V. Bijdr. 7 (1900) 85; Logs. Bot.
Jahrb. 30 (1902) 453; Back. Schoolfl. (1911)
234; Koorpb. Exk. Fl. Java 2 (1912) 523; Logs. &
ReHD. in Sargent, Pl. Wilson. 1 (1913) 485;
STEEN. Trop. Natuur 22 (1933) 175, fig.; MERR.
Sunyatsenia 1 (1934) 197; WANG, Chinese J. Bot.
1 (1936) 50; RoepKe, Trop. Natuur 25, Jub. no
(1936) 20, f. 2-3; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 118; STEEN. Bull. Jard. Bot.
Btzg III, 17 (1948) 400; BLAKELock, Kew Bull.
(1951) 268; HARA, En. Sperm. Jap. 3 (1954) 86.
—E. japonicus L. f. Suppl. (1781) 154; Jue, Pl.
Thunberg. (1918) 176.—Elaeodendron fortunei
Turcz. Bull. Soc. Nat. Mosc. 36, i (1863) 603;
Wap. Ann. 7 (1868) 582; Maxim. Bull. Ac.
Imp. Sc. St. Pétersb. 27 (1882) 460; Fors. &
Hems_. J. Linn. Soc. Bot. 23 (1886) 124.—
Elaeodendron javanicum Turcz. Bull. Soc. Nat.
Mose. 36, 1 (1863) 602.—E. radicans (MiQ.)
Sigs. ex Mia. Ann. Mus. Bot. Lugd. Bat. 3 (1867)
202; HAND.-MaAzz. Symb. Sin. 7 (1933) 660.—
Cassine fortunei O.K. Rey. Gen. Pl. 1 (1891) 114.
—E. fortunei HAND.-MAzz. Symb. Sin. 7 (1933)
660; Reup. J. Arn. Arb. 19 (1938) 75, t. 218, incl.
var. radicans (MiQ.) REHD.; Man. Cult. Trees &
Shrubs ed. 2 (1940) 559; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 118; BLAKELOcK, Kew Bull.
(1951) 268; Hara, En. Sperm. Jap. 3 (1954) 84.
—Pragmotessara japonica PieRRE, Fl. For. Coch.
(1894) sub t. 309.—E. fungosus OHw1, Acta Phyto-
tax. Geobot. 5 (1936) 186, ex descr.—Fig. 4g-j.

Erect, procumbent, or scandent shrub, or small
tree, up to 8 m. Branchlets terete, if procumbent
or scandent bearing rootlets. Leaves chartaceous,
thin-coriaceous to coriaceous, ovate or broad-
ovate, obovate or obovate-oblong, elliptic to
elliptic-oblong, rarely broad-elliptic, 2-914 by
1144-4 cm; base obtuse, or acute; apex acute or
obtuse, rarely acuminate; margin dentate-crenate;
nerves 2-6 pairs, rather fine, obliquely spreading
and loosely anastomosing near the margin;
petiole 2-13 mm. Cymes axillary and/or extra-
axillary, 1-12 cm long. Peduncle 14-8 cm. Bracts
triangular, lanceolate, 14-4 mm long, caducous.
Pedicels 3—5 mm. Flowers greenish white, 4-merous,
c. 8 mm g. Calyx lobes reniform or suborbicular,
1-114 mm long, entire or slightly erose, the outer
pair smallest. Petals subrotund, broad-ovate or
-elliptic, 314-324 by 214-314 mm, entire, some-
times erose, slightly recurved. Disk fleshy, flat
or slightly concave, slightly 4-angular or subround-
ed, usually covered with sparse papillae, sometimes
with fleshy processes especially towards the margin,
or smooth (extra-Mal. material). Stamens inserted
on the margin, sometimes in shallow notches,
c. 2Y%4 mm long; anthers broad-ovoid, 74-1 mm
long, slightly apiculate, rarely obtuse. Pistil 1-214
mm above the disk, sometimes slightly 4-angular
towards the base; style distinct, cylindric; stigma
obscure. Fruits -+- globose, smooth, green, c. 1
cm 9; in dehiscing the endocarp gets loose from the
exocarp and on both sides of each cell it has
a thickened corky-cartilaginous portion, white
and crescent-shaped in cross-section; during the
shrinkage of the more or less fleshy exocarp the
seed, pendulous from the top of the central axis, is
pushed out of the cell.

Distr. Japan, Korea, China, Ryukyu, and
Malaysia: West Central Sumatra (Kerintji),
Java (from Mt Papandajan eastwards), Lesser

Fig. 6. Distribution of Euonymus japonicus THUNB.
in Malaysia.

Dec. 1962]

CELASTRACEAE (Ding Hou)

ae

Sunda Is. (Sumbawa, Port. Timor), SW. and NE.
Celebes, Philippines (Luzon: Proy. Laguna on
Mt Banahao). Fig. 6.

Ecol. On edges of rain-forests, on ridges,
(600—)1000-—2950 m.

Uses. Roepke (1936, /.c.) found it used for
hedges in Central Java where it was the host of
an endemic butterfly. In temperate countries
frequently used as an ornamental. For the various
varieties distinguished see REHDER, J. Arn. Arb.
19 (1938) 75-80; Man. Cult. Trees. & Shrubs
ed. 2 (1940) 558-559; Bibl. Cult. Trees (1949)
405-411; BLAKELOCK, Kew Bull. (1951) 262-270;
Hara, En. Sperm. Jap. 3 (1954) 84-88.

Notes. THUNBERG based this species on J/so
Kuroggi described in KAEMPFER’s Amoen. Exot.
fasc. 5, p. 790, giving it the short diagnosis ‘flo-
ribus quadrifidis; foliis ovatis, serratis (1780). In
1784 he copied this brief diagnosis, citing KAEMP-
FER’s work, and adding a detailed description
(Fl. Jap. 100), but omitting his evaluation of 1870.
E. japonicus L. f. of 1781 is typified by THUN-
BERG’S specimens and is technically a later ho-
monym.

Elaeodendron javanicum Turcz. was based on
ZOLLINGER 2958 from Java and has curiously been
omitted in Javanese floras. Prof. ZEROv, Kiew,
kindly sent a photograph of the type which
exactly matches EF. fortunei (TURCZ.) HAND.-
MaAzz.; later an isotype was found in the Geneva
Herbarium.

This species is extremely variable, which is
partly due to its large range and partly to the
existence of many cultivars. However, its fruit
type is very characteristic as stated in the de-
scription.

There are two main groups of forms. The first,
including the typical form, is erect with obovate,
obtuse, rather coarsely crenate, glossy, dark-green
leaves. It occurs chiefly in Japan. The second
form is usually procumbent or climbing by
small rootlets produced by the stem and its leaves
are smaller, rather elliptic, acute, serrulate, dull,
and pale green. This grows chiefly in China and
was described as Elaeodendron fortunei TURCZ.
= Euonymus fortunei (TURCZ.) HAND.-MAZz..
REHDER, BLAKELOCK, and Hara still distinguish
this as a species distinct from £. japonicus,
but the differences are mainly vegetative and
because there are many intermediate specimens,
I fail to see how these two species can be main-
tained.

The root-climbing habit is not at all important
as a character. VAN STEENIS studied this in detail
in the Javanese mountains (1933, /.c.): in open
habitats the plants are perfectly erect, but in the
adjacent shaded mountain forest the seedling
grows up as a very thin, hardly branched, many
metres long root-climber clinging to often mossy
tree trunks. Under these shady conditions it
flowers seldom, but if such an inconspicuous plant
becomes exposed, as he found on a scree near the
summit of Mt Jang (E. Java), it starts to branch
profusely with abundance of flowers and fruit.
In 1953 he found both forms erect and climbing,

both flowering and fruiting, together on the
summit of Mt Perdido, Port. Timor. The climbing
form may grow into a fairly thick woody liana,
with the stem a finger thick. The use of climbing
versus erect made by e.g. REHDER (Man. Cult.
Trees & Shrubs ed. 2, 1940, 554) and BLAKE-
LOCK in their keys seems to be futile.

All Malaysian specimens, whether erect or climb-
ing, differ in another slight character from those of
China and Japan — except one specimen found in
Hupei, WiLson 502, K — in that the disk in the
Malaysian specimens is usually covered with
scattered papillae or very rarely somewhat larger
fleshy processes, the disk being smooth in the
Japanese and Chinese specimens. Under the
microscope both papillae and processes appear
to be minute emergencies, i.e. outgrowths of the
epidermal and subepidermal cell layers; the pro-
cesses are occasionally branched and sometimes
even bear a stoma at the apex (cf. also BERKELEY,
J. Elisha Mitchell Sc. Soc. 69, 1953, 191). This
is most significant, as these processes are essenti-
ally the same as those found in other species to
grow out into spines. Of course the occurrence of
spines renders the fruit an extremely showy
character and the echinate-fruited species have
even been arranged in a special section by NAKAI
(J. Jap. Bot. 17, 1941, 617) and accepted by
BLAKELOCK. However, both the spiny and
spineless fruits are in other plant groups some-
times found in a single species, for example in
species of Datura, Galium (hooked hairs), Den-
tella (emergencies), efc. Besides emergencies
occur on the disk of certain strains of E. japonicus
and are clearly visible at the base of the fruit
(cf. E. fungosus = E. japonicus and FB 7878, US).

In the light of the observations made above,
however, it is possible that the significance of the
occurrence of spines has been much overevaluated.
It is not impossible that spiny fruits and unarmed
ones may occur in the same species; in this way
one would get two series of parallel ‘pseudospecies’.
Therefore we should avoid keying out species
on this single character; in absence of other
additional clear differences it cannot serve for
specific distinction.

11. Euonymus benguetensis Merr. Publ. Gov.
Lab. Philip. 29 (1905) 26; En. Philip. 2 (1923)
480; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
119; BLAKELOCK, Kew Bull. (1951) 274; Jacoss,
Ann. Bog. 3 (1958) 65, as a Euonymus.

A scandent or decumbent shrub. Branchlets
light greenish, terete or slightly angular, usually
warty, occasionally bearing rootlets (ELMER 8729,
L). Leaves membranous to chartaceous, ovate,
elliptic, sometimes broad-ovate or -elliptic, rarely
obovate, 3-714 by 1'4-4 cm; base acute or
cuneate, sometimes obtuse; apex acute to acu-
minate, rarely obtuse; margins serrulate or cre-
nulate, sometimes entire at the lower part; nerves
3-5 pairs; petiole 2-3 mm. Jnflorescences axillary,
sometimes extra-axillary, 2-41 cm long. Peduncle
1-3'4 cm, usually 3—10-flowered, sometimes di-
vided into two very short branches at the apex.

254

FLORA MALESIANA

[ser. I, vol. 6?

Bracts at the base of the peduncle and around it,
triangular to lanceolate, 3-8 mm long, slightly
erose. Bracteoles at the base of the pedicels tri-
angular, c. 14 mm long, short fimbriate, persistent.
Pedicels 7-9 mm. Flowers greenish white, or
yellowish, 4-merous. Calyx lobes semi-orbicular,
or deltoid, 1-2 mm long, the margin reddish
brown, denticulate. Petals obovate to slightly
obovate-oblong, 4-414 by 2-3 mm, short-fimbriate,
sometimes revolute. Disk obscurely 4-angular,
fleshy, 2-3 mm in @, covered with papillae.
Stamens each attached at the margin of the disk
in a shallow notch; filaments c. 3 mm, subulate;
anthers broadly ovoid or subrotund, c. 34 mm
long, introrse, slightly apiculate. Pistil emerging
c. 214 mm from the disk, + cylindric, gradually
enlarged and + 4-angular towards the base, the
basal part densely papillose in very young flowers
later growing out into fleshy, subulate processes
gradually increasing in length after anthesis.
Fruits globose, densely covered with prickles,
up to 13 mm incl. the spines (Sumatra). Endocarp
in each cell on both sides thickened and cartilagi-
nous, loosening from the shrinking exocarp.

Distr. Malaysia: Sumatra (Mt Kerintji) and
Philippines (Luzon: Benguet and Bontoc).

Ecol. Montane rain-forests, 1200-2300 m, once
recorded on limestone cliffs.

Vern. Tabkang, lg.

Note. It is rather remarkable that a Philippine
mountain species is found also in Central Sumatra.
Though the specimen (JAcoss 4343) is in fruit,
BLAKELOCK in sched. identified it as ‘near E.
benguetensis’; 1 can find no differences between
the two. Unfortunately no mature fruit is known

from the Philippines and no flowers from the
Sumatran specimen.

12. Euonymus moluccensis BLAKELOCK, nov. sp.
in sched.

Arbor parva, c. 6 m alta. Folia chartacea,
elliptica vel elliptico-oblonga, raro ovata, 6-9,
cm longa, 3-434, cm lata, basi cuneata, apice
acuminata, margine subintegra vel leviter serrulata,
nervis utrinque 4-6; petiolus 5-10 mm longus. In-
fructescentiae cymosae simplices, pedunculo 7-12
mm longo. Pedicellus 4-5 mm longus. Capsula
immatura clavata, c. 16 mm longa, 7 mm lata, 4-
carinata, apice obtusa.—Typus MAIN & ADEN
1040, L, isotypes A, Bo, SING.

Small tree, c. 6 m. Branchlets terete. Leaves
chartaceous, slightly ash-coloured, elliptic to
elliptic-oblong, rarely ovate, 6144-914 by 3-434
cm; base cuneate; apex acuminate; margin sub-
entire or remotely slightly serrulate; nerves 4-6
pairs; ++ obliquely spreading towards near the
margin and then turning upwards; petiole 5-10
mm. Jnfructescences axillary simple cymes, some-
times on the top of a very short branchlet. Pe-
duncle 7-12 mm. Bracts deltoid, c. 4% mm long
and wide, short-ciliate. Pedicel 4-6 mm. Flowers
unknown. Fruits clavate (young), 4-ridged, c.
16 by 7 mm, apex obtuse and slightly apiculate.

Distr. Malaysia: Moluccas (Morotai: Mt
Sangawo), once collected.

Ecol. Forest, 800 m. Fr. May.

Note. This species is very characteristic by the
clavate, 4-celled fruits which are very different
in the other Malaysian Euvonymus spp.

5. GLYPTOPETALUM

THwaAITEs in Hook. J. Bot. Kew Misc. 8 (1856) 267, t. 7B; Logs. in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 125.—Fig. 7.

Shrubs or small trees. Branchlets terete or 4-angled (G. quadrangulare), some-
times slightly compressed or flattened at the nodes. Stipules small, caducous.
Leaves decussate or opposite, sometimes subopposite at the upper part of the
young branchlets, entire or crenulate. Cymes simple to 2-3 times forked, axillary
or extra-axillary, quite often also on the internodes, with a distinct peduncle and
pedicels; pedicel of the lateral flowers usually shorter than that of the central
ones, often with 2 small bracteoles just below the articulation. Flowers bisexual,
4-merous. Calyx lobes spreading, the inner pair larger than the outer one. Petals
rather fleshy, usually smooth, sometimes with a small appendage or 2 depressions
on the inner side. Disk fleshy, flat, 4-angular or slightly 4-lobed, or covering the
ovary and confluent with it. Stamens 4, inserted on the disk or on the united body
of disk and pistil; filaments very short, persistent, connective dilate; anthers
divergent, dehiscent at the top or introrse (extra-Mal. sp.). Ovary immersed in
the disk, 4-celled; style obscure; stigma obscure or obtuse. Ovule one in each
cell, pendulous from the inner angle of the top, anatropous. Capsule loculicidal,
when dehiscing the valves splitting from the central axis leaving a persistent
columella, 4-1-celled, 4— or 3-1-seeded by abortion, globose or subglobose, if

Dec. 1962] CELASTRACEAE (Ding Hou) 255

Fig. 7. Glyptopetalum quadrangulare PRAIN ex KiNG. a. Habit, < 2/4, b. flower, petals and stamens removed,

6, c. frontal and dorsal view of stamen, x 12, d. petal, x 6, e-f. ovary in sections, 12, g. fruit, nat.

size. — G. marivelense (ELM.) MeERR. h. Dehisced fruit, nat. size, i-k. seed, lateral, apical, and basal views,

1. seed in section, all x 2.— G. zeylanicum Tuw. var. brevipedicillatum D1NG Hou. m. Habit, = 7A, n.

flower, petals and sepals removed, x 12, o. petal, <6, p. stamen, X12 (a-f SF. 35405, g CORNER S.n.,
h EDANO 34166, i-/ Ebalo 584, m-p RIDLEY 2652).

256 FLORA MALESIANA [ser. I, vol. 5?

containing only 2 seeds transverse-oblong. Seeds with incomplete, fleshy aril
covering the lower 14 or 1, the raphe running down on one side and branched
into 3 to 6 bands and turning at the morphological base of the seed, the bands

ascending towards the micropylar end of the seed on the other side.

Distr. About 20 spp. in Ceylon, India, Burma, Siam, Indo-China, China (only Hainan), and Malaysia
(Malay Peninsula, Borneo, Philippines, and Celebes).

Ecol. Forests from the lowland up to 1400 m. The species are, G. quadrangulare excepted, all rare and
local in Malaysia and mostly known from a few specimens and of some either the fruits or the flowers are
not or inadequately known, therefore specific distinction may change in future.

Notes. Glyptopetalum is closely allied to Euonymus; Kurz (J. As. Soc. Beng. 44, 1i, 1875, 259) thought
they should not be separated and BAILLON (Hist. Pl. 6, 1877, 1, footnote) treated them as one genus.
BENTHAM & HOOKER (Gen. PI. 1, 1862, 361) accepted them as distinct but mentioned only slight differ-
ences, and so did BEDDoME (FI. Sylv. 1, 1874, t. 102) and Lawson (in Hook. FI. Br. Ind. 1, 1875, 612).

PRAIN (J. As. Soc. Beng, 60, ii, 1891, 207) pointed out, however, that in G/yptopetalum, besides one
ovule per cell (against at least 2 in Euonymus) the dorsal raphe does not terminate at the base of the seed
but there divides into 3-4 laciniate segments of the same appearance and structure as the raphe itself,
differing from it only in being slightly branched and not quite reaching the hilum: ‘they form a closely
adherent arillar structure with meridional segments differing in colour from the testa that it overlies’.
A third difference with Evonymus is the persistent axis (columella) in the fruit, from the apex of which
hang the seeds, the valves being suspended on threads splitting downwards from it. A good illustration
of the situation is given by TARDIEU (FI. Gén. I.-C. Suppl. 1948, 784, f. 94: 7-8).

CuuNn & How described from Hainan Evonymus fengii CHUN & How (Act. Phytotax. Sin. 7, 1958,
44, f. 1, t. 15, 2). This is clearly a G/yptopetalum; in floral characters it is closely allied to G. zeylanicum
from Ceylon, G. calocarpum from the Andaman and Nicobar Is., and G. calyptratum from Indo-China
by the pyramidal pistil, but differs from these three species by obovate, entire leaves, introrse anthers,
and especially by the remarkable round, cushion-like thickening of the connective at the insertion of the
filament. It should be called Glyptopetalum fengii (CHUN & How) DinG Hou, comb. nov. The genus is
new to China.

KEY TO THE SPECIES

Leaves with apices obtuse to rounded sometimes slightly notched.
Leaves broad-elliptic, ovate, or obovate, 414-6 by 314-314 cm. Pedicels of the lateral fruits of each
cyme at most 1 mm long above the articulation with the bracteoles . 1. G. euonymoides
2. Leaves obovate to obovate-oblong, 10-14 by 5-8 cm. Lateral fruits c. 6 mm pedicelled.
2. G. palawanense
1. Leaves with apices acute to acuminate.
3. Branchlets sharply 4-angular. Leaves usually bullate
3. Branchlets terete, sometimes also associate with slightly 4- angular o ones.
4. Leaves lanceolate to narrow-lanceolate, 20-23 by 4-7 cm; veins and veinlets obscure or invisible
on both surfaces; margin remotely denticulate 5 . . 4. G. acuminatissimum
4. Leaves usually elliptic to elliptic-oblong, ovate-oblong, rarely lanceolate, 4-17'4 by 2-634 cm; veins
and veinlets distinctly reticulate on both surfaces.
5. Infructescences c. 2 cm long. Leaf margin subentire, repandous, or with obscure, small black teeth.
5. G. loheri
5. Inflorescences or infructescences usually longer, up to 10 cm long. Leaf margin usually crenulate.
6. Petals bifoveolate at the upper part inside. Pistil evidently united with the disk, short-conical.
6. G. zeylanicum var. brevipedicellatum
6. Petals smooth inside. Disk distinctly fleshy, flat; Sey immersed in the disk and slightly above it.
7. Fruits small, subglobose, c. 8 mm o Sie) cy 7. G. euphlebium
7. Fruits larger, depressed-globose, c. 8. G. marivelense

3. G. quadrangulare

15 mm 9. .

1. Glyptopetalum euonymoides Merr. Philip. J.
Sc. 12 (1917) Bot. 278; En. Philip. 2 (1923) 481.

Shrub c. 2 m. Branchlets terete. Leaves char-
taceous to subcoriaceous, broad-elliptic or ovate,
sometimes obovate, 444-6 by 314-314 cm; base
cuneate; apex obtuse, or rounded, sometimes
slightly notched; margin entire, sometimes re-
motely crenulate; nerves 4 to 7 pairs; veins and
veinlets obscure on both surfaces; petiole 5-12 mm.
Cymes usually at the basal part of the flush, 2-3
times forked, 3-6 cm long. Peduncle 144-3 cm.
Pedicels none or very short: lateral flowers of the

ue sessile, the central ones subsessile or up to

1144 mm pedicelled. Ca/yx lobes reniform or semi-
orbicular, 1 by 144-2 mm, slightly concave with
obscured longitudinal veins. Peta/s subreniform or
suborbicular, 134-3 by 2%,—4 mm, rather fleshy.
Disk 4-angular or slightly 4-lobed, c. 2 mm 9.
Stamens c. %4 mm long, filament very short.
Pistil slightly emerging from the disk. Fruits
(young) subglobose or broadly ovoid, c. 8 mm
long, usually only one seed developed. Seeds
ellipsoid.

Distr.

Malaysia: Philippines (Luzon: Ilocos

Dec. 1962]

CELASTRACEAE (Ding Hou)

Norte, B.S. 27546, A, BM, K, US), once collected.
Ecol. On slope in thicket at low altitude.

2. Glyptopetalum palawanense Me_rr. Philip. J. Sc.
26 (1925) 466; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 126.

Small tree c. 5 m. Branchlets - compressed.
Leaves coriaceous, shining, 10-14 by 5-8 cm;
base acute to cuneate, sometimes attenuate; apex
obtuse or rounded; margins remotely crenulate
at the upper part; nerves 5-7 pairs, slender; pe-
tiole 8-12 mm. Jnfructescences 5 cm, axillary
and also on the internodes, 2-3 times forked.
Peduncle 1-3 cm. Pedicels 6 mm. Fruits globose to
depressed-globose, 1—4-celled, 8 mm by 1!4 cm.

Distr. Malaysia: Philippines (Palawan), thrice
collected.

Ecol. In primary forest at low altitude.

Vern. Panablayan.

3. Glyptopetalum quadrangulare PRAIN ex KING,
J. As. Soc. Beng. 65, ii (1896) 345; Rip. Fl. Mal.
Pen. 1 (1922) 446; SyminGTON, J. Mal. Br. R. As.
Soc. 14 (1936) 350; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 126.—Fig. 7a—g.

Shrub or small tree up to 5 m. Branchlets
distinctly 4-angled, sharply winged. Leaves
subcoriaceous to coriaceous, elliptic to elliptic-
lanceolate, 9-2914 by 3-13!4 cm; base rounded
or cuneate; apex shortly acuminate to acuminate
(acumen up to 114 cm); margin irregularly
crenulate especially in the upper half; nerves
(5—)8—12 pairs; nerves and veins depressed above,
making the leaves subbullate; petiole 5-10 mm.
Cymes 1(—2) axillary, sometimes on the internodes,
rarely terminal, up to 10 cm long (in fruit up to
12 cm), usually forked near the apex. Peduncle
114-814 cm. Bracts small. Pedicels 2-3 mm,
articulated at the base. Flowers greenish yellow.
Calyx almost divided to the base, lobes + reni-
form, 114-2 by 7-10 mm. Petals suborbicular,
314 mm 9, rather fleshy, thinner and wavy near
the edge. Disk flat, square, c. 2 mm @. Stamens
c. 1 mm, inserted near the base of the ovary.
Pistil c. 5 mm above the disk, pyramidal; style
and stigma obscure. Fruits slightly depressed-
globose, 114-2 by 1-114 cm, 3-4 celled, slightly
sulcate. Seeds broadly-ellipsoid, 8 by 6 mm,
reddish brown.

Distr. Burma (S. Tenasserim) and Malaysia:
Central West Sumatra, Malay Peninsula(common),
and Borneo (Sarawak: Kuching).

Ecol. Rain-forests, from the lowland up to
600 m.

Vern. Cha teng, pérupong, p. bukit, poko
kahava, p.restong, séminyh, M.

Note. This is the only species which seems to
be rather common. Its leaves remind of those of
G. euphlebium which has, however, not the
sharply angled branchlets.

4. Glyptopetalum acuminatissimum Merr. Philip.
J. Sc. 29 (1926) 481; Logs. in E. & P. Pfl. Fam. ed.
2, 20b (1942) 126.

Shrub up to 3 m. Leaves chartaceous, lanceolate

to narrow-lanceolate, 20-23 by 4-7 cm; base
cuneate, obtuse or sometimes rounded; apex
caudate-acuminate; margin irregularly, remotely
denticulate at the upper two-thirds; nerves 9-12
pairs, spreading, arching 2-7 mm from the edge;
veins and veinlets obscure or invisible on both
surfaces; petiole 34-1144 cm. Infructescences
axillary, c. 3 cm long, simple or once branched
cymes, usually found at the upper part of the
branchlet. Peduncle 14-2 cm. Bracts small.
Pedicels 5-8 mm. Fruits depressed-globose, 11-2
cm 9. Seeds subellipsoid or slightly oblong, 8-10
by 7 mm.

Distr. Malaysia: Philippines (Luzon: Isabela
Prov.), 3 collections.

Ecol. In forests, 100-300 m.

Vern. Guisguis, Ilocamo.

5. Glyptopetalum loheri Merr. Philip. J. Sc. 10
(1915) Bot. 321; En. Philip. 2 (1923) 481.

Shrub or small tree. Branchlets terete. Leaves
subcoriaceous, elliptic- or ovate-oblong, 6-14 by
3-614 cm; base acute to cuneate; apex acuminate;
margin subentire, repand, or with obscure small
black teeth; nerves 5—7, fine, spreading and slightly
anastomosing-reticulate near the margin; veins
and veinlets slightly elevated, finely reticulate;
petiole 3-4 mm. Cymes simple or once branched,
c. 2. cm long, axillary and extra-axillary. Peduncle
8-10 mm. Bracts small, ovate, c. 14% mm long.
Pedicels 3 mm. Calyx lobes subreniform or sub-
orbicular, 1 by 114-214 mm, reddish brown near
the margin. Fruits globose, 6-10 mm @, smooth,
sometimes slightly furrowed, 1—4-seeded. Seeds
subglobose, c. 6 mm g, reddish brown.

Distr. Malaysia: Philippines (Luzon: Rizal
and Cavite Prov.) and SE. Celebes (Kendari,
once found).

Ecol. Forests, lowland up to c. 400 m.

6. Glyptopetalum zeylanicum THw. in Hook. J.
Bot. Kew Misc. 8 (1856) 267, t. 7B.

var. brevipedicellatum DING Hou, var.
Fig. 7m-p.

Arbor parva. Folia chartacea, elliptica, 7-10
cm longa, 34-5 cm lata, basi cuneata, apice
acuta vel breviter acuminata, nervis utrinque 6-8,
venulis laxe reticulatis; petiolus c. 5 mm longus.
Cymae axillares, laxe, usque ad 8% cm longae,
pedunculo 4-5 cm longo. Pedicellus 2-3 mm longus.
Flores albi. Calycis lobi subrotundi, c. 1 mm longi.
Petala subrotunda, c. 2 mm longa, superne 2-
foveolata. Stamina supra discum inserta. Ovarium

nov.—

discum adnatum, brevissime conicum.—Typus
RIDLEY 2652, SING.

Small bushy tree. Branchlets terete. Leaves
chartaceous, elliptic, 744-1014 by 344-5 cm;

base cuneate; apex acute to short-acuminate;
margin remotely crenulate; nerves 6-8 pairs;
petiole c. 5 mm. Cymes divaricate, up to 814 cm,
once or twice forked. Peduncle 4-5 cm. Bracts
triangular, c. 1 mm long; flower stalk c. 10 mm,
articulated at about the upper 1/5. Pedicel (above
articulation) 2-3 mm. Flowers (young) white.
Calyx lobes suborbicular, c. 1 mm 9, slightly

258

erose on the margin. Petals suborbicular, slightly
concave, c. 2 mm 9, bifoveolate at the upper part
inside. Stamens 1 mm long, inserted on the lower
part of the pistil. Pistil evidently united with the
disk, short-conical, c. 144 mm long and wide,
slightly narrowed at the apex; style and stigma
obscure.

Distr. Malaysia: Malay Peninsula (Pahang:
Kota Glanggi), once found.

Notes. The Malayan variety differs from the
Ceylonese var. zeylanicum by the more lax,
spreading dichotomously branched inflorescences
and flower stalks which are articulated at about
the upper one fifth, the pedicel proper being only
2-3 mm. In var. zeylanicum the flower stalk is
articulated in the lower one fifth, the pedicel
proper being 9-12 mm.

KING cited the type (RipLEy 2652) under
Euonymus wrayi (J. As. Soc. Beng. 65, ii, 1896,
344) but his description of the 5-merous flowers
of that species does not apply to RIDLEY’s spec-
imen which has 4-merous flowers.

7. Glyptopetalum euphlebium (MEeERR.) MERR.
Philip. J. Sc. 12 (1917) Bot. 280; En. Philip. 2
(1923) 481.—G. marivelense var. euphlebium MERR.
Philip. J. Sc. 10 (1915) Bot. 321.—G. remoti-
nervium Merr. ibid. 12 (1917) 280; En. Philip. 2
(1923) 481.

Shrub or small tree up to 5 m. Branchlets
terete. Leaves firmly chartaceous, elliptic- or
ovate-oblong, 14-17 by 514-614 cm; base acute
to cuneate; apex acute and _ short-acuminate;
margins slightly recurved, obscurely and sometimes
distinctly crenulate; nerves 5-6 pairs; petiole
6-8 mm. Cymes axillary or extra-axillary, few-
flowered, sometimes depauperate. Peduncles 1/4—
41% cm. Pedicels c. 114 mm (c. 414 mm in fruit).
Flowers (young): calyx lobes reniform or trans-
verse-oblong, 1 by 2-234 mm, slightly erose, with
several obscure longitudinal veins. Petals sub-
reniform, 2 by 3 mm. Stamens small, c. 34 mm
long, inserted near the base of the ovary. Disk
4-lobed, c. 2 mm @. Pistil emerging c. 44 mm

FLORA MALESIANA

[ser. I, vol. 6?

from the disk, conical. Fruits white, pink, or red,
subglobose, c. 8 mm @, usually 1-seeded. Seeds
red, reddish brown when dry, subglobose, c. 7 mm
o.
Distr. Malaysia: Philippines (Palawan and
Luzon: Zambales Proy.).

Ecol. Primary forests, 300-1400 m.

Vern. Nou, Tagb.

Note. The leaves are very similar to those of
G. quadrangulare.

8. Glyptopetalum marivelense (ELM.) MErRR. Philip.
J. Sc. 10 (1915) Bot. 321; En. Philip. 2 (1923)
481.—Euonymus marivelensis Etm. Leafl. Philip.
Bot. 7 (1915) 2580.—G. reticulatum MeRR.
Philip. J. Sc. 12 (1917) Bot. 277; En. Philip. 2
(1923) 482.—Fig. 7h-k.

Shrub or small tree up to 7 m. Branchlets terete,
the ultimate internodes rarely 4-angular, sometimes
slightly compressed near the node. Leaves char-
taceous to coriaceous, elliptic-oblong to -lanceo-
late, or lanceolate, 4-1714 by 2-634 cm; base
cuneate or acute, very rarely obtuse; apex acu-
minate; margin subcrenate to remotely crenulate;
nerves 6—10 pairs; veins densely reticulate, distinctly
prominent on both surfaces, sometimes obscure
beneath; petiole 3-18 mm, sometimes subsessile.
Cymes axillary or extra-axillary, sometimes on a
short axillary branch, 2!4—7 cm long (up to 10 cm
in fruit), once or twice forked. Peduncle 114-344
cm. Bracts deltoid or lanceolate. Pedicels c.
3-5 mm. Calyx lobes semi-rounded or subreniform,
34-2 by 114-234 mm. Disk slightly 4-angular, with
obscure papillae (B.S. 75176). Petals suborbicular
or broad-ovate, c. 4 mm @. Stamens inserted at
the base of the ovary, c. 34 mm long. Pistil emerg-
ing c. 1 mm from the disk, short-conical. Fruits
reddish when fresh, depressed-globose, some-
times slightly wrinkled, c. 114 cm @, 3—4-seeded,
slightly furrowed. Seeds broad-oblong, 9-10 by
7-8 mm, dark-brown.

Distr. Malaysia: Philippines (Mindoro, Luzon,
and Catanduanes).

Ecol. Forests, from the lowland up to 1400 m.

6. KOKOONA

THWAITES in Hook. J. Bot. Kew Misc. 5 (1853) 379; KING, J. As. Soc. Beng. 65,
ii (1896) 346; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 164; STEEN. Sarawak
Mus. J. 8 (1958) 437._Fig. 8.

Trees, (in Mal.) evergreen, sometimes buttressed. Branchlets flat at the nodes,
almost always dark to black. Stipules very small. Leaves decussate (occasionally
in some branches sometimes subopposite or even alternate), the midrib frequently
convex causing the blades to fold when pressed for herbarium purposes. Pa-
nicles, or sometimes racemes, axillary. Flowers bisexual, usually on articulated
pedicels, with 2 to 5 small bracts at the articulation. Calyx 5-lobed. Petals 5,
contorted, slightly concave, subcoriaceous to coriaceous, sometimes punctate.
Disk fleshy, cupular, subentire, corrugated, rarely 5-lobed. Stamens 5, inserted
inside the inner edge of the disk; disk-lobes alternate with the petals; filaments

Dec. 1962] CELASTRACEAE (Ding Hou) 259

\

\
Re G ‘

Fig. 8. Kokoona ovatolanceolata Rv. a. Habit, < 24, 6. bud, c. flower, in section, d. diagram, e. stamen,

all <6, f. one fruit valve with 2 rows of imbricate seeds, g. seed, both x 74.—K.reflexa (LAws.) DING

Hou. h. Flower, petals and one anther removed, i. petal, j. stamen, all * 6.—K.ochracea (ELM.) MERR.

k. Bud, /. flower in section, m. stamen, all 6 (a-e ANDERSON 7910, f-g SAR 9291, h-j ACHMAD 945, k-m
ELMER 21881).

usually fusiform, gradually or abruptly narrowed towards the apex and trans-
parent at the upper end, very rarely terete; anthers usually with an apical, pustular,
lengthened connective, rarely this obscure or obtuse. Ovary superior or some-
times partly immersed in the disk, 3-celled, gradually narrowed into an obscure
style; stigma thick and short-cylindric, + capitate, or orbicular and flat, obtuse
or sometimes obscurely 3-lobed. Ovules 6-16 in each cell, in two series, attached

260 FLORA MALESIANA [ser I voll?

to the axis, superposed and ascending. Capsule oblong, bluntly 3-angular, 3-
celled, loculicidally dehiscing, 3-valved. Seeds 6—10(—16?) in each cell, imbricate,
erect, exalbuminous, with a conspicuous wing at the apical end, the wing very

broad, oblong, truncate or blunt.

Distr. Species 8, one in Ceylon and southern India, one in Burma, and six in Malaysia (Sumatra,
Malay Peninsula, Borneo, and the Philippines). Fig. 9.

Ecol. Lowland rain-forests, dryland, swampy or peat, rarely up to 1500 m.

Uses. THwaltes (/.c. 380) said that the pounded yellow bark of K. zeyvlanica is used by the Singhalese
as a kind of cephalic snuff, being mixed with ghee (buffalo milk butter) and introduced into the nostrils
in order to relieve severe headache.

The bark of many (all?) species contains oil, seems easy to burn and is sometimes used for tinder.

Notes. The species of this genus seem well distinct and can easily be identified if flowers and fruits
are present. Detached fruits or fruiting specimens are difficult to place; sterile material cannot be named.

For the differences with Lophopetalum, see under that genus.

KEY TO THE SPECIES

1. Anthers with distinctly prolonged connective.
2. Flowers distinctly pedicelled, after falling off leaving a distinct,
2. Ovules 6-10 per cell.
3. Connective c. 114 mm longer than the anther (c. 1 mm). Stigma broad-oblong, cylindric.
1. K. ochracea
3. Connective 14-1 mm, usually shorter than, rarely as long as, the anther (1-174 mm). Stigma
capitate or subglobose.
4. Flower bud broad-ovoid, subglobose or globose, about as long as wide. Calyx lobes semi-orbicular,
reniform, or ++ transverse-oblong, the apex rounded or + truncate . 2. K. littoralis
4. Flower bud broad-ellipsoid, longer than wide. Calyx lobes triangular, the apex acute.
3. K. coriacea
. Flowers sessile, after falling off eae an annular, cushion-like scar. Bracteoles 3-5. Ovules 14-16
per cell ata ‘J 4. K. sessilis
2 uathiers without or with obscure « or - very shortly prolonged connective.
. Connective obscure or very shortly prolonged. Filaments fusiform, thickened in the lower part,
transparent at the upper end. Stigma broad-oblong . Dare 5. K. ovatolanceolata
5. Connective not prolonged. Filaments terete, thin and not transparent at the upper end. Stigma
flat and orbicular c Shternts 6. K. reflexa

cylindric stalk. Bracteoles usually

1. Kokoona ochracea ee es En. Philip. 2
(1923) 484 (as Kokoonia); Pl. Elm. Berm. (1929)
171; Logs. in E. & P. Pfl. Dae ed. 2, 20b (1942)
165: STEEN. Sarawak Mus. J. 13 (1958) 438.—

stout, c. 114 mm long. Pistil conical, c. 14 mm,
gradually narrowed upward into the obscure
style; stigma broad-oblong cylindric, c. 74 mm
long, slightly 3-lobed or obtuse at the apex.

Ardisia ochracea E_m. Leafl. Philip. Bot. 5 (1913)
1819.—Fig. 8k—m.

Tree 25-40 m by 20-40 cm g, rarely with
buttresses up to c. 3 m tall (cf. KOSTERMANS 5792).
Leaves subcoriaceous to coriaceous, elliptic- or
ovate-oblong, oblong-lanceolate, or lanceolate,
714-134 by 314-6 cm; base attenuate, obtuse or
rounded; apex acute to acuminate, recurved
downward; margins recurved, entire or repand,
sometimes remotely crenulate; nerves 5-8 pairs,
slightly elevated on both surfaces, obliquely
spreading towards the margin and then upward;
petiole c. 1 cm. Panicles up to 12 cm, solitary or
paired. Peduncle UB: to 4 cm. Pedicels very short,
sometimes up to c. 2 mm, articulated at the base.
Bracts small, deltoid and acute. Flowers yellowish
to pale brown-orange. Calyx lobes semi-orbicular
or + transverse-oblong, 4 by 1 mm, slightly
erose. Petals fleshy, ovate rarely elliptic, 314-414
by 214-3 mm, the margin thin and transparent.
Filaments c. 1 mm; anthers including the connec-
tive c. 2'4 by 1 mm, the anther proper subglobose,
cordate at the base, the lengthened connective

Ovary with (8—)10 ovules in each cell.

Distr. Malaysia: Malay Peninsula (Johore),
Borneo (Brunei, N. and E. Borneo and Nunukan
I.), and Philippines (Palawan).

Ecol. Lowland forests, below 120 m, in Tawao,

Fig. 9. Distribution of the genus Kokoona THw.,
number of species indicated.

Dec. 1962]

CELASTRACEAE (Ding Hou)

261

North Borneo, once found in the forests near
Nypa swamps.
Vern. Borneo: ? batubagalang, Pleihari, kaju
minjak, kélapatiung, Kutei, ubar, Brunei.
Note. The bark contains inflammable oil.

2. Kokoona littoralis LAws. in Hook. f. Fl. Br.
Ind. 1 (1875) 617.—Lophopetalum dubium LAws.
Lc. 616; Rip. Fl. Mal. Pen. 1 (1922) 450.—
K. scortechinii KinG, J. As. Soc. Beng. 65, il
(1896) 347; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 165.—Lophopetalum maingayi RIDL.
Fl. Mal. Pen. 1 (1922) 450 (new name for K.
scortechinii)—Lophopetalum littoralis Ripv. Lc.
—Solenospermum littorale Lors. Notizbl. Berl.-
Dahl. 13 (1936) 223.—K. lanceolata RiDL. Kew
Bull. (1938) 237.

Tree up to 45 m by 75 cm 9, with small but-
tresses. Young twigs red. Leaves chartaceous to
coriaceous, elliptic to very narrowly elliptic,
broad-elliptic, ovate, or lanceolate, 414-1514 by
134-7 cm; base cuneate rarely obtuse; apex
acuminate, rarely blunt or acute; margins cartila-
ginous, slightly recurved, entire or repand, some-
times remotely very shallowly crenate; nerves
5-8 pairs; petiole 34-134 cm. Panicles or racemes
axillary and sometimes extra-axillary, 11-20
cm long. Peduncle up to 6 cm. Pedicels 1-214 mm.
Bracteoles small, deltoid, c. 24 mm long, acute,
denticulate. Flowers yellowish or whitish. Calyx
lobes semi-orbicular, or reniform, 1% by 114-2
mm, slightly denticulate. Petals broadly ovate,
elliptic, sometimes suborbicular, 314-6 by 234-4
mm; filaments 1-114 mm; anthers including the
connective | 14-2 14 mm, connective 14-1 mm, poin-
ted at the tip. Pistil c. 2mm. Ovary + triangular;
style obscure; stigma capitate, c. 4/5 mm, obtuse
sometimes slightly 3-notched at the top. Ovules
(6—)8-10 in each cell. Fruits 13-18 by 3-514 cm.
Seeds including the wing 714-1214 by 2!4 cm.

Distr. Malaysia: Sumatra (Morsala I., Palem-
bang, and Indragiri), Malay Peninsula (through-
out, and incl. Penang I.), and Borneo (Sarawak
and Brunei).

Ecol. Primary dryland forest, from lowland up
to 450-600 m, in the Cameron Highlands up to
1500 m. Monkeys eat the fruit (CURTIS).

Vern: Babi kurus, mata ulat, ménchali, M,
Mal. Pen., sépalis, M, Palemb., bajan garéjak,
Iban.

Notes. I have examined a sheet of the authentic
material of K. littoralis LAws. (MAINGAY 396/2,
lectotype, K), which has leafy branchlets with
five detached fruits. I have also seen the type
collection of Lophopetalum dubium LAws. (MAIN-
GAY 1436, K), which has three flowering branchlets
with detached leaves. There are two specimens cited
in the original description of K. scortechinii
KING, i.e. CurTIS 1576 (Bo, KEP) and SCORTECHINI
2042 (lectotype K, isotypes Bo, G, Kep). All the
specimens mentioned above are very similar to
each other and have been accepted as conspecific.

The leaves of the present species are very va-
riable in shape, texture and size, but the floral
characters are rather homogeneous and constant.

3. Kokoona coriacea KiNG, J. As. Soc. Beng. 65,
ii (1896) 347; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 165.—Lophopetalum coriacea RIDL.
Fl. Mal. Pen. 1 (1922) 450.

Tree, 9-15 m tall. Branchlets terete. Leaves
coriaceous, ovate to ovate-oblong, 11-12'4 by
5-71 cm; base cuneate; apex subacute; margins
slightly recurved; nerves 6 or 7 pairs, slightly
curved towards the margin; petiole c. 1 cm.
Panicles up to 12 cm long, distinctly peduncled.
Pedicels very short (c. 1 mm). Bracts -+ triangu-
lar, c. 1% mm long and wide, acute. Calyx lobes
triangular, c. 4 mm long and wide, acute, Calyx
lobus triangular, c. 1/4 mm long. Petals broad-oyate,
414 by 3mm, obtuse. Stamens c. 2mm long; fila-
ments thickened at the base, transparent at the apex;
anthers with connective c. 2 mm long, the latter
c. 4/5 mm. Free part of the pistil c. | mm, broad-
ovoid, gradually narrowed into an obscure style;
stigma subglobose, obscurely 3-notched at the apex.

Distr. Malaysia: Malay Peninsula (Perak),
once collected.

4. Kokoona sessilis DING Hou, sp. nov.

Arbor usque ad 20 m alta. Folia coriacea,
elliptico- vel ovato-oblonga, raro late ovata, 9-14
by 514-7 cm, basi obtusa, apice acuta vel obtusa,
nervis utrinque 5-8; petiolus 1% cm. Flores
sessiles, basi 3—5-bracteolatae. Flores parvi.
Calycis lobi orbiculares vel reniformes, 1-2 mm
longi. Petala late ovata vel subrotunda, 2-3 mm
longa. Stamina breviora, antheris late ovoideis,
c. 1% mm longis, appendicibus Y3—Y% mm longis.—
Typus S.F. 36296, SING, isotypes K, L.

Monopodial tree with elongate, cylindric crown,
not buttressed, up to 20 m. Leaves coriaceous,
elliptic- or ovate-oblong, rarely broad-ovate,
914-14 by 514-7 cm; base obtuse; apex acute or
obtuse; margins cartilaginous, recurved, entire;
nerves 5-8 pairs; veins and veinlets reticulate,
visible on both surfaces rarely obscure beneath;
petiole 114 cm. Panicles hanging, up to 15 cm
long. Peduncles up to 8 cm. Bracteoles 3-5 at
the base of the flower, semi-orbicular, or + reni-
form, 4-34 mm long. Flowers (rather young)
sessile, not fragrant. Calyx almost divided to the
base, lobes semi-orbicular or reniform, 1—2 by
2-3 mm, slightly erose, the outer two slightly
smaller. Petals dull yellow, broad-ovate to sub-
orbicular, 214-3 by 134-24 mm; filaments c.
1 mm; anthers broad-ovoid, including connective
c. 134 by 1 mm, the connective %4—¥ times as
long as the anther. Crenulate disk and ovary
orange-red. Pistil c. 114 mm long, conical; style
obscure; stigma cylindric c. 74 mm, obtuse.
Ovules 14-16 in each cell.

Distr. Malaysia: Malay Peninsula (Johore),
once collected.

5. Kokoona ovatolanceolata RipL. Kew Bull.
(1938) 236 (‘ovato-lanceolata’).—K. scortechinit
(non KiNG) STEEN. Sarawak Mus. J. 13 (1958)
438.—Fig. 8a-g.

Tree up to 45 by 14 m @. Bark finely fissured
below, smoother above. Buttresses up to c. | m
high. Leaves coriaceous, shining above, rather

262

FLORA MALESIANA

[ser. I, vol. 6?

dull beneath, ovate, ovate-oblong to ovate-
lanceolate, 714-11 by 3—5!4 cm; base obtuse to
rounded, sometimes acute; apex acuminate;
margin cartilaginous, + entire, sometimes re-
motely shallow-crenulate; nerves 5-7 pairs, ele-
vated on both surfaces, obliquely spreading and
curving upward; veins and veinlets rather densely
reticulate, slightly elevated on both surfaces;
petiole 34-134 cm. Panicles up to 14 cm long.
Peduncle up to 424 cm. Bracts small, deltoid.
Pedicels c. 1 mm. Flowers yellowish, fragrant.
Calyx lobes semi-orbicular or reniform, 1 by
1144 mm. Petals ovate, 344-324 by 2-214 mm;
anthers ovoid, 114 by 1 mm, acute, with a very
short and obscure connective; filaments c. 11 mm.
Pistil 114-2 mm long; ovary broad-ovoid, gradually
narrowed into a rather distinct style c. 4 mm;
stigma -+ broad-oblong, obtuse. Ovules 8-10 in
each cell. Fruit valves narrow-oblong, 10-17 by
3-5 cm, leathery, c. 5 mm thick. Seed (includ-
ing the wing) 7-11 by 2-224 cm, seed proper 12
by 14 mm.

Distr. Malaysia: Borneo (throughout).

Ecol. In primary forests, primary peat-swamp
and sometimes freshwater-swamp forests from
sea-level to low altitude. F/. June—Sept., /r.
Sept.-Nov.

Vern. Akil, dian atau barak, majan, Sarawak,
anakan, W. Born., badang, kayu api, mata ulat,
sabong api (Iban), Brun., énsabung, kayu kayan,
M, kérandji, Sampit, kulian, bekumpai, SE. Born.,
sépetir paya, N. Born.

Use. The yellowish outer bark is used by na-
tives for tinder. The timber has no commercial
use.

Note. The inner side of the outer bark is
bright orange which is said to be diagnostic (fide
SMyYTHIES BRUN 0822).

6. Kokoona reflexa (LAws.) DING Hou, comb. nov.
—Lophopetalum reflexum Laws. in Hook. f.
Fl. Br. Ind. 1 (1875) 616; Kina, J. As. Soc. Beng.
65, il (1896) 352; RoLFe, Kew Bull. (1918) 48;
Rip_. Fl. Mal. Pen. (1922) 449.—Hippocratea
maingayi LAws. in Hook. f. Fl. Br. Ind. 1 (1875)
625.—Fig. 8h-i.

A tree up to 48 m by 90 cm g, sometimes with
buttresses up to 114 m high and c. 14 m over the
ground. Leaves chartaceous, elliptic-oblong, 514—-
11 by 214-4 cm; base cuneate; apex acuminate;
margins remotely crenulate, rarely subentire,

slightly recurved; nerves 5 pairs; petiole 4/5—114
cm. Panicles 2-18 cm long, axillary as well as
extra-axillary. Peduncle up to 514 cm. Pedicel
c. 1 mm. Flowers light green, or dirty yellowish
white. Calyx lobes c. 114 by 4 mm. Petals broad-
ovate, or broad-elliptic, 3 by 214 mm. Stamens
114 mm; filaments terete, thin; anthers oblong,
1 by 34 mm, obtuse at the apex, connective not
produced. Pistil c. 114 mm long; ovary gradually
narrowed into an obscure style; ovules 10 in each
cell; stigma orbicular, flat. Fruits c. 12 cm long;
valves c. 3 cm wide. Seeds including the wing
9-11 by 214 cm.

Distr. Malaysia: Sumatra (Indragiri, Asahan,
Palembang; also Simalur I.), Malay Peninsula
(Kedah, Perak, Dindings, Selangor, and Singa-
pore), and Borneo (W. Borneo: Melawi; E.
Borneo: Samarinda; SE. Borneo: Pleihari).

Ecol. Lowland dryland forests, sometimes up
to c. 250 m, common but always in + scattered
individuals.

Vern. Sumatra: useu-useu uding, Simalur,
kémpas sakam, Indragiri, pasir, Batak, négris
hitam, n. pinang, sépalis, Palembang, résak, sajap,
M; Borneo: barjau, sabong api, Iban, batu baga-
lang, M, Pleihari, bintan, Sampit, kaju minjak,
Samarinda.

Notes. The type of Lophopetalum reflexum
Laws. (Herb. MAtNGAy 393/2, K) from Malacca
is a rather poor specimen which consists of three
defoliate branchlets and a fourth one bearing
still a leaf. There are a few detached flower buds.
The characteristic floral characters (petals con-
volute, disk cupular, and stamens inserted inside
the disk) make it easy to recognize it as a Kokoona.
Its chartaceous and greyish leaves, terete fila-
ments, obtuse anthers, and triangular disk lobes
represent the characters of a distinct and rather
wide distributed species, K. reflexa.

The type of Hippocratea maingayi LAWs. was
correctly placed by Ro.re (/.c.) under L. reflexum
of which I have examined the type.

Excluded

Kokoona luzoniensis MERR. Philip. J. Sc. 27 (1925)
32. I have not yet seen the authentic material
(LoHER 12754 & 13391). According to the char-
acters of the inflorescences (paniculate) and
stamens (3) indicated in the original description,
this seems to belong to Loeseneriella A. C. SM.

7. LOPHOPETALUM

WIGHT ex ARN. Ann. Mag. Nat. Hist. I, 3 (1839) 150; Wicut, I11. 1 (1840)
177; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 160.—Solenospermum ZOLL.
Nat. Tijd. Ned. Ind. 14 (1857) 168; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
162.—Lophopetalum subg. Solenospermum (ZOLL.) VAL. Ic. Bog. 1 (1901) 43.—

Fig. 10, 12-13.

Small to tall (in Mal.) evergreen trees. Buttresses sometimes present and up to
8 m high. Branchlets mostly dark, terete (except L. sessilifolium) and mostly

Dec. 1962] CELASTRACEAE (Ding Hou) 263

| | AM

a

Galas as
a a, ays \

Fig. 10. Lophopetalum beccarianum PirRRE. a. Habit, < 24, b-c. bud and open flower, seen from above,

x 6, d. flower, in section, petals removed, e. ovary in section, both ™ 8, f. seed surrounded by wing, * 74—

L. macranthum (Lors.) Dinc Hou. g. Flower, petals and sepals removed, A. pistil in section, i. petal,

from inside, all * 6.—L. pallidum Laws. j. Flower, petals and anthers removed, k. stamen, /. petal, from

inside, all * 6.—L. rigidum Rip. m. Section of petiole at distal end, * 6 (a-e SAN 16022, f bb 10797, g-i
GseELLERUP 701, j-/ KL 1566, m HAVILAND 2235).

flattened at the nodes. Leaves decussate, or opposite, sometimes subopposite
(occasionally a few leaves spiral on the upper part of a branchlet), distinctly
sometimes very shortly petioled, rarely + sessile, blade rarely subpeltate; margin
entire, sometimes cartilaginous. Stipules a tuft of hair-like processes, caducous.
Thyrses axillary, 1(—2-3), sometimes branched from the very base, divaricate.
Peduncle distinct sometimes very short or + absent. Bracts deltoid or lanceolate,
usually short-ciliate or -fimbriate. Pedicels distinct, usually articulated at the

264 FLORA MALESIANA [ser. I, vol. 6?

base. Flowers bisexual, 5-merous (except the ovary). Calyx dish-shaped, lobes
rounded or triangular, usually spreading, sometimes inflexed or reflexed. Petals
imbricate, inner surface usually partly covered with cristate, lamellate, or fim-
briate appendages, rarely bearing a tuft of fleshy papillae at the central part
(L. beccarianum), or naked. Disk usually fleshy and + flat, in bud usually slightly
concave, sometimes thin and patelliform, upper surface smooth, or denticulate,
sometimes covered with fleshy, subulate processes (extra-Mal. spp.), rarely only
at the base of filaments (L. pallidum), 5-angular, -+- rounded, or distinctly 5-
lobed and the angles or lobes episepalous, or epipetalous. Stamens inserted on the
disk, usually between pistil and edge, sometimes quite near the margin, rarely
each of them in a small pit (L. pallidum); filaments filiform, dorso-centrally
fixed; anthers usually broad-ovoid or -ellipsoid, versatile, + introrse, apex obtuse
or short-apiculate, free at the lower 14 or 14. Ovary usually partly immersed in
the disk, trigonal or pyramidal, pradually narrowed into a cylindric, short style,
3-celled; stigma obscure. Ovules 4-18 in each cell, arranged in two series. Capsule
oblong, or slightly + spindle-shaped, 3-lobed, -winged, or -angular, loculicidally
dehiscing. Seeds oblong, flat, attached in the middle, the wing surrounding the
seed; albumen absent, or scanty.

Distr. About 18 spp., in India, Burma, Siam, Indo-China, and throughout Malaysia (not found in
the Lesser Sunda Is. and East Java). Fig. 11.

Ecol. Frequently in lowland rain-forests, some species in dryland forests, others in peat-swamp or
freshwater-swamp forests, sometimes up to 1500 m.

Notes. The sole difference between Solenospermum and Lophopetalum is that in the first the disk is
pentagonal with alternipetalous lobes, while in the second the disk is lobed and the lobes are epipetalous.
LOESENER kept them therefore as distinct genera although VALETON /.c. had treated them already as
subgenera and LoeseNneER himself indicated Solenospermum macranthum LOEs. as an intermediate species.
I have examined most of the species described under those two genera and can not separate them by
any character without overlapping. The disks can be distinctly lobed, angular, or -- orbicular and may
vary in some species even in one specimen.

KuRz suggested to reduce Kokoona to Lophopetalum (J. As. Soc. Beng. 39, ii, 1870, 73) and said that
“the genus might be divided into 2 natural groups, the one with fimbriate or lamellate petals and large
flowers (Lophopetalum), the other with naked petals and small flowers (Kokoona)’’. However, the floral

and seed characters of Lophopetalum and Kokoona are quite different and they should be kept as two
distinct genera.

Sterile specimens, especially those from New Guinea, and detached fruiting material can hardly or not
be identified with certainty, to the species.

The bark is inflammable as in Kokoona.

The type species of the genus is L. wightianum ARN., as the second species mentioned with the generic
diagnosis, L. grandiflorum ARN., based on Euonymus grandiflorus WALL., is really a Euonymus; cf.
Starr & BALLARD in Curt. Bot. Mag. t. 9183 (1930).

BAILLON (Hist. Pl. 6, 1877, 3) regarded Lophopetalum and Glyptopetalum as sections of Euonymus,
but this lumping seems not justified.

KEY TO THE SPECIES

1. Flowers buds + flat or wider than high. Petals without appendage on the inner surfaces (except some
papilla-like ones sometimes occurring in L. beccarianum).
. Branchlets distinctly 4-angular. Leaves large, 17-38 by 514—12'4 cm. Inflorescences usually very
long, 18-45 cm. Pedicels 7—-15(-20) mm .. .... 1. L. sessilifolium
2. Branchlets usually terete. Leaves smaller, usually 4-151 by 2 oF cm. Inflorescences shorter, less
than 1314 cm long. Pedicels c. 2 mm.
3. Disk dish-shaped, distinctly 5-lobed, the edge turning slightly upward and forming a rim. Petiole
usually flat or furrowed above and round beneath, vascular strand appearing arc-shaped with the
arms bent upward and inward on a transverse section through the distal end.
4. Leaves often olivaceous above, rather yellowish green beneath when dry; apex usually obtuse,
acute, sometimes short-acuminate . . .... 2. L. beccarianum

4. Leaves often reddish brown on both Surtees! when dry. apex acuminate.
5. Petiole distinct, 5-8 mm. Inflorescences densely rusty puberulous on the young parts, usually

i)

Dec. 1962] CELASTRACEAE (Ding Hou) 265

glabrescent. Calyx patents at anthesis, distinctly larger than the disk, apex of the lobes usually

flat and obtuse . 5 3. L. floribundum

5. Petiole very short, c. 2 mm. ‘Inflorescences glabrous. ‘Apex of calyx lobes turning upcurved at
anthesis; calyx hardly larger than the disk; calyx lobes acuminate and + pointed.

4. L. glabrum

3. Disk + flat, suborbicular or obscurely 5-angular. Petiole usually round, vascular strand appearing
cylindric and enclosing 1-3 medullary bundles on a transverse section through the distal end.
(Leaves usually coriaceous, rigid, subsessile.) . oars .. 5. L. rigidum

. Flower buds short conical or subglobose, higher than wide or pisomeHines as s high as wide. Petals with

distinctly cristate, lamellate, or rarely with fimbriate appendages on the inner surfaces.

6. Disk 414-914 mm @ at anthesis.

7. Disk distinctly or sometimes obscurely 5-lobed or -angular, the lobes or angles epipetalous; anthers
apiculate. Flowers generally 10-18 mm @ at anthesis. Petiole usually round, sometimes furrowed
near the distal end above.

8. Petals slightly reniform, c. 6 by 814 mm, one small appendage at the base of each petal. Disk
7-914 mm @. Leaves coriaceous, usually covered with a layer of wax-like material above, densely
covered with papillae beneath. (Petiole usually inserted at the base of the leaf blade.)

6. L. pachyphyllum

8. Petals suborbicular, 3-4 mm in @, appendages distinctly cristate, lamellate, or rarely fimbriate,
usually 3-lobed and covering the upper 74 of the petal. Disk c. 5 mm 9. Leaves subcoriaceous, the
upper and lower surfaces not like above. (Leaves usually subpeltate.) . 7. L. wightianum

7. Disk + round, or sometimes obscurely 5-angular and the angles usually episepalous; anthers
acute or short-acuminate. Flowers c. | cm @ at anthesis. Petiole flat and slightly furrowed above,
round beneath . 5 8. L. macranthum

6. Disk smaller, 1—-3(-4) mm 9 at lauthesiss

9. Calyx almost divided to the base. Disk smooth. ate . . 9. L. micranthum

9. Calyx not divided to the base. Disk usually minutely papillose ¢ or “covered with fleshy, subulate
processes (smooth in L. forricellense and usually also in L. ledermannii).

10. Disk with fleshy, subulate processes around the base of the filaments. Leaves chartaceous to

thin-coriaceous, usually pale ash-coloured when dry 4 10. L. pallidum

10. Disk usually minutely papillose or smooth. Leaves usually coriaceous and brown to reddish

brown when dry.

11. Inner surface of the petal with a conspicuous, lobed appendage.
12. Petals smooth on the dorsal surface, sometimes thinner near the margin, the overlapping
margins just flatly pressed on the others. Nerves usually 5—8 (rarely more) pairs; veins reti-

culate, slightly elevated beneath, obscure above. (Petiole 1144-2 cm.) .

11. L. javanicum

12. The inner two or three petals with 2 distinct grooves on the dorsal surface, the overlapping
margins fitting in these grooves. Nerves usually 10-15 pairs; veins densely reticulate, elevated
beneath, usually obscure sometimes distinct above. (Petiole 114-3 cm.)

12. L. multinervium

11. Appendage small, simple, sometimes obscure on the upper surface of the petal.
13. Leaves ovate, elliptic, ovate-oblong to lanceolate, acute or acuminate.
14. Anthers twice as long as wide, acuminate. Disk papillose. Veins of the leaves usually invisible

or obscure beneath

13. L. ledermannii

14. Anthers slightly longer than wide, ‘obtuse, apiculate. Disk smooth. Veins of the leaves usually

distinct beneath,

sometimes on both surfaces

14. L. torricellense

13. Leaves mostly subobovate, apex blunt or slightly emarginate, short-apiculate, the pointed
part usually slightly concave and + sac-like. Anthers obtuse, about as long as wide. Disk

papillose

1. Lophopetalum sessilifolium RipL. Kew Bull.
(1931) 37.

Small tree up to 5 m. Branchlets distinctly
4-angular, sometimes slightly winged, reddish
brown. Leaves chartaceous, elliptic-oblong to
-lanceolate, 17-38 by 5!4-124% cm; base at-
tenuate, sometimes obtuse; apex acuminate;
nerves 10-14 pairs, slightly elevated on both
surfaces; petiole very short to obscure. Thyrses
profusely branched, sometimes almost from the

base, 18-45 cm. Peduncle very short, sometimes
up to 7 cm. Pedicels 7—15(-20) mm. Flower
buds flat, distinctly 5-angular, open flowers

patent, 7 or 8 mm 9g. Calyx lobes triangular, 114
by 1 mm, acuminate, the apex curved upward,

15. L. suboboyatum

margins short-fimbriate. Petals suborbicular or
deltoid, c. 3 mm long, acute or round at apex,
entire or sometimes slightly wavy. Disk distinctly
5-angular, acute or obtuse. Stamens small, c. 74 mm
long; anthers obtuse. Ovary almost immersed in
the disk. Fruits (young) c. 7 cm long, tuberculate,
stalk BD to 214 cm. Seeds (including wing) 514
by 114 cm, the wing c. 2 mm wide around the
seed proper. Lectotype HAvILAND 1744, K.

Distr. Malaysia: W. Borneo (Sarawak and
Sg. Landak).

Ecol. Once noted, peat-swamp forest (AN-
DERSON).

Vern. Lipeh, Sarawak.

266 FLORA MALESIANA [ser. I, vol. 6?
EE B a) Sigel oars mudah, Bajau, kapas, pérupok, sisilao, Sarawak,
i f kapas kapas, Sungai, kékan bukit, Brunei, kérupuk

f20 Vas SS

Fig. 11. Distribution of the genus Lophopetalum

WiGut ex ARN. For each subarea the number of

species has been indicated, endemic above the
hyphen, non-endemic below the hyphen.

2. Lophopetalum beccarianum Pierre, FI. For.
Coch. 4 (1894) sub t. 307; Merr. En. Born.
(1921) 354.—L. scortechinii Kinc, J. As. Soc.
Beng. ii (1896) 350; Rm. Fl. Mal. Pen. 1 (1922)
448.—L. havilandii RL. Kew Bull. (1931) 37.—
Fig. 10a-f.

Tree up to c. 30 m by 40 cm g. Leaves chartace-
ous to subcoriaceous, elliptic-oblong to -lanceolate,
oblong, sometimes ovate-oblong or lanceolate,
rarely broad-elliptic, obovate or obovate-oblong,
(7—-)11-1514(—30) by 314-7(-12) cm; base cuneate;
apex usually obtuse, acute, sometimes short-
acuminate; nerves 5-8 pairs, obliquely spreading
upwards, elevated and ridged beneath, depress-
ed sometimes plane above; petiole 34-114 cm.
Thyrses up to 18 cm long, (1—)2-5 in an axil,
sometimes branched quite from near the base,
usually glabrous, very rarely puberulous on the
young parts. Peduncle up to 4 cm, sometimes
very short. Pedicels c. 2 mm. Flowers yellowish,
c. 444 mm 9, patent. Calyx lobes triangular or
suborbicular, c. 1 mm long, entire sometimes
slightly erose, rarely short-fimbriate, sometimes
with small papillae on the outer surface. Petals
suborbicular or broad-ovate, 114-214 by 134-2
mm, naked, sometimes with villa-like appendages
on the inner side, rarely on both surfaces, some-
times with distinct venation. Disk rather thin,
dish-shaped, c. 2-3 mm 9g, 5-lobed, lobes obtuse
or rounded. Stamens short, sometimes the tissue
of the disk cushion-like thickened round the
base of the filament; anthers suborbicular, small,
c. ¥; mm long, obtuse, slightly acute, or very
short-apiculate; filaments usually reflexed after
anthesis. Pistil partly immersed in the disk, the
free part pyramidal, c. 1% mm; style and stigma
obscure. Ovules 3—4(-8) in each cell. Fruits 11 cm
long, pericarp leathery. Seeds (including the wing)
4% by 1% cm.

Distr. Malaysia: Malay Peninsula (Perak) and
Borneo (N. Borneo, Sarawak, Brunei, Bulungan,
W. Kutai, and Sangkulirang; Labuan I.).

Ecol. Primary forest, frequently on crests of
hills, on sandstone, sand, or tuff, once noted on
clay, from the lowland up to c. 400 m.

Vern. Borneo: aras, memagahas, Dusun,
bulalangabuk, Kujau, dual bukit, Kedayan, kandis

gunung, Nunukan I., sinlolosu, Tengara, winong
kora, Dusun & Tambato.

Notes. I have chosen ScoRTECHINI 1941, K,
as lectotype of L. scortechinii KING.

The leaf-tips of all the specimens available are
mostly damaged.

3. Lophopetalum floribundum WiGur, Ill. 1 (1840)
178; Laws. in Hook. f. Fl. Br. Ind. 1 (1875) 616.
—L. reflexum (non LAws.) KinG, J. As. Soc.
Beng. 65, ii (1896) 352, pro specim. Curtis 1502.

Tree 15 m tall, 40(—80) cm o. Leaves chartaceous
to thin-coriaceous, elliptic-oblong, or lanceolate,
acuminate, entire, 7-10 by 214-314(—S) cm; base
acute to attenuate; nerves 5—8 pairs, spreading and
gradually curving upwards, slightly elevated on
both surfaces; petiole 5-8 mm. Panicles solitary
sometimes branched from the base, up to 8 cm,
densely rusty puberulous (uniseriate, multi-
cellular hairs) when young, usually glabrescent;
peduncles O(-2) cm. Pedicels c. 2 mm. Calyx
usually patent at anthesis, lobes deltoid, c. 114 by
1144 mm, exceeding the disk lobes, ciliate. Petals
suborbicular, c. 3 mm 9, entire sometimes slightly
erose. Disk dish-shaped, c. 244 mm g, edge
slightly curved upward, 5-lobed, lobes deltoid
and obtuse. Stamens before the disk lobes, some-
times the tissue of the disk slightly depressed around
the base of the filaments; filaments c. 44 mm;
anthers broad-ovoid or suborbicular, c. 14 mm
long and wide, obtuse. Pistil partly immersed in
the disk, free part pyramidal, very short; style
and stigma obscure. Ovules 4-6 in each cell.
Fruits c. 8 cm long; pericarp rather thin, c. 1 mm
@. Seeds including wing c. 6 by 1 44 cm, seed proper
c. 4 by 24 cm.

Distr. Burma (Mergui: GriFFITH 620, 864, K)
and Malaysia: Malay Peninsula (Penang I.; once
in Pahang).

Ecol. Rain-forests, 150-360 m.

Vern. Kongkor, Pahang.

4. Lophopetalum glabrum DinG Hou, nov. sp.

Arbor parva, c. 6 m alta. Folia chartacea,
elliptico-oblonga, 8-13 cm longa, 34, cm
lata, basi anguste cuneata, apice acuminata,
margine integra, nervis utrinque S—7, venulis laxe
reticulatis; petiolus c. 2 mm longus. Inflorescentiae
usque ad 9 cm longae, glabrae. Pedicellus 5-7 mm
longus. Flores flavescentes. Calyx 5-lobatus, lobis
deltoideis, c. 1 mm longis, acuminatis. Petala
subrotunda, c. 2 mm longa. Discus patelliformis,
5-lobus. Stamina breviora. Pistillum pyramidatum.
Ovula in loculis 4.—Typus RUTTEN 83, U, isotypes
Bo.

Small tree c. 6 m by 20 cm g. Branchlets terete
sometimes slightly angular. Leaves chartaceous,
elliptic-oblong, 8-134 by 3-414 cm; base narrow-
ly cuneate; apex acuminate; nerves 5-7 pairs,
elevated below and slightly depressed above,
obliquely spreading towards near the margin and
then turning upward; petiole very short, c. 2 mm,
furrowed above, round beneath. Panicles up to

Dec. 1962]

CELASTRACEAE (Ding Hou)

267

9 cm long, usually branched from the base.
Peduncle very short. Pedicels 5-7 mm. Flowers
yellowish. Calyx almost as large as or slightly
exceeding the disk; lobes deltoid, c. 1 mm long
and wide; margins short-fimbriate, apex acuminate,
+ pointed and turning upwards at anthesis.
Petals suborbicular, c. 2 by 2 mm, entire some-
times slightly erose. Disk dish-shaped, the margin
slightly curved upwards, 5-lobed, lobes deltoid,
c. 1 mm long and wide. Stamens very short; an-
thers suborbicular, c. 14 mm g, obtuse. Pistil c.
14 mm emerging from the disk, pyramidal; style
and stigma obscure. Ovules 4 in each cell.

Distr. Malaysia: Borneo (Brunei; E. Borneo;
Bulungan).

Ecol. On ridge, lowland forest.

5. Lophopetalum rigidum Ript. Kew Bull. (1931)
38.—L. subsessile RD. /.c. 37.—Fig. 10 m.
Small tree up to 12 m by 10 cm 9, sometimes
with swollen stem-base (in swamp!), bark smooth.
Branchlets terete, sometimes slightly angular or
compressed, light to dark-brown, usually swollen
at the node. Leaves usually coriaceous, rigid, rarely
subcoriaceous, ovate-oblong, 414-124 by 2-6
em; base cuneate or obtuse; apex acuminate;
nerves 4-11 pairs, elevated or flat on both sur-
faces, sometimes obscure beneath, obliquely
spreading and slightly curved to the margin;
petiole short, c. 3(-8) mm, + terete, stout.

Panicles up to 10 cm, usually stout, solitary, some-
times branched almost from near the base.
Peduncle very short, sometimes up to 314 cm.
Pedicels 2-3 mm. Flowers yellowish green, 31-6
mm g. Calyx lobes and petals with small, papilla-
like processes outside. Calyx lobes ovate or
triangular, c. 114-114 by 1 mm, apex acuminate,
++ pointed and turning upward at + right angles,
short-ciliate. Petals triangular or suborbicular,
1-214 by 1-2 mm. Stamens c. 1 mm long; anthers
small, very short-apiculate. Disk -++ flat, sub-
orbicular or obscurely 5-angular, 114-342 mm 9,
smooth, or sometimes slightly tuberculate on the
upper surface. Pistil c. 1 mm emerging from the
disk, pyramidal, at the apex contracted into a
short but distinct cylindrical style. Ovules 4(—6)
in each cell. Fruits up to 11 cm long, short-
tuberculate on the outer surface. Seeds including
wing 514 by 114 cm.

Distr. Malaysia: Borneo (N. Borneo, Brunei,
Sarawak, and Buntok).

Ecol. Understorey tree of freshwater-swamp
forests, also on dryland, in heath forest; all in the
lowland, thrice found on hill top at 650, 1000, and
1380 m.

Vern.
Dyak.

Galagah, parupuk, Born., kérupok,

6. Lophopetalum pachyphyllum Kina, J. As. Soc.
Beng. 65, ii (1896) 348; Ript. Fl. Mal. Pen. 1
(1922) 448; Logs. in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 162.

In general small buttressed tree, up to 36 m
by 40 cm g. Bark greyish white, fissured. Branchlets
terete. Leaves coriaceous, greyish waxy above,

pale olivaceous beneath by a dense cover of
minute unicellular papillae, ovate-, sometimes
elliptic-oblong, 11-17 by 5-10 cm; base obtuse or
rounded; apex acute to short-acuminate, usually
damaged in the herb.; margin cartilaginous and
slightly recurved; nerves 7-9 pairs, obliquely
curving and ascending; petiolel—2 cm, almost terete,
not grooved above; blade sometimes subpeltate.
Panicles up to 21 cm long, 1(—2-3) in an axil.
Peduncle very short, sometimes up to 6 cm.
Pedicels 8-15 mm. Flowers yellowish, large, c.
18 mm @. Calyx almost as large as the disk,
slightly lobed, lobes round. Petals reniform, c.
6 by 814 mm, entire, the appendage at the base
inside, small, sometimes obscure. Disk -+ or-
bicular, slightly convex towards the center near
the ovary, 7-91, mm go, obscurely 5-lobed.
Stamens inserted near the ovary; filaments c.
2144 mm; anthers ellipsoid or slightly ovoid, 114
by 1 mm, slightly apiculate. Ovary emerging c.
2 mm from the disk, narrowed towards the apex;
ovules 16 in each cell.

Distr. Malaysia: Sumatra (Djambi) and Malay
Peninsula (Perak, Dindings, Selangor, Negri
Sembilan, and Johore).

Ecol. Dryland forests, on hill slopes, and
limestone cliffs, from the lowland up to 450 m.

Vern. Térupuk, M.

Note. As lectotype I have selected KING’s
coll. 7525, L (by error cited as 7325).

7. Lophopetalum wightianum ARN. Ann. Nat.
Hist. 3 (1839) 151; WiGur, Ic. 1 (1839) t. 162;
Ill. 1 (1840) 178; Bepp. Fl. Sylv. 1 (1869) t. 145;
Laws. in Hook. f. Fl. Br. Ind. 1 (1875) 615;
Prerre, Fl. For. Coch. 4 (1894) t. 307B, incl. var.
macrocarpum PIERRE; PITARD, Fl. Gén. I.-C. 1
(1912) 876, f. 109; Logs. in E. & P. Pfl. Fam. ed.
2, 20b (1942) 161, f. 45.—L. fimbriatum WicuHrt,
Ill. 1 (1840) 178; Laws. in Hook. f. Fl. Br. Ind.
1 (1875) 615; KiNG, J. As. Soc. Beng. 65, ii (1896)
349; Ripi. Fl. Mal. Pen. 1 (1922) 448; Logs.
in E. & P. Pfl. Fam. ed. 2, 20b (1942) 162; Tar-
DIEU, Suppl. Fl. Gén. I.-C. (1948) 807; Not.
Syst. 14 (1950) 48.—Euonymus fimbriatus BALL.
ex Laness. Pl. Util. Col. Frang. (1886) 312.—
L. winkleri Lors. Notizbl. Berl.-Dahl. 13 (1936)
221; in E. & P. Pfl. Fam. ed. 2, 20b (1942) 162.

Tree up to 50 m by 65(-195) cm go, sometimes
buttressed. Branchlets terete, sometimes slightly
4-angular. Leaves subcoriaceous, often elliptic to
elliptic-oblong, sometimes ovate to ovate-oblong,
rarely obovate or broad-obovate, 8-25 by 4-10
cm; base obtuse to rounded, sometimes cuneate,
usually subpeltate; apex acute, sometimes acumin-
ate (acumen up to | cm), rarely apiculate, usually
dried and breaking off in the herb.; nerves 6—12
pairs; petiole terete, rarely sulcate above near the
distal end, 114-214 cm. Panicles up to 12 cm long,
sometimes branching from near the base, branch-
lets rather stout, obliquely spreading. Peduncle
obscure, sometimes up to 7 cm. Pedicels S—9 mm.
Flowers 10-13 mm @. Calyx almost as large as the
disk, distinctly 5-lobed, lobes triangular, acute,
short-ciliate. Peta/s broadly ovate or suborbicular

268

FLORA MALESIANA

[ser. I, vol. 6?

with undulating margins, yellow, 3-4 mm long
and wide, appendage transverse cristate or la-
mellate, rarely fimbriate, usually manifest (some-
times small especially on the specimens from
Borneo), attached at the lower half or sometimes
at about the middle, sometimes divided into 3,
and the middle one almost reaching the apex of
petals. Disk red, concave and cup-shaped in the
bud, flat or dish-shaped at anthesis, c. 5 mm g, dis-
tinctly 5-lobed to almost orbicular, lobes epi-
petalous. Stamens inserted near the ovary; fi-
laments c. 244 mm; anthers oblong or slightly
ovoid, apiculate, c. 144 by | mm. Pistil red, c.
2 mm emerging from the disk. Ovary triangular
and narrowed upwards into the style; ovules 12-18
in each cell. Fruits 10-15 cm long. Seeds (incl.
wing) 644 by 114 cm, seed proper 434 by 1 cm.

Distr. India (Deccan, from the Concan south-
wards to Cape Comorin, and Assam), E. Pa-
kistan (Chittagong), Burma (Tenasserim, Am-
herst), Cochinchina, and Malaysia: Sumatra
(Palembang and Banka), Malay Peninsula (Kedah
Dindings, Selangor, Negri Sembilan, Malacca,
Penang, Langkawi, and Singapore), and Borneo
(E. Kutai, K. Panya, Pleihari, and Hayup).

Ecol. Everwet lowland forests, in riverine
forest, temporarily inundated forests, sometimes
near the shore just behind high-water mark, but
also on sandy dryland and coral limestone; in
India ascending to 900 m (BEDDOME, /.c.).

Vern. Sumatra: bau langit, East Coast, pérupuk,
p. talang, térupuk talang, M, trupuk, Banka;
Mal. Pen.: kéruié, Lakai, mata ulat, Kedah,
médang assam, Malacca, pérupok, Langkawi;
Borneo: bura, Dyak, nasi-nasi, Brunei, pérupuk
unung, M.

Notes. L. fimbriatum, based on GRIFFITH'S
collection from Mergui, Burma, would be distinct
from L. wightianum by the thinner and membranous
leaves, the fimbriated crest of the petals, as well
as the flowers being scarcely half the size. In the
authentic material of L. fimbriatum (K) the in-
florescences have mostly flower buds with a few
open flowers which are rather wrinkled. From the
specimens available, the differential characters
indicated by WiGuT are variable and intermediate
forms are commonly found sometimes even on
the same specimen.

Specimens from India, Burma, and Indo-China
usually have disks with distinctly broad-oblong
lobes and petals bearing manifest appendages,
while those from Borneo usually have the disk
obscurely lobed, angular, or + orbicular and
petals with sometimes small and obscure appen-
dages. Judging from the variations and intermediate
forms of the specimens available, the three names
mentioned above belong to one widely distributed
species.

8. Lophopetalum macranthum (Logs.) DinG Hou,
comb. noy.—Solenospermum macranthum Logs.
Notizbl. Berl.-Dahl. 13 (1936) 222; in E. & P.
Pfil. Fam: ed. 2, 20b (1942) 162.—Fig. 10g-i.
Tree up to 12 m. Branchlets terete, sometimes
subterete. Leaves coriaceous, ovate-oblong, rarely

oblong-lanceolate, 9-16 by 4-6 cm; base obtuse or
-+ rounded; apex obtuse, or obscurely short-
acuminate; nerves 6—7 pairs, obliquely ascending;
veins reticulate, prominent on both surfaces; petiole
7-10 mm. Panicles solitary, up to 18 cm long,
divaricate, lax, branched almost from the base.
Peduncle very short. Pedicels 5-6 mm. Flowers
yellowish white, fragrant, c. 1 cm @. Calyx
lobes semi-orbicular, or + reniform, 1 by 214 mm,
+ entire. Petals ovate or broadly ovate, sometimes
even suborbicular, 4414 by 3-4 mm, wavy on the
margin, appendage very small, inserted at the
middle or lower half, triangular with a very broad
base. Disk fleshy, -- rounded to obscurely 5-
angular, the obscure angles episepalous or epi-
petalous, c. 4144 mm 9, smooth. Filaments inserted
+ between pistil and edge of disk, c. 244 mm;
anthers broad-ovoid, 114 by 4/5 mm, acute or
short-acuminate. Pistil emerging c. 2 mm above
the disk. Ovary slightly 3-angular, narrowed into
a cylindric style. Ovules 8 in each cell.

Distr. Malaysia: New Guinea (Hollandia),
once collected; possibly a few sterile sheets from
Papua.

Ecol. In lowland rain-forest on hill, 25 m.

9. Lophopetalum micranthum Logs. Nova Guinea
8 (1910) 279.—Solenospermum micranthum Logs.
Notizbl. Berl.-Dahl. 13 (1936) 225; in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 162.

Tree. Branchlets terete. Leaves coriaceous,
obovate to slightly obovate-oblong, rarely elliptic-
oblong, 3-614 by 114-344 cm; base cuneate to
attenuate; apex acute; nerves 5—7; veins slightly
elevated beneath, invisible above; petiole c.
2 mm. Panicles up to 3 cm long, few-flowered.
Peduncle short, sometimes up to 1 cm. Pedicel
c. 144 mm. Flower yellowish green. Calyx almost
divided to the base, lobes deltoid or semi-orbi-
cular, 1 by 1-114 mm, obtuse, short-fimbriate.
Petals ovate to broadly ovate, 124-2 by 14-114
mm, acute or obtuse, + entire, appendage simple,
small. Disk fleshy, -- orbicular or obscurely
obtuse-5-angular, c. 144 mm o. Stamens inserted
on the edge of disk; anthers ovoid, c. 4/5 by 44 mm,
obtuse or very short-apiculate; filaments c. 5 mm.
Pistil emerging c. | mm from the disk; ovary
triangular at base, narrowed into a short style;
stigma obscure. Ovu/es 8 in each cell. Fruit
unknown.

Distr. Malaysia: New Guinea (Hellwig Mts:
Nepenthes Hill), once found, possibly also Mt
Arfak at 2200 m.

10. Lophopetalum pallidum Laws. in Hook. f.
Fl. Br. Ind. 1 (1875) 615; Rip. Fl. Mal. Pen. 1
(1922) 449, incl. var. curtisii (KING) RIDL.—
L. curtisii KiNG, J. As. Soc. Beng. 65, ii (1896)
351.—Solenospermum pallidum Loes. Notizbl.
Berl.-Dahl. 13 (1936) 225.—Fig. 10j-1.

Tree up to c. 40 m by 60 cm og, sometimes
buttressed. Branchlets terete. Leaves chartaceous
to thin-coriaceous, ash-coloured above, elliptic
to elliptic-oblong, sometimes ovate to ovate-
oblong, 7-12 by 3144-54 cm; base obtuse or

Dec. 1962]

CELASTRACEAE (Ding Hou)

269

cuneate; apex acute to short-acuminate; nerves
8-10 pairs; petiole 114-2 cm, grooved above.
Panicles sometimes extra-axillary, 3-7 cm long;
peduncle 1-3 cm. Pedicels 244-3 mm. Flowers
yellow. Calyx lobes semi-orbicular, reniform, or
triangular, 1 by 114-2 mm, entire. Petals deltoid,
214-3 by 2!4-3 mm, entire or slightly erose, with
fleshy, lobed processes + in the central part.
Stamens inserted between pistil and disk margin in
pits surrounded by fleshy, subulate processes
(4-4 the length of the filament); filaments
2/4114 mm; anthers broadly ovoid, 74-1 mm long,
acute. Disk dish-shaped, red, 5-angular, obtuse,
3-4 mm @g, usually covered with papillae. Pistil
triangular, emerging c. 1 mm from the disk,
gradually narrowed into a cylindric, short style.
Oyules 10 in each cell. Fruits c. 154% cm long;
pericarp hard, c. 4 cm thick. Seeds including wing
9 by 214 cm, seed proper 514 by 1 cm.

Distr. Malaysia: S. Sumatra (Palembang),
Malay Peninsula (Kedah, Selangor, Negri Sem-
bilan, Malacca, and Penang) and Borneo (Sang-
gau, Pasir, and Balikpapan).

Ecol. Dryland primary rain-forest, often on
hillsides or in sandy country, lowland up to 200 m.

Vern. Dorojolang, Pasir, kéroi, Temuan,
kéruie, Lakai, kélémpait, kérueh, manau, pélantan
gunung, tiada tahu, tjangel, M.

Use. Used as dart poison ingredient in Malaya.

Notes. I have examined both the types of L.
pallidum (MaiNGAy 393, K) and L. curtisii
(Curtis 1577, K) in the Kew Herbarium and there
is no doubt that they are conspecific.

The most characteristic characters of this species
are the fleshy, subulate processes around the base
of the filaments and the ash-coloured leaves.

It is quite closely related to the continental SE.
Asian L. celastroides LAws. and L. wallichii KURZ.
In these the disk is entirely covered with fleshy,
subulate processes and the leaves are wider, ovate
or broad-elliptic and usually brown to reddish
brown when dry. I have seen both the types of L.
celastroides and L. wallichii and several other
flowering specimens in the Kew Herbarium collect-
ed in Burma, Siam, and Indo-China; they are
similar to each other, and in my opinion L.
celastroides Laws. should be reduced to L.
wallichii Kurz.

11. Lophopetalum javyanicum (ZoLL.) Turcz. Bull.
Soc. Nat. Hist. Mosc. 36, i (1863) 598, as javanum;
K. & V. Bijdr. 7 (1900) 95; VALETON, Ic. Bog. 1
(1901) 43, t. 90; Koorp. Exk. Fl. Java 2 (1912)
§23.—Solenospermum javanicum ZOLL. Nat. Tijd.
Ned. Ind. 14 (1857) 169; Logs. Notizbl. Berl.-
Dahl. 13 (1936) 223; in E. & P. Pfil. Fam. ed. 2,
20b (1942) 162, f. 46.—L. fuscescens Kurz, J.
As. Soc. Beng. 44, ii (1875) 202, ex descr.; KiNG, J.
As. Soc. Beng. 65, ii (1896) 352; RipL. Fl. Mal.
Pen. 1 (1922) 448.—L. fimbriatum (non W1GHT)
F.-ViLL. Noy. App. (1880) 46; VIDAL, Sinopsis
(1883) 20, t. 31, f. G.—Hippocratea maingayi
(non LAws.) VIDAL, Lc. f. F.—L. oblongum KING,
J. As. Soc. Beng. 65, ii (1896) 350; RipL. Fl. Mal.
Pen. 1 (1922) 448; Craip, Fl. Siam. En. 1 (1926)

Fig. 12. Lophopetalum sp. Habit of a large tree
(Palembang, THORENAAR, 1924).

270

FLORA MALESIANA

[ser. I, vol. 6?

282.—L. oblongifolium KiNG, J. As. Soc. Beng.
65, ii (1896) 351; RipL. Fl. Mal. Pen. 1 (1922)
449: Logs. Notizbl. Berl.-Dahl. 13 (1936) 225.—
L. toxicum Loner, Ic. Bog. 1 (1897) 55, t. 16;
Merr. En. Philip. 2 (1923) 481.—L. celebicus
Koorp. Minah. (1898) 623.—L. intermedium
RIpL. Fl. Mal. Pen. 1 (1922) 449.—L. paucinervium
Me_rr. Philip. J. Sc. 20 (1922) 402.—Solenosper-
mum paucinervium Lors. Notizbl. Berl.-Dahl. 13
(1936) 223.—Solenospermum toxicum Loes. lI.c.;
in E. & P. Pfl. Fam. ed. 2, 20b (1942) 162.—
Solenospermum oblongifolium Loes. in E. & P.
Pfi. Fam. ed. 2, 20b (1942) 162.

Tree up to 45 by 1 m @, sometimes slightly
buttressed when growing in swamps. Leaves
subcoriaceous to coriaceous, elliptic-oblong or
elliptic, ovate-oblong to lanceolate, rarely obovate
or ovate, 514-18 by 214-10 cm; base acute to
cuneate, sometimes obtuse; apex acute to short
acuminate, very rarely obtuse and apiculate; nerves
5-8 pairs (very rarely more); petiole 114-2 cm.
Panicles up to 19 cm, sometimes branched almost
from the very base; occasionally furfuraceous,
glabrescent; peduncle very short, sometimes up to
414 cm. Pedicels 3-314 mm. Flowers white, light
green or yellowish green, c. 9 mm @. Calyx lobes
almost patent or sometimes the apices curved
upwards, rarely reflexed, slightly or c. 4% mm
exceeding the margin of the disk lobes, acute,
short ciliate. Petals broad-ovate, or -oblong,
2-3 by 134-214 mm, obtuse or slightly erose, some-
times slightly fimbriate at the apex, appendages
lobed or dentate, usually -+- 3-lobed, the central
lobe often prominent and longer than the lateral
ones. Disk fleshy, thick, flat sometimes concave in
the bud, obscurely 5-angular or + rounded, 214-3
(-4) mm g, usually minutely papillose on the
upper surface. Stamens inserted between ovary
and edge of disk or sometimes quite near the
margin; filaments c. 11 mm; anthers broad-
ovoid, acuminate, c. | by 24 mm, apiculate.
Pistil emerging c. 114 mm from the disk. Ovary
slightly triangular, narrowed into a cylindric
style; stigma obscure. Ovules S—8 in each cell.
Fruits 644-11 cm long; pericarp leathery, rather
thin, smooth or sometimes minutely tuberculate.
Seeds with a wing c. 6! by 114 cm.

Distr. Siam (Puket, sec. CRAiB) and Malaysia:
Sumatra (Tapus, West Coast, Palembang, Lam-
pongs, Simalur I., Nias, Banka, and Riouw),
Malay Peninsula (Perak, Kedah, and Penang),
Java (rare, mainly Central Java), Borneo (com-
mon), Philippines (Luzon, Mindoro, Samar, Sulu
Is., and Palawan), Celebes (Muna, Minahassa,
Malili, Bone, Aanaiwoi), Moluccas (Morotai and
Taliabu Is.), and New Guinea (Japen and Hol-
landia).

Ecol. Usually in dryland rain-forests in the
lowland, several collections found on loam soil
and limestone rocks, rarely in temporarily in-
undated forest or peat swamp, sometimes also
found at higher altitudes, e.g. 1000 m (Central
Java) and 1500 m (Mt Kinabalu).

Vern. Sumatra: béhu, Nias,
bungo, téradih uding, Simalur,

bintol langsa
dilau— rumba,

émpidingan, kumbang, marpitis, M, madang-
gambici, Batak, madan landjut, Pariaman; Mal.
Pen.: kachang rimba, Kedah; Java: mandalaksa,
J; Borneo: médang bora, m. tolei, Balikpapan,
agar-agar, dampal, djéréndjang, pérupuk, pp.
gunung, séraya puteh, takau, tulang, M, dual,
Dusun & Kedayan, kaju api, E. Kutei, kayu
malam pérampuan, ranau, Sungei, kédjé woos,
kétapang, maréndai, Kutei, bura, maratémon,
péndjolawat, sang, Dyak, pisang pisang, tapatung,
Dusun, mémagahar, tanggom apui, N. Born.;
Philip.: sampol, Bis., abtab, abitab, bokbok,
butifigi, dayanddng, Tag., buyun, Sul., dita, Neg.,
kalibambangan, Mand., puti-i-babde, puti-i-lalaki,
Lan., sudkad, P. Bis.; Celebes: bongkorio, kulilawa
puté, Muna, kabalo, Malili, kaléleng, Bone,
totorintek, Minahasa; Moluccas: mému, Sula;
New Guinea: ra ai, Ambai, tatokwa, Hollandia.

Notes. ZOLLINGER cited two collections 779
(non vidi) and 3254 (lectotype G, Fi); TuRCZANI-
Now cited the latter number by error as 3654.

The inflorescences of some specimens show
witches’ broom malformations, e.g. KOSTERMANS
9580.

12. Lophopetalum multinervium RipL. Kew Bull.
(1931) 39.—Lophopetalum sp., THORENAAR, Trop.
Natuur 16 (1927) 76; ENpERT, Verslag M.O.
Born. Exp. (1927) 221.—Solenospermum aquatile
RipL. Kew Bull. (1938) 236.—Fig. 13.

Tree up to 45 m by 70 cm o. Buttresses up to
80 cm high, extending 8 m over the ground, some-
times with knee-roots. Leaves coriaceous, ovate to
ovate-oblong, elliptic to elliptic-oblong, 1014-18
by 414-8 cm; base obtuse or rounded, sometimes
cuneate; apex acuminate; nerves 10-15 pairs;
petiole (1—)134-3 cm. Panicles up to 12 cm long,
sometimes scurfy on the young parts. Peduncle
up to 6 cm long. Pedicels 3!,—4 mm. Flowers light
or yellowish green, rarely white, c. 7 mm @.
Calyx triangular, 34 by 144 mm, acute, short-
ciliate. Petals broad-ovate, 24-3 by 134-2 mm,
acuminate, appendage with broad base inserted
at the lower 1, lobes rather small, the inner two
or three always with 2 grooves on the dorsal side.
Disk fleshy, flat, c. 3 mm 9, obscurely S-angular,
densely covered with minute papillae. Stamens
inserted between the ovary and the disk margin;
filaments c. 134 mm; anthers broad-ellipsoid
or -ovoid, 1 by 24 mm. Pistil emerging c. 144 mm
from the disk. Fruits 7-814 cm long; pericarp
leathery, hard, furfuraceous outside. Seeds (in-
cluding the wing) c. 5 by 114 cm.

Distr. Malaysia: Sumatra (East Coast, In-
dragiri, Riouw and Palembang), Banka, Malay
Peninsula (Perak, Pahang and Singapore), and
Borneo (common).

Ecol. In lowland forests, usually occurring in
peat swamps (on shallow peat) and inundated
forest, rarely found at higher altitude (Mt Ki-
nabalu at 1350-1500 m).

Galls. There are small, club-shaped or ellipsoid
galls, c. 1 mm long, occurring on the upper sur-
faces of the leaves; sometimes a few subglobose,

Dec. 1962]

CELASTRACEAE (Ding Hou)

ail

fruit-like galls c. 1 cm o are found on the in-
florescences.

Vern. Sumatra: Pérupuk, p. talang, Palembang,
pupu, Bengkalis; Borneo: bako, Dyak, dual, N.
Born., pasana, pérupuk, M, Iban, & Banka.

Note. The type of Solenospermum aquatile,
MoTLey 861 (K), has been recorded as growing in
water (cf. RipL. 1938, /.c.). The specimen is a
rather young branchlet and has rather young
flowers and fruits. In addition to the ecological
habit, it matches L. multinervium very well.

Fig. 13. Lophopetalum multinervium RipL. Pneu-
matophores in swamp forest of Sugut For. Res.,

c. 39 miles N of Sandakan (MeEwer, 1961).

13. Lophopetalum ledermannii (Loes.) DiInG Hou,
nov. comb.—Solenospermum ledermannii Logs.
Notizbl. Berl.-Dahl. 13 (1936) 224; in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 162.

Tree up to 26 m by 56 cm g. Branchlets terete.
Leaves coriaceous, rigid, opaque, ovate-oblong to
lanceolate, sometimes ovate, 5-10 by 214-414 cm;
base acute or obtuse; apex acuminate or acute;
nerves 5-8 pairs; petiole 7-10 mm. Panicles up to
9 cm, solitary, branched almost from the base.

Peduncle very short, sometimes up to 114 cm.
Pedicels 2-3 mm. Flowers white, 6-7 mm o@.
Calyx patent or slightly reflexed in bud, lobes
deltoid to broad-triangular, 4% by 4-114 mm,
acute or obtuse, sparsely, very short-fimbriate.
Petals ovate or deltoid, 2—3 by 134—2 mm, obtuse;
appendage small, lanceolate, in the centre. Sta-
mens inserted usually between the edge of the
disk and the base of the ovary, sometimes quite
close to the margin; anthers ovoid-oblong, 1 by 14
mm, short-acuminate; filaments c. 24 mm. Disk

rounded, fleshy, c. 2 mm @, smooth, sometimes
with sparse papillae. Pistil pyramid, 1-14 mm
above the disk; style and stigma obscure. Ovules
7-8 in each cell. Fruits unknown.

Distr. Malaysia: Moluccas (Morotai) and
New Guinea (also in Mios Noem and Japen I.).

Ecol. Primary, rarely secondary forests, from
the lowland up to 850 m.

Vern. Sewaidjakas, Manikiong, sidomokoe,
Galela, tenggarenop, Papua, wajarora, Wanapi,
weekal or weekar, Tor.

Note. There are three specimens cited in the
original description of which I have selected
DocTEeRS VAN LEEUWEN 9622 as lectotype (L;
isotypes Bo & K).

14. Lophopetalum torricellense Logs. in K. Sch. &
Laut. Nachtr. (1905) 303.—Solenospermum tor-
ricellense Lors. Notizbl. Berl.-Dahl. 13 (1936)
224, ex descr., incl. var. opacum Logs.; in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 162.

Tree up to 29 m by 36 cm o. Bark greyish,
fairly rough. Branchlets terete sometimes slightly
4-angular. Leaves coriaceous, rigid, opaque, ovate,
or broad-ovate, rarely elliptic-oblong, or obovate,
(3-)414-8 by (1144-)24%4-414 cm; apex acute to
short acuminate, rarely obtuse; base obtuse,
cuneate, or round; nerves 4-6 pairs; petiole 3-7
mm. Panicles solitary. Peduncle very short, some-
times up to 8 mm. Pedicels 3-4 mm. Flowers
yellowish green, 6-7 mm @. Calyx lobes triangular,
13-14 by 1 mm, short-fimbriate, in bud the lobes
patent sometimes slightly inflexed at the apex.
Petals triangular 244 by 124 mm, obtuse; append-
ages very small, inserted almost at the centre.
Stamens c. 1‘ mm, inserted almost on the margin
of the disk; anthers broad-ellipsoid or -ovoid,
4/, by 24 mm, obtuse and short-apiculate. Disk
+ rounded, fleshy, c. 2mm @, smooth, sometimes
sparsely short-papillose. Pistil emerging c. 1 mm
from the disk. Ovules 6 in each cell. Fruits 6-8 cm
long. Seeds (incl. wing) 34% by 1 cm.

Distr. Malaysia: North New Guinea.

Ecol. Primary forests usually c. 1000-2200 m,
rarely occurring at 600 m.

Vern. Hemouw, Manikiong.

Note. The available specimens (BRAss &
VERSTEEGH 11901, 11905, 12564; Brass 12326) are
very homogeneous. They match very well the
isotype of Solenospermum torricellense var. opacum
(LEDERMANN 10059, L) and agree also with the
description of Lophopetalum torricellense (type:
SCHLECHTER 14507, not seen).

272

FLORA MALESIANA

[ser Ie vollnG?

15. Lophopetalum subobovatum Kina, J. As. Soc.
Beng. 65, ii (1896) 349, as ‘sub-obovatum’; RIDL.
Fl. Mal. Pen. 1 (1922) 448.—Solenospermum
apiculatum RipL. Kew Bull. (1938) 235.

Tree up to 39 m by 70 cm o. Bark rough,
fissured, greyish sometimes dark-brown. Branch-
lets terete. Leaves coriaceous, subobovate some-
times elliptic, 444-13 by 214-7 cm; base cuneate;
apex obtuse and apiculate, the pointed part usually
slightly folded upwards and_ shallow-sac-like,
sometimes obscurely acute, rarely notched; nerves
4-6 pairs; petiole 14-114 cm. Panicles solitary, up
to 10 cm. Peduncle very short, sometimes up to

usually covered with minute papillae, c. 3 mm g.
Stamens inserted between pistil and edge of disk;
filaments c. 1144 mm; anthers slightly deltoid, c.
2, mm long and wide, obtuse. Pistil c. 144 mm
emerging from the disk, triangular at the base,
gradually narrowed into a cylindric style. Ovules
6 in each cell. Fruits c. 7 cm long, usually minutely
tuberculate. Seeds including the wing 434-6 by
1-114 cm.

Distr. Malaysia: Sumatra (Riouw, Indragiri),
Malay Peninsula (Penang, Pahang, and Johore),
and Borneo (common).

Ecol. Dryland lowland rain-forests, below 400

c. 4 cm. Pedicels 3-6 mm. Flowers white. Calyx m, twice recorded from 600-700 m in North
lobes patent or the apex slightly curved upwards, __ Borneo.

triangular, 14-24 by 1 mm, slightly ciliate. Petals Vern. Duol, dual, Brunei, kadjo, Dyak,
triangular, or slightly ovate-oblong, 3-314 by kungkur, mata ulat, paropo, pérupok, tjauto putih,
2-214 mm, slightly erose, appendage rather small, M.

at the upper half. Disk slightly angular, fleshy,

8. MICROTROPIS

WALL. [Cat. (1830) n. 4337-40, nomen] ex MEISN. Pl. Vasc. Gen. Tabul. Diagn.
(1837) 68, nom. gen. cons. prop.; Comment. (1837) 49, non E. Mey. 1836; cf.
Taxon 11 (1962) 226; MerR. & FREEM. Proc. Am. Ac. Arts Sc. 73 (1940) 276; Logs.
in E. & P. Pfl. Fam. ed. 2, 20b (1942) 126.—Microtropia ReicHB. Nomencl.
(1841) 190.—Paracelastrus Mig. Fl. Ind. Bat. 1, 2 (1859) 590; F. N. WILLIAMs,
Bull. Herb. Boiss. II, 5 (1905) 224.—Fig. 14.

Shrubs or small trees. Terminal node of each flush usually with one, sometimes
two pairs of subpersistent incipient bracts. Stipules minute, early caducous, or
exstipulate (?). Leaves decussate or opposite, glabrous (except in one extra-Mal.
sp.), entire. Flowers sessile or subsessile, in axillary or extra-axillary dichotomous
or paniculate cymes, sometimes condensed to sessile clusters, at the base of the
flush, bisexual (sometimes unisexual by abortion in extra-Mal. spp.), 5- or 4-
merous. Calyx deeply lobed, lobes almost free, persistent, imbricate, unequal in
size, the outer 2 or 3 usually smallest. Petals slightly united at the base, some-
times free (M. bivalvis), imbricate, erect. Stamens usually dorsifixed; filaments
subulate, usually united at the base into (? inserted on) a ring or short tube (the
ring sometimes interpreted as a disk) (except in Mal. in M. tenuis and M. te-
trameris), the united part free from the petals or sometimes adnate to them (in
M. tenuis stamens even inserted in the mouth of the corolla); anthers broad-oyvoid
or ovoid, introrse, rarely extrorse (M. discolor). Ovary free, completely or in-
completely 2-celled. Ovules 2 in each cell, erect, collaterally attached to the axis
towards the base at the inner angle (see below); style very short or cylindric;
stigma obscure, or discoid, sometimes slightly 2-4-lobed. Capsule short-apiculate
to beaked, rarely obtuse, lengthwise striate, laterally split along one side, sustained
by a persistent calyx. Seed usually 1, erect, on a knob-like thickened placenta,
albuminous, enveloped by the aril, usually wrinkled, testa similar to aril, smooth,

soft, red or red-brown.

Distr. About 70 spp., distributed in Central America (4 spp.), SEE. Asia, and Malaysia,
from Ceylon and India (Deccan Peninsula, Silhet, Assam) eastward to S. China (Yunnan, Szechuan,
Kwantung, Fukien), Hainan, Formosa, central Japan, Riukiu, southward through Burma, Siam and
Indo-China to Malaysia (Sumatra, Malay Peninsula, West Java (once found), Borneo, and the Phi-
lippines). Fig. 15.

Dec. 1962] CELASTRACEAE (Ding Hou) 273

Fig. 14. Microtropis tetrameris DING Hou. a. Habit, = 74, b. flower, c. diagram, d. corolla with stamens,

e. pistil, all < 6, f-g. ditto, section, < 12, h. fruit, nat. size, i-j7. sections of fruit, pericarp removed, obvious-

ly the remaining placenta containing 2 seeds, * 3.—M. elliptica KiNG. k. Flower, /. corolla with stamens

on ring-shaped adnate disk, m. pistil, all x 6, n. young fruit * 2 (a-g KOSTERMANS 7312, h, j ditto 4482, 7
ditto 4399, k-m RAHMAT sI Toroes 5591, m ISMAEL 60).

Ecol. Forests, from the lowland up to 2700 m.

Embryol. I have not succeeded in observing the exact situation of the ovules and seeds.
Mr W. VAN HEEL has made a series of microtome sections of flower buds of M. curraniiand M. tetrameris.
It appears that in each cell the placentas are distinctly axile and 2-lobed, but show no trace of distinct
ovules. In one slide one of the placental lobes would seem to contain an incipient embryo-sac, but the
insertion or immersion of the ovules on or in the placental tissue must wait pending the examination of
sufficient fixed material. Obviously the formation of ovules occurs rather late in the development. In
the longitudinal sections it appeared that in bud the septum is not complete at the apex, but closes later.

274 Lens FLORA MALESIANA [ser. I, vol. 6?

In sections of mature flowers it looks as if the ovules are immersed in placental tissue.

Stipules. In the past it was stated in generic descriptions that there are no stipules in this ge-
nus. However, I have found stipules on a very young branchlet of M. platyphylla (cf. CLEMENS 31809, L).
Frequently at the tip of the branchlet, there is a pair of stipule-like, small aciculiform bodies which have
been interpreted as ‘incipient leaves’ by MERRILL & FREEMAN (/.c. 275). They are resembling bracts;
sometimes two opposite pairs (cf. CLEMENS 29518 of M. kinabaluensis) appeared on the internode towards
the base of a young shoot or inflorescence; they are called incipient bracts in this treatment.

Notes. There is a drawing of Microtropis longifolia WALL. published by Biswas (J. Ind. Bot. Soc. 19,
1940, 143, f. 3) showing a 5-merous flower (except the ovary) which has a flat disk (in the description
on page 144 stated ‘disk annular, more or less cup-shaped, with minutely fimbriate rim’) with 5 stamens
inserted on it and the basal part of the ovary immersed in the disk. The free part of the ovary is + pyrami-
dal, c. 1 mm long. Unfortunately Biswas did not make a drawing of the transverse or longitudinal
section of it. It seems to me this flower belongs to Euonymus instead of Microtropis. The flowering spe-
cimens cited by him are MAUNG Ba PE 13105, BURKILL 30313, and P. T. RussELL 41, and these obviously
belong to Microtropis. MERRILL and FREEMAN (/.c. 298) have based a new species, Microtropis pachy-
phylla, on MG. Po CHIN 6553 (type) and BuRKILL 30313. They have stated clearly that the filaments are
short, thick, and inserted on the margin of the disk; ovary narrow-ovoid, 2-celled.

Specific discrimination is very difficult in this genus. This is partly caused by the fact that so many
species have been described on inadequate material and that specimens are very seldom represented with
both flowers and mature fruit. Furthermore the majority of the species seem to be rare and collections
are few. The main key characters used by MERRILL & FREEMAN are whether the inflorescences are ample
or condensed, whether the flowers are 4- or 5-merous, and the leaf-shape. However, in many species
4- and 5-merous flowers occur together in one specimen; the inflorescences occur condensed merely in
degree; the leaf-shape, size, and texture are very much depending on the local ecology and altitude.
Consequently, MERRILL & FREEMAN had great difficulty in framing a satisfactory key: in 14 cases one
species occurs twice in their key, in 2 cases even thrice.

I have tried to delimit species by means of qualitative characters and this has led to a substantial
reduction in their number.

KEY TO THE SPECIES

1. Anthers extrorse. Petals always conspicuously keeled inside, the innermost 1 or 2 slipper-shaped
even in open flowers ... . . . 1. M. discolor
1. Anthers introrse or sometimes -+ latrorse. Petals smooth inside, very rarely slightly keeled in some
flowers in M. bivalvis, the innermost 1 or 2 not slipper-shaped.
2. Petals united in the lower half; filaments not united in a ring, inserted on the corolla tube.
3. Flowers at anthesis c. 414 mm long. Stamens inserted at the mouth of the corolla tube. Pistil +
flask-like, c. 244 mm long. . F . . 2, M. tenuis
3. Flowers ot anthesis c. 244 mm long. Stamens inserted at ‘the base of the corolla tube. Pistil short-
conical, 1 mm long. . . .. 3. M. tetrameris

Petals oni slightly united at the base; filaments united at ‘the base c or inserted on a ring or short tube.

4. Pistil -- cylindric, often slightly constricted in the middle; apex discoid or truncate, as wide as the
base or sometimes even slightly wider.

5. Calyx lobes suborbicular, 244-3 mm o. Anthers 1-114 by 1 mm. Fruits up to 20 by 12 mm,
usually furfuraceous near the apex. Inflorescences usually ps aaa and rachis rather
Stouty sven 3 . 4. M. valida

5. Calyx lobes suborbicular or : subreniform, Me 1Y, by 1- 2% mm. "Anthers C. 4, by 14 mm. Fruits
12-15 by 8-10 mm, furfuraceous, glabrescent. Inflorescences dichotomous cymes, peduncles and
rachis thin and slender . . cos ss | Sap IM bivalvis:

4. Pistil ovoid, ovoid-oblong, or short- onical (de cylindric i in 'M. evar more or less gradually
narrowed towards the apex; apex obtuse, slightly notched or 4(—6)-toothed, narrower than the base.

6. Connective broad and prominent on both sides and separating the thecae. Peduncles and rachis
when present distinctly furfuraceous . . » « 2) TOIMEgellipiics

6. Connective invisible at least on the inner aide: "Peduncles ‘and ‘rachis when present glabrous.

7. Calyx lobes transparent (when boiled), from the base with 1—5 longitudinal, few-branched veins,
distinct or conspicuous on both surfaces. Fruits rounded at the apex, mucronate by the persistent
style.

8. Leaves elliptic-oblong to -lanceolate, 11-22 cm long. Pistil + flask-shaped.
7. M. kinabaluensis
8. Leaves obovate, obovate-oblong, elliptic, rarely suborbicular, 3!4—-7 cm. Pistil short-conical.
8. M. curranii

7. Calyx lobes rather dull and only transparent at the margin, venation usually invisible, sometimes
loosely reticulate and slightly visible on both surfaces. Fruits as far as known acute to short-
acuminate or rostrate, terminating in the persistent style.

9. Apex of pistil 4(—6)-toothed (sometimes obtuse when young).

Dec. 1962]

CELASTRACEAE (Ding Hou)

275

10. Petiole distinct, 1-2 cm. eee lobes suborbicular or reniform,

at the apex

10. Petiole very short (c. 2 mm) ¢ or none. Calyx lobes suborbicular, 3

acuminate at the apex
9. Apex of pistil obtuse.
11. Pistil + cylindric, smooth, c.
Fruit unknown

. Pistil + short-conical, usually longitudinally striate, Le

terete.

2 by 2-3 mm. Anthers obtuse
. 9. M. sumatrana
4 mm. Anthers short-

by 21,
Peale se. 10. M. rigida

2 by | mm. Branchlets ee aot (Leaves sessile.)

11. M. ovata
h by jal 4 mm. Branchlets usually

12. Thyrses or rarely dichotomous CYELES; sPeduncle distinct (14-2 cm), rachis or internodes
distinct to the naked eye. Fruits 114-214 by 1-114 cm; apex acute to short-acuminate, or

rostrate, furfuraceous near the apex .

12. M. platyphylla

12. Condensed cymes, fascicles or glomerules, sessile or - subsessile, rachis or internodes obscure
or invisible to the naked eye. Fruits smaller, c. 1 by 14 cm, obtuse, furfuraceous.

1. Microtropis discolor (WALL.) WALL. Cat. (1830)
n. 4337; Arn. Ann. Nat. Hist. 3 (1839) 152; Laws.
in Hook. f. Fl. Br. Ind. 1 (1875) 614; Kurz, For.
Fl. Burma 1 (1877) 251; KiNG, J. As. Soc. Beng.
65, ii (1896) 341; Ripv. FI. Mal. Pen. 1 (1922)
444: Craip, FI. Siam. En. 1 (1926) 281; MeErr.
& Freem. Proc. Am. Ac. Arts Se 73 (1940) 292;
Loes. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 129.
—Cassine discolor WALL. in Roxb. Fl. Ind. ed.
Wall. 2 (1824) 378.

Shrub or small tree. Branchlets terete. Incipient
bracts 3-8 mm long. Leaves chartaceous, elliptic-
oblong, sometimes lanceolate, 714-1414 by
2-5'%4 cm; base attenuate; apex acuminate; nerves
5-7 pairs; petiole 5-10 mm. Cymes simple or
thrice dichotomously branched, c. | cm_ long;
peduncle 3-8 mm. Bracteoles deltoid, c. Y% mm
long and wide. Calyx lobes suborbicular to
slightly reniform, 1-134 by 114 mm, erose on the
margin. Petals + oblong, distinctly keeled inside,
the inner one or two usually slipper-shaped,
2-314 by 1-144 mm. Stamens 114 mm; filaments
subulate, slightly dilated and united at the base;
anthers dorso-basifixed, slightly oblong or ellip-
soid, c. 34 mm long, extrorse, distinctly apiculate.
Pistil 1-114 mm long, slightly wrinkled, slightly
swollen at the base and gradually narrowed into
a cylindric style; stigma obscurely 2-lobed, or
obtuse. Fruits ellipsoid, 11-15 by 7-10 mm,

subfurfuraceous.

Fig. 15. Distribution of the genus Microtropis

WALL. ex MEISN.; species density in Malaysia,

endemic above the hyphen, non-endemic below
it; in Asia c. ? 40 spp.

13. M. wallichiana

Distr. India (Sikkim, Khasia, Assam, and
Bengal), Burma (rather common), Siam (Betong,
Kaw Chang, and Lasau), Indo-China (Bienhoa
and Lakhon), S. China (Yunnan) and Malaysia:
Malay Peninsula (Penang).

Ecol. Rain-forests, 300-1260 m.

2. Microtropis tenuis SyMINGTON, J. Mal. Br.
R. As. Soc. 14 (1936) 350, t. 19; MerR. & FREEM.
Proc. Am. Ac. Arts Sc. 73 (1940) 304.

Small tree. Branchlets subterete. Leaves charta-
ceous, elliptic-oblong to -lanceolate, 444-914 by
2-414 cm; base attenuate or cuneate; apex
acuminate; nerves 6-10 pairs; petiole 6-10 mm.
Inflorescences simple or elongate-paniculate
cymes, rarely 1-flowered; peduncle 1-2! cm.
Bracteoles at the base of the flowers deltoid,
transparent, the margin reddish brown, short-
fimbriate, c. 114 mm long. Calyx lobes suborbicul-
ar or broadly obovate, 244-3'4 mm @, the lower
half rather thick and slightly wrinkled outside,
the upper half irregularly splitting, erose or
short-lacerate. Corolla whitish, c. 44% mm long,
divided at about the upper half, lobes almost
oblong, obtuse. Stamens inserted at the mouth
of the corolla; filaments free for c. 114 mm;
anthers dorso-basifixed, broad-ovoid, 4/, by 4
mm, obtuse, the lower half free. Pistil c. 244 mm
long; ovary ovoid and gradually narrowed into a
cylindric style; stigma discoid or slightly capitate.

Distr. Malaysia: Malay Peninsula (Pahang:
Mt Tapis).
Ecol. In dipterocarp forest, 750 m.

Note. Known only from the authentic col-
lection (SF 28818, StNG, L) which was printed
erroneously as ‘‘S’p. 28878” in the original de-
scription.

3. Microtropis tetrameris DING Hou, oy.
Fig. 14a-j.

A M. tenui SYMINGTON, cui affinis, foliis ellip-
ticis raro ovatis, floribus subsessilibus, minoribus,
staminibus basi in tubum insertis, ovario conico,
circiter 1 mm longo differt——Typus KOSTERMANS
7312, L, isotypes Bo, K.

Small tree up to 15 m by 30 cm go. Branchlets
terete. Incipient bracts 2-7 mm. Leaves charta-
ceous to subcoriaceous, elliptic rarely ovate, 7-11
by 314-5 cm; base attenuate; apex short-acuminate

Sp.—

276

FLORA MALESIANA

[ser. I, vol. 6?

to acuminate, sometimes acute; nerves 4—7 pairs;
petiole 5-8 mm. Inflorescences paniculate-cymose,
up to 414 cm long, few- to many-flowered; pe-
duncle 7-14 mm. Flowers green or greenish-
white; bracteoles broad-ovate, c. 1 mm long,
sometimes keeled outside. Calyx lobes suborbicu-
lar, 1-114 mm g, two opposite pairs, denticulate.
Corolla c. 24% mm long, the lobes very broad-
oblong, or -- quadrangular, 1-114 mm long,
obtuse. Stamens c. 1144 mm long; anthers sub-
orbicular, free at the lower 14, c. ?/; mm long,
obtuse or slightly apiculate, usually perpendi-
cularly bent towards the center; filaments subulate,
fleshy, c. 1 mm, inserted at the base of the corolla
tube and usually the lower half adnate to it;
anthers dorso-basifixed. Pistil short-conical, c. 1
by 1 mm, obtuse, slightly furrowed. Fruits el-
lipsoid or slightly obovoid, 12-15 by 8-10 mm,
obtuse, furfuraceous; persistent calyx lobes patent.

Distr. Malaysia: Borneo (Balikpapan: Kos-

TERMANS 7312, 4242, 4399, 4482; Pleihari: bb
10372).

Ecol. Primary forest, from lowland up to
700 m.

Vern. Masintan, Born., ratubégalang, M.

Note. The specimens cited above are very
homogeneous and bear both flowers and fruit.
M. tetrameris is very closely allied to M. tenuis
especially by the leaf-shape, inflorescences, 4-
merous flowers, and stamens not united into a tube.
It differs, however, from it by the characters
indicated in the key, by the nervation, and also
by the geographical distribution. The structure
of the seeds (fig. 14i-j) is not clear.

4. Microtropis valida Rip. J. Str. Br. R. As. Soc.
n. 75 (1917) 19; Fl. Mal. Pen. 1 (1922) 445;
Merr. & FREEM. Proc. Am. Ac. Arts Sc. 73 (1940)
304.—M. bicolor Merr. & FREEM. l/.c. 298.—
M. pauciflora BOERL. ex MERR. & FREEM. /.c. 303.

Shrub or small tree up to 5 m. Branchlets terete.
Incipient bracts 3-8 mm. Leaves subcoriaceous to
coriaceous sometimes chartaceous, ovate to
lanceolate, elliptic-oblong or elliptic, 11-2714 by
414-12 cm; base rounded, acute, or cuneate to
attenuate; apex acute to acuminate, sometimes
obtuse or short-apiculate; nerves 6-14 pairs;
petiole 1-2 cm. Inflorescences usually paniculiform,
114-6 cm, rarely a simple cyme or fasciculate less
than 1 cm; peduncles 1-4 cm, sometimes very
short. Bracteoles slightly ovate or suborbicular,
144-2 mm long. Flowers 4(—5)-merous. Calyx
lobes suborbicular, 2!4—3 mm @, concave, slightly
wrinkled outside, margin transparent, slightly
erose. Petals broad-elliptic or -oblong, 2144-3 by
134-214 mm, obtuse. Stamens c. 1¥%. mm long;
anthers very broadly oblong, c. 1-114 mm long,
obtuse and short-apiculate, connective distinct,
broad on the dorsal side; filament flat, c. 1 mm,
distinctly united at the lower 1/5. Pistil -- cylindric,
slightly constricted at the middle, slightly fur-
rowed, c. 2 mm long, truncate or discoid at the
top, rarely notched. Fruits ellipsoid, c. 21 by 11
mm, acute to acuminate and furfuraceous at the
apex, terminating in a very short, persistent style.

Distr. Malaysia: Sumatra (no precise locality),
Malay Peninsula (Perak, Pahang, and Johore),
and Borneo (Sarawak).

Ecol. Rain-forests from lowland up to 1800 m,
once collected in mossy forest (KEP 36566).

Notes. The flower of M. bicolor is unknown.
In the characters of the leaves and the persistent
calyx lobes this species is similar to M. valida
with the exception that the fruits (two collections)
are fasciculate. There is a _ sterile specimen,
Beccari (PB 1488, K), which has large leaves up
to 27 by 814 cm with rather obscure venation
underneath. There are galls on the upper surfaces,
which are pale, corky bodies lobed but globose
in outline, c. 3-4 mm @o. This specimen matches
the type collection, BECCARI (PB 2617, K) which
also bears these characteristic galls.

The authentic material of M. pauciflora
was collected by KorTHALS (s.n., L) in West
Central Sumatra. The specimens are rather poor
with detached flower-buds and young fruit. One
specimen had two flowers still attached to the
inflorescence; the one I dissected was 5-merous
as indicated in the original description. The
vegetative and floral characters are quite similar
to those of the M. valida with the exception that
the flowers as far as we know are 5-merous.

5. Microtropis bivalvis (JACK) WALL. List (1830)
n. 4340; Laws. in Hook. f. Fl. Br. Ind. 1
(1875) 614; Kina, J. As. Soc. Beng. 65, ii (1896)
341; Riv. FI. Mal. Pen. 1 (1922) 444; Merr. &
FreeM. Proc. Am. Ac. Arts Sc. 73 (1940) 301;
Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 129.
—Celastrus bivalvis JacK, Mal. Misc. 1, 5 (1821)
19; Roxs. Fl. Ind. ed. Carey & Wall. 2 (1824)
399.—Paracelastrus bivalvis Mig. Fl. Ind. Bat.
1, 2 (1859) 591; F. N. Witiams, Bull. Herb.
Boiss. II], 5 (1905) 224.—M. filiformis KING,
J. As. Soc. Beng. 65, ii (1896) 342; Rrpv. Fl. Mal.
Pen. 1 (1922) 444; BurK. & HENDERSON, Gard.
Bull. S.S. 3 (1925) 360; Crais, Fl. Siam. En. 1
(1926) 281; Merr. Pap. Mich. Ac. Sc. 1938,
24 (1939) 79; MeRrR. & FREEM. Proc. Am. Ac.
Arts Sc. 73 (1940) 300; Logs. in E. & P. Pfl. Fam.
ed. 2, 20b (1942) 129, f. 35.—M. peduncularis
Rip_. Kew Bull. (1924) 262; Fl. Mal. Pen. 5
(1925) 298; Crars, Fl. Siam. En. 1 (1926) 282;
MerrR. & FREEM. Proc. Am. Ac. Arts Sc. 73
(1940) 300.

Small tree or shrub, up to 7 m. Branchlets
terete. Incipient bracts small, c. 2 mm long.
Leaves chartaceous to subcoriaceous, elliptic to
elliptic-oblong, ovate to lanceolate, (6—)12—20\4
by 214-8 cm; base acute to attenuate; apex acumi-
nate; nerves 6-10 pairs; petiole 5-8 mm. Cymes
2-10 cm, 2-5 times dichotomously branched,
rarely simple, few- to many-flowered; peduncle
1-714 cm. Flowers pale-greenish-yellow or yellow-
ish; bracteoles deltoid, c. 1 mm long and wide.
Calyx lobes suborbicular to subreniform, 74-11
by 1-214 mm, very short-fimbriate or slightly
erose and transparent on the margin. Pefals free,
oblong or oblong-ovate, sometimes slightly
contracted at the base, the inner one or two con-

Dec. 1962]

CELASTRACEAE (Ding Hou)

277

cave or laterally folded, 214-334 by 144-134 mm.
Stamens c. 114 mm long; filaments flat, basi-
fixed, c. 4/; mm, united at the lower half or lower
¥,; anthers oblong, c. 74; mm long, obtuse.
Pistil c. 114 mm long, cylindric, slightly contracted
at the middle, slightly ridged; stigma discoid.
Fruits ellipsoid, 114-114 by 4/,—-1 cm, furfuraceous,
glabrescent; style persistent.

Distr. Siam (Kasum), Burma (Tenasserim),
and Malaysia: Sumatra (Asahan and Pajakum-
buh) and Malay Peninsula (Kedah, Perak,
Dindings, Kelantan, Trengganu, Pahang, Se-
langor, Langkawi Is., and Penang).

Ecol. Chiefly in primary forests, rarely in
secondary forests, 300-1800 m.

Vern. Kaju bile-bile, k. borhum, sungai buloh,
Asahan.

Notes. The chief differences between M.
bivalvis, M. filiformis and M. peduncularis are
found in the length of inflorescences and the size
of leaves and flowers. These differences hold only
for the type specimens but in additional speci-
mens one can find the intermediate sizes some-
times even on a single specimen.

According to MERRILL & FREEMAN (/.c. 272 &
301), WALLICH n. 4340 (BM, G, K, and L) is
made up of material collected in Penang by JACK
and Porter and it is reasonably safe to assume
that this WALLICH number does represent the type
collection of Celastrus ? bivalvis Jack. They have
also pointed out that Jack, who described his
species with ‘corolla none’, concluded to this
from specimens with fallen petals; this inadequacy
led Miquet (/.c.) to propose for it the new gene-
tic name Paracelastrus, based solely on JACK’s
original description of Ce/astrus bivalvis.

6. Microtropis elliptica KiNG, J. As. Soc. Beng. 65,
ii (1896) 340; Ript. Fl. Mal. Pen. 1 (1922) 444;
Merr. & FREEM. Proc. Am. Ac. Arts Sc. 73 (1940)
296; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
129.—M. ophirensis Riv. J. Str. Br. R. As. Soc.
n. 35 (1901) 10; Fl. Mal. Pen. 1 (1922) 444; Merr.
& FREEM. Proc. Am. Ac. Arts Sc. 73 (1940) 297.
—M. vinculans BoerLt. & Koorb. in Koord.-
Schum. Syst. Verz. 2 (1911) 33; Logs. in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 129.—M. bivalvis
(non WALL.) Koorp. Exk. Fl. Java 2 (1912) 524;
Koorpb.-ScHuM. Syst. Verz. 1 (fam. 158) (1912) 3;
Koorp. Atlas 1 (1913) t. 138.—M. javanica MERR.
& Freem. Proc. Am. Ac. Arts Sc. 73 (1940) 295.
—WM. longirostris MerRR. & FREEM. /.c. 303.—
Fig. 14k-n.

Shrub or small tree up to 15 m. Branchlets
terete. Incipient bracts 14-1 cm. Leaves charta-
ceous to coriaceous, elliptic to elliptic-oblong,
sometimes ovate to lanceolate, 414-22 by 2-11
cm; base cuneate; apex acute to acuminate; nerves
5-9 pairs; petiole 3-13 mm. J/nflorescences usually
in several leaf-axils along the branchlet, some-
times extra-axillary near the base of a young
shoot, rarely ramiflorous, -- condensed or short-
to distinct-peduncled, rather laxly cymose, 1—S cm
long, few-flowered; peduncles when present
Y%-114 cm, densely furfuraceous. Bracteoles

deltoid, c. 4 mm long. Flowers white, 5-merous,
occasionally a few flowers 4-merous. Calyx lobes
+ semi-orbicular, sometimes subreniform, 1-114
by 114-214 mm, in full developed flowers 3-41,
mm. Petals elliptic- or ovate-oblong, or slightly
oblong, 3-4 by 114-2 mm, up to 5 by 3 mm.
Stamens c. 2-21 mm, finally 344 mm _ long;
filaments flat, united at the lower 14-24; anthers
broad-oblong, 4/,-1 mm long, obtuse; connective
broad and prominent on both sides, shortly
sometimes distinctly protruding beyond the
anther-cells. Pistil ovoid-oblong, 2-3 by 34-1 mm,
slightly, gradually narrowed to the apex; apex
obtuse, sometimes obscurely notched. Fruits
ellipsoid or obovoid, irregularly, obscurely striate
in the dry state, 114-2 by 34-114 cm, furfuraceous;
apex short-acuminate, sometimes forming a dis-
tinct beak 5-12 mm.

Distr. Malaysia: Sumatra (Tapanuli, Taram,
and Djambi), Malay Peninsula (Perak, Treng-
ganu, Pahang, Malacca, Johore, and Penang),
and West Java (Bantam: Mt Pulasari, Udjong
Kulon).

Ecol. In primary forests, 90-1500 m, once
found on sandstone near Tjampo R. in W. Su-
matra (coll. ISMAEL).

Notes. The type of M. vinculans was collected
by Koorpers (212798, Bo, not 21297f as cited by
MERRILL & FREEMAN) at Pang Kalan-Dula,
Central Sumatra. There is only one sheet of the
type with a few detached leaves, a piece of root,
and some fragments of flowers. From these
fragments left I could unfortunately not verify
any floral character. The leaves are similar to
those of M. elliptica.

The type of M. javanica was collected by
Koorpbers (7. 9921, B, L) at Mt Pulasari, Bantam,
W. Java, at 1250 m, in June 1892; it has extra-
axillary, short-peduncled, few-flowered, con-
densed inflorescences and very young 5-merous
flowers. The flowers were apparently associated
with some 4-merous ones as indicated by VA-
LETON’S note on the type. Only in Nov. 1960 rich
additional material was collected by KOSTERMANS
at Mt Pajung, Udjong Kulon, c. 100 km south-
west of the type locality at 300-470 m. Two of these
specimens have very well preserved and developed,
distinctly peduncled, dichotomous cymes (up to
4 times branched and c. 314 cm long). There
are many flowers in different stages of develop-
ment; I have dissected more than twenty of them
and so far I have seen only 5-merous ones. The
characters of leaves, inflorescences and flowers
fall within the range of M. elliptica.

As lectotype of M. elliptica | have chosen
MAINGAY 945/2, in K, isotype sheets in L; there
are several good sheets of this collection at Kew,
bearing both flowers and fruits.

7. Microtropis kinabaluensis Merr. & FREEM.
Proc. Am. Ac. Arts Sc. 73 (1940) 304, incl. var.
acuminata.—M. sterrophylla MERR. & FREEM. l.c.
305.

Shrub or small tree up to c. 3 m. Branchlets
terete. Incipient bracts up to 114 cm long. Leaves

278

FLORA MALESIANA

[ser IF volaie2

chartaceous to subcoriaceous, elliptic-oblong to
-lanceolate, 11-22 by 3144-914 cm; base cuneate to
attenuate; apex acuminate; nerves 6-12 pairs;
petiole 1-214 cm. Paniculate cymes 14-24% cm
long. Flowers (young) dull white; bracteoles
1144-2 mm long. Calyx lobes suborbicular, c.
2144 mm Q, transparent, slightly concave, with
3-5 longitudinal veins, margin- slightly erose.
Petals ovate- or broad-elliptic, 244-3 by 114-2
mm, obtuse. Stamens c. 2 mm long; anthers
broadly ovoid to subrotund, 24 mm long, slight-
ly apiculate; filaments c. 1144 mm, united at the
lower 1 mm. Pistil c. 114 mm long, + flask-
shaped. Fruits yellowish orange, red when ripe,
ovoid or broad-ellipsoid, 114-2 by 1 cm, slightly
furrowed, the obtuse apex crowned by the per-
sistent style.

Distr. Malaysia: N. Borneo (Mt Kinabalu).

Ecol. Primary forests, 900-1500 m.

Notes. The distinctions between M. kinabalu-
ensis, its var. acuminata, and M. sterrophylla are
chiefly based on leaf characters. However, they
share the very characteristic calyx lobes which
are transparent when boiled, bearing 3—5 longitu-
dinal veins, and the fruit-shape, rounded at the
apex, crowned by a mucro of the persistent style,
from which I conclude that they are con-
specific.

8. Microtropis curranii Merr. Philip. J. Sc. 3
(1908) Bot. 238; En. Philip. 2 (1923) 482; Merr.
& FrReeM. Proc. Am. Ac. Arts Sc. 73 (1940) 287,
incl. var. zambalesensis et var. obovata MERR. &
FREEM.; Logs. in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 29.

Small tree or shrub up to 9 m. Branchlets terete,
or sometimes slightly 4-angular. Leaves subcoria-
ceous to coriaceous, obovate, obovate-oblong,
elliptic, rarely suborbicular, 314-7 by 114-334
cm; base cuneate to attenuate, sometimes de-
current; apex acute, rarely short-acuminate,
sometimes obtuse and apiculate, the tip usually
damaged; nerves 4~7 pairs; petiole 2-10 mm.
Cymes up to 114 cm, (1—)3-flowered, occasionally
a central axis elongated above the first pair of
flowers and more-flowered; peduncle 14-114 cm.
Flowers white. Bracteoles ovate, c. 1 mm long.
Calyx lobes suborbicular to subreniform, 2-214
by 214-314 mm, erose. Petals elliptic, 334 by
2mm. Stamens c. 11/4. mm long; anthers subglobose,
c. /; by 34 mm, slightly apiculate; filaments
flat, 134 mm long, gradually dilated towards the
base, united at the lower half. Pistil short-conical,
1144 by 1 mn, slightly longitudinally striate, ob-
tuse or sometimes obscurely 2-lobed. Fruits
(young) ellipsoid, furfuraceous.

Distr. Malaysia: Philippines (Luzon: Zam-
bales, Benguet and Rizal Prov.).

Ecol. In mossy forest and on exposed ridges
on the higher mountains, 1500—2500 m.

Note. There are several specimens cited in the
original description of M. curranii, but the type
has not been designated. Since the epithet is
“curranii’ | have chosen CURRAN FB 4966 (US,
isotype in K) as lectotype. The authentic collec-

tions cited in the original description of var.
zambalesensis are RAMos BS 4698 (K) and
CuRRAN & MERRITT 8071 (K), and the former is
here selected as the lectotype. The two varieties
differ only slightly in leaf-size, texture, and shape;
in my opinion they do not deserve distinction,
especially because these differences may be due
to different altitude.

9. Microtropis sumatrana Merr. Pap. Mich. Ac.
Sc. 1933, 19 (1934) 164, t. 26; MeRR. & FREEM.
Proc. Am. Ac. Arts Sc. 73 (1940) 304.

Small tree up to 17 m. Branchlets terete. In-
cipient bracts conspicuous, 7-15 mm long. Leaves
chartaceous to subcoriaceous, elliptic- or ovate-
oblong, 12-23 by 6-11 cm; base acute; apex
acute to acuminate; nerves 6-11 pairs; petioles
1-2 cm. Thyrses and/or cymes up to 4% cm,
simple or 2-3 dichotomously branched; peduncle
1-3 cm, rarely very short. Flowers white; brac-
teoles small, 114-2 mm long, short-fimbriate.
Calyx lobes suborbicular or reniform, 2-3 by
2-314 mm, concave, stiff, smooth inside, usually
irregularly split, short-lacerate, venation visible
but not elevated. Petals oblong, 3-314 by 14-144
mm, obtuse. Stamens 1'4-134 mm long; fi-
laments dorso-basifixed, flat, gradually dilated
towards the base, united at the lower half;
anthers suborbicular, 14 mm long, obtuse, some-
times obscurely apiculate. Pistil c. 14 mm long,
gradually narrowed towards the apex; apex 4-
lobed, or obtuse (in young flowers). Fruits
ellipsoid, broad-obovoid, or ovoid-oblong, 15-17
by 8-1114 mm, furfuraceous, glabrescent, slightly
Trugose, acuminate at the apex.

Distr. Malaysia: Sumatra (Simalur and Batu
Is., Sum. East Coast Res.: Marbau) and Borneo
(Sipitang and Balikpapan).

Ecol. In lowland and mossy forest (800-1050
m); in Sumatra at low altitude, once noted on
sandstone.

Vern. Anuntus, délok, kudang pajo, lala-lalar
délok, soe’ ah-baseum, surin, téradih, Sum.

Note. MERRILL & FREEMAN (/.c.) made an
erroneous correction of an error in the original de-
scription in which ‘stipules’, i.e. incipient bracts,
are not 7 cm but 7 mm long.

10. Microtropis rigida RipL. Kew Bull. (1931)
36; Merr. & FREEM. Proc. Am. Ac. Arts Sc. 73
(1940) 297.

Leaves coriaceous, elliptic-oblong, 7-12 by
214-6 cm; base obtuse; apex acuminate; nerves
c. 5 pairs; petiole very short. Thyrses up to 214
cm long; peduncle c. 1 cm. Bracteoles ovate,
c. 2 mm long. Calyx lobes suborbicular, c. 3 by
214 mm, sometimes irregularly split, smooth
inside, slightly wrinkled outside, very short
fringed. Petals oblong, obtuse, 2 by 1 mm, the
lower part adnate to the disk. Stamens c. 2 mm
long; anthers ovoid c. 24 mm long, short-apicu-
late; filaments 114 mm, subulate, the lower 24
united, dorso-fixed, the connective distinct on the
dorsal side. Pistil c. 244 mm long; ovary + glo-
bose, c. 2 mm g, narrowed to the apex; style very

Dec. 1962]

CELASTRACEAE (Ding Hou)

279

short; stigma obscurely 4-6 lobed. Fruits (im-
mature) oblong.

Distr. Malaysia:
collected.

Borneo (Sarawak), twice

11. Microtropis ovata MeRR. & FREEM. Proc. Am.
Ac. Arts Sc. 73 (1940) 297.

Shrub up to 3 m. Branchlets 4-angular. Leaves
coriaceous, ovate to ovate-oblong, 4!4-10 by
2-7 cm; base rounded, sometimes obscurely
cordate; apex acute; nerves 5-9 pairs; petiole
very short or none. Cymes (rather young) con-
densed, less than 1 cm; peduncle very short.
Bracteoles broad-ovate, c. 11 mm long, + keeled
outside. Calyx lobes suborbicular to subreni-
form, 214-2'4 by 2!4-3 mm, short-lacerate.
Petals broadly ovate or ovate-oblong, 2'4-3 by
2 mm. Stamens c. 144 mm; anthers transverse
broad-oblong, 74 by 4/; mm, notched at the
apex; filaments c. 1 mm, united at the lower half.
Pistil ovoid-oblong, c. 2 by 1 mm. Fruits un-
known.

Distr. Malaysia: Borneo (Mt Kinabalu).

Ecol. Forests, 1200-1500 m.

12. Microtropis platyphylla Merr. Philip. J. Sc.
10 (1915) Bot. 319; En. Philip. 2 (1923) 482;
Merr. & FREEM. Proc. Am. Ac. Arts Sc. 73 (1940)
306, incl. var. ellipticifolia MERR. & FREEM.;
Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 129.—
M. philippinensis Merr. Philip. J. Sc. 13 (1918)
Bot. 306; Merr. & FREEM. Proc. Am. Ac. Arts
Sc. 73 (1940) 304.—M. rostrata MerR. Philip. J.
Sc. 17 (1920) 275; En. Philip. 2 (1923) 482;
Merr. & FREEM. Proc. Am. Ac. Arts Sc. 73 (1940)
306; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
129.—M. fasciculata Quis. & MerR. Philip. J.
Sc. 37 (1928) 162; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 129.—M. chartacea MERR. & FREEM.
Proc. Am. Ac. Arts Sc. 73 (1940) 303.—M. rubra
Exo. [ex Merr. En. Philip. 2 (1923) 482, in obs.,
pro syn.) ex Merr. & FREEM. lc. 305.—M.
basilanensis Merr. & FREEM. l/c. 306.—M.
borneensis MERR. & FREEM. /.c. 296.

Shrub, 2-7 m tall. Branchlets terete or some-
times slightly angular. Incipient bracts 4-12 mm
long. Leaves chartaceous to subcoriaceous, el-
liptic to elliptic-lanceolate, ovate to lanceolate,
sometimes obovate, rarely broad-elliptic, 414-24
by 414-11 cm; base acute to cuneate; apex acute
to short-acuminate; nerves 6-12 pairs; petiole
114-2 cm. Paniculate and/or sometimes dicho-
tomous cymes up to 314 cm long, rarely fascicles
and cauline; peduncles usually present, rather
stout, 1-114 cm, sometimes very short or none.
Bracteoles suborbicular or ovate, 1-2 mm long,
slightly erose. Flowers 5(-4)-merous. Calyx lobes
suborbicular or subreniform, 2-314 by 2-3'4 mm,
slightly concave and wrinkled outside, erose.
Petals elliptic or broad-ovate, 244-3 by 134 mm,
obtuse. Stamens 134-234 mm long; anthers
broad-oblong, -ovoid, or subglobose, 1—*/,; mm
long, obtuse, or short-apiculate; filaments 114-134
mm, united at the lower 34-1 mm. Pistil short-
conical, 114-134 mm long; apex obtuse rarely

notched. Fruits ovoid or ellipsoid, 1-21 by 1-114
cm; apex acute, short-acuminate, or rostrate,
furfuraceous near the tip.

Distr. Malaysia: Philippines (Luzon: Proy.
Nueva Ecija, Rizal, Tayabas, Camarines Norte
and Camarines Sur, and Alabat I.; Catanduanes,
Panay, Basilan I. and Siargao I.) and N. Borneo
(Mt Kinabalu).

Ecol. In forests at low and medium altitudes
up to 500 m; a mountain form found in mossy
forest of Mt Kinabalu, 1500-2700 m.

Notes. The authentic material of the present
species and all its synonyms cited here had been
collected from the central Philippines (except one
sheet from Basilan and several from Mt Kinabalu
respectively) at low and medium altitudes not
higher than 500 m (except several mountain form
from Mt Kinabalu at 1500-2700 m, described
as M. borneensis). All species described from the
Philippines are similar to one another, as was
already remarked by MerrRILL & FREEMAN.
The characters which they used to distinguish
them are chiefly the leaf-shape, size and texture,
the structure of the inflorescences or infructes-
cences (cymose, paniculate-cymose, or fasciculate),
the number of floral parts (S- or 4-merous), and
the apex of the fruit (rostrate or not). After studying
the authentic material and additional collections,
the differences appear to be only quantitative and
gradual; in my opinion only one species is con-
cerned.

13. Microtropis wallichiana WIGHT ex THWAITES,
En. Pl. Zeyl. (1858) 71; Laws. in Hook. f. Fl. Br.
Ind. 1 (1875) 613; Trimen, Fl. Ceyl. 1 (1893)
269; MerrR. & FREEM. Proc. Am. Ac. Arts Sc. 73
(1940) 283; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 127.—M. ramiflora (non WIGHT)
Tuwaites, En. Pl. Zeyl. (1858) 72; Stapr, Trans.
Linn. Soc. II, Bot. 4 (1894) 140; Merr. En. Born.
(1921) 354.—Paracelastrus wallichianus F. N.
WILLIAMS, Bull. Herb. Boiss. II, 5 (1905) 224.—
M. zeylanica MerrR. & FREEM. Proc. Am. Ac.
Arts Sc. 73 (1940) 282.—M. suborbiculata MERR.
& FReEEM. /.c.

Shrub or small tree. Branchlets terete or slightly
4-angular. Incipient bracts 3-8 mm. Leaves sub-
coriaceous to coriaceous, elliptic to elliptic-
lanceolate, sometimes broad-elliptic or -obovate,
3-18 by 1-6 cm; base cuneate to attenuate, or
rounded to subtruncate, rarely cuneate; apex
acuminate, or rounded, sometimes acute, rarely
shallowly notched; nerves 4-9 pairs; petiole
obsolete to 114 cm. Condensed cymes less than
1 cm long, few- to many-flowered; peduncle very
short or none, sometimes up to 2-3 mm. Flowers
yellowish. Bracteoles small, ovate, 1-2 mm long.
Calyx lobes subreniform, c. 1144-2 by 2-3 mm,
erose or short-lacerate. Pera/s rather fleshy, el-
liptic-oblong, broad-ovate, -elliptic, or -obovate,
214-314 by 124-2 mm, obtuse. Stamens c. 2 mm
long; anthers broad-oblong or ovoid, c. */,—*/, by
3/,-4/, mm, obtuse or slightly notched, acute or
slightly apiculate; filaments flat, 11-144 mm, unit-
ed at the lower !—24. Pistil short-conical, 11 by |

280

FLORA MALESIANA

[ser 1 voleG?

mm, obtuse, striate-furrowed, sometimes slightly
contracted at the base. Fruits slightly oblong-
ellipsoid or oblong-obovoid, c. 1 by % cm,
furfuraceous, obtuse and crowned by the persistent
style.

Distr. Ceylon and Malaysia: Sumatra (Atjeh)
and Borneo (Mt Kinabalu).

Ecol. In forest, 1250-2500 m.

Notes. This species has two altitudinally
differentiated forms, which is clearly demonstrated
in a series of excellent specimens collected by
VAN STEENIS in Atjeh, N. Sumatra. The floral and
fruiting characters show no variation in these
forms, but the leaves are variable especially in
shape and venation. Specimens collected from
1250—c. 2000 m have leaves which are elliptic to
elliptic-lanceolate, cuneate to attenuate at the
base, distinctly petioled, and with rather dense-
reticulate venation; leaves of specimens collected

between c. 2000 and 2500 m are commonly sub-
orbicular, usually rounded to subtruncate at the
base, more or less sessile and with rather loose-
reticulate venation. This phenotypic variation in
leaf-shape with altitude agrees with the general
scheme outlined by vAN STEENIS in this Flora,
cf. vol. 5, p. clxxx, fig. 3-4.

Excluded
Microtropis ? coriacea WALL.[Cat. (1831) n. 4338,
nomen] ex EtTtINGsH. Denkschr. Ak. Wiss. Wien
Math.-Nat. Cl. 13 (1857) 64, t. 4 f. 12, descr. fol.;
Merr. & FrReEM. Proc. Am. Ac. Arts Sc. 73
(1940) 306, based on specimens collected by
PorTER in Penang = Salacia sp.

Microtropis lanceolata Borrt. & Koorp. in
Koord.-Schum. Syst. Verz. 2 (1911) 33 is accord-
ing to MERRILL & FREEMAN (Proc. Am. Ac. Arts
Sc. 73, 1940, 307) = Linociera sp. (Oleaceae).

9. BHESA

Ham. ex ARN. Edinb. New Phil. J. 16 (1834) 315; DinG Hou, Blumea Suppl. 4
(1958) 150.—Kurrimia WALL. [List (1831) n. 4334-4336, 7200, nomen, pro maj.
part.| ex ARN. Noy. Act. Ac. Caes. Leop.-Car. 18 (1836) 328; Logs. in E. & P.
Pfl. Fam. 3, 5 (1892) 210; ed. 2, 20b (1942) 158, non Kurrimia WALL. ex MEISN.
Pl. Vasc. Gen. 1 (1837) 67; ibid. 2 (1837) 48, quae est [tea (Saxifr.).—Pyrospermum
Mig. Sum. (1861) 402.—Nothocnestis Mig. Sum. (1861) 530.—Trochisandra
Bepp. Fl. Sylv. 1 (1871) 120, t. 120.—Fig. 16.

Buttressed evergreen trees. Branchlets terete, light to dark-brown, their tips
enclosed by caducous, convolute stipules. Stipules finely lengthwise veined, usually
with colleters at the base inside, leaving large scars. Leaves spiral, coriaceous,
entire, midrib and nerves prominent on both sides, with distinct crossbar veins;
petiole terete or slightly furrowed above, long, knee-like thickened at the upper
end underneath. Racemes solitary or paired. Panicles axillary, sometimes crowded
at the tips of twigs sustained by stipules only and pseudoterminal (B. paniculata),
but a terminal bud always present between them. Pedicels with an articulation.
Bracts small, caducous. Flowers 5-merous, occasionally some flowers 4-merous,
whitish to greenish, fragrant. Calyx deeply lobed, lobes imbricate, sometimes ++
valvate. Petals contorted. Stamens inserted on the disk or just beneath the outer
margin; anthers lengthwise dehiscent, introrse, latrorse or extrorse, basifixed,
obtuse or short-apiculate; cells free for the lower 14-24. Disk fleshy, subentire
or lobed, glabrous, sometimes puberulous (B. robusta). Ovary free, usually with
a tuft of hairs at the top, 2-celled; styles 2, filiform, free or slightly united at the
base; stigma small. Ovules 2 in each cell, erect, anatropous. Capsule entire or
2-lobed, 1—2-celled, lengthwise dehiscing loculicidally by 2 valves or on one
side, mostly 1—2-seeded. Seeds completely or partly covered by the fleshy aril;

albumen abundant, fleshy; cotyledons linear-oblong.

Distr. Species 5, four of them found in Malaysia, a fifth endemic in Ceylon, distributed from Ceylon
to SE. Asia (India, Pakistan, Burma, Siam, and Indo-China); in Ma/aysia: Sumatra, Malay Peninsula,
Borneo, the Philippines, and New Guinea as far east as the Louisiades (Sudest I.). Fig. 17.

Ecol. Rain-forests at low and medium altitudes, sometimes found up to 2150 m.

Uses. See under B. paniculata.

Note. Sterile specimens show a resemblance to those of Ventilago (Rhamnac.) especially by the ve-
nation; the latter lack, however, the knee-like thickened apex of the petiole as found in Bhesa.

Dec. 1962] CELASTRACEAE (Ding Hou) 281

Fig. 16. Bhesa archboldiana (MERR. & PERRY) D1NG Hov. a. Habit, in fruit, 24, b. stipule, from inside,
3, c. flower, X 4, d. ditto in section, pistil removed, * 4, e. pistil, f-g. ditto in sections, 8, h. seed with
aril, i. seed in section, both nat. size.—B. indica (BEDD.) D1NG Hou. j. 2-lobed fruit, both lobes dehiscing,
k. seed enveloped by aril, both nat. size. —B.paniculata Arn. /. Flower, * 4, m. ditto in section, pistil
removed, <8, n. pistil, «8, o. fruit, one lobe barren, nat. size.—B. robusta (RoxB.) DING Hov. p.
Flower, 4, g. ditto in section, pistil removed, * 8, r. pistil, x 8, s. fruit, nat. size (a Brass 7754, b-g
Brass 28084, h-i BrAss 7754, j Kep 10158, k SF. 4177, /-n ASHTON 2613, 0 CLEMENS 27396, p-r KOSTER-
MANS 10319, s ditto 9716).

282

FLORA MALESIANA

[ser. I, vol. 6*

KEY TO THE SPECIES
(based on flowering material)

1. Flowers in panicles.

2. Disk deeply 5-lobed. Filaments inserted between the lobes of the disk. Anthers extrorse. Ovary

always with a tuft of woolly hairs at the top . ee
. Disk subentire. Filaments a continuation of the disk. Anthers introrse. Ovary glabrous or sometimes

N

slightly hairy at the top .
1. Flowers in racemes.

3. Calyx lobes broad-ovate or subrotundate, 114-2

1. B. paniculata
2. B. indica

mm long. Petals 214-3 by 74-114 mm. Anthers

latrorse. Disk usually puberulous. Filaments inserted just below the outer disk margin. Ovary with
a tuft of woolly hairs at the top. Styles entirely free. Flowers subsessile to shortly pedicelled (c.

1 mm)

3. B. robusta

3. Calyx lobes obovate, c. sy mm long. Petals 4-4 by 2 mm. 1. Anthers introrse. Disk usually glabrous.
Filaments a continuation of the disk. Ovary glabrous. Styles shortly united at the base. Flowers

distinctly pedicelled (3-7 mm)

4. B. archboldiana

KEY TO THE SPECIES
(based on fruiting material)

apex (often unequally) 2-lobed.

i)

Ne

Fruits 214

. Infructescences paniculate. Fruits obovoid, obcordate, broadly ellipsoid to suborbicular, the blunt

. Fruits 1-134(—2) cm long, acute to attenuate at the base. Aril covering most of the seed.

1. B. paniculata

—314(-5'4) cm long, obtuse at the base. Aril covering only the lower 14 or 14 of the seed.

2. B. indica

1. Infructescences simple racemes. Fruits ovoid-oblong or elliptic-oblong, attenuate and pointed to the

apex, not pores
3. Pedicels (2!
of nerves
3. Pedicels 3-7 mm. ‘Leaves ity

1. Bhesa paniculata Arn. Edinb. New Phil. J. 16
(1834) 315; Wap. Rep. 1 (1842) 538 (‘Rhesa’);
Dinc Hou, Blumea Suppl. 4 (1958) 151.—
Kurrimia paniculata WALL. [List (1831) n. 4336,
nomen] ex ARN. Noy. Act. Ac. Caes. Leop.-Car.
18 (1836) 328; Kurz, J. As. Soc. Beng. 39, ii
(1870) 73; Laws. in Hook. f. Fl. Br. Ind. 1 (1875)
622; KING, J. As. Soc. Beng. 65, ii (1896) 355;
Merr. En. Born. (1921) 354; Rpt. Fl. Mal. Pen.
| (1922) 452; Merr. En. Philip. 2 (1923) 483;
Crais, Fl. Siam. En. 1 (1926) 286; CorNeER,
Ways. Trees (1940) bith t. 43, f. 49; Logs. in
E. & P. Pfl. Fam. ed. 2, 20b (1942) 160; BROWNE,
For. Trees Sar. & Brun. (1955) 78. —Schmidelia

conferta BLANCO, Fl. Filip. ed. 2 (1845) 217
(sphalm. Schmidellia).—Pyrospermum calophyl-
lum Mia. Sum. (1861) 402.—Kurrimia luzonica
VIDAL, Rey. Pl. Vasc. Filip. (1886) 88; Logs.

in E. & P. Pfl. Fam. ed. 2, 20b (1942) 160.—
Kurrimia minor Rwy. Kew Bull. (1938) 235, excl.
BECCARI 2624, quae es
161-0.

Tree up to 35 m by 90 cm g, buttresses up to 2 m.
Bark smooth, light brown, thin. Stipules lan-
ceolate, 134-3 cm. Leaves elliptic-oblong to
-lanceolate, rarely ovate-oblong, 6-39 by 2-15 cm;
base obtuse or cuneate; apex shortly acuminate or
obtuse; nerves 11-20 pairs; petiole 1-10 cm.
Panicles crowded at the ends of the twigs, up to
38 cm long; rachis puberulous, glabrescent.
Pedicels 2-3 mm. Calyx lobes deltoid, sometimes
broadly-oblong, 24-1 by 24-1 mm, puberulous

14—)1 mm or less. Leaves 2-31 2 times as ee as broad, with aa )13—15(-19) pairs

3. B. robusta

times as ; long « as ‘broad, with 9- I ‘pairs of 1 nerves.

4. B. archboldiana

outside. Perals oblong, broad-ovate, 134-2 by
1-114 mm, obtuse, puberulous inside. Stamens
114-2 mm long; anthers triangular, 4% by 4 mm,
obtuse. Disk 5-lobed, lobes broad-oblong, trun-
cate. Ovary ellipsoid, 11-2 by 1 mm; styles free,
about half as long as the ovary. Fruits 1-134 (-2)
by 4/,-114 cm, usually 2-lobed, turning yellow,
then pink, red or dark red. Seeds 2-4, broad-
ellipsoid, subglobose, 9-11 by 6-8 mm, pale brown,
usually largely covered by the pink aril.

Distr. Common and widely distributed in
southern India, S. Siam, and Malaysia: Sumatra,
Malay Peninsula, Borneo, and the Philippines.

Ecol. Primary and secondary forests on dry-
land or in peat swamp, or periodically inundated
forests, from the lowland up to 450 m, a few
specimens collected from 900-1500 m (Perak and
Mt Kinabalu). F/. March-Dec., fr. Jan.—Dec.

Vern. Sumatra: arang, Benkoelen, damas,
djung, kaju si darang daja, East Coast, hajodolok
rawang, kalumpan, Batak., kaju djambu, k. tulang,
pimpoéh, Palembang, Jagan bunga, Pematang,
kérindjing rénak, mérlantaén rawang, sétomuhila,
M, ponau, séngafoh-balah, s. délok, s. étén bungo,
s. uding, tutum séngafoh, Simalur; Billiton:
kérangji, M; Banka: mélabung; Mal. Pen.: bénga
médang ayér, Temuan, biko-biko, bunak, M,
médang tandok, m. tijoh, Negri Sembilan, sapan,
Selangor; aha tung, bintan, madang bura, médang
latak, nga, tjanggam, Dyak, asam pau, S. Born.,
duhat burung, réngas, Balikpapan, kayu minyak,
résak buntun, sulang sulang, Sarawak, médang

Dec. 1962]

CELASTRACEAE (Ding Hou)

283

|
|
|
|

Fig. 17. Distribution of the genus Bhesa Ham.
ex ARN. Species density, endemic above the hyphen,
non-endemic below it.

kuning, pangil-pangil, sarunai, Brunei, rarasan
tatahon, sampaka, Dusun, ruwas, M, sanggam,
W. Born., sémita, Boeloengan, simun, Iban.

Uses. The dull-brown wood is rather hard and
is used for house-building purposes; it is durable
under the roof, is not attached by insects, and is
not liable to cracking. The black, acid fruits are
eaten similarly as Nephelium (HeyYNeE, Nutt. PI.
1927, 984; BuRKILL, Dict. 1, 1935, 1288).

Note. Sterile herbarium material of B. panicu-
lata and B. robusta can not be distinguished with
certainty. The stipular scars are usually oblique in
B. robusta and + transverse in B. paniculata.
However, this is not a constant character.

2. Bhesa indica (BEDD.) DING Hou, Blumea Suppl.
4 (1958) 152.—Trochisandra indica Bebb. Fl.
Sylv. 1 (1871) 120, t. 120.—Kurrimia bipartita
Laws. in Hook. f. Fl. Br. Ind. 1 (1875) 622.—
Kurrimia indica GAMBLE, Fl. Madras 2 (1918)
207; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
160.—Fig. 16j-k.

Tree up to 20 m by 60 cm g. Stipules lanceolate,
2-3 cm. Leaves oblong-ovate to -lanceolate, 11-40
by 4-15 cm; base rounded or obtuse; apex acute
to short-acuminate; nerves 16-21 pairs; petiole
up to 11 cm. Panicles up to 15 cm, rachis and
stalks puberulous when young; peduncle up to
4 cm. Pedicels c. 2 mm, puberulous. Calyx lobes
elliptic-lanceolate, c. 1 by 24 mm. Petals slightly
obovate-oblong or sometimes elliptic-oblong, c.
24% by 1144 mm, obtuse. Disk flat, subentire.
Stamens inserted on the margin of the disk and
continuous with it. Ovary broad-oboyate or sub-
globose, c. 114 by 2 mm, glabrous, or sometimes
slightly hairy at the top; styles almost divided to
the base. Fruit 2-lobed to the lower 1/,—/;,
sometimes one of the lobes small and abortive,
24%4-314(-S'4) cm long. Seeds ellipsoid, 214-214
by 114-134 cm, sometimes slightly flattened, the
aril covering its lower 14 or 14.

Distr. S. India, Burma (Mergui), Lower Siam
(Kopal, Sarat, and Kao Nom Sao), and Malaysia:
Malay Peninsula (Kedah, Selangor, and Malacca).

Ecol. Dense, moist forests usually at 800-2150
m, rarely also occurring at low altitude (KERR
12263 and SF 4177).

Vern. Bénak, biku biku, buah chéndara, hashim,
M, malayray, Temuan.

Note. This is a new record for Malaysia. It is
closely allied to, but quite distinct in flower and
fruit from, B. paniculata to which Lawson (i.c.)
reduced it.

3. Bhesa robusta (Roxs.) DING Hou, Blumea
Suppl. 4 (1958) 152.—Celastrus robustus Roxs.
Fl. Ind. 2 (1824) 395; ed. Carey 1 (1832) 626.—
Kurrimia pulcherrima WALL. [List. (1831) n. 4334,
nomen] ex LAws. in Hook. f. Fl. Br. Ind. 1 (1875)
622, nomen illegit. pro cit. prior.; KING, J. As. Soc.
Beng. 65, ii (1896) 354; Rip. Fl. Mal. Pen. 1
(1922) 452; KANJILAL, DAs & PurRK. Fl. Assam 1,
2 (1937) 270; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 159; Tarpieu, Fl. Gén. I—C. Suppl. 1
(1948) 807; Not. Syst. 14 (1950) 46.—Kurrimia
calophylla WALL. [List (1831) n. 4335, nomen].—
B. moja HAM. ex ARN. Edinb. New Phil. J. 16
(1834) 315; WALP. Rep. 1 (1842) 538 (sphalm.
Rhesa).—Nothocnestis sumatrana MiaQ. Sum.
(1861) 531.—Kurrimia robusta Kurz, J. As. Soc.
Beng. 39, ii (1870) 73; ScHeFF. Nat. Tijd. N. I.
34 (1874) 98; Kurz, For. Fl. Burm. | (1877)
253; Pierre, Fl. For. Coch. 4 (1893) t. 296B,
incl. var. roxburghii PIERRE et var. thorelii PIERRE;
PiTARD, FI. Gén. I.-C. 1 (1912) 893, f. 112, 3-8;
Crais, Fl. Siam. En. 1 (1926) 286; CoRNER,
Ways. Trees (1940) 190, f. 49; Merr. J. Arn.
Arb. 35 (1954) 141.—Kurrimia maingayi LAws.
in Hook. f. Fl. Br. Ind. 1 (1875) 622; Logs. in
E. & P. Pfl. Fam. ed. 2, 20b (1942) 159.—Fig.
16p-s.

Tree up to 40 m by 62 cm o. Bark rather rough,
brown, peeling off profusely in strips 2-3 cm wide
and 2 mm thick. Buttresses up to 4 m high, c.
34 m out. Stipules lanceolate, 5-10 mm. Leaves
elliptic or oblong-elliptic, sometimes ovate-oblong
or obovate-oblong, 6-16 by 214-814 cm; base
obtuse or cuneate; apex acute to short-acuminate;
nerves (11—)13—15(—19) pairs, slightly elevated or
flat above; petiole 1-3 cm. Racemes up to 15 cm
long, 1(—2) in a leaf-axil; peduncle short. Flowers
subsessile. Calyx lobes broad-ovate or sub-
rotundate, 114-2 by 1-144 mm, obtuse. Petals
oblong-elliptic, 244-3 by 74-144 mm, obtuse.
Stamens c. 2 mm long, attached beneath the
outer margin of the disk; anthers almost deltoid,
c. Y mm long and wide, obtuse, free for the lower
2/,, latrorse. Disk cupular, subentire or obscurely
notched, the rim usually puberulous. Ovary
subglobose c. 1 mm g, with a tuft of hairs at the
apex; styles free, longer than the ovary. Fruits
ovoid-oblong to lanceolate, with 2 vertical grooves,
much tapered to the apex, 3-34 by 1-144 cm,
usually 1l-seeded. Seeds + oblong, usually on a
knob-like thickened placenta, completely or some-
times only the lower half enveloped by the aril.

Distr. NE. India, Bhotan, E. Pakistan (Chit-
tagong Hills), Burma (Martaban, Pegu, Te-
nasserim), Andaman Is., Siam (Chantaburi, Sarat,
and Sriracha), Indo-China (Tonkin, Laos, Cam-
bodia), and Malaysia: Sumatra, Malay Peninsula,
and Borneo.

284

FLORA MALESIANA

[sera I voliiG2

Ecol. Chiefly in primary lowland forests, some-
times in periodically inundated forests, rarely up
to 1075 m. Fi. Feb.-Aug., fr. Jan.—Dec.

Vern. Sumatra: adjan, béngkinang, M, balam
budju, katian, tjabé, Palembang; Billiton: djanting,
M; Banka: djurung laki, M; Mal. Pen.: boko-boko,
médang gidap, m. tijoh, pauh rusa.

Use. The wood is used for house building and
beams.

Note. The young racemes sometimes start as
strobilaceous buds, the lower part of the raceme
being covered by numerous, imbricate, small pairs
of tardily caducous stipules (KOSTERMANS 7172,
10319, ENDERT 123E 1P 789).

4. Bhesa archboldiana (MeRR. & PERRY) DING
Hou, Blumea Suppl. 4 (1958) 152.—Kurrimia
archboldiana Merr. & Perry, J. Arn. Arb. 20
(1939) 335; Logs. in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 159.—Fig. 16a-i.

Tree up to 30 m by 50 cm g. Buttresses up to
3 m. Bark pale brown, hard, fissured. Branchlets
light brown. Stipules lanceolate, 4-11 mm.
Leaves elliptic to elliptic-oblong, sometimes ovate-
oblong, 8-16 by 314-8 cm; base rounded or acute;
apex acute to short-acuminate; nerves 9-11 pairs;
petiole 1-414 cm. Racemes 1(—2) in a leaf-axil;
peduncle very short. Pedicels 3-8 mm. Flowers

greenish pink, usually 2 or 3 in a cluster. Calyx
lobes obovate, 3 by 2 mm, obtuse. Petals elliptic-
or obovate-oblong, 414 by 2 mm, obtuse. Stamens
c. 24% mm long; anthers broad-ovoid, c. 1 by
2/, mm, free for lower 24, introrse, slightly api-
culate; filament a continuation of the disk. Disk
shallow-cupular, usually glabrous. Ovary subglo-
bose, c. 114 mm g; styles 2, almost as long as the
ovary, slightly united at the base. Fruits yellow,
ellipsoid, with 2 vertical grooves, gradually
narrowed towards both ends. Seeds ellipsoid,
18 by 9 mm, almost completely enveloped by the
orange aril.

Distr. Louisiades (Sudest I.), D’Entrecasteaux
Is. (Normanby I.), and Malaysia: New Guinea
(Waigeo I., Manokwari, Japen I., Hollandia,
W. Division, E. Division, and Koitaki).

Ecol. Primary forest, from the lowland up to
530 m.

Vern. Diik, Amberbakan, djéra, Papua, kar,
Selogof, onggotu, Depapre, seborereko, Ma-
nikiong.

Excluded
Kurrimia gracilis VipaL, Rey. Pl. Vasc. Filip.
(1886) 89, is according to MERRILL, En. Philip. 2

(1923) 521 = Ventilago dichotoma (BLANCO)
Merr. (Rhamnac.).

10. CASSINE

LINnE [Gen. Pl. (1737) 338]; Sp. Pl. (1753) 268; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 176.—Elaeodendron Jaca. f. ex Jaca. Ic. Pl. Rar. 1? (1782) t. 48;
Murray, Syst. ed. 14 (1784) 241, as Elaeodendrum; Jaca. f. Nov. Act. Helvet.
1 (1787) 38, f. 2; SPRAGUE, Kew Bull. (1929) 43; Logs. in E. & P. Pfl. Fam. ed. 2,
20b (1942) 172.—Fig. 18.

Shrubs or trees. Stipules small, caducous. Leaves decussate (occasionally some
leaves alternate), or alternate (Afr. spp.), subcoriaceous or coriaceous, entire or
crenulate. Cymes axillary or extra-axillary, distinctly peduncled. Flowers bi-
sexual (unisexual in some extra-Mal. spp.), 4-5-merous. Calyx lobes imbricate.
Petals imbricate, spreading. Stamens inserted on or slightly under the outer margin
of the disk; filaments subulate; anthers subglobose, or deltoid, versatile, introrse.
Disk fleshy, flat, orbicular to lobed. Pistil short-conical, or + flask-like, the base
slightly united with the disk, or partly immersed in it. Ovary 2-celled (3—4-celled in
extra-Mal. spp.); style very short or obscure; stigma obscure or slightly 2-lobed.
Ovules 2 in each cell, erect, attached at the base. Fruits indehiscent, 1—2-celled,
the exocarp thin or fleshy, the endocarp firm-leathery, or mostly a stone, very

rarely the mesocarp corky (C. viburnifolia). Seeds 1-2, exarillate, albuminous.

Distr. About 80 spp. (if the African genus Mystroxylon with spiral leaves is included), cosmopolitan
in the tropics of both hemispheres but the bulk of the species in Africa; in Malaysia only 2 spp.; not yet
found in New Guinea. Fig. 19.

Ecol. The two species in Malaysia behave very different. C. viburnifolia is a typical mangrove plant,
C. glauca is a characteristic, though rare, constituent of the monsoon forest.

Notes. The genus Elaeodendron is mostly cited from 1787 but fig. 2 of that reference has also been
published in 1782 as t. 48.

There has been some disagreement about the status of the distinction between Cassine, Elaeodendron,
Mystroxylon, and two other genera. SONDER (FI. Cap. 1, 1860, 451-452) had them as five different gen-

Dec. 1962] CELASTRACEAE (Ding Hou) 285

Fig. 18. Cassine viburnifolia (Juss.) DING Hou. a. Habit, in fruit, < 24, b. flower, in section, = 8, c. section
of ovary, < 16, d. fruit, e. ditto in section, both nat. size. —C. glauca (RottB.) O.K. var. cochinchinensis
Pierre. f. Fruit, g. ditto in section, both 24 (a, d-e SAN A 2895, b-c SAN 10390, f-g KOORDERS 30189),

era; in Cassine the drupe was defined as juicy with a thin crust-like putamen, in Elaeodendron it being
rather dry, with a very hard ligneous putamen. BENTHAM & Hooker (Gen. Pl. 1, 1862, 363, 367) kept
Cassine and Elaeodendron separate, but BAILLON (Hist. Pl. 1, 1877, 33) recognized only the latter. In
1892 LoEsENER reduced Elaeodendron to Cassine and distinguished them as two different sections of
Cassine subg. Elaeodendron, adding that sect. Elaeodendron would have vessels with scalariform, rarely
also simple perforations, and Cassine simple, round or elliptic perforations. Later, however, he reinstated
Cassine, Elaeodendron, and Mystroxylon as distinct genera (in E. & P. Pfl. Fam. Nachtr. 1897, 223;
Bot. Jahrb. 28, 1900, 154; in E. & P. Pfl. Fam. ed. 2, 20b, 1942, 110).

In 1927 Davison (Bothalia 2, 289) merged Elaeodendron and some other genera with Cassine concluding
that there are no generic differences between them. PERRIER DE LA BATHIE, though agreeing that these
two genera cannot be distinguished, arranged all species of Madagascar under Elaeodendron (Not. Syst.
10, 1942, 196-200). Recently also BLAKELOCK followed Davison (Kew Bull. 1956, 556), especially because
MeTCALFE & CHALK (Anat. Dic. 1, 1950, 393) had found that the anatomical characters of the vessels
do not hold in conjunction with the other characters. The recognition of only one genus, Cassine, seems
therefore to be final.

The structure of the pericarp is variable, in some Australasian species both endocarp and exocarp
are rather thin, in others the endocarp is a bony, thinner or thicker stone, and this may be covered with
a pulpy (when fresh probably juicy) exocarp, or the stone may be covered with a thin dry pericarp;

286

FLORA MALESIANA

[ser. I, vol. 6?

in C. viburnifolia there is, between the thin exo- and endocarp on two sides, a thick pithy corky mesocarp.
Also in this genus sterile or fruiting material is very difficult to identify, if at all.

KEY TO THE SPECIES

1. Flowers 4-merous (occasionally some flowers 5S-merous). Fruits obovoid-oblong, often rhombic in
cross-section, c. 12 by 6 mm, the exocarp very thin, mesocarp swollen and soft-corky on both sides,
endocarp rather bony, less than 1 mm thick. Leaves cy discolorous; cymes usually large, up to

11 cm long .

1. C. viburnifolia

1. Flowers 5-merous. Fruits broad- ellipsoid, terete in cross- .-section, 20-25 by 12- 20 mm, the exocarp
coriaceous, endocarp stony, 5S—7 mm thick. Leaves concolorous; cymes usually small, less than 3 cm

long

1. Cassine viburnifolia (Juss.) DING Hou, comb.
nov.—Aegiphila viburnifolia Juss. Ann. Mus. Hist.
Nat. Paris 7 (1806) 76.—Euonymus viburnifolius
Merr. Philip. J. Sc. 9 (1914) Bot. 312, pro comb.,
excl. specim. quae est Euonymus cochinchinensis.—
Elaeodendron viburnifolium Merr. Philip. J. Sc.
16 (1920) 449, t. 1; En. Philip. 2 (1923) 484;
Pl. Elm. Born. (1929) 170; Logs. in E. & P. Pfi.
Fam. ed. 2, 20b (1942) 173.—Elaeodendron subro-
tundum KinG, J. As. Soc. Beng. 65, ii (1896) 356,
excl. RIDLEY 100la, quae est Euonymus cochin-
chinensis; Merr. J. Str. Br. R. As. Soc. n. 76
(1917) 93; En. Born. (1921) 354; Rip. Fl. Mal.
Pen. 1 (1922) 453, f. 44; Crats, Fl. Siam. En. 1
(1926) 287.—Fig. 18a-e.

Small tree or shrub, up to 10 m. Leaves charta-
ceous to subcoriaceous, often slightly discolorous
when dry, obovate, obovate-oblong, sometimes
broad-elliptic, or suborbicular, 4-10!4 by 2-6 cm;
base cuneate; apex acute, obtuse, or rounded,
rarely acuminate; margin remotely crenulate or
subentire; nerves 4-6 pairs; petiole 8-13 mm.
Cymes up to c. 11 cm, usually at the upper parts
of the twigs. Peduncle up to c. 6 cm. Pedicels
1-2 mm. Flowers white, 4-merous (occasionally
some 5-merous). Calyx lobes almost free, broad-
ovate or suborbicular, c. 1 mm g. Petals oblong
or slightly oblong-ovate, 134-2 by 1-114 mm,
obtuse. Disk c. 144 mm o. Stamens just inserted
beneath the outer disk margin, erect, c. 144 mm.
Pistil ovoid. Fruits oboyoid-oblong, often rhombic
on cross-section, sometimes slightly compressed,
the lower 24 gradually narrowed towards the base,
c. 12 by 6 mm; mesocarp thick corky and soft on
both sides, endocarp rather bony; usually 1-
seeded. Seeds obovate-oblong, 6 by 3 mm.

Distr. Siam (Puket, ex Crais), Andaman Is.
and Malaysia: NE. Sumatra (Belawan; also in
Banka and Billiton), Malay Peninsula (Kedah,
Perak, Selangor, Johore, and Singapore), Borneo
(common), Philippines (Sulu Arch.? cf. MERRILL
1923), and Central Celebes (Mengkoka Mts:
Talala).

Ecol. In or along the mangrove swamps and on
the edge of tidal rivers. In the genus the fruit
structure is unique and is clearly adapted to
dispersal by water (RIDL. Disp. 1930, 120, 267,
357, 426).

Vern. Borneo: Barak laut, barat barat, landing-
landing, mémpénai or méta pélandok, rambai laut,
M, api-api, Kedayan and M, damak-damak,

2. C. glauca var. cochinchinensis

Fig. 19. Area and localities of Cassine viburnifolia
(Juss.) Dinc Hou (@) and C. glauca (RoTTB.)
O.K. var. cochinchinensis PIERRE (+).

Kedayan, changlin, gurah, Brunei, pungsu, Brunei
and N. Born., kachang-kachang, Tidung.

Use. The husk of the fruit is said to be used to
stupefy fish (fide NoorDIN SAN 10574, N. Born.).

2. Cassine glauca (Rotts.) O.K. Rev. Gen. Pl. 1
(1891) 114.—Mangifera glauca Rotts. Nye Samm.
Vid. Selsk. Skrift. 2 (1783) 534, t. 4 f. 1.—Celastrus
glaucus VAHL, Symb. Bot. 2 (1791) 42, non R. Br.
(1814), nomen.—Elaeodendrum glaucum PERS.
Syn. 1 (1805) 241; K. & V. Bijdr. 7 (1900) 100.—
Neerija dichotoma Roxs. Fl. Ind. 1 (1820) 646.—
Elaeodendron roxburghii W. & A. Prod. (1834)
157.—Elaeodendron ellipticum Decne, Nouv. Ann.
Mus. Hist. Nat. Paris 3 (1834) 478; Mia. FI. Ind.
Bat. 1, 2 (1859) 591.—C. elliptica O.K. Rev.
Gen. Pl. 1 (1891) 114.

var. cochinchinensis PieRRE, Fl. For. Coch. 4
(1893) t. 296A, ex descr.—Elaeodendron ellipticum
Decne.—Elaeodendron glaucum var. macrocarpa
K. &. V. Bijdr. 7 (1900) 101-102; Back. Schoolfl.
(1911) 235; Bekn. Fl. Java (em. ed.) 6 (1948)
fam. 133, p. 7.—Fig. 18f-g.

Tree, 18-25 m by 53-95 cm go. Leaves sub-
coriaceous, elliptic to elliptic-oblong, obovate,
broad-obovate, ovate, or ovate-oblong, 514-15
by 214-6 cm; base cuneate, obtuse; apex obtuse
or rounded, sometimes short-acute; margins
crenulate, or subentire; nerves 4-9 pairs; petiole
c. 4-10 mm. Cymes few-flowered, usually at the

Dec. 1962]

CELASTRACEAE (Ding Hou)

287

upper part of the branchlets, up to 3 cm. Peduncle
1-214 cm. Pedicels 2-5 mm. Flowers 5-merous.
Calyx lobes slightly reniform, 4% by 1144 mm.
Petals oblong, c. 3 by 144 mm. Disk c. 2 mm @.
Stamens inserted on the outer margin of the disk,
c. 1 mm; pistil partly immersed, the emerging part
short-conical, c. 1 mm long. Fruit a stone covered
by a thin leathery exocarp + mesocarp, broad-
ellipsoid, 2-214 by 1144-2 cm, usually with 2
longitudinal furrows on the outside; exocarp +
mesocarp coriaceous, thin, endocarp stony, up
to 7 mm thick, usually 1-celled and 1-seeded.
Seeds oblong-elliptic, obtuse at both ends, 114 by
1mm.

Distr. Cambodia (Prov. Dongnai) and Ma-
laysia: SE. Java (Puger), Lesser Sunda Is. (W.
Timor), and S. Moluccas (Tanimbar and Kai Is.),
four localities only.

Ecol. In East Java found in periodically very
dry places, weathered coral chalk ground in the
teak-forest and in heterogeneous primary forest
with numerous deciduous tree species below 300 m.

Notes. Elaeodendron ellipticum DECNE is known
only from the type (RIEDLE s.n., P) collected in

Timor. Its vegetative and floral characters are
similar to those of the Javanese specimens.

The type variety, var. glauca (incl. var. rox-
burghii (W. & A.) PieRRE and var. montana
(THw.) PierreE.—Neerija dichotoma Roxs.—
Elaeodendron roxburghii W. & A.) occurs in
Ceylon and India and can be distinguished by
longer, lax, many-flowered cymes (4-6 cm),
larger flower (petals 444 by 3 mm; stamens c.
2 mm long, and disk c. 3 mm @) and smaller fruits
(12-18 mm long).

Cassine australe (VENT.) O.K.—Elaeodendron
australe VENT. Jard. Malm. (1805) t. 117, from
Australia (Queensland and N.S. Wales), is allied
but differs in having drupes with a rather thick
fleshy exocarp and unisexual, 4-merous flowers.

Excluded

Elaeodendron pauciflorum TULASNE, Ann. Sc. Nat.
IV, 8 (1857) 109. In Ind. Kew. this species has
erroneously been referred to as from Malaya.
The type, BERNIER 6182, came from St. Marie,
Madagascar (cf. PERRIER DE LA BAtTuHig, FI.
Madag. fam. 116, 1946, 54).

11. PLEUROSTYLIA
W. & A. Prod. 1 (1834) 157; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 180.—

Fig. 20.

Shrubs or trees. Leaves decussate, subcoriaceous to coriaceous. Stipules very
small, caducous. Cymes in the axil of leaves or bracts, (1—)few-flowered. Flowers
bisexual, 5— (occasionally with a few 4-)merous flowers. Calyx lobes imbricate.

Fig. 20. Pleurostylia opposita (WALL.) ALSTON. a. Habit, x 24, b. flower, <6, c. ditto in section, 6, d.
ovary in section, X 12, e-f. frontal and dorsal view of stamen, x 12, g-h. frontal and lateral view of fruit,
i. fruit in section, showing endocarp, all 3 (a-f Ramos BS. 22350, g-i BRASS 6257).

FLORA MALESIANA

288 [ser. I, vol. 6?

Petals imbricate. Disk cupular, fleshy. Stamens inserted outside the disk; anthers
subbasifixed, introrse, with thick dorsal connective. Pistil conical or flask-shaped.
Ovary free or slightly united with the disk at the base, 2-celled or usually only 1-
celled by abortion; style short, terminal, slightly dilated at the top; stigma capitate
or slightly peltate. Ovules 2(—8 in extra-Mal. sec. Logs.) in each cell, erect. Nuts
1-(also 2?-)celled, with a prominent persistent hardened style on the lateral side;
exocarp + mesocarp thin-coriaceous, endocarp crustaceous, thin, inside glossy
and irregularly ridged, easily separated from the meso- + exocarp, with several
irregular clefts at the base around the hilum of the seed. Seeds 1(—2). Aril evidently

absent.

Distr. About 6 spp. in the tropics and subtropics of Africa, Madagascar, Mascarenes, Ceylon,
Indo-Malaysia, Queensland, and New Caledonia; in Malaysia one species.

Note. Wicut & Arnott (/.c.) stated the 1-celled ovary and fruit to be due to abortion and that the
rudimentary one can be scarcely traced. In the Malaysian material I have examined there is only one cell
in the ovary and fruit; there is a faint indication of a septum (fig. 20d) in cross-section. I have not traced

a single mature, undamaged developed seed!

1. Pleurostylia opposita (WALL.) ALSTON in Trimen,
Fl. Ceyl. 6 (Suppl.) (1931) 48; Merr. & METCALF,
Lingn. Sc. J. 16 (1937) 394.—Celastrus opposita
WALL. in Roxb. Fl. Ind. ed. Wall. 2 (1824) 398.
—P. wightii W. & A. Prod. (1834) 157; WiGHT,
Ic. (1839) 155; Laws. in Hook. f. Fl. Br. Ind. 1
(1875) 617; TRIMEN, FI. Ceyl. 1 (1893) 271;
Logs. Bot. Jahrb. 39 (1907) 171, incl. var. neo-
caledonica Loes.; MerR. Philip. J. Sc. 12 (1917)
Bot. 281, pro var. neocaledonica; RipL. Fl. Mal.
Pen. 1 (1922) 453, as Pleurostylis; MERR. En.
Philip. 2 (1923) 484, pro var. neocaledonica;
Crais, Fl. Siam. En. 1 (1926) 283; Logs. in E. & P.
Pfl. Fam. ed. 2, 20b (1942) 180, f. 31, J.—P.
heynei W. & A. Prod. (1834) 157.—Elaeodendron
microcarpum WHITE & FRANCIS, Proc. R. Soc.
Queensl. 37 (1926) 154, t. 3.—Elaeodendron
mindanaense Merr. Philip. J. Sc. 12 (1917) Bot.
277; MeRR. & Perry, J. Arn. Arb. 29 (1939) 36.

—Fig. 20.
Tree, up to 15 m tall, sometimes a shrub.
Branchlets subterete or obscurely 4-angular.

Leaves chartaceous, ovate to obovate-oblong,
rarely suborbicular, 3-8 by 114-5 cm; base
cuneate; apex obtuse, acute, rarely acuminate, very
rarely retuse; margin entire, sometimes slightly
recurved; nerves about 6 pairs; petiole 2-5 mm.
Cymes 1(—2), axillary, sometimes also terminal,
rarely on the internodes in opposite pairs, few-
flowered, sometimes 1-flowered. Peduncle very
short, 2-3 mm. Pedicel 1-2 mm, articulation
towards the base of stalk. Flowers green. Calyx
lobes rounded or subreniform, 4 by 34-1 mm.
Petals elliptic, or broadly ovate, 1144-11 by 1 mm,
reflexed at anthesis, obtuse or slightly acute.

Disk fleshy, cup-shaped, margin slightly crenate.
Stamens 1-11 mm, attached just beneath the
margin of the disk; filaments subulate, fleshy,
slightly flattened; anthers slightly ovoid, c. ¥4 mm
long, slightly apiculate. Pistil flask-like, the base
adnate to the disk; style very short; stigma ca-
pitate. Fruits ellipsoid or sometimes slightly
obovoid, 5-7 by 4-5 mm, obtuse, 1(—2)-seeded,
sustained by the usually persistent floral parts.

Distr. Ceylon, S. India, Siam (Puket, fide
Crats), China (Hainan), through Malaysia: Malay
Peninsula (Penang, Langkawi Is.), Philippines
(Luzon and Mindanao), and New Guinea to
Queensland and New Caledonia.

Ecol. In lowland primary and secondary light
rain-forest, up to 650 m (cf. MERRILL, 1917).

Uses. The very pretty wood is used in S. India
to make combs; it makes a very beautiful furniture
wood. It contains a moderately poisonous al-
kaloid (cf. Burk. Dict. 2, 1935, 1771; Boorsma,
Bull. Dépt. Agric. Ind. Neéerl. 16, 1908, 5).

Notes. Crarp already suggested the con-
specificity of Celastrus oppositus and Pleurostylia
wightii. In 1931 ALstoN made the transfer but
referred erroneously to ‘Wall. ex Carey in Roxb.
Fl. Ind. ed. II (1832) 389° where the species is not
mentioned.

MERRILL & Perry (J. Arn. Arb. 20, 1939, 336)
identified two specimens collected by BRAss (6257,
8685) in New Guinea as Elaeodendron mindanaense
Merr. They have, however, a cup-shaped disk, a
one-celled ovary with 2 ovules, a thin pericarp,
and entire and reticulate leaves; these characters
agree very well with those of Pleurostylia.

12. PERROTTETIA

H.B.K. Nov. Gen. & Sp. 7 (Dec. 1824) 73, t. 622, non DC. Ann. Sc. Nat. I, 4
(Jan. 1825) 95; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 191.—Caryospermum
BL. Mus. Bot. Lugd. Bat. 1 (1850) 175.—Fig 21.

Shrubs or small trees. Branchlets smooth, usually zigzag. Leaves alternate,

Dec. 1962] _ CELASTRACEAE (Ding Hou) ie 289

(

e

Fig. 21. Perrottetia alpestris (BL.) Logs. ssp. moluccana (BL.) DING Hou. a. Habit, x 24, 6. flower, * 6,

c. ditto, in section, * 8, d. young stamen, frontal and dorsal view, 12, e-f. fruit and its cross section,

3, g-h. seed and its cross section, * 5.—P. alpestris (BL.) LokEs. ssp. philippinensis (VIDAL) DING Hou. i.

Flower, 6, j. leaf margin, 3.—P. alpestris (BL.) Logs. ssp. alpestris. k. Leaf with domatiae, x 7A, /.

leaf margin, = 3, m. domatia enlarged (a RoBBINs 368, b-d HOOGLAND & SCHULTE 7385, e-h ZIPPELIUS
64/d, i-7 STEINER 1941, k-m MEER 1616).

sometimes spiral (extra-Mal.), midrib prominent, sometimes with domatia in the
axils of midrib and nerves. Stipules small, caducous. Inflorescences cymose, axil-
lary in small divaricate thyrses generally shorter than the leaves. Flowers bisexual,
rarely unisexual. Calyx lobes 5 or 4, very rarely 6-8, short, erect, triangular,
valvate or with slightly imbricate margins, very rarely distinctly imbricate.
Floral envelopes persistent. Petals 5 or 4, erect, similar to the calyx lobes in shape
and equal in size or somewhat larger, but not in colour, sometimes ciliate, more
or less distinctly keeled, valvate or slightly overlapping. Disk flat, cup- or ring-
shaped, entire or fine-undulate. Stamens 5 or 4, very rarely 6-8, inserted on the
margin of the disk; filaments subulate; anthers subglobose or ovoid, lengthwise
dehiscent, -_ introrse. Ovary semi-immersed in the disk, mostly 2-celled, or
seemingly 4-celled at the base (cf: Logs. /.c.). Ovules 2 in each cell, basally
attached, erect. Berry globose, 2-4-seeded. Seeds erect, subglobose, thin arillate;
testa thick, muricate-foveolate, or tuberculate, crustaceous, fleshy outside;
endosperm thin; embryo small.

290 FLORA MALESIANA [ser. I, vol. 6?

Distr. About 15 spp., in Central China (E. Szechuan and W. Hupeh, | sp.), Formosa (1 sp.), through-
out Malaysia to NE. Queensland and Solomons (2 spp.), the Hawaiian Is. (1 sp.), and Central America
(Mexico to Columbia, c. 10 spp.). It is remarkable that the genus has not been recorded in continental
SE. Asia between Sumatra and Central China. Fig. 22.

Ecol. In Malaysia chiefly in primary and secondary rain-forest and thickets, from the foothills
through the montane zone up to 2640 m, sometimes also in the lowland, not found in East Java and the
Lesser Sunda Is. and obviously bound to an everwet climate.

Taxon. The Malaysian material falls apart into three groups which are, however, distinguished only
by two minor though constant characters while their geographical distribution shows exactly replacing
areas. These taxa have, therefore, here been distinguished as geographical races and have been designated
as subspecies. All of them have a rather considerable altitudinal range in Malaysia and show the normal
morphological response of woody plants by reduction of leaf-size, thicker leaf texture, and more con-
densed habit in proportion to increase of altitude.

The Chinese species was in Ind. Kew. wrongly attributed to Japan with the erroneous basionym Ce-
lastrus racemosus TuRCZ. from Java (not Japan). Its proper synonymy runs as follows:

P. racemosa (Outv.) Loes. Bot. Jahrb. 24 (1897) 201, 200; in E. & P. Pfl. Fam. Nachtr. 1 (1897) 224;
Bot. Jahrb. 29 (1900) 447; ibid. 30 (1901) 474.—Ilex racemosa OLtv. in Hook. Ic. Pl. 19 (1889) t. 1863.
It differs from P. alpestris by glabrous sepals, ciliate petals, very long exserted stamens (similar to some
American species!), and less nerves; it has 5-merous flowers.

The Formosan species, P. arisanensis HAYATA is distinct by very narrow petals and sepals and is said
to be deciduous; it has 4-merous flowers as the Central Malaysian P. alpestris ssp. philippinensis.

The heavy reduction in number of Malaysian species accepted here may also be necessary in the New
World species when these are critically compared and revised; CUATRECASAS already observed that
‘they differ from each other by minute characters’ (Lloydia 11, 1948, 223).

Attention should be given to the tendency towards unisexual flowers and monoecism, or even dioecism,
in comparing floral characters.

It is remarkable that such a block of very closely related species shows such enormous disjunctions in

its colossal geographical range.

1. Perrottetia alpestris (BL.) Lors. in E. & P.
Pfl. Fam. 3, 5 (1892) 220.—Celastrus alpestris
BL. Bijdr. (1826) 1145.

The synonyms have been arranged under the
subspecies

KEY TO THE SUBSPECIES

1. Flowers 5-merous.

2. Leaf margins distinctly and rather closely
glandulose-serrulate or -crenulate, the upper
part of the teeth callose; base usually rounded.
ssp. alpestris
. Leaf margins entire, or with sparse, remote,
minute, pointed teeth; base acute or cuneate,
rarely obtuse nen ssp. moluccana

1. Flowers 4-merous. Leaves serrulate.
ssp. philippinensis

tN

ssp. alpestris.—Celastrus alpestris BL. Bijdr. (1826)
1145.—Caryospermum serrulatum Mig. Fl. Ind.
Bat. 1, 2 (1859) 592; K. & V. Bijdr. 7 (1900) 97.
—Caryospermum alpestre O.K. Rey. Gen. Pl. 1
(1891) 113.—P. alpestris Logs. in E. & P. Pfl.
Fam. 3, 5 (1892) 220; Nachtr. (1897) 224; Bot.
Jahrb. 24 (1897) 200; K. & V. Ic. Bog. 2 (1904)
137, t. 127; Back. Schoolfl. (1911) 235; Koorp.
Exk. Fl. Java 2 (1912) 526, f. 80; Rip. Fl. Mal.
Pen. 1 (1922) 454; Koorp. Fl. Tjib. 2 (1923)
146; Burk. & HENDERSON, Gard. Bull. S.S. 3
(1925) 360; BarTLeTT, Pap. Mich. Ac. Sc. 6
(1926) 53; HENDERSON, Gard. Bull. S.S. 4 (1927)
95; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942) 193.
—Maesa perakensis Ripv. J. Fed. Mal. St. Mus.
4 (1909) 45.—Fig. 21k—m.

An evergreen shrub or a small tree up to 8 m
by 10 cm g. Branchlets puberulous when young,

glabrescent. Leaves chartaceous, nerves and veins
on both surfaces usually covered with appressed
hairs, sometimes the upper surfaces glabrescent;
Ovate-oblong, lanceolate, or sometimes ovate,
614-23 by 214-9 cm; base obtuse or rounded,
sometimes slightly cordate; apex acuminate (acu-
men up to 1!4 cm); margin glandulose-serrulate,
or -crenulate, the upper part of the teeth callose,
usually black, sometimes yellowish, blunt; petiole
7-15 mm. Inflorescences usually 1-2 cm long,
sometimes even shorter, rarely up to 5 cm, some-
times branched almost near the base; peduncle
very short, sometimes up to 2 cm. Bracts small,
c. 2 mm long. Pedicels short, 14-114(-24%4) mm.
Flowers white, or light greenish, small, c. 3 mm 9.
Calyx lobes 5, deltoid, 14-24 mm long, sparsely
puberulous on both surfaces, sometimes only on
the inside especially towards the upper part.
Petals 5, triangular, or ovate, 14-114 by 74-1 mm,
sparsely puberulous on both surfaces, sometimes
only puberulous on the margins. Disk shallow
cupular, c. 144 mm @. Stamens c. 74 mm long;
filaments much longer than the disk. Pistil slightly
united with the disk at the base, conical, c. /% mm
long, narrowed into a short style; stigma obscurely
2-lobed. Fruits globose or subglobose, 214-3'4
mm, reddish black, 3—4-seeded, glabrous. Seeds
subglobose, or slightly obovate, 114-2 mm long,
sometimes cuneate at the base, foveolate-rugulose
or tuberculate.

Distr. Malaysia: commonly distributed in
Sumatra, Malay Peninsula, and Java (as far E
as Mt Lawu). Fig. 22.

Ecol. Everwet primary and secondary forests
and thickets in the colline and montane zones,
500-2500 m, chiefly above 1000 m.

Dec. 1962]

CELASTRACEAE (Ding Hou)

291

Fig. 22. Old World distribution of the 3 species of

the genus Perrottetia; in the Malaysian P. al-

pestris (BL.) Logs. there are three replacing sub-
species, separated by dotted lines.

Vern. Sumatra: bérbah, Bencoolen, kaju
attarasa pidong, k. si lando, Asahan, kaju musang,
W. Coast; Mal. Pen.: kayu tungas, Perak; Java:
kibélut, ki-bémok, ki dagé, kihiris, ki-hurang, ki-
kiris, ki purut, kipiit, kitarasi, S, kémalon, keé-
maluhan, wadéran, J.

ssp. moluccana (BL.) DING Hou, comb. nov.—
Vertifolia rubra RumPH. Herb. 3 (1741) 100, t. 67.
—Caryospermum moluccanum BL. Mus. Bot.
Lugd. Bat. 1 (1850) 176.—Caryospermum ar-
borescens F. vy. M. Fragm. Phyt. Austr. 6 (1868)
202; ibid. 8 (1874) 279; BaILeEy, Queensl. Fl. 1
(1899) 258.—P. moluccana (BL.) Logs. in E. & P.
Pfl. Fam. 3, 5 (1892) 220; Nachtr. (1897) 224;
Pfi. Fam. ed. 2, 20b (1942) 193.—P. arborescens
(F. v. M.) Logs. Bot. Jahrb. 24 (1897) 200; in
E. & P. Pfl. Fam. ed. 2, 20b (1942) 193.—P.
grandifolia Rip. Trans. Linn. Soc. II, Bot. 9
(1916) 30; Merr. & Perry, J. Arn. Arb. 20 (1939)
336; Logs. in E. & P. Pfl. Fam. ed. 2, 20b (1942)
193.—P. nervosa Ript. Trans. Linn. Soc. II,
Bot. 9 (1916) 30.—P. Jlauterbachiana_ Logs.
Notizbl. Berl.-Dahl. 12 (1934) 36, incl. f. B ma-
crophylla Loers.; in E. & P. Pfl. Fam. ed. 2, 20b
(1942) 193.—P. schlechteri Loes. Notizbl. Berl.-
Dahl. 12 (1934) 37; in E. & P. Pfl. Fam. ed. 2,
20b (1942) 193.—P. traumatophylla MeERR. &
Perry, J. Arn. Arb. 22 (1941) 261.—Fig. 21a-h.

Treelet 5—10 m, rarely tree up to 24 m. Leaves
at the base cuneate to attenuate, rarely obtuse.
Peduncles 1-5 cm, sometimes almost none.
Fruits 214-6 mm 9g.

Distr. Solomon Is. (Bougainville, Isabel, and
Guadalcanal Is.), Australia (NE. Queensland,
rare), and Malaysia: Moluccas (Ceram, Ambon,
and Buru) and New Guinea (rather common;
also Fergusson I.). Fig. 22.

Ecol. Forests, from lowland up to 2640 m.

Vern. Moluccas: kasubuwa, Ceram; New
Guinea: aimbeh, Wahgi, bopeh, Chimbu, fonanitur,
Onjob lang., gamaha, Asaro, ihira, Mairi, kun-
guna, Hagen, togump, Togaba, fsi/tsibi, Tomba,
waljamahp, walumap, Enga lang.

ssp. philippinensis (VIDAL) DING Hou, comb. nov.
—Caryospermum philippinense VIDAL, Rev. Pl.
Vasc. Filip. (1886) 89.—P. philippinensis Logs.
in E, & P. Pfl. Fam, 3, 5 (1892) 220; Bot. Jahrb.
24 (1897) 200; in E. & P. Pfi. Fam. ed. 2, 20b
(1942) 191.—P. alpestris var. philippinensis STAPF,
Trans. Linn. Soc. II, Bot. 4 (1894) 141; Logs.
in E. & P. Pfl. Fam. Nachtr. (1897) 224; MERR.
Philip. J. Sc. 5 (1910) Bot. 200; Gisss, J. Linn.
Soc. Bot. 42 (1914) 64.—P. alpestris (non (BL.)
Logs.) Koorp. Minah. (1898) 396; Merr. En.
Born. (1921) 354; En. Philip. 2 (1923) 484.—
Fig. 21i-j.

Shrub or small tree up to 16 m. Leaf-base
rounded, sometimes cuneate. Inflorescences (1—)
5—12(—21) cm long. Fruits c. 3 mm o.

Distr. Malaysia: Philippines (Luzon, Mindoro,
Panay, Camiguin de Misamis, and Mindanao),
Borneo (N. Borneo, Sarawak, and Kutai), and
Celebes (Minahasa to SW. Celebes). Fig. 22.

Ecol. Rain-forests and thickets, (350—)700—
2700 m.
Vern. Borneo: maesa, Sarawak; Philip.:

balakbakan, Buk., bubayug, Ig., dafgalis, Bag.,
tigaundako, C. Bis.; Celebes: kaju-werang, ma-
hawerang, mukuwerung, Minahasa, lotong matjula,
Polewali.

Excluded

Perrottetia caudata Ripi. Trans. Linn. Soc. II,
Bot. 9 (1916) 31, from New Guinea, leg. BODEN
Koss, in K, BM, is, according to kind information
of Mr L. L. ForMAN, Kew = Rhus cf. linguata
W. Sis (Anacardiaceae).

of

Lie
ny

7

LOGANIACEAE (P. W. Leenhouts, Leyden)

Trees, shrubs, woody climbers, or herbs. Hairs simple, stellate, or glandular-
capitate; colleters often present in the axils of the leaves, stipules, and sepals
(among Mal. genera absent in Buddleja only). Leaves nearly always opposite,
entire or nearly so, penninerved, rarely 3—7-plinerved (Strychnos) or curvinerved
(Mitrasacme); stipules interpetiolar (in many genera reduced to a stipular line)
and in some genera moreover intrapetiolar. Flowers in cymose to thyrsiform
(rarely racemose or spicate) inflorescences or solitary, 5-(rarely 4-, in Anthocleista
up to 16-)merous, nearly always bisexual, actinomorphic (in some genera slightly
zygomorphic). Disk sometimes present (not in Mal. spp.). Sepals united or free.
Corolla gamopetalous, very rare with a corona. Stamens isomerous in Mal. spp.
(in 2 extra-Mal. genera less), alternating, inserted on the corolla tube (with one
exception in Buddleja), included or exserted; anthers basifixed or sometimes slightly
dorsifixed (in the Spigelieae), slightly to deeply bifid at base, lengthwise dehiscent.
Ovary superior (in Polypremum, Cynoctonum, and Mitrasacme p.p. semi-inferior),
(1—)2(-4)-celled, placentas axile (parietal if 1-celled), often peltate; ovules 1-c©
per cell, amphitropous or anatropous; style usually one. Fruit always superior,
capsular, baccate, or drupaceous. Seeds 1-co, with copious endosperm; embryo
minute, straight, cotyledons small.

Distribution. About 28 genera with some 600 spp., almost confined to the tropics of both eastern
and western hemispheres, a few genera extending to the warm-temperate regions, mainly towards the
south. In Malaysia 11 genera with 80 spp.

Ecology. The majority of the genera and species are confined to the everwet tropical lowland. A
few are indifferent to climate (e.g. Cynoctonum mitreola, Fagraea racemosa, and F. fragrans); a few
others are characteristic for areas subject to distinct periodical drought, e.g. Strychnos lucida (fig. 29),
Mitrasacme (M. elata, fig. 41, and a few others). As to altitude almost all representatives occur below
2000 m, Buddleja asiatica and a few species of Fagraea and Geniostoma are ascending to the lower border
of the subalpine zone, but none is found above 3000 m.

Almost all species occur in dryland sites; Fagraea crenulata is confined to temporarily or permanently
inundated swamps (fig. 18), several Neuburgias occur in damp alluvial forest, Fagraea racemosa can
also stand marshy conditions.

The role played by Loganiaceae in vegetation is small as none occurs gregariously. Only very few species
are sizeable trees, notably old specimens of Fagraea fragrans, F. elliptica, F. crenulata, both Norrisias,
and further a few Neuburgias. A temporary predominance in seral vegetation types is found with Buddleja
asiatica which is a characteristic shrub of disturbed terrain and Fagraea fragrans which is rather fire-
resistant (fig. 20).

Flowers are usually whitish and frequently fragrant, mainly attracting insects; some obviously nocturnal
(certain Fagraeas); it has been suggested that some large-flowered Fagraeas are visited by birds, or even
bats, but this matter is insufficiently investigated.

The seeds from species with capsular fruits are mostly very light and often winged (Norrisia, Gelse-
mium, Buddleja); they are dispersed by wind; the small to rather big baccate fruits of some genera are
swallowed by birds and bats who will be responsible for the dispersal of the seeds. Neuburgia seems to
possess buoyant fruits, possibly dispersed by water.

Wood-anatomy. DEN BerGer, Determinatietabel houtsoorten van Malesié, Veenman, Wageningen
(1949) 20, 21, 27, 60; Descu, Mal. For. Rec. 15! (1941) 302 (hand lens); CHALK & CHATTAWAY, Trop.
Woods 50 (1937) 1; METCALFE & CHALK, Anat. Dic. 2 (1950) 928; Mo_t & JANSSONIUS 4 (1926) 652;
PEARSON & BROWN, Commercial timbers of India 2(1932) 738. According to CHALK (METCALFE & CHALK
/.c.) the woods of the family suggest more than one distinct group, but according to them the material
available for examination was insufficient to justify any more detailed conclusions. JANSSONIUS (MOLL &
JANSSONIUS /.c.) studying Geniostoma rupestre Forst. (= G. haemospermum StEvuD., G. miquelianum
K. & V., G. oblongifolium K. & V.), Buddleja asiatica Lour. and Fagraea racemosa JACK ex WALL. (=
F. morindaefolia Bu.) and F. blumii G. DON(= F. obovata-javana BL.) has claimed that they belong to
two families, one including Geniostoma and Buddleja, the other Fagraea. It seems to me that the differ-
ences between the genera are not large enough to justify such a conclusion and that for the present, there
is no evidence against LEENHOUTS’ subdivision into tribes.—C.A.R.—G.

(293)

294 FLORA MALESIANA [sera dy vole?

Phytochemistry. From this family the great pantropic genus Strychnos has attracted the attention of
many phytochemists. It was known early that most species are toxic and contain alkaloids. Today many
pure Strychnos alkaloids are known and all of them proved to be indol derivatives. Most South Ameri-
can species contain highly complex curarizing quaternary bases in the barks. The most recent reviews
of these alkaloids are those of BERNAUER (Fortschr. Chemie Org. Naturstoffe 17, 1959, 183; Planta
Medica 9, 1961, 340). On the other hand Asiatic and Australian Strychnos species contain convulsive
alkaloids of the strychnine-type. Strychnine, brucine, and closely related bases were demonstrated to be
present in many species from Asia, Malaysia, and Australia. The alkaloids of the African species of
Strychnos have not yet been investigated thoroughly. Indole-alkaloids occur also in the African genus
Mostuea and in the Chinese and North-American genus Ge/semium. In other genera alkaloids have been
demonstrated to be present but their chemistry is unknown. Complex indole-alkaloids are restricted, as
far as we know, among Angiosperms to the families of Loganiaceae, Apocynaceae, and Rubiaceae. This
may well indicate that they are all phylogenetically closely related as suggested by HUTCHINSON.

Besides alkaloids some Loganiaceae accumulate pseudo-indicans, i.e. glycosides producing blue
colours under certain conditions. Loganin and loganic acid were found in many species of Strychnos
(A. DENOEL ef a/., Contribution a l’étude chimique des Strychnos du Congo Belge, Bruxelles 1953;
publ. par Ministére des Colonies, Direction de l’Agriculture). Loganin has also been isolated from Me-
nyanthes trifoliata (Gentianaceae). In the genus Buddleja loganin is replaced by the related glycoside
aucubin, which is a characteristic compound of many families of Tubiflorae (CHASLOT, Thése [Pharm.]
Univ. Paris 1955). This may indicate rather clear relationships between Buddlejeae and some families of
Tubiflorae. This assumption is strengthened by the fact that still other types of glycosides are common to
Buddlejeae and different taxa of Tubiflorae.The flavonoid compound buddleo-flavonoloside is known to be
identical with linarin from Linaria (Scrophulariaceae) and according to a recent suggestion of HARBORNE
& CORNER (Biochem. J. 81, 1961, 242) the third heteroside (= buddleoside) known to occur in Buddleja
is probably identical with orobanchoside from several species of Orobanche (Orobanchaceae). The
recent elucidation of the structures of loganin, aucubin, and other pseudo-indicans (for instance as-
peruloside from Rubiaceae) and related compounds (gentiopicrin and gentianin from Gentianaceae)
seems to be highly interesting for taxonomy (for structures compare R. THoMas, Tetrahedron Letters
1961, 544-553). It is very probable that many constituents isolated from species belonging to families
of the orders Loganiales, Contortae, Tubiflorae, and Rubiales are biochemically intimately related.

As a whole, the Loganiaceae have hitherto not been studied intensively by phytochemists. Most of the
research has been concentrated on the alkaloid-bearing species of Strychnos. Chlorogenic acid, some triter-
penes, rubber (fruit of Fagraea), bitter principles (Fagraea) and saponins were demonstrated to be pres-
ent in some species. The distribution and chemistry of these compounds is not yet sufficiently well
known to permit their use forchemotaxonomical speculations. On the other hand, as already mentioned,
Loganiaceae are biochemically related to Apocynaceae and Rubiaceae by the glycosides and alkaloids
of the subfamily of Loganioideae and to Scrophulariaceae and derived families by the different types of
glycosides present in the subfamily of Buddlejoideae-——R. HEGNAUER.

Morphology. Stipules. The leaf-bases are probably all developed into vaginae connatae. They can be
developed as a stipular line only, can form a sheathlike connection between the petioles (vaginae inter-
petiolares; fig. 23b), or can be moreover connected in the leaf-axils as an ocrea (fig. 10b). In the last case
either the interpetiolar part may be strongest developed (interpetiolar stipules, fig. 31a) or the axillary
part is predominant (axillary scales’ of Fagraea, fig. 3, 6, 20f). The stipules resemble those of the Ru-
biaceae; they are persistent except in Mostuea where the upper portion withers. Some species of Fagraea
have auricles at the base of the petiole outside of (and sometimes hiding) the ocrea; they may be taken
at first sight as stipules, but are of distinctly laminar origin (fig. 3c-d). Most genera of the Buddlejeae
show a (sometimes faint) stipular line. In Buddleja the lamina is often decurrent along these stipular lines;
in that case there may be a pair of auricles at the leaf-base, or these may be combined into an ‘interpetiolar
stipule’ on either side of the node, or the leaves may be perfoliate. In some Buddlejas the interpetiolar
stipules develop into a second pair of (slightly smaller) leaves decussate to and inserted between the normal
leaf pair (analogous to those of Galium and Asperula in the Rubiaceae). Whether the stipular line of the
Buddlejeae is comparable with the vaginae connatae of the other Loganiaceae deserves further study. It
seems to me that there is a gradual series from the stipules of the Loganiaceae to those of the Rubiaceae,
the latter usually being better developed organs. In the related Apocynaceae axillary scales similar to those
found in Fagraea are also known, for example in Conopharyngia, but they also occur in other families,
for instance Guttiferae. HASSELBERG (Symb. Bot. Upsal. 2, 3, 1937, 1-170) gave an extensive account of
the morphology of the stipules in the Loganiaceae.

Aestivation of the corolla-lobes. The aestivation of the corolla-lobes can be valvate, imbricate, or
contorted. This character seems to be of some systematical importance; in most of the tribes, and in all
genera with only one exception, we find only one type of aestivation:

Potalieae: contorted.

Buddlejeae: mostly imbricate, valvate in Peltanthera and Nuxia.

Antonieae: valvate.

Gelsemieae: imbricate.

Strychneae: valvate.

Dec. 1962] LOGANIACEAE (Leenhouts) 295

Loganieae: Logania imbricate, Geniostoma imbricate or contorted, Labordia contorted.

Spigelieae: Spigelia and Mitrasacme valvate, Cynoctonum and Polypremum imbricate.

Delimitation and subdivision. Loganiaceae are considered to be a heterogenous family. Especi-
ally the position of Buddleja and related genera has given rise to controversial opinion. To gain a better
insight in this matter I felt compelled to study all genera, from almost all of which herbarium material
was available and has been analyzed; in few cases my work was supplemented by examining published
descriptions and plates. Furthermore, I have used data derived from anatomy, cytology, palynology,
embryology, and phytochemistry.

From these studies emanated the following subdivision into tribes, to which have been added the names
of genera which I have accepted:

A. Potalieae: (1) Potalia, (2) Anthocleista, (3) Fagraea.

B. Buddlejeae: (4) Peltanthera, (5) Sanango, (6) Nuxia, (7) Androya, (8) Gomphostigma, (9) Buddleja,
(10) Emorya, (11) Adenoplusia, (12) Adenoplea.

C. Antonieae: (13) Bonyunia, (14) Antonia, (15) Norrisia, (16) Usteria.

D. Gelsemieae: (17) Gelsemium, (18) Mostuea.

E. Strychneae: (19) Strychnos, (20) Gardneria, (21) Neuburgia.

F. Loganieae: (22) Geniostoma, (23) Labordia, (24) Logania.

G. Spigelieae: (25) Polypremum, (26) Cynoctonum, (27) Spigelia, (28) Mitrasacme.

Genera dubia: Refzia, Desfontainea.

I agree with SOLEREDER (in E. & P. Nat. Pfl. Fam. 4, 2, 1892-95, 19-50) that the Loganiaceae represent
a well-circumscribed family, and that at least some of the tribes show clear mutual relationships. From
this follows that I disagree with HUTCHINSON (Fam. FI. Pl. ed. 2, 1, 1959, 370-378) who raised nearly
all tribes to family rank, keeping only the Loganieae and Gelsemieae in Loganiaceae in the restricted sense;
he largely maintained the suprageneric taxa of BENTHAM & Hooker. His delimitation of some of the tribes
differs from that of SOLEREDER; therefore, his families are sometimes less homogeneous and less sharply
defined than SOLEREDER’s tribes. For the rest, the raising of tribes to the rank of families leads, it seems
to me, not to scientific gain if these families are mutually kept in the same position within an order.

Critical remarks in more detail follow below; all generic names mentioned by either SOLEREDER or
HUTCHINSON are included, as well as some more recently ascribed to the family.

A. Potalieae. For a survey, see LEEUWENBERG, Act. Bot. Neerl. 10 (1961) 1-53.

(1) Potalia. Monotypic; tropical South America. Shows distinct relationships to both other genera.

(2) Anthocleista. 14 spp.; tropical Africa. For a revision see LEEUWENBERG, /.c. Closest allied to Fagraea.

(3) Fagraea. c. 35 spp.; SE. Asia, Malaysia, tropical Australia, and the southwestern Pacific. Distinctly
related to both other genera.

B. Buddlejeae. This is a group of mutually distinctly related genera, the systematic position of which
is still a source of controversial opinion. In fact this all comes to the same thing: its position with either
the Scrophulariaceae or the Loganiaceae, as a tribe or a subfamily, or as a family of its own. Up to and
including EICHLER (Bliithendiagr. 1, 1875, 210) this generic assemblage was mostly treated as a tribe of
the Scrophulariaceae. BENTHAM & HOOKER (Gen. PI. 2, 1876, 787) were apparently the first who included
it in the Loganiaceae (as a subtribe of the tribe Euloganieae), probably on account of the presence of
stipules. SOLEREDER, /.c., raised it to the rank of a subfamily, and opposed it to the other Loganiaceae,
mainly on anatomical grounds, remarking that, apart from the stipules, they could as well be included
in the Scrophulariaceae. WILHELM (Die Samenpfil. 1910, 90) was apparently the first who raised
this group to family rank. These Buddlejaceae were usually placed near the Loganiaceae, but sometimes
near the Scrophulariaceae. Recently HARTL (Oest. Bot. Z. 103, 1956, 185-242) included them again as a
tribe in the Scrophulariaceae, in which he was followed by some German authors (WAGENITZ, TROLL).

The main differences between the Buddlejeae and the other Loganiaceae mentioned are:

(i) Intraxylary phloém. Absent in the Buddlejeae and Polypremum, present in all other Loganiaceae.

(ii) Glandular-capitate hairs. Present in Buddlejeae except Peltanthera, in Mostuea (on the style),
in Logania (on the ovary), and in Mitrasacme (on the calyx, sec. KLetr in Mez, Bot. Arch. 5, 1924,
327), absent in all other Loganiaceae.

(iii) Stellate hairs. Present in the Buddlejeae (except Peltanthera and Sanango) and in Spigelia, absent
in all other Loganiaceae.

(iv) Unicellular or uniseriate hairs. Absent in most of the Buddlejeae, present in Peltanthera and
Sanango of the Buddlejeae and in the other Loganiaceae.

(v) Pericyclic cork in the Buddlejeae except Peltanthera and probably Sanango, superficial cork in the
other Loganiaceae.

(vi) Colleters. Absent in the Buddlejeae and Polypremum, present in the other Loganiaceae, at least
in the leaf axils.

The genera usually included in the Buddlejeae (those cited in the survey above under B, plus Poly-
premum) are thus still marked off from the other Loganiaceae by the absence of intraxylary phloem and
of colleters. Data derived from cytology, palynology, embryology, and phytochemistry sustain in general
a relationship with the Scrophulariaceae. However, these data are very incomplete and comparisons often
based on one or a few genera which are only distantly related (for example Spigelia and Buddleja). These
arguments may therefore loose significance if all genera are examined, as for example has already appear-

296 FLORA MALESIANA [ser. I, vol. 6?

ed in palynology and chromosome numbers which have yielded a very varied pattern within the Loga-
niaceae. A comparison in these fields of Pe/tanthera with the Antonieae and of Polypremum with the Spi-
gelieae would be highly desirable.

Two other facts are in favour of a closer relationship between the Buddlejeae and the Loganiaceae.
The genus Pe/tanthera (and the closely related Sanango) is on the one side closely related to Nuxia, but
shows on the other side a distinct alliance to the Antonieae, and was in fact included by HUTCHINSON
in the Antoniaceae. Polypremum has been included mostly in the Buddlejeae, but HUTCHINSON included
it in the Loganiaceae sensu stricto; in my opinion it should be included in the Spigelieae on account of the
herbaceous habit, the membranous, sheath-like interpetiolar stipules, the partly inferior ovary, and the
nearly basally inserted peltate placentas, all characters which are unusual in the Buddlejeae. Inclusion of
Polypremum in the Spigelieae as accepted here entails the breakdown of the only two characters by which
the Buddlejeae deviate from the other Loganiaceae.

Concluding, the affinity of the Buddlejeae is clearly with both the Loganiaceae and the Scrophulariaceae.
But on account of the unsharp delimitation against the Loganiaceae and especially the distinct relation-
ships with the Antonieae I prefer to include them in the Loganiaceae. In the Scrophulariaceae they would
represent an isolated basal offshoot; within the Loganiaceae they are definitely less isolated than the Po-
talieae and the Spigelieae.

(4) Peltanthera. Monotypic; tropical Central and South America. Especially related to Sanango and
to Nuxia, furthermore to the Antonieae (included by HUTCHINSON in the Antoniaceae).

(5) Sanango. Monotypic; tropical South America. Related to Peltanthera.

(6) Nuxia (syn. Lachnopylis). About 20 spp.; South Africa and Madagascar. Close to Peltanthera and
to Sanango.

(7) Androya. Monotypic; Madagascar. Closely related to Nuxia but sufficiently different. Originally
described in the Oleaceae.

(8) Gomphostigma. A few spp.; South Africa.

(9) Buddleja (syn. Chilianthus and Nicodemia). More than 100 spp.; in the tropics and subtropics,
worldwide with the exception of the western half of Africa, Australia, and the Pacific. Emorya, Adeno-
plusia, and Adenoplea are closely related and should probably be united with it.

(10) Emorya. Monotypic; southwestern North America. Probably to be included in Buddleja.

(11) Adenoplusia. A few spp.; Madagascar and East Africa. Closely related to Adenoplea; both should
possibly better be sunk in Buddleja.

(12) Adenoplea. A few spp.; Madagascar. Should probably be combined with Adenoplusia, and both
possibly with Buddleja.

C. Antonieae.

(13) Bonyunia. About 2-5 spp.; tropical South America. Close to the next two genera.

(14) Antonia. Monotypic; tropical South America. Close to both Bonyunia and Norrisia.

(15) Norrisia. Two spp.; Malaysia. Close to the foregoing two genera.

(16) Usteria. Monotypic; tropical Africa. Somewhat isolated.

D. Gelsemieae.

(17) Gelsemium. Three spp.; southern North and northern Central America and SE. Asia (also in
Malaysia). Shows also relationship to the Antonieae.

(18) Mostuea (syn. Coinochlamys). Eight spp.; tropical Africa and northern South America. Re-
vision: LEEUWENBERG, Med. Landbouwhogesch. Wageningen 61, 4 (1961) 1-31. Shows also relationship
to the Strychneae.

E. Strychneae.

(19) Strychnos (syn. Scyphostrychnos; see LEEUWENBERG, Act. Bot. Neerl. 11, 1962, 47-50). About
150-200 spp.; worldwide in the tropics and subtropics.

(20) Gardneria (syn. Pseudogardneria). Five spp.; SE. and E. Asia, Malaysia. Revision: LEENHOUTS,
Bull. Jard. Bot. Brux. 32 (1962) 431-439.

(21) Neuburgia (syn. Couthovia and Crateriphytum). About 10-12 spp.; Malaysia and the southwestern
Pacific. Shows also affinities to the Loganieae.

F. Loganieae.

(22) Geniostoma. About 20-40 spp.; Malaysia and the Pacific. Close to Labordia.

(23) Labordia. About 20 spp.; Hawaii. Hardly generically different from Geniostoma.

(24) Logania. About 20-30 spp.; Australia (also Tasmania and New Zealand) and New Caledonia.

G. Spigelieae.

(25) Polypremum. Monotypic; subtropical North and South America. By SOLEREDER and most
subsequent authors included in the Buddlejeae, by HUTCHINSON in the Loganiaceae sensu stricto. Dis-
tinctly related to Mitrasacme and Cynoctonum; compare also the notes to the Buddlejeae.

(26) Cynoctonum (syn. Mitreola). Six spp.; tropical and subtropical America, Madagascar, SE. Asia,
Malaysia, and North Australia.

(27) Spigelia (syn. Pseudospigelia). About 50 spp.; tropical and subtropical America, 1 species na-
turalized in Africa and Malaysia.

(28) Mitrasacme. About 40 spp.; Australia (also Tasmania and New Zealand), SE. and E. Asia,
Malaysia, the Carolines, and New Caledonia.

Dec. 1962] LOGANIACEAE (Leenhouts) ; 297

Mitrasacmopsis, a monotypic genus from Madagascar, described by Jover in the Loganiaceae-Spigelieae
and accepted as such by HUTCHINSON, should be excluded from this family. The leaves show abundant
raphides, a character which is absent in the Loganiaceae, but characteristic of many Rubiaceae, as Dr
C. E. B. BREMEKAMP informed me. It should be included in the Rubiaceae-Hedyotideae and seems to be
especially close to the African genus Diotocranus.

Desfontainea. Monotypic; Andine South America. Included by HUTCHINSON in the Potaliaceae, also
occasionally placed in or near the Solanaceae.

Retzia. Monotypic; South Africa. Relationships uncertain.

Systematic affinities. The relationships of the Loganiaceae are threefold:

Especially the Poralieae are distinctly related to the Apocynaceae-Tabernaemontaninae. The main
characters in which they differ from the Apocynaceae are the absence of laticiferous vessels (but Fagraea
has a laticiferous tissue under the epidermis of the fruits) and of the more complicated type of stigma
characteristic of the latter family.

The Spigelieae show a distinct relationship to the Rubiaceae-Hedyotideae. As a whole, superior versus
inferior ovary is a good distinguishing character between Loganiaceae and Rubiaceae. In this case the
character is, however, not sharp: in the Spigelieae the ovary is superior or partly inferior, in the Hedyoti-
deae it is often only halfway inferior or even less. The best distinguishing characters are the absence of
raphides and the always superior fruit in the Spigelieae.

The Buddlejeae show an indubitable affinity to the Scrophulariaceae, as discussed above. The main
difference with the latter family is the stipular line which is present in most genera.

Possibly there is some relationship with the Oleaceae which must then be found with the Antonieae
and Gelsemieae.

The morphological relationships are mainly confirmed by the data provided by the auxiliary sciences.

I agree with HUTCHINSON in accommodating the Loganiaceae in an order Loganiales, from which the
orders Contortae (Gentianales), Tubiflorae (Scrophulariales), and Rubiales can be derived. The Oleaceae
should possibly be included in the Loganiales, as proposed for example by SOLEREDER and by HUTCHINSON.

Uses. Some species are reputed for their alkaloids. Best known of these are strychnine and brucine
in the seeds of several Strychnos spp., especially S. ignatii and S. nux-vomica; furthermore the very poison-
ous gelsemin from the roots of Gelsemium. Some species of Fagraea and the Norrisias are good timber-
trees. A few species, mainly of Fagraea and Buddleja, are planted as ornamentals.

Notes. Many years ago Dr C. A. BACKER started a revision of this family but had to give up on
account of failing eyesight; his MSS were put at my disposal. I have further cooperated with Dr A. J. M.
LEEUWENBERG, Wageningen, who is revising the African Loganiaceae, by exchange of MSS.

KEY TO THE GENERA
1. Woody plants.
2. Flowers 4-merous. At least underside of leaf always tomentose; hairs stellate or glandular-capitate.
2. Buddleja
2. Flowers 5-merous. Plants mostly -- glabrous; underside of leaf not tomentose; hairs neither stellate
nor glandular-capitate.
3. Leaves 3-5-plinerved. Tendrils often present .............. . 5. Strychnos
3. Leaves penninerved. No tendrils.
4. Inflorescences terminal, sometimes also lateral.
5. Corolla-lobes valvate in bud.
6. Corolla outside densely tomentose, without a hair-ring in the mouth. Capsule 2-valved, densely

hairy. Seeds many, minute, spindle-shaped .............. .- 3. Norrisia
6. Corolla outside glabrous, inside with a hair-ring in the mouth.Fruit drupaceous, indehiscent,
glabrous. Seeds 1-2, large, spindle-shaped, remaining in a stone . . . .. 7. Neuburgia

5. Corolla-lobes imbricate or contorted in bud.
7. Corolla-lobes contorted. Trees, shrubs, or epiphytes. Fruit baccate, almost never dehiscent.
Seeds many, angular, embedded in pulp eA ee Oiaeter Ho) o Loko neMtO 1. Fagraea
7. Corolla-lobes imbricate. Liana or straggling shrub. Fruit a dry capsule, 2-valved. Seeds c. 8,
elliptic to bean-shaped, winged all around .............. 4 Gelsemium
4. Inflorescences exclusively lateral.
8. Fruits baccate, indehiscent.
9. Corolla-lobes contorted in bud. Seeds small, angular ......... . 1. Fagraea
9. Corolla-lobes valvate in bud. Seeds fairly large (14-1 cm), orbicular or elliptic. 6. Gardneria
8. Fruits capsular, with 2 caducous valves. Seeds smal!, embedded in a red or orange placenta.
8. Geniostoma
1. Annual, rarely perennial herbs.
10. Flowers 4-merous. Leaves l-nerved or curvinerved. ........... - Il. Mitrasacme
10. Flowers 5-merous. Leaves penninerved.
11. Stem with some pairs of small leaves and at the base of the inflorescence a pseudowhorl of 4
lancernleavessminiiorescences mn branched! sa. sem einci semen eileen inner . 10. Spigelia
11. No whorled larger leaves at the base of the inflorescence. Inflorescences dichasially branched.
9. Cynoctonum

298 _______FLora MALESIANA [ser. I, vol. 6?

Fig. 1. A strangling Fagraea sp. at Pondok Patjet, Mt Singalang, Central Sumatra (W. MEYER,
1956).

Dec. 1962] LOGANIACEAE (Leenhouts) : 299

1. FAGRAEA

TuHuns. Vet. Acad. Handl. Stockh. 3 (1782) 132, t. 4; LeeNH. Bull. Jard. Bot.
Brux. 32 (1962) 418-431.—Bertuchia DENNST. Schluess. Hort. Malab. (1818) 30,
nom. inval—Cyrtophyllum REINw. [ex BL. Cat. (1823) 47; Isis 1 (1823) 313-314,
nom. nud.] Syll. Pl. Ratisb. 2 (1826) 8.—Kuhlia ReINw. [ex BL. Cat. (1823) 51,
nom. nud.] Syll. Pl. Ratisb. 2 (1826) 6, nom. illeg., non H.B.K. (1825).—Picrophloeus
BL. Bijdr. (1826) 1019.—Utania G. Don, Gard. Dict. 4 (1838) 663.—Kentia
SteuD. Nomencl. ed. 2, 1 (1840) 845, nom. illeg —Flemingia HUNTER in Ridl. J.
Str. Br. R. As. Soc. n. 53 (1909) 83, nom. inval., non Roxs. ex Air. (1812).—
Fig. 1-23.

Terrestrial, epiphytic, or hemi-epiphytic, often scrambling shrubs, woody
climbers, or small to fairly large trees, glabrous in all parts. Stipules connate into
an ocrea which usually early splits (interpetiolarly) into 2 axillary scales, these
free or partly to entirely adnate to the base of the petiole; axillary colleters all
around the node. Leaves petioled or sometimes sessile, coriaceous or more or less
fleshy, nearly always entire (crenulate in F. crenulata), base usually decurrent and
sometimes auriculate; penninerved, nerves often, veins nearly always inconspic-
uous to invisible. Flowers solitary, in twos, or in 3- to many-flowered cymose
(rarely glomerulous, thyrsoid, or by reduction racemose or spicate) inflorescences,
nearly always terminal, usually with a pair of strong basal branches in the upper
leaf-axils; peduncle nearly always quadrangular in section, slightly compressed,
sometimes terete, branches usually distinctly compressed. Bracts (except lower
ones) small and scale-like. Bracteoles usually present, mostly similar to the bracts,
but smaller, in some species large and enveloping the calyx, in that case often
2(-3) pairs present (fig. 21). Calyx thick-fleshy to coriaceous, in some species even
almost woody, lobes imbricate, nearly always rounded, thick with a thin margin,
inside with colleters at the base. Corolla fleshy, sometimes very thick, creamy-
white (outside sometimes greenish or pinkish), the second day turning to yellow or
orange, tube consisting of a tubular, thin(ner)-walled basal part which may be
very short and included in the calyx but can form the greater part of the tube, and
a thicker-walled upper part which is either tubular and only slightly widened to-
wards the mouth, or narrowly to widely funnel-shaped; lobes contorted, over-
lapping to the right, variable in length but always shorter than the tube, rounded.
Stamens inserted in the throat (between the basal and the upper part of the tube),
in some species on a thickened ring (fig. 23b); filaments broadened at and geni-
culate just above the base, strap-shaped, sometimes filiform; anthers basifixed,
either deeply bifid at the base and blunt-ellipsoid, or shallowly bifid and acute-
linear; cells dehiscing lengthwise, introrse. Ovary ellipsoid, tapering into a filiform
to cylindric (in sicco deeply lengthwise grooved) style about as long as the tube
or distinctly exserted; stigma capitate, obconical, peltate, or 2-lobed; ovary either
1-celled with 2 parietal placentas, or 2-celled with axile placentas; placentas peltate,
elliptic, with c ovules. Fruit: berry, but in some large-fruited species 4-valved;
usually globular to ellipsoid, crowned by the style-base, pale greyish-green or
whitish or turning via yellow and orange to bright red; under the epidermis with
a sticky white latex; calyx not or slightly enlarged (caducous in some forms of
F. blumei). Seeds ~, irregularly angular, c. | mm long, minutely warty, brown.

300 FLORA MALESIANA [ser. I, vol. 6?

Distr. About 35 spp., from Ceylon and the Malabar coast through SE. continental Asia to South
China, Hainan, and the southern peninsula of Formosa, throughout Malaysia, in the Northern Territory
and NE. Queensland, and in the Pacific from the Marianas to the Marquesas and the Tubuai Is. and New
Caledonia in the southwest; distinctly centered in Malaysia. LEENH. Pac. Pl. Areas 91. Fig. 3.

Ecol. Mostly heliophilous plants, along forest edges, on river-banks, in open places, also shrubs or
trees in light forest and savannahs; from sea-level up to c. 3000 m. As epiphytic shrubs they are usually
found on the trunk of large trees, clasping the stem with their roots; the latter may reach the soil whereby
the plant becomes a hemi-epiphyte (fig. 13). Some collectors noted that the same species could be found
as an epiphyte at lower altitudes, terrestrial at a higher altitude. F. crenulata is adapted to permanent or
periodical swamp conditions. As to climate, most species grow under everwet conditions, a few only are
tolerant to seasonal conditions (for example F. fragrans).

Flowers are mainly visited by insects (especially Xy/ocopa spp. and butterflies), some large-flowered
species also by birds, possibly some species by bats (see Derx, Ann. Bog. 1, 1950, 50). The anthesis of an
individual flower apparently lasts for two days, opening is after sunset; in F. blumei, F. auriculata, and
possibly in all large-flowered species the free halves of the lobes are released within 5—10 minutes, standing
out as the blades of a turbine after which the limb expands suddenly. The flowers are protandrous
(see BuRCK, Teysmannia 3, 1892, 201-203).

Dispersal is mainly effected by birds, not rarely by bats, apparently sometimes also by ants. At least
some or several species are myrmecophilous: they possess extra-floral nectaries near the leaf-base, on the
leaf-blades, and on the calyces; in a few species with large auricles at the petiole ants live in the shelter
provided by these. For the possible protection, given by these ants, see Burck, Ann. Jard. Bot. Btzg 10
(1891) 95-98, and Mrs NiEUWENHUIS-VON UXxkULL-GULDENBANDT, ibid. 21 (1907) especially p. 252
and t. 27 f. 60 & 67.

Anat. See VON GUTTENBERG, Ann. Jard. Bot. Btzg 44 (1934) 35-41, f. 3440; for the extrafloral nec-
taries see also ZIMMERMANN, Ann. Jard. Bot. Btzg 18 (1901) 1, f. 1-7.

Uses. A few species are good timber trees, notably F. crenulata, F. elliptica, and F. fragrans. The latex
under the epidermis of the fruits is widely in use as a glue. Some of the species with large flowers are local-
ly cultivated as ornamentals.

Morph. The rebranching is mainly sympodial (also in the two species with axillary inflorescences).
The lower two or three internodes of the axillary shoots are in several species much longer than the further
ones of the same shoot; this is especially conspicuous in the species of sect. Cyrtophyllum, in which a
shoot of some strongly lengthened internodes abruptly ends in a number of densely crowded leaf-pairs
(fig. 6).

The stipules are as well intra- as interpetiolary connate and form an annulus around the twig. This
annulus is nearly always highest in the leaf-axils and often shows a suture between the leaf-bases. With
very few exceptions it soon splits into two ‘median stipules’, which I have called axillary scales (fig. 2a).

The auricles which are present in some species (fig. 2b—d) are no stipules but appendages of the base of
the blade; they are always inserted outside the stipules. In petioled leaves the blade is often decurrent
as a narrow wing along the petiole broadening at the base into the auricle, but in others the terete petiole
bears no trace of such wings. See HASSELBERG, Symb. Bot. Upsal. 2, 7. 3 (1937) 49-SO.

The ovary can be 2-celled with axile placentation, or 1-celled with 2 parietal placentas. The latter may be
the more primitive condition. The systematic value of this character cannot be thoroughly studied in the
herbarium. I tried to get a rough impression by making sections of one or two ovaries of several species,
of one ovary of several specimens of F. gracilipes (belonging to ‘elata’, ‘muelleri’, and ‘cambagei’), and of
several ovaries of one specimen of F. gracilipes. | found that the ovary in one specimen is constant,
‘elata’ and ‘cambagei’ both showed only 1-celled ovaries, but in ‘muelleri’ both types occurred. In some
other species there was also a difference between my observations and descriptions or figures published.
The tentative conclusion is that the inner structure of the ovary is variable and anyhow cannot serve for
sectional or specific discrimination. During this investigation I found several ovaries which were solid,
which may point to a tendency of dioecism, or dioecio-polygamism. This matter should be examined
in more detail with plenty of fresh material.

Taxon. The genus has been subdivided here into three sections, sect. Cyrtophyllum with 3 spp.,
sect. Racemosae with F. racemosa as only species, and sect. Fagraea with 27 spp.

A further subdivision of sect. Fagraea into sharply separated infrageneric taxa is not well possible.
The flowers provide still the best characters, especially the form of the stigma, 2-lobed or not, seems to be
important. There is a group of species characterized by a 2-lobed stigma associated with linear anthers
and inside the corolla with a ring on which the stamens are inserted (fig. 23). Unfortunately F. annulata
has the ring but not the other two characters and F. gardenioides has linear anthers but no ring and no
2-lobed stigma.

The remaining species have a globular, obconical, or truncate stigma, which is, however, sometimes
grooved with a tendency to be slightly 2-lobed. There are two groups of species, one of which is charac-
terized by auricles at the leaf-base, the other lacking this character. Though this character is constant for
the species it cannot serve for further infrageneric subdivision as it occurs in various degree of develop-
ment and seems to be of minor importance systematically.

As to other specific characters the shape of the stipules is fairly constant.

Dec. 1962] LOGANIACEAE (Leenhouts) 301

In general it has been found necessary to adopt a rather wide specific delimitation because it was often
difficult to find reliable characters; especially the size of the corolla shows great but grading variation.
F. auriculata, F. berteriana, F. blumei, F. ceilanica, and F. gracilipes are such species with wide circum-
scription.

Nomenclature. The name Bertuchia DENNST. is considered to be invalid as it is not fully in accordance
with Art. 42 of the Montreal Code.

The name Kuhlia REINW. is illegitimate, being a later homonym of Kuhlia H. B. K.; Utania and Kentia
were both published as substitutes, the latter superfluously.

Flemingia HUNTER is not validly published as RIDLEy’s publication of the old MS of HUNTER’s was
only for historical reasons, and not to validate any name.

Broce

Fig. 2. Different kinds of stipules and auricles in Fagraea. a. Axillary scales and very faint auricles in

F. eymae BAcK. (BW. 8908), b. not-reflexed auricles in F. fastigiata BL. (KOORDERS 4329), c. small re-

flexed auricles in F. tacapala LEENH. (EYMA 2473), d. well developed auricles in F. auriculata JACK
(SINCLAIR s.n.). All * 34.

KEY TO THE SPECIES

1. Leaves minutely crenulate. Stem and branches prickly. .......... . 12. F. crenulata
1. Leaves entire. Stem and branches unarmed.

2. Inflorescences exclusively axillary.

3. Inflorescences thyrsoid, oe conus: mostly many-flowered; peduncle slender. Stamens and

style far exserted .. . . 2, F. fragrans
3. Inflorescences condensed racemose, umbelliform, few= flowered; peduncle short and thick. Stamens
and style hardly exserted. ... . . 3. F. umbelliflora

2. Inflorescences terminal (the lowest pair of strong branches often i in the upper leaf-axils).
4. Flowers and fruits small (calyx 2-3 mm, corolla-tube 314-8 mm, fruits 5-8 mma). Inflorescences
many-flowered, corymbose 48 ee edeeenelliptica:
4. Flowers and fruits distinctly larger. Inflorescences rarely corymbose.
5. Leaves with distinct auricles (reflexed or not) at or slightly above the base (fig. 2b—d).
6. Auricles reflexed. Young leaves not connate. Inflorescence without an involucrum of reduced

leaves.
7. Auricles on the petiole distinctly above the axillary scale (fig. 2b). Calyx 7 mm long, sepals
connate for more than 24 of their length. . . . . 11. F. fastigiata

7. Auricles at the very leaf-base, clasping the axillary scale (fig. 2c- ‘d). Calyx (8—)10 mm or more,
sepals for their greater part (mostly nearly completely) free.
8. Flowers in many-flowered inflorescences. Bracteoles one pair, attached to the lower half of the
pedicel. Corolla-tube 2—3 cm.

9. Leaves blunt to acute at the apex; nerves 8-10 pairs. Inflorescences about corymbose
(branches erecto-patent and not of the same length). Bracteoles (14—)1—11!4 cm. Calyx 114-
2% cm long, sepals nearly free. Corolla-tube widely funnel-shaped. . . . 14. F. tacapala

9. Leaves rounded (apart from the acumen) at the apex; nerves 7-8 pairs. Inflorescences py-
ramidal (branches transverse to the rachis and of nearly the same length). Bracteoles 214-3
mm. Calyx 8-10 mm, sepals connate for 14—!4. Corolla-tube slenderly funnel-shaped.

15. F. woodiana

8. Flowers solitary or inflorescences up to 5(—7)-flowered. Bracteoles 1—3 pairs, attached to the

upper half of the pedicel. Corolla-tube 4144-15 cm.

10. Calyx enveloped by 2(—3) pairs of decussate bracteoles, the upper of which being at l2ast

half as long as the calyx (fig. 21). Flowers solitary.
11. Nerves hardly visible. Sepals rounded (fig. 18b) . . . .... . . 419. F. involucrata

302, FLORA MALESIANA [ser. I, vol. 6?

11. Nerves distinctly prominent on the lower side. Sepals long and acute (fig. 21e).
20. F. macroscypha
10. One or two pairs of small appressed bracteoles at the base of the calyx, much smaller than

the latter. Inflorescences 1—3(—7)-flowered . . - . . 18. F. auriculata
6. Auricles not reflexed. Leaves connate when young. Inflorescence with an involucrum of reduced
leaves.

12. Corolla-tube tubular, slender, c. 11 cm. Sepals nearly halfway up connate. 16. F. longiflora
12. Corolla-tube funnel-shaped, 3-314 cm. Sepals connate for about 14 of their length.
17. F. carstensensis
5. Leaves without distinct auricles (fig. 2a).

13. Stipules, even in full-grown leaves, connate around the twig into an ocrea which is hardly longer
in the leaf-axils than between the petioles (fig. 10b). Inflorescences distinctly and mostly fairly
long-peduncled, racemiform to spiciform, usually with many spaced, decussate, rather short,
cymose glomerules (sometimes corymbiform or the whole inflorescence reduced to one fairly
long-stalked glomerule) (fig. 9) . . . . 4. F, racemosa

13. Stipules soon interpetiolary (nearly) fully split, distinctly lengthened ‘ in the axils (fig. 23f).
Inflorescences not distinctly peduncled; flowers either solitary, or in few-flowered sessile glomeru-
les, or with a pair of strong basal branches in the upper leaf-axils.

14. Full-grown stigma distinctly 2-lobed (fig. 23d). Stamens inserted on a fleshy ring in the corolla-
tube; anthers linear, only slightly bifid at the base (fig. 23b).

15. Nervation seemingly dense, as between every two nerves an intermediate vein is nearly as
strongly developed as these, together c. 20 pairs (nervation not very conspicuous). Leaves
relatively large and especially broad (9—23 by 4!4—-14 cm). Inflorescences usually widely and
laxly branched . . . As 31. F. berteriana

15. Nervation (nearly) invisible, : nerves ‘spaced, 4- ile) pairs. “Leaves usually smaller and often
narrower (4-15 by 2!4-7!4 cm). Inflorescences rather dense.

16. Leaf-base subcordate to rounded, rarely broadly cuneate, distinctly set off from the petiole.
Bracteoles, if present, apical, appressed to the calyx . . : . . . 28. F. salticola
16. Leaf-base acute, decurrent. Bracteoles attached about halfway the pedicel.

17. Corolla-tube tubular or narrowly funnel-shaped, 1 —4 cm. Axillary scale usually distinctly
broader than the base of the petiole, 3-10 mm long. Leaf apex often shortly and broadly
acuminate. Fruits broadly ellipsoid to subglobose, contracted into a strong conical beak.

29. F. bodenii

17. Corolla-tube long-tubular, 7-11 cm. Axillary scale inconspicuous, 3-5 mm long. Leaf-apex

blunt to rounded, not acuminate. Fruits oblong-ellipsoid, acute . . . 30. F. gitingensis
14. Full-grown stigma capitate, obconical, or peltate (in the latter case sometimes slightly 2-lobed).

Stamens not inserted on a ring (F. annulata excepted); anthers oblong to elliptic, about halfway

bifid (F. gardenioides excepted).

18. Corolla-tube 6-14 cm, cylindric or almost so.

19. Calyx 14-1 cm long; anthers 4 mm. Flowers in sessile glomerules, rarely solitary.
8. F. tubulosa
19. Calyx 114-5 cm; anthers 7 mm long or more. Flowers solitary or in up to about 5-flowered
cymes.

20. Calyx enveloped by an involucrum consisting of two pairs of large bracteoles.

21. F. resinosa

20. Calyx not enveloped by such an involucrum.

21. Anthers linear, bifid at the very base only. Stipules conspicuous. . 23. F. gardenioides
21. Anthers oblong to elliptic, bifid till about the middle. Stipules not very conspicuous.
22. Flowers in c. 5-flowered cymes. Bracteoles small, attached about halfway the pedicel, pa-
tent. Calyx 114-134 cm high. Anthers 7 mm long. Fruits subglobular, 5 cm o. Axillary
scale truncate to slightly emarginate. The Ceylonese form ... . 5. F. ceilanica
22. Flowers solitary or rarely in twos. Bracteoles 14-1 cm long, attached just below and
appressed to the calyx. Calyx (114—)2-5 cm high. Anthers 1-134 cm long. Fruits ellipsoid,
4-7 cm long. Axillary scale rounded... ..:::.... =. . 22H ecamosa
18. Corolla-tube less than 6 cm, funnel-shaped.
23. Flowers sessile, in 3—7-flowered glomerules.

24. Calyx 34-114 cm high. Axillary scale inconspicuous, rounded, appressed to the twig. Leaves
7-24 by’ 334-10 cm; midrib slender, rounded beneath, nerves invisible. 7. F. acuminatissima

24. Calyx 214-3 cm high. Axillary scale conspicuous, 2-lobed, adnate to the petiole. Leaves
20-38 by 8-18 cm; midrib bold, keeled beneath, nerves conspicuous. 13. F. truncata

23. Flowers either solitary or distinctly pedicelled in few-flowered, or in widely branched and

many-flowered inflorescences.

25. Inflorescences warty-lenticellate ... see 2 DONE blomei

25. Inflorescences not warty-lenticellate (at least during anthesis!).

26. Calyx partly enveloped by an involucrum consisting of 1 or 2 pairs of large bracteoles.

Dec. 1962] LOGANIACEAE (Leenhouts) 303

27. Involucrum composed of one pair of confluent bracteoles (fig. 14c). 5. F. ceilanica

27. Involucrum composed of two pairs of free bracteoles . ... . . 25. F. calcarea
26. Calyx without involucrum.
28. Nerves distinctly prominent beneath . 9. F. ridleyi

28. Nerves either impressed, or inconspicuous to invisible.

29. Stamens inserted on a distinct thickened ring about halfway the corolla-tube.

6. F. annulata

29. Stamens not inserted on a distinct thickened ring.

30. Leaves subsessile, rounded to subcordate at base, faintly auriculate (fig. 2a); nerves
5-6 pairs. Inflorescences usually 2-flowered . ; 27. F. eymae
30. Leaves nearly always petioled, acute to decurrent at base; ‘mostly more nerves (one
form of F. ceilanica has a rounded to subcordate leaf-base, but is not auriculate and has
4-15-flowered inflorescences).
31. Axillary scale conspicuous, rounded, truncate, or emarginate at the apex. Inflorescen-
ces up to 5-flowered. Pedicels robust. Anthers 7/4 mm long or more. Stigma peltate.
32. Leaves often slightly auriculate at the base; midrib keeled beneath; nerves incon-
spicuous, up to 20 pairs. Sepals connate for half their length or less. Fruits ellips-
oid, 6-7 cm long, not distinctly beaked . 18. F. auriculata ssp. parviflora
. Leaves never auriculate; midrib rounded beneath; nerves about invisible, up to 10
pairs. Sepals connate for half their length or more. Fruits subglobular, up to 5 cm
long, with a long and slender beak . 5 24. F. curtisii
31. Axillary scale inconspicuous, rounded, rarely truncate at the apex. Inflorescences
mostly more than 5- flowered. Pedicels nearly always slender. Anthers up to 744 mm
long. Stigma capitate, obconical, or more exceptionally peltate.

33. Usually a shrub or climber. Midrib flat or prominulous above, nerves usually invisi-
ble. Calyx 1-234 (-4)cm long (in the form ‘khasian@ 6-7 mm). Anthers usually more
than 5 mm long. Stigma obconical to peltate . . . 5. F. ceilanica

33. Usually a tree. Midrib mostly grooved above, nerves delicately grooved on both
sides. Calyx 14-34 (—114) cm long. Anthers up to 5 mm long. Stigma capitate to sub-
obconical 5 on ey Re peectetc eicene ater Ma 26. F. gracilipes

wo
iw)

1. Section Cyrtophyllum

(REINW.) BL. Rumphia 2 (1838) 34.—Cyrtophyllum REINW. 1826.—Picrophloeus
BL. 1826.—Fagraea subg. Cyrtophyllum Mia. FI. Ind. Bat. 2 (1857) 375.—Fagraea
sect. Parviflorae Bru. J. Linn. Soc. Bot. | (1856) 100.—Flemingia HUNTER in
Ridl. 1909, nom. inval.

Leaves not auriculate. Inflorescences corymbose, with many small flowers;
corolla-tube always tubular; stamens and style far exserted (except in F. umbelli-
flora); fruits globular, small.

1. Fagraea elliptica Roxs. [Hort.
84, nom. nud.| Fl. Ind. ed. Wall. Q
ed. Carey 1 (1832) 462; DC. Prod. 9 (1845) 30; 2
Mia. FI. Ind. Bat. 2 (1857) 376; K. & V. Bijdr.9 | ,
(1903) 84; Atlas 2 (1914) f. 330; Koorp. Fl. o z
Tjib. 3 (1918) 48; CAmMMmerRL. Bull. Jard. Bot. ‘4

Btzg III, 5 (1923) 316; HeyNne, Nutt. Pl. (1927)
1270; BAKH. f. in Back. Bekn. FI. Java (em. ed.) 7
(1948) fam. 170, p. 13.—Picrophloeus javanensis
Bi. Bijdr. (1826) 1020; Don, Gard. Dict. 4
(1837) 66; DC. Prod. 9 (1845) 32.—Cyrtophyllum it
speciosum BL. Bijdr. (1826) 1022; DC. Prod. 9 a1
(1845) 31; Riv. Fl. Mal. Pen. 5 (1925) 322, L— a Se

Beng. (1814) = ae a ae a

2 (1824) 32;

fo]
ol=
ol-

Obs

ol=\;

incl. var. montanum.—Willughbeia elliptica SPRENG.
Syst. Veg. 4 (1827) Cur. Post. 71.—F. speciosa BL.
Rumphia 2 (1838) 35, t. 81; Mus. Bot. 1 (1850)
172; non Riw.. J. Str. Br. R. As. Soc. n. 50 (1908)
122 (= F. fragrans); MerR. Philip. J. Sc. 11 (1917)
Bot. 306.—F. picrophloea BL. Rumphia 2 (1838)
36, nom. illeg.; Mus. Bot. 1 (1850) 173; Mua.
Fl. Ind. Bat. 2 (1857) 377.—F. kimangu BL. Mus.

Fig. 3. Distribution of Fagraea in and around

Malaysia. The number above the hyphen refers

to the total number of species, that below the

hyphen to the number of endemic species in each
island or district.

Bot. 1 (1850) 173; Walp. Ann. 3 (1852) 76,
sphalm. kimanga; Mia. Fl. Ind. Bat. 2 (1857) 377.

304

FLORA MALESIANA

[ser. I, vol. 6?

—F. valida Mia. FI. Ind. Bat. 2 (1857) 376.—F.
sumatrana Mia. Fl. Ind. Bat. 2 (1857) 377;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923) 318.
—F. aurantiodora S. Moore, J. Bot. 66 (1928)
105.—F. pseudoelliptica KAN. & Hatus. Bot.
Mag. Tokyo 56 (1942) 161, f. 5.—F. javanensis
BAKH. f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 170, p. 12; Blumea 6 (1950) 382.—F. pu-
silliflora BAKH. f. [in Back. Bekn. Fl. Java (em. ed.)
7 (1948) fam. 170, p. 13, Dutch descr.] Blumea 6
(1950) 383.—Fig. 4.

Fig. 4. Fagraea elliptica Roxs., Sandakan (North
Borneo) (MEYER, 1960).

Tree up to 45 m by 0.15-114 m @, sometimes
with buttresses up to 1 m high, or shrub, rarely a
vine. Leaves usually lanceolate or oblong to
obovate, sometimes broadly obovate, 71/—24(—32)
by 134-15 cm, thinly to thickly coriaceous, base
acute to slightly attenuate, apex mostly short- to
long-acuminate, less often obtuse, rarely broadly
rounded or subretuse; nerves 6—20 pairs, faintly
to distinctly conspicuous, often slightly impressed
above, beneath usually distinctly prominulous or
(in thick leaves) hardly visible; petiole varying
from rather thin to robust, 1-4 cm long. Stipules
connate into a 114—7!4 mm long ocrea which
may split into rounded axillary scales, partly
adnate to the petiole. Inflorescences terminal and
often also in the topmost leaf-axils, up to 30 cm
wide, often much smaller, very many-flowered;
peduncle 1-13 cm; pedicels during anthesis 1-4
mm (later up to 6 mm), often provided with 2
minute bracteoles inserted about halfway or some-
what lower down. Ca/yx campanulate, 2—3 mm

long, divided + halfway. Corolla: tube narrow,
cylindric, 314-6(-8) mm. Stamens erect or later
reflexed; anthers oblong, 114 mm long, cells free
in the basal half. Style (accrescent during anthesis
?) varying from 1144 mm and included to 134 cm
and far exserted; stigma small, capitate, very
obscurely 2-lobed. Berry globose, 14-34 cm g,
tipped by a minute circular style-rest, orange-
coloured to brick-red.

Distr. Malaysia, except Central and East Java,
the Lesser Sunda Is., and the Philippines.

Ecol. In very different sites, both on dry and on
marshy or temporarily inundated soils, mostly on
sand, often along rivers, in humid forests, open
forests, shrubberies, heath forest, grass wastes,
rocky strand forest, also as a shrub on open dry
hill-tops, from sea-level up to 1800 m. Fi. (mainly
April-June), fr. Jan.—Dec.

Uses. The brownish to pale yellow, medium to
very hard, very durable wood is used for house-
building, for bridges, and for padi pounders. Also
used medicinally against stomach-ache.

Vern. Kaju badjam, k. galumbang batu, k.
kadjang, k. kisa, k. labih, k. rama-rama, k. sabo,
k. si margalugur, k. si margapuk, randa tiung,
ruku_ biruwang, sélura, sépukan, sibatukon, simar
tarasa, témbésu kétam, t. rawah, t. rawang, t.
talang, Sum., rubi, ténggél dotan, tutun ténggeél
délok, Simalur, témbésu, Nias & Billiton, kaju
bujuk, k. sabo (or sobo), teémbésu samsu, Banka,
(jénis) témbasu, Mal. Pen., ki mangle, ki mangu,
ki minjak, ki tandu, ki térong badak, rangkot
tandus (or tando), Java S., bintulu, brut-brut,
témasu (kbini), tembaso bansang, témbusu, Born.,
tariwan, Cel.; Moluccas: bientaus, Sula, kau,
wadil, Buru, atieso, Ceram, tonki tonki, Ambon;
niperen, New Guinea.

Note. F. picrophloea BL. is illegitimate, being
based upon Picrophloeus javanensis; the epithet
javanensis should, and could, have been used.

2. Fagraea fragrans Roxs. [Hort. Beng. (1814)
84, nom. nud.) Fl. Ind. ed. Wall. 2 (1824) 32; ed.
Carey 1 (1832) 461; Don, Gard. Dict. 4 (1837)
68; BL. Mus. Bot. 1 (1850) 172; Bru. J. Linn.
Soc. Bot. 1 (1857) 100; Mra. Fl. Ind. Bat. 2 (1857)
375; ScuerF. Nat. Tijd. N.I. 31 (1870) 22; Kurz,
Fl. Burm. 2 (1877) 205; CLARKE in Hook. f. FI.
Br. Ind. 4 (1883) 85; VIDAL, Sinops. (1883) t. 69
f. A: Watt, Dict. 3 (1890) 312; AHERN, Compil.
Timber Tree P. I. (1901) 88, pl.; K. & V. Bijdr. 9
(1903) 86; Kina, J. As. Soc. Beng. 74, 1i (1908)
611; Rip. J. Str. Br. R. As. Soc. n. 50 (1908) 121;
Ei. Leafl. Philip. Bot. 2 (1909) 597; Dorp, FI.
Gén. I.-C. 4 (1914) 177; K. & V. Atlas (1914)
f. 332; ENDERT, Tectona 13 (1920) 132; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 318; Foxw.
Mal. For. Rec. n. 3 (1927) 157; HEYNE, Nutt. PI.
(1927) 1270; Ho_trum, Gard. Bull. S. S. 9 (1935)
73; EssENBURG, Tectona 28 (1935) 606, fig.;
DE GRAAF, Tectona 28 (1935) 611; VAN DER
Voort, Trop. Natuur 28 (1939) 207, f. 9-10;
CoRNER, Ways. Trees (1940) 424, t. 122-123;
Kerr in Craib, Fl. Siam. En. 3 (1951) 55; BROWNE,
For. Trees Sar. & Brun. (1955) 244, t. 32. —

Dec. 1962]

LOGANIACEAE (Leenhouts) 305

<n RE eR Aap eR eR

Ane

Se

Fig. 5. Fagraea fragrans Roxs. at Tanglin Barracks, Singapore (CORNER).

306 FLORA MALESIANA [ser. I, vol. 6?

b. opened corolla with the long-

Fig. 6. Fagraea fragrans ROXB. a. Habit, with flowers and fruits, x 7/4,
x 2 (a after

exserted stamens, X 2, c. pistil, 2, d. small part of twig, showing the axillary scales,
Rumphia 2, t. 80, b-d BvuMe in herb. L 908.127—224).

Dec. 1962]

LOGANIACEAE (Leenhouts)

Fig. 7. Burned-over savannah with Fagraea fragrans Roxs., Padang Bolak, Sumatra East Coast Res.
Corky bark charred by fire (VAN DER VoorT, 1939).

Cyrtophyllum peregrinum Reinw. [ex BL. Cat.
(1823) 47, nom. nud.; Isis 1 (1823) 313 & 314,
nom. nud.) Syl. Pl. Ratisb. 2 (1826) 9; BL. Bijdr.
(1826) 1022; DC. Prod. 9 (1845) 31; Ript. FI.
Mal. Pen. 2 (1923) 421; Ho_trum, Gard. Bull. S.S.
5 (1931) 189.— Willughbeia fragrans SPRENG. Syst.
Veg. 4 (1827) Cur. Post. 71.—F. peregrina BL.
Rumphia 2 (1838) 34, t. 80; Mus. Bot. 1 (1850)
172; ScHerF. Nat. Tijd. N. I. 31 (1870) 22.—
Cyrtophyllum fragrans DC. Prod. 9 (1845) 31;
Ript. Trans. Linn. Soc. Bot. 3 (1893) 323.—
Cyrtophyllum lanceolatum DC. Prod. 9 (1845) 31;
Ript. Fl. Mal. Pen. 2 (1923) 421.—F. lanceolata
WALL. [Cat. (1829) mn. 1599, nom. nud.) SCHNIZL.
Iconogr. 2 (1851) t. 131, fig. 1, nom. illeg., non
BL. 1826; Mia. Fl. Ind. Bat. 2 (1857) 376; Burk.
Dict. (1935) 995.—F. wallichiana Bru. J. Linn.
Soc. Bot. 1 (1856) 98; CLARKE in Hook. f. FI. Br.
Ind. 4 (1883) 85; Kina, J. As. Soc. Beng. 74, ii
(1908) 607; CorNeR, Ways. Trees (1940) 426.—
F. speciosa (non Bu.) Riv. J. Str. Br. R. As. Soc.
n. 50 (1908) 122.—Flemingia fragrans HUNTER
in Ridl. J. Str. Br. R. As. Soc. n. 53 (1909) 83,
nom. inval.—F. caudata Riv. J. Str. Br. R. As.
Soc. n. 79 (1918) 97.—F. gigantea RIDL. l.c. 98;
Foxw. Mal. For. Rec. n. 3 (1927) 157, pl.; Burk.
Dict. (1935) 995; CorNER, Ways. Trees (1940)
425; Browne, For. Trees Sar. & Brun. (1955)
245.—Cyrtophyllum caudatum Rivv. J. Str. Br.
R. As. Soc. n. 79 (1918) 98, nom. altern., illeg.—
Cyrtophyllum wallichii Rip. L.c., nom. altern.,

illeg —F. cochinchinensis A. CuHeEv. Cat. Pl. J.
Bot. Saigon (1919) 65, pro specim., excl. basionym;
Brown, Min. Prod. Philip. For. 3 (1921) 220;
Merr. En. Philip. 3 (1923) 314; Burk. Dict.
(1935) 994; Brown, Useful Pl. Philip. 3 (1950)
223.—Fagraea sp. ENDERT, Tectona 13 (1920)
142.—F. sororia J. J. SmitH in Cammerl. Bull.
Jard. Bot. Btzg III, 5 (1923) 319, f. 5; HEYNE,
Nutt. Pl. (1927) 1270; Streup, Tectona 24 (1931)
1132—1133; Trop. Natuur 22 (1933) 110.—
Cyrtophyllum giganteum Riv. [J. Str. Br. R. As.
Soc. n. 79 (1918) 98, nom. altern., inval | Fl. Mal.
Pen. 2 (1923) 421.—F. ridleyi GANDOGER, Bull.
Soc. Bot. Fr. 70 (1924) 921, nom. illeg., non
K. & G. 1908.—Fig. 5-8.

Tree (sometimes a shrub), 8—25(—55) m tall,
up to 1.35(-214) m @, sometimes with buttresses
up to 214 m. Leaves petioled, oblong-lanceolate to
lanceolate or obovate-oblong, 4-15 by 114-6 cm,
mostly thinly, rarely firmly coriaceous, sometimes
more or less bullate, the base acute or shortly
attenuate, the apex rarely blunt to acute, usually
short- to long-acuminate, sometimes to caudate,
obtuse; nerves 4-9 pairs, rarely more or fewer,
sunken above, slightly prominent or invisible
beneath; petiole thin, 1-2'4 cm; axillary scale
rounded, c. 1-2 mm long, partly free from the
petiole. Inflorescences in the higher leaf-axils,
(1—)3- to oo-flowered, often rather dense; peduncle
thin, 114-714 cm; pedicels thin, 14-2! cm, often
with 2 minute bracteoles in or below the middle

208

Fig. 8. Fagraea fragrans Roxs., cult. in Botanic Gardens, Singapore (HENDERSON, 1949)

(in few-flowered inflorescences sometimes with a
second pair near the base). Calyx campanulate,
214-8 mm long, divided over 14-34 of its length.
Corolla-tube narrowly funnel-shaped, 34-214 cm.
Anthers oblong-linear to oblong-elliptic, 134—3 mm
long, cells free in the basal half. Style 114-6 cm
long; stigma small, capitate or obconical, faintly
2-lobed. Berry broadly ellipsoid, tipped by the
short but distinct style-rest, 34-1 cm g, red or
orange.

Distr. Bengal (near Calcutta, wild?), Lower
Burma (from Rangoon southward), the An-
daman Is. (CLARKE, /.c.), Siam, South Indo-China
(N. to Tourane, Central Annam), and Malaysia:
Sumatra, Malay Peninsula, W. Java (naturalized),
Bawean, Borneo, SW. Philippines (Balabac,
Palawan, Busuanga, Culion, Mindoro), Celebes,
and Japen I. near NW. New Guinea.

According to MIQUEL, /.c., also in the Moluccas;
I do not know on what collection this assertion
was based. The statements ‘‘China”’ and “‘Japan”
in older literature go back to ROxBURGH and
BLuME, who gave this as the possible origin of
cultivated specimens.

Ecol. Humid, often seasonally but not con-
stantly swampy, light forest, secondary forests,
lalang fields, in the Malay Peninsula also along the
beach; it does very well in poorly aerated, com-
pact or swampy soils, but also on poor sands or
podsols; from sea-level up to c. 800 m. Fi. mainly
April—June, fr. mainly July-Noy. Flowering (and
consequently fruiting) is distinctly periodical:
HoLttum (Gard. Bull. S. S. 9, 1935, 73-78)

found in Singapore that ‘‘...general flowering
is in progress about four months after the onset of
the dry weather.” and ‘‘...the time from the
stimulus to flower-development to the ripening of
fruit is about seven months and a half” (p. 75).
Rip.tey (Trans. Linn. Soc. Bot. 3, 1893, 323)
cited as flower-visitors, mainly in the evening,
“innumerable butterflies, of the Hesperidae sec-
tion, and humming-bird hawk-moths (Macro-
glossa luteata and M. insipida).” The fruits are
mainly eaten by bats (Pteropus edulis and Cy-
nopterus spp. according to RIDLEY, Disp. 1930,
348), furthermore by birds (Otocompsa analis,
Rip. /.c. 481), and possibly also by fruit-eating
ants (Polyrachis sp., Riv. /.c. 526). A light-
demanding tree, often a pioneer on burned-over
areas (VAN DER Voort, /.c.) (fig. 7).

Uses. This beautiful tree is often planted,
especially along roads and as a shade tree,
mainly in Siam, Sumatra, Singapore, and locally
in Java. Furthermore it is commonly planted for
reforestation, as the timber is highly valued,
being hard and very durable — even in the soil or
in water — though not very beautiful. A decoction
of the bark is used as a febrifuge, e.g. for malaria,
a decoction of the leaves and twigs is drunk in
Kedah, Malaya, for passing blood in stools,
such as happens in dysentery.

Vern. Jronwood, E. The common Malayan
name is rémbésu (lémésu, sémésu, tamasu, tamésu,
témasu(k), témbési, teémbusu, témé(n)su, témusu)
in many combinations. Kayu musuh-musuh, k.
tammusuh padang-padang, k. témbusu, pohon su-

Dec. 1962]

LOGANIACEAE (Leenhouts)

309

meéjar, témbésu bukit, t. kapur, t. langkanang, t.
lilin, t. paya, t. pématang, t. rawang, t. rénah,
t. talang, Sum., kayu témbusu, pako (or poko(k))
témbusu, p. tummoésoh paya, rériang, riang-riang,
tamsao, témbésu bukit, t. hutan, t. kémpang (or
kampong), t. luar, t. padang, t. talang, t. témbaga,
t. tikus, Mal. Pen., ki badak, Java S., (s)ambinaton
(or binaton or tombiaton), loang, mahoei, ma-
rambungkam, pérépat, séranai, tambiaton, témbésu
butat, t. danau, t. hutan, t. laut, t. padang, t. pasir,
t. rébong, t. rusa, t. tanduk, tibuan, Born.; Philip.:
délo, dilo, tiling, Tagb., susulin, Tag., téka, uring,
trung, Kuy.; anrali, bitjoro, nosu, (pohon) kulahi
(or kolahi), Cel., mana hurudu, manderi, Japen.

Notes. On the whole this species is fairly
uniform. Only two of the synonyms were based
upon clearly distinguishable forms, viz F. wal-
lichiana (= F. lanceolata SCHNIZL. = Cyrto-
Phyllum lanceolatum DC.) and F. caudata, the
former comprising all the specimens known from
P. Penang near the Malay Peninsula, the latter
described from Borneo. Both are characterized by
(1-)3-flowered inflorescences with relatively large
flowers and by lanceolate leaves, tapering at both
ends, distinctly caudate in the latter form.

Furthermore, foresters, especially in Malaya,
distinguish between F. fragrans, being a fairly
small tree of the secondary forest, and F. gigantea,
a canopy tree of the high lowland forest; there are
also differences in the bark, which is dark-brown
and deeply irregularly fissured in the first mentioned
form, brown or reddish brown with closer, narrow-
er and more regularly longitudinal ridges and
fissures in the second one; in the leaves, being
flat and with c. 8 pairs of nerves in F. fragrans,
having a wavy, undulating margin and 5-6 pairs
of nerves in F. gigantea; the timber of F. fragrans
is also more heavy than that of F. gigantea. F.
fragrans is often mentioned to propagate freely,
F. gigantea would not propagate or only ex-
ceptionally. In my opinion these are merely eco-
types, or even only specimens of different age:
F. fragrans is a light-demander, thus a pioneer of
open country, where it can maintain itself fairly
well, being fire-resistant. The name F. gigantea
is probably applied to the few old specimens
left after the forest has restored itself, or — as
there seems to be mostly a small group of speci-
mens — which filled up a glade. And these old
specimens stand in an ecological position in which
propagation is impossible.

In old literature the inflorescences are often
said to be axillary and terminal; the latter state-
ment is apparently based upon inaccurate ob-
servation, and then copied again and again.

Sterile specimens can be distinguished from the
related F. elliptica especially by the stipules:
in F. elliptica they form together an annular
ocrea closely appressed to the twig, in F. fragrans

they are mutually nearly free and form small
cups in the leaf-axils (fig. 6d).

For the name Flemingia fragrans HUNTER see
under the notes to the genus.

The basionym of F. cochinchinensis A. CHEV. is
Aidia cochinchinensis Lour. (type in BM). This
was reduced by Merr. Comm. Lour. (1935) 365
to Randia cochinchinensis (LOUR.) MERR. (Ru-
biaceae).

3. Fagraea umbelliflora Gmc & BENED. Bot.
Jahrb. 54 (1916) 193, f. 12.

Treelet or shrub, 2-5 m tall. Leaves petioled,
broadly ovate to obovate or lanceolate, 8-18 by
3-9 cm, thin- to stiff-coriaceous, base broadly to
acutely cuneate, attenuate, apex broadly rounded,
rather abruptly terminated by a short and slender
acumen or acute and hardly acuminate; nerves
5—10 pairs, inconspicuous to invisible, sometimes
prominulous beneath; petiole slender, 214-5 cm;
axillary scale for the greater part adnate, 3-5 mm
long, cup-shaped, rounded at the apex. Inflores-
cences axillary, umbelliform, nearly sessile or
with an up to c. 1 cm long, thick peduncle, c.
4-10 fairly long-pedicelled flowers, pedicels
slender, without bracteoles, 1-114 (in fruit —2)
cm. Calyx campanulate, 4-6 mm long, divided
somewhat less than halfway down. Corolla-tube
nearly tubular, 2-214 cm, slightly widened towards
the mouth. Anthers oblong-sagittate, 2-3 mm long,
cells free in their basal half. Stigma small, narrowly
cuneate, truncate. Fruits subglobular, c. 1 cm 9;
pedicels hardly thickened, calyx appressed.

Distr. Malaysia: New Guinea (Japen I.,
Vogelkop Peninsula, Sepik region).

Ecol. In dense, very humid high-stemmed
primary forest, 60-1200 m. Fi. July, Nov., fr.
Nov.

Vern. Niejap.

Notes. The type is cited as LEDERMANN 9714;
the only original specimen I did see, which is
apparently an isotype (in L) bears the number
9614.

In principle the inflorescences seem to be um-
belliform condensed racemes, terminal on short
axillary shoots or knobs, but with the exception of
some minute bracts these shoots are quite leafless.
Therefore it comes very near to a really axillary
inflorescence, as we find in this genus only in
F. fragrans.

Though the affinity of F. wmbelliflora is not fully
clear I have tentatively included it in sect. Cyrto-
Phyllum; it differs from the other species amongst
other characters, by the stamens and style which
are only very slightly exserted. Within sect. Cyr-
tophyllum it seems to be closest to F. fragrans.
Outside the section it may have some affinity with
F. gracilipes.

2. Section Racemosae

Bru. J. Linn. Soc. Bot. 1 (1856) 99 & 73.—Kuhlia REINW. 1826, non H.B.K.
1825.—Utania G. Don, 1838.—Kentia StEuD. 1840, nom. illeg—Fagraea sect.

310 FLORA MALESIANA ___[ser. 1, vol. 6

mas
N
N
N
Ny
N
N
N
N
N
iS
N
iS
N
N
N
i.
N
i

1959

Fig. 9. Fagraea racemosa WALL. a. well-developed normal inflorescence, < 14, b. part of infructescence,

< 1, c. inflorescence of ‘F. pendula’, = 4, d. inflorescence of ‘F. spicata’, * 1 (a partly after Rumphia

2, t. 79, the leaves from WOMERSLEY NGF 3870, 6b WoMERSLEY NGF 3870, c CLEMENS 21546, d AGAMA BS
553).

Dec. 1962]

LOGANIACEAE (Leenhouts)

311

Pseudoracemosae SOLER. in E. & P. Nat. Pfl. Fam. 4, 2 (1892) 43.

Leaves not auriculate; stipules connate in an ocrea. Inflorescences peduncled,
racemiform, with a number of distant decussate pairs of small cymes (some-
times basal cymes long-stalked, sometimes all cymes sessile and close together,
sometimes the whole inflorescence reduced to a long-stalked, few-flowered glo-
merule). Stamens and style not or only slightly exserted.

4. Fagraea racemosa JACK ex WALL. in Roxb.
Fl. Ind. 2 (1824) 35; DC. Prod. 9 (1845) 29,
incl. var. grandis [WALL. Cat. (1829) n. 1601-2,
nom. nud.]; BENTH. J. Linn. Soc. Bot. 1 (1856)
99; Mia. Ann. Mus. Bot. Lugd.-Bat. 2 (1866)
218, incl. var. coarctata; BENTH. Fl. Austr. 4 (1869)
367; Kurz, Fl. Burm. 2 (1877) 205; CLARKE in
Hook. f. Fl. Br. Ind. 4 (1883) 84; K. Sco. &
Hote. Fl. Kais. Wilh. Land (1889) 109; F. M.
Bait. Queens]. Fl. 3 (1900) 1023; Kina, J. As.
Soc. Beng. 74, ii (1908) 608, incl. var. pauciflora;
Ript. J. Str. Br. R. As. Soc. n. 50 (1908) 118;
Dop, Fl. Gén. I.-C. 4 (1914) 175; Burk. J. Str.
Br. R. As. Soc. n. 73 (1916) 215, 226, 259; GILG &
BeNeD. Bot. Jahrb. 54 (1916) 184; Merr. Sp.
Blanc. (1918) 306; En. Philip. 3 (1923) 315;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
310; Rip. Fl. Mal. Pen. 2 (1923) 418; Burk. &
HeENpDeRS. Gard. Bull. S. S. 3 (1925) 399; LANE-
Poorer, For. Res. (1925) 134; Burk. Dict. (1935)
996; CORNER, Ways. Trees (1940) 425; Baku. f.
in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam.
170, p. 10; F. S. WALKeR, For. Br. Sol. Is. (1948)
137; Brown, Useful Pl. Philip. 3 (1950) 224,
f. 87; KERR in Craib, Fl. Siam. En. 3 (1951) 57;
Merr. J. Arn. Arb. 33 (1952) 225; BROWNE,
For. Trees Sar. & Brun. (1955) 246.—F. volubilis
WALL. in Roxb. FI. Ind. 2 (1824) 36; Don, Gard.
Dict. 4 (1837) 68; DC. Prod. 9 (1845) 30; Mia.
Fl. Ind. Bat. 2 (1857) 367; VIDAL, Sinopsis (1883)
t. 69 f. B.—Kuhlia morindaefolia REtNw. [ex BL.
Cat. (1823) 51, nom. nud.) Syll. Pl. Ratisb. 2
(1826) 7, nom. illeg.; BL. Bijdr. (1826) 777.—
F. malayana Mart. Nov. Gen. Sp. 2 (1826) 91;
Don, Gard. Dict. 4 (1837) 68; DC. Prod. 9 (1845)
30.—Willughbeia racemosa SPRENG. Syst. Veg. 4
(1827) Cur. Post. 71.—Willughbeia volubilis
SprENG. /.c.—F. morindaefolia BL. Rumphia
2 (1838) 32, t. 73 f. 2, t. 79, incl. var. robusta;
DC. Prod. 9 (1845) 29; BL. Mus. Bot. 1 (1850)
169; CLARKE in Hook. f. Fl. Br. Ind. 4 (1883)
84; K. & V. Bijdr. 9 (1903) 74; Em. Leafi.
Philip. Bot. 2 (1909) 596; K. & V. Atlas (1914)
f. 331, ‘morindifolia’.—F. cordifolia BL. Rumphia
2 (1838) 33; Mus. Bot. 1 (1850) 171; Mia. Fl. Ind.
Bat. 2 (1857) 368.—F. coarctata BL. Rumphia 2
(1838) 33; Mus. Bot. 1 (1850) 170; Mia. FI.
Ind. Bat. 2 (1857) 368, incl. var. ligustrina; VAL.
Bull. Dép. Agr. Ind. Néerl. 10 (1907) 45.—
F. ligustrina BL. Rumphia 2 (1838) 33; Mus. Bot.
1 (1850) 171, incl. var. disparifolia; BENTH. J. Linn.
Soc. Bot. 1 (1856) 100, incl. var. brachystachya;
CLARKE in Hook. f. Fl. Br. Ind. 4 (1883) 85;
Kina, J. As. Soc. Beng. 74, ii (1908) 609; RIDL.
Fl. Mal. Pen. 2 (1923) 420.—Utania morindaefolia
Don, Gard. Dict. 4 (1838) 663.—Kentia morin-

daefolia SteuD. Nomencl. ed. 2 (1840) 845, nom.
illeg —F. scholaris BLco, Fl. Filip. ed. 2 (1845)
93; ed. 3, 1 (1877) 171.—F. appendiculata BL.
Mus. Bot. 1 (1850) 169; Walp. Ann. 3 (1852)
76.—F. cuspidata Bit. Mus. Bot. 1 (1850) 170;
Walp. Ann. 3 (1852) 76; Mia. Ann. Mus. Bot.
Lugd.-Bat. 2 (1866) 218; Merr. J. Str. Br. R. As.
Soc. n. 77 (1917) 237; En. Philip. 3 (1923) 314.—
F. robusta Bu. Mus. Bot. 1 (1850) 170; Walp.
Ann. 3 (1852) 76.—F. subreticulata BL. Mus.
Bot. 1 (1850) 171; Walp. Ann. 3 (1852) 76;
Baku. f. in Back. Bekn. Fl. Java (em. ed.) 7
(1948) fam. 170, p. 10.—F. crassipes Bru. J. Linn.
Soc. Bot. 1 (1856) 99; Mia. Fl. Ind. Bat. 2 (1859)
1080; Gisss, J. Linn. Soc. Bot. 42 (1914) 111.—
F. latifolia Mig. Fl. Ind. Bat. 2 (1857) 369.—F.
thwaitesii F. v. M. Fragm. 2 (1861) 137.—F.
maingayi CLARKE in Hook. f. Fl. Br. Ind. 4 (1883)
84; Kinc, J. As. Soc. Beng. 74, i1 (1908) 608;
Ript. Fl. Mal. Pen. 2 (1923) 419; Foxw. Mal.
For. Rec. n. 3 (1927) 159.—F. spicata BAKER,
Kew Bull. (1896) 25; Merr. J. Str. Br. R. As. Soc.
n. 77 (1917) 237.—F. rodatzii K. Sco. & LAUT.
Fl. Schutzgeb. (1901) 499.—F. congestiflora ELM.
Leafl. Philip. Bot. 8 (1915) 2741.—F. grandifolia
Me_rr. J. Str. Br. R. As. Soc. n. 77 (1917) 231;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923) 338.
—F. cymosa Merr. J. Str. Br. R. As. Soc. 1. 77
(1917) 234; CAMMeERL. Bull. Jard. Bot. Btzg III,
5 (1923) 338.—F. stenophylla Becc. [For. Born.
(1902) 524, f. 65(1), nom. nud.) ex Merr. J. Str.
Br. R. As. Soc. n. 77 (1917) 236; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 337.—F.
pauciflora Ript. Fl. Mal. Pen. 2 (1923) 419,
f. 110; Burk. & HeNpeERS. Gard. Bull. S. S. 3
(1925) 399; Burk. Dict. (1935) 996.—F. eucalyp-
tifolia CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
312, f. 2.—F. teysmannii CAMMERL. I. c. 314,
f. 3.—F. gracilis CAMMERL. /.c. 316, f. 4.—F.
pendula Merr. Pl. Elm. Born. (1929) 251.—F.
minor (non BL.) Herne, Pfl. Clemens Kinabalu
(1953) 91. — Fig. 9-12.

Tree, 2—10(—16 or even —35?) m, up to c. 30cm @,
shrub or sometimes a straggling climber. Leaves
varying from broadly ovate via elliptic, obovate-
oblong and oblong to oblong-lanceolate or rarely
even linear, 5-50 by 1—23 cm, thinly to thickly
coriaceous or almost fleshy, sometimes bullate
between the nerves; base cuneate, obtuse, rounded
or often slightly to distinctly cordate; apex acute,
obtuse, rounded or very often shortly to rather
long acuminate; nerves usually 4-12 pairs, sunken
or less often flat to very slightly prominent above,
distinctly prominent beneath; reticulations lax,
more or less distinctly visible; petiole 14-5 cm,
thin to stout; stipules connate into an annular,

312

FLORA MALESIANA

[ser. I, vol. 6”

Fig. 10. Fagraea racemosa WALL. a. Fruiting twig

of ‘F. ligustrina’, X 24, b. ocrea, X 114, c. leaf
of ‘F. teysmannii’, < 24, d. leaf of ‘F. eucalypti-
folia’, X 2/4; (a Ker 15425, b WHITEHOUSE in herb.
Bri 016414, c PURSEGLOVE 4500, d BROOKE 10668).

up to 34 cm high ocrea which clasps the twig.
Inflorescences erect, nodding, or very often
drooping, 2-60 cm long (incl. the 1-30 cm long,
slender to robust peduncle); usually thyrsoid with
decussate, spaced, nearly sessile cymes of c. 7
(the higher ones less) long-stalked flowers; by
reduction spiciform if the internodes are short
combined with sessile cymes and short-stalked
flowers, corymbose if all cymes or at least the lower
ones are long-stalked, laxly branched, and many-
flowered, racemiform if the cymes are sessile and
few-flowered or in the apical part even 1-flowered,
or glomerulous if the whole inflorescence is reduced
to 5—7 flowers crowded on a long stalk; pedicels
(0—)34-3 cm, thin to stout, 2-bracteolate at the
base. Calyx campanulate to subglobose, 4-114
cm, divided about halfway down or slightly deeper.
Corolla-tube 2-4 cm, funnel-shaped. Anthers
rather thick oblong, c. 314 mm long, cells free in
their basal half. Stigma (broadened) obconical,
up to c. 144-2 mm broad, subtruncate, faintly
2-lobed. Berry globose-ellipsoid-ovoid, tipped by
the very short but distinct style-base, up to 214 cm
long, said to be sky-blue (or red?) when ripe.

Distr. SW. Indo-China, Siam, South Burma,
the Andamans and Nicobars, throughout Malaysia
(except the eastern half of Java, Madura I., and
the Lesser Sunda Is.), Solomon Is., and North
Australia (Northern Territory: Victoria River,
Providence Hill; Queensland: Cape York Penin-
sula).

Ecol. Light to rather dense, often secondary
forests on swampy to dry soil, on muddy river-
banks, podsolized sand, sometimes in savannahs
and lalang fields, 0-2000 m. F/. mainly April-
Sept., fr. July—Novy.

The flowers have a smell of butter or butter-
milk, as they produce diacetyl (see DeRx, Ann.
Bog. 1, 1950, 49-52), a substance which is charac-
teristic of bat-visited flowers; there are, however,
no observations that Fagraea flowers are visited
by bats. According to RIDLEY (Disp. 1930, 421)
the fruits are probably dispersed by bats. For
galls see Gisss, J. Linn. Soc. Bot. 42 (1914) 111,
and DocTers VAN LEEUWEN, Zoocecidia (1926)
464, f. 887.

Uses. The soft to moderately hard timber is
used for construction and as firewood. Decoctions
of the leaves, bark, and roots play a role in Ma-
layan medicine, mainly as tonics (see BURK.

4, Dict. /.c.); in the Philippines the bark and the

flowers are used as an antidote for snake bites.

Vern. False Coffee-Tree, E, gluguh babi, kaju
buliga, k. isop-isop dahojong, k. kapis, k. kopi-kopi,
k. obi, k. rupun-rupun, k. si markopi-kopi, léngugu,
rampisi, tuma taveuh pajo, Sum., tuma tafeu,
Simalur, kayu naga, Enggano, coffee utan, dada
kora (or kura), daun pépulit, d. puleh, glam tikus

jantan, kahwa hutan, (kaju) lidah rusath), (k.)

rumpo-rumpo, k. skobang, katiduran puak, kela
(payam), kémusang, kopi (h)utan, lambisi (or
limbisii) (pajah), mémbéra gading, méngkudu
badak, m. hutan, mumpuleh, (or mémpuleh, or
mumpulih), pépulit, (poké) pakan payd, p. sttibal
paya, p. suruas jantan, (p.) sépuleh, riang-riang

to
—_
ww

Dec. 1962] _ LOGANIACEAE (Leenhouts)

Fig. 11. Fagraea racemosa WALL. in secondary vegetation, Brunei (Borneo) (ASHTON, 1960).

314 FLORA MALESIANA

[ser. II, Vols 6°

Fig. 12. Fagraea racemosa WALL., close-up from the tree of fig. 11 (ASHTON, 1960).

Dec. 1962] LOGANIACEAE (Leenhouts)
gélugot, sapooli, sébérotut, sékobang, sépulis, long thyrsoid inflorescences, and medium-sized
sérawas, séruas, sétébal, siittibal, tahi musang, flowers (fig. 9a). An extremely coarse form, with

témbusu (or tumpusu), t. bukit, t. jantan, t. paya
(or gajah, or kaya), wa nam, Mal. Pen., kendung
badak, ki tjankuda, kopi métado, mangu leweng,
tjangkudu (or tjangkuda) badak, t. gunung, t.
léwéng, t. utan, tjang-tjangkuduan, S, baros, me-
lingu, J, dawun sukung, engkudu hutan, gémpar,
kadampuhok, kafi hutan, kalam kubak, kasadapan,
katiduran puak, kayu badal, k. ranau, kélagikong,
lankonis, legkau, lundus, mambunal, manggapuak,
nya gohong, puak bulan, sankaniap, sukong (or
sukung), tadapan puak, tapa labie, témhusu (or
témasuk) gaja, ténggilan (or tinggiran) puah (or
puak), (t)(o)dopon (or dapan) puak, ukudu ayer,
Born.; Philip.: baagu, Bag., bagontapai, porigabu,
Mbo., bakau, balat-buaya, kabal, libadkan, talob-
dlak, Tag., bogosald, S. L. Bis., bulobudya, ham-
budia, himbubiiya, lambudya, malabudya, salak,
P. Bis., cudman, gatibuka, makatibuga, sinalas,
Sub., kukodmon, Bik., kakdo-ita, magusayak,
Sul., kibudia, Laguna, malabago, Cebu; J/eraa,
Cel., kore haru, Halmaheira, barua, batteriengien,
berean, bisip, bogabogoia, dabe, fohkipeh, galud,
kankanis, kubugup, mahobaatje, makereko, malar,
misal, omborupa, sibéh, simbe, New Guinea,
gwaret, Aru Is.

Notes. Wide-spread, polymorphous species,
described under a host of names. The extreme
forms look so different that one might feel warrant-
ed to keep them wide apart, but on close exami-
nation of a sufficient number of specimens they
prove to be connected by numerous intergrades
which render a sharp delimitation impossible.
Notwithstanding the differences all specimens have
a certain habit in common so that the species can
nearly always be recognized at first sight.

The most common form is fairly coarse, with
thick twigs, relatively large, ovate, thick-coriace-
ous leaves, rounded to subcordate at the base,

larger and thicker leaves, nerves less conspicuous,
and large flowers was described from the Malay
Peninsula as ‘F. maingayi’. Another coarse form,
characterized by corymbose inflorescences with
short rachis and long lower branches, is the
Bornean ‘F. pendula’ (fig. 9c). ‘F. spicata’ differs
from the typical form only by its fairly short,
contracted inflorescences with sessile cymes or
flowers (Borneo) (fig. 9d). Other Bornean forms,
characterized by narrow leaves, are ‘F. teysmannii”
with lanceolate leaves (fig. 10c), and ‘F. eucalypti-
folia’ and *F. stenophylla’ with linear leaves, ta-
pering at both ends (fig. 10d); these narrow-
leaved forms have moreover small few-flowered
inflorescences with small flowers; they occupy
a rheophytic habitat (gravelbeds of stream-banks).
‘F. pauciflora’ (Mal. Pen.) differs from the normal
form by being more slender in al! parts, and espe-
cially in the inflorescence which has less and re-
latively small flowers. The most extreme form is
represented by ‘F. ligustrina’ (Mal. Pen. to
Moluccas, not uniform), with slender twigs,
small, elliptic leaves, tapering at base and apex,
nervation not very conspicuous, glomerulous
inflorescences, and flowers with a very small
calyx and a slender corolla; in this form the fruit
is also distinctly smaller (fig. 10a).

The names Kuhlia morindaefolia BL. and Kentia
morindaefolia StEup. are illegitimate as these
generic names were illegitimate.

Martius erroneously described F. malayana,
of which I saw the type in the Brussels Herbarium,
with a 6-merous corolla and 6 stamens and wrong-
ly assumed it to be related to F. e/liptica. BENTHAM
J. Linn. Soc. Bot. 1 (1856) 100, who only knew
the deficient description, took it for possibly
synonymous with Tabernaemontana corymbosa
Roxs. (Apocynaceae).

3. Section Fagraea

Sect. Fagraeae verae & sect. Eufagraea BL. Mus. Bot. 1 (1850) 163 & 169 resp.
—Sect. Corymbosae Bru. J. Linn. Soc. Bot. 1 (1856) 97.—Subg. Eufagraea MiQ.
Fl. Ind. Bat. 2 (1857) 367.—Sect. Pseudocorymbosae SOLER. in E, & P. Nat. Pfl.

Fam. 4, 2 (1892) 42.

Leaves auriculate or not. Inflorescences corymbose, dichasial, glomerulous,
or flowers solitary; always terminal and usually with a pair of strong branches in
the upper leaf-axils, therefore the inflorescence as a whole mostly sessile. Stamens

and style hardly or not exserted.

5. Fagraea ceilanica THUNB. Vet. Acad. Handl.
Stockh. 3 (1782) 132, t. 4; Nov. Gen. Pl. (1782)
35; BL. Rumphia 2 (1838) 29, t. 78 f. 2 (‘zeylanica’);
Hook. Bot. Mag. (1874) t. 6080; CLARKE in Hook.
f. Fl. Br. Ind. 4 (1883) 83, incl. var. brevituba;
TRIMEN, Fl. Ceyl. 3 (1895) 170; GAMBLE, FI.
Madras 5 (1923) 865; LeENH. Bull. Jard. Bot.
Brux. 32 (1962) 420.—Modagam RUuHEEDE,
Hort. Mal. 4 (1673) 119, t. 58.—F. obovata WALL.

in Roxb. Fl. Ind. 2 (1824) 33, non BL. (1826),
quae est F. blumei; DC. Prod. 9 (1845) 29; Hook.
Bot. Mag. (1846) t. 4205; Grirr. Notul. 4 (1854)
35; Ic. Pl. Ind. Or. 4 (1854) t. 382; non Mia.
Sum. (1861) 226 & 550, incl. var. latifolia (= F.
blumei); Kurz, Fl. Burm. 2 (1877) 205; CLARKE
in Hook. f. Fl. Br. Ind. 4 (1883) 83, p.p., incl. var.
gardneri; RipL. Trans. Linn. Soc. Bot. 3 (1893)
323; TRIMEN, FI. Ceyl. 3 (1895) 171; non F. M.

316

FLORA MALESIANA

[ser. I, vol. 6?

BAIL. Queensl. Agr. J. 3 (1898) 157, incl. var.
papuana (= F. berteriana); KiNG, J. As. Soc.
Beng. 74, ii (1908) 606, p.p.; ELm. Leafl. Philip.
Bot. 2 (1909) 598; 3 (1910) 857; Dop, Fl. Gén.
I.-C. 4 (1914) 174, excl. syn. p. p.; non CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 322 (= F.

Fig. 13. Fagraea ceilanica THUNB., a young spe-
cimen enclasping the trunk: of a tree with its
roots. Bogor (Java) (JAcons, 1958).

blumei); GAMBLE, Fl. Madras 5 (1923) 865; MERR.
En. Philip. 3 (1923) 315, p.p.; Ripv. Fl. Mal. Pen.
2 (1923) 418, p.p.; SASAKI, Cat. Gov. Herb.
Formosa (1930) 414; non Merr. Contr. Arn.
Arb. 8 (1934) 138 (= F. blumei); Burk. Dict. 1

(1935) 996, p. min. p.; HENDERS. J. Mal. Br. R.
As. Soc. 17 (1939) 58; KANsILAL & Das, FI.
Assam 3 (1939) 318; non BAKH. f. in Back. Bekn.
Fl. Java (em. ed.) 7 (1948) fam. 170, p. 11, nec in
Blumea 6 (1950) 382, incl. var. brevicalyx (both
= F. blumei); Kerr in Craib, Fl. Siam. En. 3
(1951) 56; non Hetne, Pfl. Clemens Kinabalu
(1953) 91 (= F. blumei).—F. litoralis Bu. Bijdr.
(1826) 1021; Rumphia 2 (1838) 28, t. 74, incl. var.
amboinensis; DC. Prod. 9 (1845) 30 (‘littoralis’);
Bi. Mus. Bot. 1 (1850) 166; Mia. Fl. Ind. Bat. 2
(1857) 374; Ann. Mus. Bot. Lugd.-Bat. 2 (1866)
217, incl. var. moluccana & forstenii; RACIBORSKI,
Ann. Jard. Bot. Btzg 17 (1900) 43, f. 23; K. & V.
Bijdr. 9 (1903) 82; CAMMERL. Bull. Jard. Bot.
Btzg III, 5 (1923) 328; Doct. VAN LEEUWEN,
Zoocecidia (1926) 463, f. 886; BAKH. f. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 11.—
F. lanceolata Bu. Bijdr. (1826) 1021, non WALL.
(1829), nec Mia. (1857); Rumphia 2 (1838) 31,
t. 77; DC. Prod. 9 (1845) 29; BL. Mus. Bot. 1
(1850) 167; Mia. Ann. Mus. Bot. Lugd.-Bat. 2
(1866) 218; K. & V. Bijdr. 9 (1903) 81; K. & G.
J. As. Soc. Beng. 74, ii (1908) 607; Koorpb. FI.
Tjib. 3 (1918) 47; CAmMMeRL. Bull. Jard. Bot.
Btzg III, 5 (1923) 329; Riv. FI. Mal. Pen. 2
(1923) 420; non HeNnpers. Gard. Bull. S.S. 4
(1927) 99 (= F. carnosa); Baku. f. in Back.
Bekn. FI. Java (em. ed.) 7 (1948) fam. 170, p. 12.—
F. minor REINW. ex BL. [Cat. (1823) 47, nom.
nud.| Bijdr. (1826) 1021; Rumphia 2 (1838) 27,
t. 73 f. 1; DC. Prod. 9 (1845) 30; BL. Mus. Bot. 1
(1850) 166; Mia. Fl. Ind. Bat. 2 (1857) 370;
non HEINE, Pfl. Clemens Kinabalu (1953) 91 (= F.
racemosa).—F. malabarica BL. Rumphia 2 (1838)
29.—F. rostrata BL. Rumphia 2 (1838) 31, t. 76
f. 2; DC. Prod. 9 (1845) 30; BL. Mus. Bot. 1 (1850)
168; Mia. Fl. Ind. Bat. 2 (1857) 374; SCHEFF.
Ann. Jard. Bot. Btzg 1 (1876) 38.—F. crassifolia
Bi. Rumphia 2 (1838) 31, t. 78 f. 1, mon WALL.
(1829) (= F. ridleyi); DC. Prod. 9 (1845) 30; BL.
Mus. Bot. 1 (1850) 166; Bru. J. Linn. Soc. Bot. 1
(1856) 98; Mia. Fl. Ind. Bat. 2 (1857) 373; Ann.
Mus. Bot. Lugd.-Bat. 2 (1866) 217.—F. coro-
mandelina WiGut, Ic. 4 (1850) t. 1316.—F. ma-
labarica WiGut, l.c. t. 1317, nom. illeg., non BL.
(1838).—F. amboinensis BL. Mus. Bot. 1 (1850)
166; Walp. Ann. 3 (1852) 75; Merr. Int. Rumph.
(1917) 424, excl. syn. Rumph. (p.p. 2); CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 329.—F. forstenii
Bt. Mus. Bot. 1 (1850) 166; Walp. Ann. 3 (1852)
75; non Koorp. Minah. (1898) 540 and Koorp.—
ScHum. Syst. Verz. 3 (1914) 105 (= F. truncata).
— F. celebica Bu. Mus. Bot. 1 (1850) 167; Walp.
Ann. 3 (1852) 75.—F. heterophylla BL. Mus. Bot. 1
(1850) 168; Walp. Ann. 3 (1852) 75; Mia. Fl. Ind.
Bat. 2 (1857) 375.—F. congesta BL. Mus. Bot. 1
(1850) 168; Walp. Ann. 3 (1852) 75.—F. fuscescens
Bi. Mus. Bot. 1 (1850) 168; Walp. Ann. 3
(1852) 75.—F. splendens Bu. Mus. Bot. 1 (1850)
168; Walp. Ann. 3 (1852) 75.—F. khasiana Bru.
J. Linn. Soc. Bot. 1 (1856) 99; CLARKE in Hook.
f. Fl. Br. Ind. 4 (1883) 84; BraAnpis, Ind. Trees
(1906) 476; KANNILAL & Das, Fl. Assam 3 (1939)
319.—F. oxyphylla Mig. Fl. Ind. Bat. 2 (1857)

Dec. 1962]

LOGANIACEAE (Leenhouts)

317

371, nom. illeg—F. gardneri THWaAtTESs, En.
(1860) 200; Beppome, FI. Sylv. (1869) 164.—
F. ternatana Mia. Ann. Mus. Bot. Lugd.-Bat. 2
(1866) 217; CAMMeERL. Bull. Jard. Bot. Btzg III,
5 (1923) 333; non HotTH. & LAM, Blumea 5
(1942) 230 (= F. gitingensis).—F. oblonga K. & G.
J. As. Soc. Beng. 74, ii (1908) 612; Ripv. J. Str.
Br. R. As. Soc. n. 50 (1908) 121; Fl. Mal. Pen. 2
(1923) 417; Burk. & Hot. Gard. Bull. S.S. 3
(1923) 61; Burk. & HENDeRS. Gard. Bull. S.S. 3
(1925) 399.—F. sasakii HayaTA, Ic. Pl. Formos.
3 (1913) 151, t. 29; KANEH. Formosan Trees,
rey. ed. (1936) 623, f. 581.—F. gardeniaeflora
WERNH. Trans. Linn. Soc. Bot. 9 (1916) 111.—F.
macrodendron GitG & BENED. Bot. Jahrb. 54
(1916) 187.—F. birmanica GANDOGER, Bull. Soc.
Bot. Fr. 65 (1918) 58.—F. prainii GANDOGER,
l.c.—F. lutea CAMMERL. Bull. Jard. Bot. Btzg III,
5 (1923) 330, f. 8; non Merr. Contr. Arn. Arb. 8
(1934) 138 (= F. blumei).—F. chinensis MERR.
Philip. J. Sc. 23 (1923) 261.—F. loheri MERR.
ibid. 27 (1925) 49.—F. pyriformis S. Moore,
J. Bot. 63 (1925) Suppl. p. 70.—F. sparei HENDERS.
Gard. Bull. S.S. 7 (1933) 114, t. 28 A.—F. rahmatii
Merr. Pap. Mich. Ac. Sc. 23 (1938) 189.—F.
angiensis KANEH. & Hatus. Bot. Mag. Tokyo
56 (1942) 160, f. 3. —F. archboldiana MeErR.
& Perry, J. Arn. Arb. 23 (1942) 412.— Fig.
13-15.

Epiphytic or (more rarely) terrestrial shrub,
climber, or small tree, up to 15 m. Leaves very
variable as to shape and size, varying from ovate
to obovate (to spathulate in the Deccan and
Ceylon) via oblong or oblong-obovate to narrowly
lanceolate, 414-35 by 114-9 cm, fleshy or co-
riaceous, base attenuate and decurrent, cuneate,
or rounded to subcordate, apex rounded to acute,
usually (and mostly gradually) acuminate, acumen
short to long and blunt to acute; nerves 4—8(—12)
pairs, inconspicuous or mostly invisible (only in
the typical form from Ceylon and S. Deccan
slightly prominent beneath); petiole slender to
robust, 14-314(—5) cm, exauriculate, the axillary
scale appressed to the twig or (more rarely) partly
or entirely adnate to the petiole, mostly inconspic-
uous, rarely somewhat broader than the petiole,
Y%4—-34(-1) cm long, usually rounded, rarely
truncate or emarginate, often slightly confluent
between the leaves. Jnflorescences dichasial,
branched from the base, usually rather dense,
sometimes laxly thyrsoid and up to 15 cm long,
(1—)3-17-flowered; pedicels slender (then usually
thickened towards the calyx) to stout, 144-314 cm
long; bracteoles 2, inserted from about halfway
the pedicel to just below the calyx, very small to
c. 24 cm long (usually the more apical the larger),
sometimes confluent at the base and together
enclasping the calyx. Ca/yx campanulate to slightly
urceolate, 1-234(-4) cm long (in Assam 6-714
mm), connate for 14 to more than 14. Corolla
widely to slenderly funnel-shaped, tube 2-5 cm
(in the typical form from Ceylon and SW. Deccan
8144-10 cm). Anthers oblong to ovate, (314-)
5-7'4(-1214) mm long, cells free at the base for
14-14. Stigma obconical, slightly cup-shaped to

peltate, 1-5 mm @o. Fruits ovoid or ellipsoid to
subglobular, 3-5 cm long (specimens from Assam
134 cm), slightly to strongly beaked, glaucous-
white (sometimes mentioned as being coffee-
brown or black); calyx often warty-lenticellate,
lobes appressed, spreading, or reflexed.

Distr. Ceylon, SW. Deccan, Assam, Burma,
Siam, Indo-China, SW. China (SW. Yunnan),
Hainan, Hongkong, Formosa (S. peninsula only),
and throughout Malaysia.

Cited by Sasaki, lc. (as F. obovata) from
Ponape (Carolines); I saw no material from there;
confusion with small-flowered F. berteriana seems
probable.

Ecol. Occupying a wide range of habitats, in
open localities and along forest edges but also in
primary and secondary forests, both on dry and on
marshy or periodically inundated soils, under
everwet as well as under seasonal conditions, from
sea-level up to 2500 m. F/. mainly in the dry season,
fr. in the rainy period.

Uses. Sometimes cultivated as an ornamental
shrub.

Vern. Kaju djottik, Sum., angiladn bilu, bani
bani luam, olor arélah, Simalur, ara burong,
Mal. Pen., kikunteh, kitérong, S, kémrungén,
képitu, J, rauai kajarok, Sumba, salang mapit,
Born.; Philip.: bankalon, Bag., busalsal, Sbl.,
kotofgogan, Mbo., malatunog, C. Bis., mamagon,
Bik., panagang, Tagb., pospusug, pukut, Ig.;
papatjeda bangan, p. utan, pasi gunung, Ternate,
tonki utan, Ambon, méngando, Moluccas (?),
kulaken (or kuleka), N. G.

Notes. In the circumscription given here an
extremely variable species. Both inside and out-
side Malaysia it comprises several local forms,
some of which have very conspicuous characters;
it is quite understandable that they were described
as distinct species, sometimes not even thought
to be closely related. After a careful study of a
large number of specimens from all parts of the
area I feel convinced that all these forms belong to
one species. Most of the more extreme ones are in
fact not sharply delimited but grade into other
forms. In a few cases where there is a really sharp
demarcation this depends on characters which
are ephemeral and of no systematic importance
in this species. Though it would not be impossible
to subdivide this species into a (fairly large)
number of forms and varieties I have refrained
from doing so. I confine myself to an enumeration
of the more important local forms in Malaysia
with the synonyms belonging to them:

‘F. litoralis’ is the most common form to which
most of the synonyms belong; it occurs throughout
Malaysia: \eaves medium-sized to small, elliptic,
acute at the base, gradually acuminate at apex,
nerves invisible; axillary scale appressed to the
twig, small, rounded; inflorescence 3—5-flowered;
bracteoles inserted about halfway the pedicel,
small; calyx 1-134 cm, sepals free for at least half
their length; anthers 4-34 cm long; stigma up
to 314 mm g, mostly about obconical; fruits
subglobular, not strongly beaked, calyx spreading.

“Kinabalu-race’ (Borneo: Mt Kinabalu): differs

318°

FLORA MALESIANA

[ser. I, vol. 62

from ‘F. /itoralis’ by lanceolate leaves, solitary
flowers, and the calyx which is appressed to the
fruit. It comes close to the type of ‘F. sasakii’
from Formosa.

‘F. pyriformis’ (Sumatra): differs from ‘F.
litoralis’ mainly in its calyx, which is 134-214 cm
long and slightly constricted in the mouth, the
sepals being halfway connate.

‘F. oblonga (Sumatra, Malay Peninsula): a
very characteristic form which differs from ‘F.
litoralis’ by long and narrow leaves (11-35 by 2-9
cm), when dried greyish green, beneath lighter or
brownish, apex usually long and slender acuminate;
nerves 10-12 pairs, visible though inconspicuous;
petiole long and slender (2-5 cm); axillary scale
slightly broader and truncate; inflorescence re-
latively many-flowered; calyx 114 cm _ long,
slightly constricted at the mouth.

‘F. sparei’ (Sumatra, Malay Peninsula): a cha-
racteristic form which differs from ‘F. litoralis’ by
small (7-10 by 314-5 cm) stiff leaves with a blunt
to slightly acuminate apex; inflorescences 1-—9-
flowered; calyx large (214-234 cm); bracteoles
inserted just below and appressed to the calyx,
c. ¥% cm long, lanceolate.

Fig. 14. Fagraea ceilanica THUNB., calyces and

bracteoles of Philippine specimens. a. Bracteoles

small and attached about halfway the pedicel,

b. bracteoles slightly larger, attached just below

the calyx, c. bracteoles connate, forming a cupule

around the calyx, all * 24 (a WILLIAMS 1316,
b BS 45700, c WENZEL 936).

“Philippine-race’ (Philippines: Mindoro, Luzon,
Catanduanes, Leyte, Negros, Mindanao) mainly
differs from ‘F. litoralis’ by a large calyx (214 cm)
which at its base is enveloped by a cupule com-
posed of the confluent, 1144 cm long bracteoles.
‘F. loheri’ is intermediate between this form and
‘F. litoralis’: it has a long calyx and high-inserted
bracteoles, but these are smaller, patent, and not
confluent at the base (Luzon) (fig. 14).

“F. ternatana (Luzon, Moluccas, New Guinea)
differs from ‘F. Jitoralis’ by somewhat broader
leaves, a blunt to rounded, slightly acuminate
apex; axillary scale adnate to the petiole, slightly
larger, truncate to emarginate; calyx large (214-
2% cm), slightly constricted at the mouth; anthers
long and narrow (1 cm); stigma broadly peltate
(5S mm Qg); fruit distinctly beaked. ‘F. angiensis’
from New Guinea differs only from ‘F. ternatana’
by its leaves, which are more similar to those of
‘F. litoralis’ but slightly obovate (fig. 15).

‘F. gardeniaeflora’ (incl. F. archboldiana; New

Guinea) differs from the closely related ‘F. terna-
tana’ and ‘F. angiensis’ by a 234-4 cm long calyx,
enveloped by a cupule composed of the strongly
enlarged (134-214 cm long) partly connate brac-
teoles; anthers 1-114 cm long.

‘F. minor (best developed in Borneo) differs
mainly from ‘F. /itoralis’ by the ovate leaves which
are rounded to subcordate at the base.

‘F. forstenii’ and ‘F. celebica’ (Celebes) differ
mainly from ‘F. Jitoralis’ by the spindle-shaped
fruits, the latter furthermore by the broad, almost
deltoid axillary scale.

The affinity of F. ceilanica is especially with 6.
F. annulata and 10. F. blumei (see under these
species for differences and further remarks).

The pseudo-axillary inflorescences, as mentioned
by CAMMERLOHER in a note to his description of
F. lutea, are apparently only the normal, axillary
lower branches of a furthermore broken terminal
inflorescence.

According to MERRILL’s description, in F. loheri
the tube of the corolla would be 11 mm long, the
lobes 16 mm; in fact, the measurements are exactly
the reverse.

The leaves are always exauriculate, but in F.
gardeniaeflora on both sides of the broadened
petiolar base a distinct wart (probably a gland)
can be found, which may look like a rudimentary
auricle.

Funis toaccae Rumputus, Herb. Amb. 5 (1747)
481, t. 179 has been referred by MERRILL, Int.
Rumph. (1917) 424 to F. amboinensis BL. but as
he rightly remarked, the fruits doubtless belong to
an Apocynacea; the latex mentioned to occur in the
twigs points to the same family. See also HEYNE,
Nutt. Pl. (1927) 1292, under Apocynacea. On the
other hand the large flowers (according to the fig-
ure they must be about 10 cm) with 5 free stamens
may represent some Fagraea (though not the
present species).

F. oxyphylla MiQ. was published as a new more
appropriate name after combination of three
earlier species and is therefore illegitimate.

6. Fagraea annulata Hiern, Nova Guinea 8 (1909)
202; CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
Biss 1, Al);

Tree, 6-8 m tall. Leaves petioled, (oblong-)ovate
to (oblong-)obovate, 5-14 by 244-514 cm, firmly
herbaceous or thinly coriaceous, base acute, often
subcontracted and decurrent, apex very shortly
acute-acuminate; nerves c. 10 pairs, very thin
and hardly visible above, invisible beneath, veins
inconspicuous except sometimes some intermediate
ones; petiole slender, 34—2!4 cm; exauriculate;
axillary scale adnate to the petiole, 44-14 cm
long, fairly broad (much broader than the
base of the petiole), truncate to faintly 2-lobed.
Inflorescences cymose, 2—15-flowered, dense; pe-
dicels thick, 14-12 mm long, often 2-bracteolate
above the middle and then thickened above the
bracteoles. Ca/yx campanulate, often with recury-
ed lobes, 34-1 cm long, divided about halfway.
Corolla-tube funnel-shaped, 234-31 cm, inside
about halfway with a distinct horizontal rim bear-

Dec. 1962]

LOGANIACEAE (Leenhouts)

Fig. 15. Fagraea ceilanica THUNB., a flowering specimen of ‘F. angiensis’, Mt Gwamongga at 2450 m
(W. New Guinea) (SLEUMER, 1962).

ing the stamens. Anthers oblong, c. 34 cm long, to
about halfway bifid. Stigma subpeltate, c. 214 mm
9. Fruits (unripe) ellipsoid-oblong, c. 114 by 34 cm.
Distr. Malaysia: SW. New Guinea (Noord
River), once collected.
Ecol. On river-bank at low altitude. F/. June.

Note. Doubtless very close to F. ceilanica; in
fact the only difference with that species is the
thickened annulus in the corolla-tube bearing the
stamens. This character it has in common with the
small group of species with a 2-lobed stigma (F.
berteriana and allies). As only one specimen is
known it is not impossible that it is of hybrid ori-
gin, the more so as in New Guinean specimens of
F. ceilanica there seems to be a slight introgression

of berteriana-characters,
the leaves.

especially concerning

7. Fagraea acuminatissima Merr. J. Str. Br. R.
As. Soc. n. 77 (1917) 232; CAMMERL. Bull. Jard.
Bot. Btzg III, 5 (1923) 330, f. 9.—F. obovata (non
WALL.) KING, J. As. Soc. Beng. 74, ii (1908) 606,
p.p.; Riwi. Fl. Mal. Pen. 2 (1923) 418; Burk.
Dict. (1935) 996.—F. longicuspis GANDOGER, Bull.
Soc. Bot. Fr. 65 (1918) 58.

Usually a shrub or small tree, sometimes a
climber. Leaves petioled (suborbicular to) oblong
to lanceolate or subobovate, 7-24 by 334-10 cm,
thickly coriaceous (fleshy when fresh); base acute
and decurrent, apex obtuse or rounded, abruptly
contracted into a narrowly triangular, acute,

320

FLORA MALESIANA

[ser. I, vol. 6?

3%4(—2) cm long acumen; nerves invisible; petiole
robust, 114-5 cm long, exauriculate; axillary
scale appressed to the twig, inconspicuous, round-
ed. Jnflorescences subsessile, subglomerulous,
3—5-flowered; pedicels thick, 44-34 cm long,
about halfway with a pair of small, appressed to
patent bracteoles. Calyx campanulate, 34-114 cm
long, deeply divided (about 24—34 of its length).
Corolla-tube funnel-shaped, 214-4'4 cm long.
Anthers oblong, thick, ?/;-34 cm long, cells free
in their basal half. Stigma broadly obconical to
peltate, c. 144 mm 9. Fruits ellipsoid to globular,
c. 1144-2 cm 9; calyx appressed.

Distr. Malaysia: Sumatra (also Banka and
Riouw Is.), Malay Peninsula, W. and N. Borneo.

Ecol. Mainly in peat-swamp forests, sometimes
also in bamboo forests, up to 750 m. Fi. Febr.—
Noy., fr. Febr., July, Sept.

Vern. Malikoh, Banka, akar tunkoo beavak,
témbusu, Mal. Pen., baruas babi, dawi-dawi, nyatu,
Born.

Notes. This species belongs to the wider
relationship of F. cei/anica (in the Malay Peninsula
it has always been identified with F. obovata).
Its most distinctive characters are the often black-
ish, round twigs, the dull- to olive-brown upper
and smooth red-brown lower side of the very stiff
dried leaves, and the small, + sessile, glomerulous
inflorescences, with the small calyces. Apart from
the flowers which are very long and slenderly
tubular in F. tubulosa there is a great resemblance
with that species; a good difference is found in
the midrib, which is rounded beneath in the
present species, strongly keeled in F. tubulosa.

8. Fagraea tubulosa BL. Mus. Bot. 1 (1850) 167;
Walp. Ann. 3 (1852) 75; Mig. Fl. Ind. Bat. 2
(1857) 373; Kina, J. As. Soc. Beng. 74, ii (1908)
604; CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
326; RipL. Fl. Mal. Pen. 2 (1923) 415.

Straggling climbing shrub. Leaves petioled,
ovate-oblong, oblong, or oblong-lanceolate, 10—
2214 by 5-10 cm, thinly coriaceous, base broadly
cuneate or obtuse, narrowly decurrent, apex
abruptly short- to long-subcaudate; nerves 5—7
pairs, invisible or almost so above, faintly visible
Or quite inconspicuous beneath; petiole 14-3 cm
long; exauriculate; the axillary scale inconspicuous,
adnate to the petiole, blunt, 3 mm long. Flowers
1-6 in sessile glomerules; pedicels very short with
1-2 pairs of small bracteoles. Ca/yx campanulate,
14—1 cm long, divided somewhat less than halfway
down. Corolla-tube tubular, very slender, 8—9 cm.
Anthers elliptic, c. 4 mm long, cells in their basal
half free. Stigma peltate, 2 mm o. Fruit unknown.

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

Ecol. In forests, c. 900 m. F/. June.

Note. A very distinct species, well characterized
by its glomerulous, long and slender, fragile
flowers. Apart from this character much alike
(and possibly related to) F. acuminatissima, which
has the midrib rounded beneath, not keeled as it
is here.

9. Fagraea ridleyi K. & G. J. As. Soc. Beng. 74,
ii (1908) 612, non GANDOGER (1924); RiDL. J.
Str. Br. R. As. Soc. m. 50 (1908) 118; Fl. Mal.
Pen. 2 (1923) 417.—F. crassifolia WALL. Cat.
(1829) n. 1602, nom. nud., non BL. (1838) (= F.
ceilanica).—Fig. 16.

Liana or big straggling shrub. Leaves petioled,
obovate to suborbicular, 13—24 by 9-18 cm, thick-
coriaceous, base acute, somewhat decurrent, apex
rounded, with or without an abrupt short and
blunt acumen; nerves 4—10 pairs, flattish or slight-
ly impressed above, prominent beneath; petiole
2-4 cm, robust, exauriculate; stipules connate in
a 4 mm high ocrea which in older leaves is split
between the petioles. Inflorescences cymose, 3—9
flowered; pedicels 14-2 cm, thick, tapering into
the calyx, usually with a pair of small patent
bracteoles somewhat above the middle. Calyx
campanulate, 2-214 cm high, tube woody, lobes
114-134 cm, thick-coriaceous. Corolla-tube funnel-
shaped, 314-4 cm. Anthers bifid to halfway, ob-
long, 344 cm long. Stigma subpeltate, 2144 mm og.
Fruits inversely pear-shaped to globular and conic-
ally beaked, c. 5 by 314 cm; calyx loosely appress-
ed.

Distr. Malaysia: Malay Peninsula,
Arch., and Borneo (Sarawak, Brunei).

Ecol. In the Malay Peninsula on rocky sea-
cliffs and along rivers, in Borneo in primary
forests on sandstone hills, 0-300 m. Fi. May,
Aug., fr. April, May, and Sept.

Vern. Buah télan kényalang, Born. (this means:
the fruit which is swallowed by the hornbills).

Notes. Specimens from Borneo differ slightly
from those of the Malay Peninsula; the main
difference is in the fruits, which are globular with
a strong conical beak in Borneo, inversely pear-
shaped in the Malay Peninsula.

The relationship of F. ridleyi is doubtless with
F. blumei (which differs by warty-lenticellate in-
florescences and conspicuous axillary scales) and
with F. ceilanica (which differs by invisible nerves).

As lectotype I have designated RIDLEY 5845
(Smnc; dupl. BM, K).

Lingga

10. Fagraea blumei G. Don, Gard. Dict. 4 (1837)
69 (‘blumii’); SteuD. Nomencl. ed. 2, 1 (1840)
624; DC. Prod. 9 (1845) 30.—F. obovata BL.
Bijdr. (1826) 1021, nom. illeg., non WALL. (1824).
—F. obovato-javana BL. Rumphia 2 (1838) 29,
t. 75, nom. superfl., incl. var. bebeak; Mus. Bot.
1 (1850) 164; Mie. Ann. Mus. Bot. Lugd.-Bat. 2
(1866) 217, incl. var. latifolia; K. & V. Bijdr. 9
(1903) 78; Koorp. Exk. Fl. Java 3 (1912) 59; Fl.
Tjib. 3 (1918) 47; Hocnr. Candollea 6 (1936) 473.
—F. leschenaultii BL. Rumphia 2 (1838) 30;
Mus. Bot. 1 (1850) 164; Mug. Fl. Ind. Bat.
2 (1857) 370.—F. plumeriaeflora DC. Prod. 9
(1845) 29; Bento. J. Linn. Soc. Bot. 1 (1856)

98 (‘plumeriaefolia’); Mig. Fl. Ind. Bat. 2
(1857) 375; Exm. Leafl. Philip. Bot. 2 (1909)
598; ibid. 3 (1910) 857; Merr. En. Philip. 3

(1923) 315, excl. spec. Celeb.; non KoorD. Minah.
(1898) 540, nec Koorp.—ScHum. Syst. Verz. 3
(1914) 105 (both are F. truncata).—F. obovata

Dec. 1962] ___LOGANIACEAE (Leenhouts) 321

Fig. 16. Fagraea ridleyi K. & G. a. Habit (note the glands on the lower side of the leaf), b. opened corolla,
c. style and stigma, d. fruit, all * 24 (RmpDLEY 5845).

(non WALL.) Mig. Fl. Ind. Bat. 2 (1857) 369; En. Philip. 3 (1923) 315,p.p.; Doct. VAN LEEUWEN,
Sum. (1861) 226, incl. var. latifolia; CLARKE in Zoocecidia (1926) 463; Merr. Contr. Arn. Arb. 8
Hook. f. Fl. Br. Ind. 4 (1883) 83, p.p.; CAMMERL. (1934) 138; BAKH. f. in Back. Bekn. Fl. Java (em.
Bull. Jard. Bot. Btzg III, 5 (1923) 322; Merr. ed.) 7 (1948) fam. 170, p. 11; Blumea 6 (1950)

322

FLORA MALESIANA

[ser. I, vol. 6?

Fig. 17. Fagraea blumei G. Don. a. Habit,

382, incl. var. brevicalyx.—F. vaginata K. & G.
J. As. Soc. Beng. 74, ii (1908) 610; Rip. FI.
Mal. Pen. 2 (1923) 418; S. Moore, J. Bot. 63
(1925) Suppl. 69; Burk. Dict. 1 (1935) 997.—
F. negrosensis ELM. Leafl. Philip. Bot. 2 (1909)
599; Merr. En. Philip. 3 (1923) 315.—F. cuernosen-
sis ELM. Leafl. Philip. Bot. 2 (1909) 600; Merr.
En. Philip. 3 (1923) 314.—F. intermedia RIDL.
J. Fed. Mal. St. Mus. 8, n. 4 (1917) 63.—F.
spatiosa 8. Moore, J. Bot. 63 (1925) Suppl. 69.—
F. lutea (non CAMMERL.) Mere. Contr. Arn. Arb.
n. 8 (1934) 138.—Fig. 17.

Tree up to 26 m, scandent or erect, epiphytic or
terrestrial shrub, or climber. Leaves mostly obo-
vate, sometimes elliptic to oblong, rarely lanceolate,
7-60 by 2-30 cm, thin- to stiff-coriaceous, base
acute, mostly long-attenuate, apex rounded to
cuneate, nearly always acuminate, acumen taper-
ing to abrupt, short and broad to fairly long and

24, b. opened corolla, < 1, c. stigma, < 5, d. anther, x 2
(a-d BUNNEMEYER 4950).

slender; nerves 3—15 pairs, usually slightly though
distinctly prominent on both surfaces, rarely in-
conspicuous to invisible; petiole 44-4 cm, mostly
robust, often narrowly winged, exauriculate;
axillary scale for the greater part adnate to the
petiole but distinctly broader, 4 mm-—1!4 cm long,
usually 2-lobed to truncate, rarely rounded.
Inflorescences terminal with 2 strong branches in
the upper leaf-axils, cymose to corymbose, lax to
rather dense, (1—)7—c.50-flowered, sparsely to
densely warty-lenticellate (still more conspicu-
ously so in fruit); pedicels usually stout, 144-214 cm,
2-bracteolate in the lower two-thirds. Calyx cu-
pular to campanulate, very slender to rather broad,
4-20 mm long, sepals connate for 4-2/4. Corolla-
tube narrowly to widely funnel-shaped, 114-4 cm,
lobes varying from about 14 to nearly as long as
the tube. Anthers thickly ovate-oblong to lanceo-
late, 4-10 mm, to halfway bifid. Stigma capitate

Dec. 1962]

LOGANIACEAE (Leenhouts)

323

to peltate, 1-5 mm og. Fruits either subglobular
(sometimes beaked) and 3-4 cm 9, or spindle-
shaped and 5-7 by 134-2!4 cm, said to be black
(Sumatra) or yellow (Philippines); calyx with
loosely appressed to spreading or caducous lobes.

Distr. Malaysia: Sumatra, Malay Peninsula
(Perak, Pahang, Selangor), Java, Borneo, and the

Philippines; a few dubious specimens from
Celebes.
This species comprises two subspecies, as
follows:

ssp. bDlumei.—F. blumei G. Don.—F. obovata BL.—
F. obovato-javana BLu., incl. var.—F. leschenaultii
BL.—F. obovata (non WALL.) Auct., incl. var.
—F. vaginata K. & G.—F. negrosensis ELM.
—F. intermedia RipL.—F. spatiosa S. Moore.
—F. lutea (non CAMMERL.) MERR.

Axillary scale + flat, dark-brown. Leaves
obovate, up to 27 by 13 cm, thick- and stiff-
coriaceous, dark-brown when dry, base long-
attenuate, apex rounded with a minute, broadly
triangular acumen; nerves 3-6 pairs, distinctly
forked towards the margin, conspicuous beneath.
Pedicels up to 1 cm, with a pair of bracteoles
about halfway. Calyx up to 114 cm. Corolla-tube
up to 214 cm. Anthers up to 7 mm. Stigma usually
1-214 mm @. Fruits (sub)globular, calyx persistent.

Distr. Sumatra, Malay Peninsula, Java, rare in
Borneo and the Philippines (Negros).

Ecol. Preferably on a fertile wet soil and in
light shade, in and especially along primary and
secondary forests, on river-banks, efc.; usually
under everwet, in East Java also under seasonal
conditions. Rarely under 500 m (except in the
Malay Peninsula), usually 1200—1800(—2200) m.
Fl. fr. Jan.—Dec.

Uses. The rubbed leaves are used as a medicine
against fever and headache; latex from the fruits
used to make lime-twigs for catching birds. The
fruits are said to be poisonous.

Vern. Akar konjal, kaju djottik, kama lojang,
Sum., akar térong kusang, aréng kidjamudju, bebeak,
kalang mangga, ki minjak, kitérong areui, S,
djeruk utan, (én)dog-(én)dogan, kémplek, ki hurip
(or horip), ki kéndal, ki teréng, salisor, wuru, J,
njam plongolos, Md, dirung, Born.; Philip.: buton,
dolis, Bag., maganonok, Sub.

ssp. plumeriaeflora (DC.) LEENH. nov. stat.—F.
plumeriaeflora DC.

Axillary scale concave, greenish. Leaves elliptic,
17-60 by 6-30 cm, thin-coriaceous, greyish green
when dried, base not long-attenuate, apex blunt,
abruptly terminated by a slender acumen; nerves
9-15 pairs, looped and joined near the margin,
visible on both sides. Pedicels 114-214 cm, brac-
teoles at the base. Calyx 134-2 cm. Corolla-tube
314-4 cm. Anthers 6-10 mm. Stigma c. 4 mm g.
Fruits spindle-shaped, calyx caducous.

Distr. Philippines (Luzon, Catanduanes, Sa-
mar, Leyte, Panay, Mindanao).

Ecol. Humid primary forests, 500-2000 m.
Fl. fr. Jan—Dec. The very fragrant flowers are
visited by butterflies.

Vern. Butoon ta-usd, Bag., kalaynig, Buk.

Notes. This species is related to F. ceilanica,
F. fastigiata, and F. tacapala. The conspicuous
stipules, the distinctly prominent nerves, and the
warty lenticellate inflorescences are good cha-
racters to distinguish it from F. ceilanica; only some
forms from the Philippines and from Sumatra
have about the same leaves as the latter species.
F. fastigiata is doubtless very close to ssp. blumei,
but differs by the auricles inserted slightly above
the leaf-base; it could be conspecific. F. tacapala
comes near ssp. plumeriaeflora, is also different by
the presence of auricles.

Typical specimens of the two subspecies are so
different that one would prefer to distinguish them
as good species. There are, however, two series of
intermediate specimens; the type of F. cuernosensis
and several other specimens I have not referred to
a subspecies.

The first series is formed by the specimens from
Negros (among which the types of F. negrosensis
and F. cuernosensis), several specimens from
Mindanao, and one from Luzon (FB 8996).
The Negros specimens come close to ssp. blumei
and differ mainly by their ceilanica(‘litoralis’)-like
leaves; most of the Mindanao specimens have the
same kind of leaves, but the inflorescences and
flowers look more like those of ssp. plumeriaeflora,
with which they have also the spindle-shaped
fruits in common; the calyx is not caducous,
however. The latter character is the main differen-
ce between the remaining specimens and ssp.
plumeriaeflora.

The second series goes via Borneo. Among the
Bornean specimens one (BECCARI PB 3032) comes
close to ssp. blumei, from which it differs only by
the greater number of nerves. All other Bornean
specimens combine the leaves of ssp. plumeriae-
flora with the inflorescences, flowers, and fruits of
ssp. blumei.

Especially ssp. blumei shows a wide range of
variability; this mainly concerns the inflorescences
and the flowers (many- to few-flowered inflores-
cences, small to big calyces and corollas, the
former cupular to campanulate and with short to
long lobes, the latter slender to wide). This varia-
bility is gradual, and in Java nearly the full range
of specimens is found. Some local races with de-
viating characters are restricted to Central Su-
matran volcanoes. The specimens from Mt.
Merapi are mainly aberrant by the caducous
calyx under a globular fruit. Those of Mt. Kerintji
differ only from typical b/umei in the large, many-
flowered inflorescence and the small flowers
(calyx cupular, 4 mm high, less than halfway
connate, corolla-tube 114 cm long). The specimens
from Mt. Singalang have oblong, acute leaves, the
nerves of which are invisible, a very lax, few-
flowered inflorescence and the smallest flowers
(calyx tubular, 4 mm long, more than halfway
up connate, corolla-tube c. 5 mm). On account of
the leaf-shape, the inflorescences, and the small
flowers, the latter race is sometimes confused with
F. fragrans (which has, however, lateral inflores-

324

FLORA MALESIANA

[ser. I, vol. 6?

cences). Specimens from Mt. Sibajak are mainly
characterized by elliptic, acute leaves, the nerves
of which are nearly invisible, by short-stalked and
densely crowded inflorescences of 3-9 flowers,
with big (6 by 10 mm) bracteoles clasping the
calyx, and by sepals connate for only 14.

The nomenclature of F. blumei is somewhat
confused. BLUME, who was the first to recognize
and publish it in 1826, named it F. obovata, ap-
parently unaware of the fact that WALLICH had
used the same epithet already two years earlier
for another species. Later on G. Don, BLUME
himself, and STEUDEL corrected this error, using the
epithets blumii, obovato-javana, and blumei
respectively. All these names were clearly based
upon F. obovata BL.

11. Fagraea fastigiata BL. Rumphia 2 (1838) 30,
t. 76 f. 1; DC. Prod. 9 (1845) 30; BL. Mus. Bot. 1
(1850) 164; Mig. Fl. Ind. Bat. 2 (1845) 369;
K. & V. Bijdr. 9 (1903) 80; Koorp. Exk. FI.
Java 3 (1912) 59; CAmMMeERL. Bull. Jard. Bot.
Btzg III, 5 (1923) 322, pro spec. Jay., fr. excl.;
BAKH. f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 170, p. 11, p.p.; non Rip. J. Str. Br. R. As.
Soc. n. 30 (1897) 167, nec ENDERT, Tectona 13
(1920) 117 (= F. crenulata).—Fig. 2b.

Liana, epiphytic shrub, or small tree, up to
13 m by c. 10 cma. Leaves sessile, oblong-obovate,
13-27 by 6-12 cm, thin-coriaceous, base narrowly
cuneate, long-decurrent, again somewhat broaden-
ed and subauriculate slightly above the twig,
completely hiding the axillary scale which is
adnate to the midrib, + 34 cm long and rather
narrow; apex blunt to faintly blunt-acuminate;
nerves 5-7 pairs, inconspicuous above, pro-
minulous to prominent beneath. Inflorescences
corymbose, laxly branched, with c. 15—20 sub-
sessile flowers; bracteoles small, appressed, just
below the calyx. Calyx slender-campanulate, c.
7 mm, less than 14 divided. Corolla-tube widely
funnel-shaped, 3 cm. Anthers ovate-lanceolate,
7 mm, cells in their basal half free. Stigma un-
known. Fruits globular, 2 cm @; pedicels slender,
calyx-lobes reflexed.

Distr. Malaysia: Malay Peninsula
Ijok F. R.) and West and Central Java.

Ecol. Forests, c. 600-1000 m. Fr. June. For a
gall see DoctTers VAN LEEUWEN, Zoocecidia
(1926) 463, f. 885.

Vern. Lanupa, S.

Note. This species is closely related to F.
blumei ssp. blumei; it is even possible that it will
finally appear to be conspecific with that species,
from which it mainly differs by its auriculate
leaf-base. By this character it seems to link up
F. blumei with the group of species (n. 14-20) with
well-developed auricles.

(Perak:

12. Fagraea crenulata MAINGAY ex CLARKE in
Hook. f. Fl. Br. Ind. 4 (1883) 83; KING, J. As.
Soc. Beng. 74, ii (1908) 610; Dop, Fl. Gén. I.-C.
4 (1914) 176; CammMerL. Bull. Jard. Bot. Btzg III,
5 (1923) 321; RipL. Fl. Mal. Pen. 2 (1923) 420;
Foxw. Mal. For. Rec. n. 3 (1927) 160, plates;

Heyne, Nutt. Pl. (1927) 1269; DE Voocp, Trop.
Natuur 21 (1932) 60, f. 1; Burk. Dict. 1 (1935)
994; CorRNER, Ways. Trees (1940) 423, t. 121.—
F. fastigiata (non BL.) Riv. J. Str. Br. R. As. Soc.
n. 30 (1897) 167; ENpDERT, Tectona 13 (1920) 117.
—Fig. 18.

Fig. 18. Fagraea crenulata CLARKE in marshes
near Palembang (Sumatra) (DE VooGD).

Tree, up to 23 m by 50 cm 9, trunk and branches
with simple or divided prickles. Leaves sessile or
subsessile, broadly obovate, 18-45 by 14-25 cm,
thin-coriaceous to stiff-herbaceous, base narrowed,
then widened again into a pair of amplexicaulous
or recurved auricles, margin finely crenulate,
apex very broadly rounded to nearly truncate;
nerves 5-7 pairs, clearly visible above, prominent
beneath; axillary scale fully adnate to the petiole
or midrib, fairly narrow, acute, 114-2 cm, com-
pletely enveloped by the leaf-base. Inflorescences
corymbose, widely branched and many-flowered,
up to 25 cm; pedicels 1-2 cm, gradually thickened
towards the apex; bracteoles appressed, not very
large, usually about halfway the pedicel. Calyx
1 cm long, divided to quite near the base. Corolla-
tube funnel-shaped, 134-2 cm. Anthers oblong,
6 mm, cells free in their basal third. Stigma pel-
tate, 114-2 mm 9g, faintly 2-lobed. Fruits ellipsoid,
214 cm or longer; calyx more or less spreading.

Distr. South Indo-China and Malaysia: along
the east coast of Sumatra, the west coast of the
Malay Peninsula, and the southwest and south
coast of Borneo.

Ecol. Permanent or periodical swamps behind

Dec. 1962]

LOGANIACEAE (Leenhouts)

the mangrove belt, along rivers, up to 10 m. Fi.
fr. Jan.—Dec.

Uses. Furnishes a good, though cross-grained,
timber; wood very valuable for piling, resists
teredo borers when used with the bark on; further-

more used as firewood. In Bengkalis (Sum.)
planted on yards.
Vern. Cabbage tree, E, bébira. bira_ bira,

bubira, malabira, mélabira, Sum., bérah, bira(h),

malabéra, mal(a)bira, Mal. Pen., kayu bulan,
Born.

Notes. The tree reminds in habit of Terminalia
catappa L.

Young trees have densely spiny trunks (LORZING
in Herb. Bo). These sharp conical prickles form
part of the bark and are apparently corky in na-
ture.

This is the only species of which the leaves are
not entire. Its affinity seems to be with F. blumei.

13. Fagraea truncata Bit. Mus. Bot. 1 (1850)
165; Walp. Ann. 3 (1852) 75; Mia. FI. Ind. Bat. 2
(1857) 372; Ann. Mus. Bot. Lugd.-Bat. 2 (1866)
216; non Koorp. Minah. (1898) 540, Koorp.—
Scuum. Syst. Verz. 3 (1914) 105 (= F. nov. sp.
aff. longiflora)—F. plumeriaeflora (non DC.)
Koorp. Minah. (1898) 540, sphalm. plumeriae-
folia; Koorp.-ScHuo. Syst. Verz. 3 (1914) 105.—
F. forstenii (non BL.) Koorp. Minah. (1898) 540;
Koorp.-Scuum. Syst. Verz. 3 (1914) 105.

Liana, shrub, or tree, up to 12 m high. Leaves
oblong, oblong-obovate or oblanceolate, 20-38
by 8-18 cm, coriaceous, base broadly to narrowly
cuneate, apex broadly cuneate to rounded,
abruptly, up to 114 cm long, blunt- or acute-
acuminate; nerves 7-10 pairs, slightly impressed
above, prominulous beneath; petiole 2-3 cm,
narrowly winged over the whole length, ex-
auriculate; axillary scale adnate, c. 1 cm long,
broadened towards the apex, 2-lobed. Jnflores-
cences glomerulous, 3—7-flowered; pedicels 14-114
cm, with a pair of 1-2 cm long, lanceolate brac-
teoles. Calyx widely campanulate, 2!4-3 cm
long, slightly less than halfway connate. Corolla-
tube widely funnel-shaped, 4 cm. Anthers oblong,
1 cm, split halfway. Stigma obconical-peltate, c.
3 mm @. Fruits ellipsoid, + 314 by 244 cm,
orange; calyx loosely appressed.

Distr. Malaysia: Celebes (Minahasa).

Ecol. Primary forests, 700-1250 m. Fi. July,
fr. Jan., April.

Vern. Matenga, simbélan, sumering.

Notes. Doubtless related to F. fastigiata of
Java and to F. blumei ssp. plumeriaeflora from the
Philippines.

The leaves are exauriculate, but sometimes there
is a pair of gland-like structures on both sides of
the leaf-base.

14. Fagraea tacapala LeEENH. Bull. Jard. Bot.
Brux. 32 (1962) 430—F. fastigiata (non BL.)
CammeRL. Bull. Jard. Bot. Btzg III, 5 (1923) 322,
pro specim. Celeb.—¥ig. 2c.

Treelet (up to 3 m high), tall epiphytic or ter-
restrial shrub, or woody climber. Leaves (sub)

sessile or distinctly petioled, (oblong-)obovate,
rarely elliptic, 20-35 by 7-17 cm, thin-coriaceous,
base (narrowly) cuneate or attenuate, usually
decurrent, auriculate, apex blunt to broadly
cuneate or shortly acuminate; nerves 8-10 pairs,
prominulous on both sides; petiole (0O—)2—5 cm,
axillary scale adnate to the petiole, large (nearly
1-114 cm), much broader than the petiole, dis-
tinctly bilobed. Inflorescences broadly corymbose,
laxly branched, with up to c. 30 flowers; pedicels
14-2 cm, at the base to somewhat less than half-
way provided with a pair of ovate to lanceolate,
(4-)1-1% cm long bracteoles. Calyx cam-
panulate, 1144-214 cm long, confluent at base.
Corolla-tube funnel-shaped, 214 cm. Anthers ob-
long, 14-1 cm long, cells free in their basal half.
Stigma peltate, funnel-shaped or discoid, 214-4
mm @. Fruits subglobular to spindle-shaped, up
to 7 cm long; calyx more or less spreading.

Distr. Malaysia: Celebes, Moluccas (Ceram).

Notes. This species, with its three subspecies,
is more or less intermediate between F. blumei
ssp. plumeriaeflora on one side and F. woodiana
on the other. Especially ssp. gracilis comes in its
leaves very close to F. blumei ssp. plumeriaeflora;
a constant difference between these two species is
the presence of (mostly small) auricles in F.
tacapala, their absence in F. blumei.

The three subspecies, in the sequence gracilis-
ceramensis-tacapala form together a series from
slender to very coarse.

KEY TO THE SUBSPECIES

1. Leaves distinctly petioled. Calyx 114 cm,

slender eee SSD ACIS:

1. Leaves sessile. Calyx 114-214 cm, fairly wide to
wide.

2. Calyx 2'4 cm long, strongly warty-lenticellate.

ssp. tacapala

2. Calyx 1144-114 cm long, smooth or with few

lenticels Ria ssp. ceramensis

ssp. tacapala.—F. fastigiata BL. sensu CAMMERL.
p-p.

Leaves sessile; axillary scale 114 cm long, much
broader than the petiole, deeply 2-lobed. Inflo-
rescence very coarse, densely warty-lenticellate;
pedicels 1 cm, thick; bracteoles lanceolate to ovate,
1-114 cm, usually inserted about halfway the
pedicel, sometimes both or one of them nearly
apical. Calyx 214 cm long, strongly warty-
lenticellate, very broad; anthers 7!4-10 mm;
stigma discoid-peltate.

Distr. SW. Celebes.

Ecol. In primary and secondary forests, along
forest-edges and waysides; on volcanic tuff, at
1000-2000 m alt. F/. Jan., June.

Vern. Ta‘kapala.

ssp. gracilis LEENH. Bull. Jard. Bot. Brux. 32
(1962) 431.

Leaves distinctly petioled; axillary scale some-
what less than 1 cm long, relatively narrow. In-
florescence not very coarse, not warty-lenticellate;

326

FLORA MALESIANA

[ser. I, vol. 6?

pedicels slender, 1-2 cm; bracteoles oblong, c.
¥,-1 cm. Calyx slender, 1144 cm; anthers 5 mm;
stigma funnel-shaped.

Distr. Celebes.

Ecol. Rain-forests, along streams
marshy soil, 500-1200 m. F/. June—July.

Vern. Pulularé.

or on

ssp. ceramensis LEENH. Bull. Jard. Bot. Brux. 32
(1962) 430.

Leaves sessile; axillary scale nearly 114 cm long,
broadly cup-shaped, faintly 2-lobed. Inflorescence
large, fairly coarse, smooth or sparsely warty-
lenticellate; pedicels moderately slender, 1-1 cm;
bracteoles ovate, c. 34-1 cm. Calyx fairly broad,
144-1144 cm long; anthers 6 mm; stigma disk-
shaped.

Distr. Moluccas (Ceram).

Ecol. In forests, 500-800 m. Fi. Jan., May.

15. Fagraea woodiana F. v. M. Austr. J. Pharm.
(Sept. 1886) 323; Bot. Centralbl. 29 (1887) 241;
S. Moore, J. Bot. 61 (1923) Suppl. 35.—F. an-
thocleistifolia G1LG & BENED. Bot. Jahrb. 54 (1916)
192.

Tree, up to 20 m by 41 cm g, shrub, or liana.
Leaves subsessile, oblong-obovate, 25-50 by 10-20
cm, thin-coriaceous, base attenuate and decurrent,
ending in a pair of auricles c. 14-114 cm @, apex
broadly rounded and emarginate to slightly
acuminate; nerves 7-8 pairs, prominulous on both
sides, mainly underneath; axillary scale for the
greater part or entirely adnate to the petiole,
c. 1-144 cm long, very broad and deeply 2-lobed
to truncate. Inflorescences pyramidal-thyrsoid
with c. 3 pairs of horizontal branches, these up to
3 cm long, rebranched or not, at the end with a
more or less dense 5—7-flowered cyme, the total
inflorescence about 20 cm long; pedicels 1-1 cm,
bracteoles inserted at their base or up to halfway,
small and thin. Ca/yx campanulate, | cm, divided
for c. 2. Corolla-tube funnel-shaped, c. 2-3 cm.
Anthers bifid to about halfway, sagittate, 6-714
mm. Stigma peltate, faintly 2-lobed, 214-3 mm o.
Fruits ovoid, c. 2-3 cm long, yellow; pedicel warty-
lenticellate, calyx more or less spreading.

Distr. Malaysia: New Guinea.

Ecol. In primary and secondary forests, either
temporarily inundated or not, from sea-level up
to 1600 m. Fi. Aug., Oct., Dec., fr. Aug., Oct.
The flowers are fragrant like those of jasmine.

Vern. Amiundam, subekwa, wuribéding.

Notes. Apparently related to F. tacapala.

The type is wrongly cited as Fores 741, this
must be 744.

16. Fagraea longiflora Merr. Philip. J. Sc. 3 (1908)
Bot. 260; ELM. Leafl. Philip. Bot. 2 (1909) 597;
Mere. En. Philip. 3 (1923) 315.—F. macgregorii
Me_rr. Philip. J. Sc. 13 (1918) Bot. 51.
Epiphytic shrub. Leaves sessile, broadly to
narrowly obovate, 35-70(-100) by 10-32 cm,
thin-coriaceous, narrowed towards the base, at
the base widened into a pair of rounded, some-
times more or less reflexed wings at both sides of

the axillary scale, apex more or less rounded with
a distinct, small (up to 2 cm long) and narrow,
acute acumen; nerves 9-12 pairs, prominulous on
both sides; axillary scale adnate to the midrib,
114 cm long, acute at the apex, completely hidden
by the leaf-base. Inflorescences glomerulous, with
4—c. 15 subsessile flowers, enveloped by an in-
volucrum of reduced leaves. Calyx campanulate,
214 cm (-6 cm in fruit), slightly more than half-
way divided. Corolla-tube tubular, slender, c.
11 cm. Anthers narrowly ovate, 7 mm long, cells
free to slightly above the middle. Stigma peltate,
14-34 cm @. Fruits subglobular, ovate, or ellipsoid,
3-414 by 234 cm; calyx accrescent, the fruits
moreover enveloped by the long bracts.

Distr. Malaysia: Philippines (Luzon, Ca-
tanduanes); a dubious specimen from Mindanao
(BS 38650).

Ecol. In forests, 100-1400 m. F/. April, Oct.—
Dec., fr. May.

Vern. Sapiag, Klg.

Notes. Probably near to F.blumei ssp. plumeriae-
flora; distinguishable, however, at first sight by the
glomerulous inflorescences and the long corolla-
tube. Possibly also related to F. carstensensis from
New Guinea.

As lectotype I have designated FB 8026 (PNH,
+; dupl. K), the only syntype I have seen.

MERRILL erroneously cited the type number of
F. macgregorii as BS 17938, this should be BS
19738.

17. Fagraea carstensensis WERNH. Trans. Linn.
Soc. Bot. 9 (1916) 111.—Fig. 19.

Small tree (c. 3 m). Leaves sessile, broadly
obovate to obovate-oblong, 17-35 by 5-17 cm,
coriaceous, base subcordate by a broadly rounded,
probably flat wing (apparently not auriculate),
connate when young, apex rounded, shortly
acute-acuminate; nerves 5—12 pairs, conspicuous
though flat above, prominent beneath; axillary
scale fully adnate to the midrib, 114-134 by ec.
1-114 cm, quadrangular; the reduced leaves at the
base of the inflorescence are connate and much
broadened at the base, ++ perfoliate. Inflorescences
sessile and glomerulous, c. 15-flowered; bracts
and (one pair of) bracteoles relatively large (114
and 1 cm long resp.), enclasping the calyx. Calyx
c. campanulate, 2 cm long, fairly slender, sepals
free for c. 34 of their length. Corol//a-tube funnel-
shaped, c. 314 cm long. Anthers elliptic, c. 6 mm
long, thecae free in their basal third. Stigma
obconical-peltate, 3 mm o. Fruit unknown.

Distr. Malaysia: W. New Guinea (Mt Tamrau
and Mt Carstensz), twice collected.

Ecol. Forests, 200-860 m.

Note. Probably nearest related to F. longiflora
and F. woodiana.

18. Fagraea auriculata Jack, Mal. Misc. 2, 7. 7
(1822) 82; WALL. in Roxb. Fl. Ind. 2 (1824)
34 & 573; BL. Bijdr. (1826) 1020; WALL. PI. As.
Rar. 3 (1832) 16, t. 229; Jack in Hook. Comp.
Bot. Mag. 1 (1836) 254; BL. Rumphia 2 (1838)
26, t. 72, excl. syn. Valli-Modagam RHEEDE; DC.

Dec. 1962]

oS

LOGANIACEAE (Leenhouts)

Fig. 19. Fagraea carstensensis WERNH., Tamrau Mts, Vogelkop Peninsula, W. New Guinea (VAN ROYEN
& SLEUMER 7755) (VAN RoyeEN, 1961).

Prod. 9 (1845) 29; Hassk. Flora 28 (1845) 246;
Bi. Mus. Bot. 1 (1850) 165; Mig. Fl. Ind. Bat. 2
(1857) 371; Kurz, For. Fl. Burma 2 (1877) 204
(‘auricularia’); CLARKE in Hook. f. Fl. Br. Ind. 4
(1883) 83; Sorter. in E. & P. Nat. Pfl. Fam. 4,
2 (1892) 42, f. 23; RipL. Trans. Linn. Soc. Bot. 3
(1893) 322; K. & V. Bijdr. 9 (1903) 76; KiNG, J.
As. Soc. Beng. 74, ii (1908) 605; RipL. J. Str. Br.
R. As. Soc. n. 50 (1908) 119; Exo. Leafl. Philip.
Bot. 2 (1909) 599; Koorpb. Exk. Fl. Java 3 (1912)
59; Dop, FI. Gén. I.-C. 4 (1914) 173, f. 21 (5-7);
Burk. J. Str. Br. R. As. Soc. n. 73 (1916) 258;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
326; Merr. En. Philip. 3 (1923) 314; Ripv. FI.
Mal. Pen. 2 (1923) 416; BeumEr, Trop. Natuur
15 (1926) 208, cum fig.; HEYNE, Nutt. Pl. (1927)
1269; RipDL. Disp. (1930) 420; STEEN. & RUTTNER,
Arch. Hydrobiol. Suppl. 11 (1932) 319, f. 16,
20, 21; Burk. Dict. 1 (1935) 993; BAKH. f. in Back.

Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 10;
KerR in Craib, Fl. Siam. En. 3 (1951) 54; Merr.
J. Arn. Arb. 33 (1952) 224.—F. javanica REINW.
ex BL. Cat. (1823) 47, nom. nud.—Willughbeia
auriculata SPRENG. Syst. 4 (1827) Cur. Post. 71.
—F. imperialis Miq. Fl. Ind. Bat. 2 (1857) 372;
Ann. Mus. Bot. Lugd.-Bat. 2 (1866) 216, t. 5 &
6; Burck, Ann. Jard. Bot. Btzg 10 (1891) t. 8
f. 1-2; CAMMERL. Bull. Jard. Bot. Btzg III, 5
(1923) 327.—F. euneura SCHeEFF. in Hassk. Flora
52 (1869) 308; Nat. Tijd. N.I. 31 (1869?) 20;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
328.— F. borneensis ScHEFF. in Hassk. Flora 52
(1869) 309; Nat. Tijd. N.I. 31 (1869?) 21; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 326.—F. nonok
Exo. Leafl. Philip. Bot. 3 (1910) 858; Merr. En.
Philip. 3 (1923) 315.—F. epiphytica Em. Leaf.
Philip. Bot. 8 (1915) 2743.—F. jackii Em. l.c.
2744.—F. curranii MeRR. Philip. J. Sc. 13 (1918)

328

FLORA MALESIANA

[ser. I, vol. 6°

Bot. 50; En. Philip. 3 (1923) 314.—F. fastigiata
(non BL.) CAMMERL. Bull. Jard. Bot. Btzg III, 5
(1923) 322, pro specim. Simalur.—F. bracteosa
CAMMERL. /.c. 323, f. 6-7.—Fig. 2d, 20.

Epiphytic, more rarely terrestrial, shrub or
climber, with age sometimes becoming a small
to medium-sized tree (up to 20 m by c. 25 cm 9).
Twigs often sharply 4-angled, each ridge crowned
by a small acute spine (which in large specimens
may be developed into a small extra auricle).
Leaves mostly distinctly petioled, varying from
narrowly oblanceolate to oblong or obovate,
9-40(—60) by 4-25 cm, usually thickly coriaceous,
base broadly cuneate to narrowly acute, more or
less decurrent, apex rounded to acute, mostly
terminated by a narrowly triangular, acute acu-
men; nerves 5—20 pairs, often inconspicuous to
invisible; petiole robust, often about quadrangular
in cross-section, 24(—8) cm, at the base (faintly
to) distinctly auriculate, auricles about orbicular,
somewhat convex, appressed to the twig; axillary
scale for the greater part adnate to the petiole,
loosely appressed to the twig, !4-34 cm long,
fairly broad, rounded to slightly 2-lobed at the
apex. Flowers solitary or in 2—7-flowered cymes;
pedicels robust, 1-5 cm, quite near the top or
somewhat lower down with 1 or 2 pairs of ap-
pressed, small to fairly large bracteoles. Calyx
campanulate, 2-714 cm long, mostly divided to
near the base, sometimes only slightly more than
halfway down. Corolla thin to very thickly fleshy-
leathery (in the herbarium sometimes nearly
woody), tube widely to narrowly funnel-shaped,
414-15 cm, limb wide-spreading (in the fresh
state the whole corolla may be up to nearly 30 cm
long and more than 30 cm across). Anthers bifid
to about the middle or somewhat lower down,
elliptic to oblong, 34-2 cm, thick. Stigma peltate,
flat or slightly hollowed, 4-8 mm @ (in vivo up
to 2cm). Fruits oblong-ellipsoid to ovoid, crowned
by the persistent style-base, 6-15 cm long, greyish-
white when ripe, dehiscing with 4 lobes; seeds
embedded in an orange to red pulpa (which is
derived from the placentas); calyx somewhat
accrescent, mostly more or less spreading.

Distr. Lower Burma (Sandoway, Tenasserim),
Siam, South Indo-China, and Malaysia: Sumatra,
Malay Peninsula, Java, Bali, Borneo, Philippines,
and Moluccas (Halmaheira and surrounding is-
lands). In older literature also often cited for
India (Malabar), but this is based upon a wrong
interpretation of RHEEDE’s Valli-Modagam.

Ecol. In primary and secondary rain-forests,
often along clearings, river-banks, paths, efc.,
sometimes in mangrove swamps on tidal water-
ways, on coastal rocks, in alang-alang fields and
scrub, from sea-level up to 1500(-1900) m.
Fl. fr. Jan.—Dec. The flower biology of this magni-
ficent and gigantic flower is relatively well known:
it is protandrous, and visited by insects and birds
(see Ripv. J. Str. Br. R. As. Soc. n. 50, 1908, 119,
and Burck, Ann. Jard. Btzg 10, 1891, 97). The
seeds are dispersed by birds and ants (see RIDL.
ic. and Disp. 1930, 421).

Uses. The sticky fluid under the epidermis of

the fruit is used in Sumatra as a glue, especially
as a bird-lime. Forms with large flowers are
cultivated as ornamentals.

Vern. Bira-bira, kayu tulang, kénang (or
kémang) hutan, puleh, sikaso udieng, simbu badak,
téréntang langit, terong térong, Sum., pélir musang,
sagam, Mal. Pen., ki nangka, ki térong (badak),
kontol buta, mangando badak, Java S.; Philip.:
nato, tumakos, Mbo., nonok, P. Bis., piakang, Buk.

Notes. F. auriculata is nearest related to F. in-
volucrata and F. macroscypha, both especially
characterized by the involucrum of large brac-
teoles around the calyx, moreover by the slender,
terete petiole.

The auricles are usually described as being
green, once, however, they were mentioned to be
orange. They provide shelter to ants (see BURCK,
Ann. Jard. Bot. Btzg 10, 1891, 95).

SCHEFFER’S names were apparently published for
the first time in a review by HASSKARL, dated
June 9th, 1869, and published July Ist, 1869, in
Flora. Though SCHEFFER’s paper was dated July
1868, it was not yet published at August 2Ist,
1869, as appears from the annual report of the
“Koninklijke Natuurkundige Vereeniging in Ne-
derlandsch Indié’. So probably HAsskARL had
received either a proof or a pre-issued reprint.

KEY TO THE SUBSPECIES

1. Corolla widely funnel-shaped.

2. Flowers large; calyx 3!4-714 cm, corolla-
tube 5-15 cm. Auricles at the leaf-base well
developed © sous! <0 ssp. auriculata

2. Flowers small; calyx 2-314 cm, corolla-tube
c. 5 cm. Auricles at the leaf-base small or
almost absent . . . ssp. parviflora

1. Corolla slenderly funnel-shaped.

ssp. borneensis

ssp. auriculata.—F. auriculata JACK.—F. javanica
Bi.—Willughbeia auriculata SPRENG.—F. imperia-
lis Mig.—F. epiphytica ELM.

Auricles well developed, up to 2 by 2'4 cm;
leaves shortly petioled to subsessile. Inflorescences
1—3(—6)-flowered. Calyx 314-7!4 cm. Corolla-
tube 5-15 cm long, widely funnel-shaped. Fruits
8-15 cm long; calyx-lobes often appressed.

Distr. Burma, Siam, South Indo-China, and
Malaysia: Sumatra, Malay Peninsula, Java, Bali,
Borneo, and SE. Philippines (Panay, Mindanao).

ssp. borneensis (SCHEFF.) LEENH., nov. Sstat.—F.
borneensis SCHEFF.—F. nonok EtM.—F. jackii
E_mM.—F. bracteosa CAMMERL.

Auricles usually well developed, though not
very large (34-114 cm @); leaves mostly distinctly
petioled. Inflorescences 3—7-flowered. Calyx 244—
314 cm. Corolla-tube 8-9 cm long, narrowly
funnel-shaped (basal half narrowly tubular).
Fruits 6—7 cm long; calyx-lobes spreading.

Distr. Malaysia: Northern half of Borneo and
W. Philippines (Palawan, Mindoro, Sibuyan).

ssp. parviflora LEENH., mov. -ssp—F. euneura
ScHEFF.—F. curranii MERR.

Dec. 1962]

LOGANIACEAE (Leenhouts)

329

Fig. 20. Fagraea auriculata Jack; the giant flower
of ‘F. imperialis’, cult. Hort. Bog.
(Hj. JENSEN).

Auricles small or almost absent (up to 34 cm @);
leaves mostly distinctly petioled. Inflorescences
(1—)3—S-flowered. Calyx 2-314 cm. Corolla-tube
414-514 cm long, widely funnel-shaped. Fruits
6-7 cm long; calyx-lobes spreading.

Distr. Malaysia: Philippines (Luzon) and
Moluccas (Halmaheira and adjacent islands).

Note. Ssp. parviflora has been based upon
F. curranii MErr.

19. Fagraea involucrata Merr. J. Str. Br. R. As.
Soc. n. 77 (1917) 233; non var. longipetiolata MERR.
Pl. Elm. Born. (1929) 251 (=F.? macroscypha).—
F. macroscypha (non BAKER) HEINE, Pfl. Clemens
Kinabalu (1953) 91.—F. wniflora (non MERR.)
Heine, /.c. 92.—Fig. 21a—d, 22.

Epiphytic shrub or liana which later can become
a fairly slender tree. Leaves petioled, oblong to
lanceolate, (9-)12—25 by 4-8 cm, thickly coriace-
ous, base cuneate or slightly contracted, acute,
apex rather abruptly and shortly acute acuminate;
nerves c. 15 pairs, on either side nearly always in-
conspicuous to invisible, or slightly prominent on
the lower surface; petiole 114-414 cm, mostly
terete, auriculate; axillary scale largely free from

the petiole, 14-1 cm long, rounded. Flowers soli-
tary; pedicel very robust, 3¥4-214 cm, thickened
upwards, bearing in its upper part (2—)3 closely
approximate, decussate pairs of appressed, co-
riaceous, ovate, 2-6 cm long bracteoles which
enclose the lower part of the calyx (inner brac-
teoles distinctly larger than outer ones). Calyx
7-9 cm long, divided to quite near its base, seg-
ments oblong, rounded, leathery, very concave.
Corolla inside green with 5 broad cream streaks,
tube at base tubular, abruptly widened in its upper
third part, 14-15 cm, the upper part inside rather
densely tomentose. Anthers oblong, very thick,
114-134 cm, cells free to distinctly below their
middle. Stigma peltate-cupshaped, 34 cm o.
Fruits ovoid, about 414 by 4 cm; calyx spreading.

Distr. Malaysia: Borneo.

Ecol. In forests, on river-banks and along paths
up to 1800 m. Fi. fr. probably Jan.—Dec.

Vern. Kaju ara jangkit.

Note. Closely related to F. macroscypha (see
there) and to F. auriculata, especially its ssp.
borneensis, different from the latter by the solitary
flowers with much larger calyx, by the greater
number of exceptionally large bracteoles, and by
the abruptly widened corolla-tube.

5

of
J

y Bx

Fig. 21. Fagraea involucrata MERR. a. Habit, 1.
b. flower-bud, < 14, c. apical part of style and
stigma, * 14, d. young fruit, calyx partly removed,
<14.—F. macroscypha BAKER. e. Calyx with
involucrum, V4. (a SF 26733, b ENperT 4613,
c CLEMENS 30072, d SAN 16380, e CREAGH S.n.).

330

Fig. 22.

20. Fagraea macroscypha BAKER, Kew Bull. (1896)
25.—? F. involucrata MeERR. yar. longipetiolata
Merr. Pl. Elm. Born. (1929) 251.—Fig. 21e.
Epiphytic shrub, climber, or finally a tree.
Leaves petioled, oblong-lanceolate, 714-27 by
314-614 cm, thick coriaceous, base acute, apex
obtuse or rounded, tapering into a long and narrow
acute acumen; nerves 10-15 pairs, inconspicuous
above, prominulous beneath; petiole 2-3 cm,
rather robust, terete towards the base, auriculate;
axillary scale nearly free, loosely appressed to the
twig, 3 mm high, rounded. Flowers solitary, pedicel
very thick, 34-1 cm, at the apex with 2 decussate
pairs of large (2144-3 cm), appressed bracteoles,
these together embracing the base of the calyx.
Calyx campanulate, 614-9 cm long, divided to
near the base, lobes ovate-lanceolate, tapering

Fagraea involucrata MeERR., Bundu Tukan, North Borneo (W. Meyer, 1960).

into a long and slender acumen. Corolla white
to cream with a broad green medium stripe, tube
12-14 cm, tubular, slightly widened upwards.
Anthers divided to below the middle. Stigma
peltate, slightly hollowed, 5 mm or more @.
Fruits reversed pear-shaped or acutely obovoid,
c. 7 by 4cm.

Distr. Malaysia: Borneo.

Ecol. In logged area, up to 200 m. Fi. March,

Sept.
Vern. Kayu ara jangkit.
Notes. Closely related to F. involucrata,

mainly different by the long and acute sepals and
by the nerves which are prominent on the lower
surface of the leaves.

As far as I can judge from sterile specimens,
F. involucrata yar. longipetiolata comes closer

Dec. 1962]

LOGANIACEAE (Leenhouts)

331

to this species than to any other one. MERRILL,
in describing this variety, already had some
doubt about its identity.

21. Fagraea resinosa LEENH. Bull. Jard. Bot.
Brux. 32 (1962) 429.

Leaves distinctly petioled, oblong to lanceolate,
10-16 by 4-7 cm, stiffly coriaceous, base acute,
apex blunt, abruptly terminated by a short, nar-
row, and acute point; nerves c. 15-20 pairs, in-
conspicuous above, invisible beneath; petiole
214-4 cm, terete or slightly flattened above,
exauriculate, axillary scale partly adnate to the
petiole, 4-5 mm, rounded. Flowers solitary, known
only in bud, pedicel relatively slender, 1-11 cm,
at the apex with 2 pairs of bracteoles, all broadly
rounded and embracing the basal part of the calyx,
the upper 214 cm, the lower 114 cm long. Calyx
campanulate, 314 cm, deeply divided, lobes
broadly rounded. Anthers broadly oblong, 6 mm
long, slightly bifid at the base only. Stigma dish-
shaped, 214 mm @ (or more ?). Fruits ovoid,
414 by 3 cm; calyx appressed.

Distr. Malaysia: Borneo (Brunei: Bt Belalong;
W. Borneo: Mt Amai Ambit).

Ecol. Forests, c. 700 m. Fr. April.

Notes. Related to, but distinctly different from,
F. macroscypha and F. involucrata.

In its vegetative parts resembling F. carnosa
which differs by its smaller, slender calyx not
enveloped by large bracteoles, and by the smaller
number of nerves (c. 6 pairs).

22. Fagraea carnosa JACK, Malay. Misc. 2, 1. 7
(1822) 81; in Hook. Comp. Bot. Mag. 1 (1836)
254; DC. Prod. 9 (1845) 30; Mia. FI. Ind. Bat. 2
(1857) 374; Kurz, Fl. Burm. 2 (1877) 204;
CLARKE in Hook. f. FI. Br. Ind. 4 (1883) 82;
Kina, J. As. Soc. Beng. 74, ii (1908) 604; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 323; RuipL.
Fl. Mal. Pen. 2 (1923) 416; HeNperRs. J. Mal.
Br. R. As. Soc. 17 (1939) 58.—F. monantha Mia.
Fl. Ind. Bat. 2 (1857) 373; Ann. Mus. Bot. Lugd.-
Bat. 2 (1866) 218; ScHeEFF. Nat. Tijd. N.I. 31
(1870) 21.—F. rotundifolia Rip. J. Str. Br. R.
As. Soc. n. 50 (1908) 117; Fl. Mal. Pen. 2 (1923)
416.—F. uniflora Merr. J. Str. Br. R. As. Soc.
n. 77 (1917) 235; CAMMERL. Bull. Jard. Bot. Btzg
ILI, 5 (1923) 337.—F. flavidula Rw. Fl. Mal. Pen.
§ (1925) 322.—F. lanceolata (non BL.) HENDERS.
Gard. Bull. S.S. 4 (1927) 99.

Epiphytic (rarely terrestrial) shrub or climber.
Leaves petioled, (suborbicular to) elliptic to
lanceolate or oblong-obovate, 712-16 by 314-814
cm, thickly and rigidly coriaceous, base cuneate,
sometimes shortly contracted, apex acute to round-
ed or even slightly emarginate, and in both latter
cases shortly acutely acuminate; nerves 6—7 pairs,
at both surfaces inconspicuous or almost so, some-
times slightly sunken above; petiole 34-3 cm,
exauriculate; axillary scale halfway up to nearly
completely adnate to the petiole, 14-34 cm long,
obtuse or rounded, not very conspicuous. Flowers
1(—2), pedicel very thick, 14-2 cm, bracteoles 1 or
2 pairs, apical, appressed, 14-1 cm long. Calyx

slender, (1!4—)2—5 cm long, divided + halfway
down or slightly less. Corolla-tube rather slender,
tubular, 7-14 cm. Anthers oblong, 1-134 cm, cells
free to somewhat below the middle. Stigma
capitate to peltate-cupular, 4-5 mm 9. Fruits
(oblong) ellipsoid, 4-7 cm, greyish blue; calyx
appressed.

Distr. Lower Burma (Moulmein) and Malaysia:
Sumatra, Malay Peninsula, and Borneo.

Ecol. Forests on limestone, also along the
beach, from sea-level up to 2000 m. F/. mainly
Jan., fr. Jan., Aug., Nov.

Vern. Andor hapal, Sum., numpang, Born.

Notes. F. uniflora was apparently based upon
an anomalous specimen, with 6 calyx-lobes and
8-9 corolla-lobes; I have not seen the type speci-
men.

This species is best characterized by the slender
though rather large, spindle-shaped calyx, further-
more by the very stiff, usually minutely pointed
leaves and the smooth, greyish-yellow twigs.

JACK’s type specimen is apparently lost. Among
the few Sumatran specimens at my disposal none
fits his original description very well, and there is
no later collection from the type-locality, cited
as Bencoolen, S. Sumatra. The only specimen
which is well in accordance with the original
diagnosis is RosTabo s.n. from Tringganu, Bundi,
Malaya, collected Febr. 1904 (K), the type of
F. rotundifolia Riv.

23. Fagraea gardenioides Ript. J. Fed. Mal. St.
Mus. 5 (1914) 42; Fl. Mal. Pen. 2 (1923) 416.

Epiphytic shrub or possibly sometimes a treelet.
Leaves petioled, (oblong or elliptic to) obovate,
614-13 by 4-614 cm, coriaceous (in vivo probably
thick fleshy), base cuneate, attenuate, apex round-
ed or blunt, without or with a short acumen;
nerves c. 7-8 pairs, mostly invisible, sometimes
minutely grooved beneath; petiole 1-2 cm,
exauriculate; axillary scale partly adnate to the
petiole, 14-34 cm long, about square, truncate to
2-lobed, distinctly broader than the petiolar base.
Inflorescences terminal; flowers 1-5 in a sessile
cyme; pedicel 14-2 cm; bracteoles appressed to the
calyx, ovate to roundish, 14-34 cm long. Calyx
campanulate, 114-3 cm long, connate for 13-7.
Corolla-tube tubular, 6—7 cm. Anthers linear (-lan-
ceolate), 9 mm, up to c. 4 bifid at the base.
Stigma subpeltate, sometimes slightly 2-lobed,
3-5 mm @. Fruits (young) ovoid to ellipsoid, 4 cm
or more long, tapering into a conical beak; calyx
loosely appressed.

Distr. Malaysia: Malay Peninsula, Borneo.

Note. On the one side closely related to F.
curtisii, which differs by the short corolla-tube
and the longer and relatively narrower, acute
leaves. On the other side especially ssp. borneensis
shows some relationship to F. berteriana and
allies.

ssp. gardenioides.

Rather coarse. Petioles narrowly winged; axillary
scale 34 cm long; blade up to 13 by 7'4 cm, not
acuminate. Flowers up to 3; pedicel up to 2 cm,

332

FLORA MALESIANA

[ser. I, vol. 6?

thick. Calyx 214-3 cm, less than halfway up con-
nate. Corolla-tube relatively wide. Stigma ob-
conical with a membranous margin, c. 5 mm g.

Distr. Malay Peninsula.

Ecol. On hills, c. 1400-1900 m. Fi. Jan., April,
July, Aug.
ssp. borneensis LEENH. Bull. Jard. Bot. Brux.
32 (1962) 425.

Rather slender. Petioles not winged; axillary
scale 144-14 cm long; blade 7-10 by 4-614 cm,
fairly abruptly, shortly and broadly acute-acu-
minate. Flowers up to 5; pedicel up to 34 cm,
rather slender. Calyx 1144 cm, connate for 24.
Corolla-tube slender. Stigma cupular-peltate,
slightly 2-lobed, c. 3 mm @.

Distr. Borneo (Sarawak: near Kuching).

Ecol Fi. Dec:

24. Fagraea curtisii K. & G. J. As. Soc. Beng. 74,
ii (1908) 605; RipL. Fl. Mal. Pen. 2 (1923) 417;
Kerr in Craib, Fl. Siam. En. 3 (1951) 55.

Low spreading tree or (sometimes epiphytic)
shrub. Leaves petioled, elliptic, ovate-elliptic, or
oblong-obovate, 10-21 by 5-814 cm, firmly co-
riaceous, base cuneate or contracted, apex shortly
and often abruptly triangular-acuminate; nerves
7-10 pairs, invisible on both surfaces; petiole
1-114 cm, robust; exauriculate; axillary scale for
the greater part adnate to the petiole, 14-34 cm
long, narrowed towards the truncate to slightly 2-
lobed apex. Flowers in (1—)2-4-florous cymes; pe-
dicels stout, 1144-214 cm; 1-2 pairs of bracteoles
near the apex, appressed, broadly ovate, 14-1 cm
long, caducous. Calyx campanulate, 134-314 cm
long, divided + 144 to nearly halfway down.
Corolla-tube funnel-shaped, c. 414 cm. Anthers
oblong, thick, + 8 mm, bifid to about the middle.
Stigma peltate, c. 24% mm @. Fruits subglobose or
broadly (ob)ovoid, 334-5 cm long; calyx funnel-
shaped, loosely appressed.

Distr. North Burma (Kachin State) and Ma-
laysia: Malay Peninsula (Puket, Kedah, Kelantan,
Langkawi I.).

Ecol. From sea-level up to 1300 m (in Burma),
on limestone. F/. Aug.—Sept., fr. Febr.

Note. On the one side closely related to F.
gardenioides, on the other to F. calcarea. The
former differs by its long tubular corolla, the
other by its large bracteoles.

25. Fagraea calearea HENpDeRS. Gard. Bull. S.S.
710933) PUTS ite 28: Bs

Climbing shrub. Leaves petioled, elliptic to
broadly ovate or obovate, 12-17'4 by 6-9 cm,
thinly coriaceous, base cuneate, somewhat de-
current, apex rounded, without or with an abrupt
very short, triangular acumen; nerves 8-10 pairs,
very faint on both surfaces; petiole 114-3 cm,
fairly slender, exauriculate; axillary scale adnate
to the petiole, 44-1 cm long, truncate, not very
conspicuous. Flowers 1(—3); pedicels stout, 44-114
cm; bracteoles 2 pairs, the outer ovate or ovate-
lanceolate, acute, up to 3 cm long, the inner
ovate, rounded, 2 cm long, all embracing the basal

part of the calyx. Calyx ovoid, 314-4 cm long,
divided for about 24. Corolla-tube funnel-shaped,
c. 5 cm. Anthers oblong, c. 1 cm. Stigma broadly
funnel-shaped. Fruits unknown.

Distr. Malaysia: Malay Peninsula (Pahang:
B. Chintamani near Bentong), once collected.

Ecol. Climbing on a limestone rock face, at
low altitude. F/. Oct.

Note. Evidently allied to F. curtisii but differing
in the thinner, wider and more rounded leaves, the
flowers of the single specimen known being mostly
solitary (in a few inflorescences in threes), and
somewhat larger, and in the much larger and more
deeply cut calyx. Possibly also related to F. in-
volucrata.

26. Fagraea gracilipes A. Gray, Proc. Am. Ac.
Arts Sc. 4 (1859) 323; SEEMANN, FI. Vit. (1866)
165; A. C. Smitn, J. Arn. Arb. 33 (1952) 113;
LeenH. Bull. Jard. Bot. Brux. 32 (1962) 426.—
Gardneria fagraeacea F. v. M. Fragm. 6 (1868)
130.—F. muelleri Bru. Fl. Austr. 4 (1869) 368,
nom. illeg.; F. M. Batt. Queensl. Fl. 3 (1900)
1023.—F. dolichopoda GiLG & BENED. Bot. Jahrb.
54 (1916) 196.—F. fagraeacea Druce, Rep.
Exch. Cl. Brit. Is. 1916 (1917) 623.—F. amabilis
S. Moore, J. Bot. 61 (1923) Suppl. 36.
—F. cambagei Domtn, Bibl. Bot. 89 (1929) 1071;
Mere. & Perry, J. Arn. Arb. 23 (1942) 413.—
F. elata Merr. & Perry, /.c.—F. obtusifolia
Merr. & PERRY, /.c. 415; F. S. WALKER, For. Br.
Solomon Is. (1948) 137.—Fagraea sp. KAN. &
Hatus. Bot. Mag. Tokyo 56 (1942) 163.

Small to medium tree, usually up to 15 m, rarely
up to 40 m by 45 cm 9; sometimes a scrambling
shrub. Leaves petioled, broadly elliptic or elliptic
obovate to oblong, (6—)9-17(-23) by (214-)
414-814(-10) cm, coriaceous, base narrowly acute
to broadly cuneate, nearly always attenuate, apex
blunt, usually terminated by a short, broad, and
blunt acumen; midrib grooved above, towards the
base sometimes flattish; nerves 5-8(—9) pairs,
minutely grooved above, grooved to prominulous
beneath, usually inconspicuous or sometimes in-
visible; petiole 14 cm, mostly slender, exauricul-
ate; axillary scale more than halfway up to nearly
completely adnate to the petiole, but appressed to
the twig, not conspicuous, 2'4—5 mm long, blunt
to rounded. Inflorescences cymose, few- to many-
flowered, dense to lax, always relatively small, the
branches often more or less drooping, pedicels
sometimes very slender, 14-3 cm, with 1(—3) pairs
of small bracteoles, inserted from about halfway
to just below the calyx. Calyx campanulate, c.
6-7 mm(—1 1! cm), divided about halfway or some-
what more. Corolla-tube funnel-shaped (basal
half mostly tubular), slender, 214-5 cm long.
Anthers bifid to c. 4-14 from the base, oblong
(to elliptic), 3-5 mm. Stigma small, subobconical,
slightly 2-lobed or entire. Fruits broadly ellipsoid-
obovoid to -ovoid, abruptly terminated by a
strong long-conical beak, the whole fruit 3-4
cm long, orange or red; pedicels apparently some-
what lengthened, calyx appressed, slightly ac-
crescent.

Dec. 1962]

LOGANIACEAE (Leenhouts)

S88

Distr. Fiji, Santa Cruz Is., Solomon Is. (Santa
Isabel, New Georgia), Australia (NE. Queensland)
in Malaysia: New Guinea (also Aru Is.). Cf.
LEENHOuTS, Pac. Pl. Areas map 91.

Ecol. In and along the edges of rain-forests,
as well on a dry as on a swampy or temporarily
inundated soil, furthermore in secondary forests,
in palm-swamps and in mossy oak forests, 0-1530
(-2000) m. Fi. Jan.—Dec., fr. April, June—Nov.

Uses. A hard, heavy timber, very durable,
esteemed by the natives, who use it for making
combs, houseposts, canoes, efc., also suitable for
survey-marks, heavy construction, and turnery.
In Fiji tea made from the bark and leaves is used
medicinally. The fruits are sometimes pickled and
eaten.

Vern. Mulgrave plum, E, besron, boggong,
ibelaka, iedéwob, i(eg)bérakka, mofruka, nasam,
ngorarien, nietsamber, numae, New Guinea.

Notes. A variable species; the Papuan speci-
mens are rather uniform and well characterized by
the brown to blackish twigs and the thick, fleshy-
coriaceous, broadly elliptic to obovate leaves,
which are distinctly attenuate at the base and
obtuse or rounded, but always shortly and bluntly
acuminate at the apex, and with the nerves deli-
cately grooved on both surfaces. Some small-
leaved and small-flowered specimens remind of the
Australian form known as F. muelleri; a few spe-
cimens, with large oblong leaves, come close to
the other Australian form which was described
under the name of F. cambagei.

The relationships are not fully clear to me. On
the one side it reminds of F. berteriana, on the
other hand it has several characters in common
with F. umbelliflora.

The name F. muelleri is illegitimate, as it had
been based upon Gardneria fagraeacea F. v. M.,
and the epithet of the latter should have been
used.

27. Fagraea eymae BACKER in Leenh. Bull. Jard.
Bot. Brux. 32 (1962) 425.—Fig. 2a.

Shrub. Leayes subsessile to shortly petioled,
oblong to elliptic, c. 9-10 by 414-514 cm, firmly
coriaceous, base rounded or subcordate, faintly
auriculate, margin recurved, apex obtuse or mi-
nutely obtusely pointed; nerves 5—6 pairs, on the
upper surface faintly visible, beneath incon-
spicuous or almost so; axillary scale for the greater
part adnate to the petiole, 1,—34 cm long, about
quadrangular to slightly bilobed, boat-shaped,
nearly completely hidden by the leaf-base. Jn-
florescences usually 2-flowered, pedicels thick, 1-2
cm long; bracteoles near the apex, ovate to oblong,
%-14% cm long. Calyx campanulate, 2-244 cm
long, divided somewhat over halfway down.
Corolla-tube distinctly funnel-shaped, 414 cm,
lobes exceptionally short, rounded. Anthers ob-
long, thick, 8-11 mm long, cells free in their basal
half. Stigma + cupular-peltate, 214 mm @. Fruits
globular, 214-3 cm 9; calyx appressed.

Distr. Malaysia: West New Guinea (Wissel
Lake area).

Ecol. In permanently inundated, secondary

forest on peat at c. 1750 m. Fi. Jan.

Vern. Jpu or iepo.

Note. Apparently related to F. sa/ticola and to
F. monticola.

28. Fagraea salticola LEENH. Bull. Jard. Bot.
Brux. 32 (1962) 429.—Fig. 23.

Tree, up to c. 25 m high by 35 cm g. Leaves
shortly petioled, elliptic to slightly obovate, 4-11
by 214-7 cm, somewhat convex, very thickly and
stiffly coriaceous (fleshy when fresh), the base
slightly cordate to rounded, rarely broadly cu-
neate, apex rounded to blunt; nerves c. 5 pairs,
nearly invisible, sometimes slightly sunken above;
petiole 14-1 cm, thick, exauriculate; axillary scale
adnate to the petiole, distinctly broader than this
and convex, 4-7 mm long, widened towards the
2-lobed apex. Inflorescences cymose, rather dense,
with 3-10 flowers; pedicels 14 cm long, thick,
tapering towards the calyx, either without or with
a pair of bracteoles appressed to the calyx. Calyx
ellipsoid-campanulate, 114-114 cm long, divided
about halfway up. Corolla-tube funnel-shaped,
214 cm long. Stamens inserted on a thickened ring
in the inside of the corolla; anthers attached just
above the base, lanceolate, c. 7 mm. Stigma
bilobed, 3-4 mm go. Fruits ellipsoid, acute, 314
by 114 cm, orange; calyx appressed.

Distr. Malaysia: East New Guinea (Western
and Eastern Highlands).

Ecol. In mossy and Podocarpus-Libocedrus
forests at 2500-2850 m. Fi. fr. July—Aug.

Uses. Wood previously used by natives for
making spears.

Vern. Banda, mama, mband, tihl.

Notes. A very characteristic species, which
only shows a distinct resemblance to F. eymae as
to the vegetative parts. Apart from F. annulata,
F. salticola, F. bodenii, and F. berteriana — with
which it is connected by the annulus inside the
corolla and as to the latter three also by the only
very slightly bifid anthers and the bilobed stigma —
it may be related to F. monticola GiLG & BENED.
(1916), and it would even be possible that it is
identical with F. melanochlora GitG & BENED.
(1916), from which, however, it differs in some
points.

29. Fagraea bodenii WeERNH. Trans. Linn. Soc.
Bot. 9 (July 1916) 111; Merr. & Perry, J. Arn.
Arb. 23 (1942) 414.—F. jasminodora GILG &
BeNeED. Bot. Jahrb. 54 (Oct. 1916) 190.—F. ampla
S. Moore, J. Bot. 61 (1923) Suppl. 35.—F. sua-
veolens CAMMERL. Nova Guinea 14 (1924) 117,
t. 13.—F. papuana MERR. & PERRY, J. Arn. Arb.
23 (1942) 414.

Tree, up to 30 m by 50 cm, or erect shrub, some-
times epiphytic. Leaves petioled, elliptic- to oblong-
obovate to oblong, 5-15 by 214-7'4 cm, rigidly
coriaceous, base acute or shortly contracted, apex
obtuse or rounded, often terminated by a short
and broad, obtuse and often recurved point;
nerves c. 6-10 pairs, not or hardly visible on the
upper surface, very faintly visible or obsolete
beneath; petiole firm to slender, 34-3 cm long,

334

FLORA MALESIANA

[ser. I, vol. 6?

Fig. 23. Fagraea salticola LEENH. a. Habit with flowers, =<

d. stigma, = 3, e. twig with fruits,

< 24, f. leaf-bases, showing the axillary scale, =

x 5c. pistols
1 (a-f HOOGLAND

24, b. opened coroila,

& PULLEN 5623).

exauriculate; axillary scale adnate to the petiole,
usually distinctly broader than this, 143-1 cm
long, rounded or obtuse. /nflorescences cymose,
mostly short and dense, 2—15-flowered; pedicels
thick, 34-344 cm; bracteoles halfway. Calyx
narrowly campanulate, 34-114 cm long, divided
slightly less to somewhat more than halfway down.
Corolla deep-olive outside, cream-coloured to
white inside, tube tubular or narrowly obconical,
114-4 cm. Stamens inserted on a thickened ring
in the tube, anthers linear-lanceolate, 34-114 cm,
slightly bifid at base. Stigma distinctly 2-lobed,
2-3 mm o. Fruits broadly ellipsoid-subglobose,
contracted into a long conical, robust beak, c.
4 by 2'4 cm, orange; calyx appressed.

Distr. Malaysia: New Guinea.

Ecol. Forests and shrubberies on slopes, on
limestone hills, etc., 80-2840 m. Fi. Oct.—Febr.,
May, Aug., fr. Oct.—Febr.

Notes. Distinctly related to F. gitingensis and
in less degree to F. annulata.

Specially characterized by the conspicuous,
boat-shaped stipules and the strongly beaked
fruit on a short, stiff pedicel. F. gracilipes has
also beaked fruits, but its pedicels are usually

distinctly longer and more slender, even drooping;
F. salticola has the same kind of stipules, but they
are less conspicuous.

30. Fagraea gitingensis ELM. Leafl. Philip. Bot. 3
(1910) 859; Merr. En. Philip. 3 (1923) 315.—
F. ternatana (non Mig.) HoLttH. & LAM, Blumea
5 (1942) 230.

Gnarled tree or subarboreous shrub, 3-10 m,
trunk up to 25 cm g. Leaves petioled, (obovate-)
oblong to (obovate-)lanceolate, 744-12 by 214-4
cm, thickly coriaceous, base tapering, decurrent,
apex very obtuse or rounded; nerves 5-12 pairs,
faintly visible above, subinconspicuous beneath;
petiole 114-3 cm, rather slender, narrowly winged
towards the lamina; exauriculate; axillary scale
adnate to the petiole, 1-14 cm long, truncate or
slightly emarginate. Inflorescences 3—9-flowered
cymes, pedicels not very thick, 4-144 cm;
bracteoles about halfway. Ca/yx narrowly cam-
panulate, 1-2 cm long, divided + halfway down.
Corolla-tube long-cylindric, narrow, slightly wid-
ened in the upper part, 7-11 cm. Anthers narrow-
ly linear-lanceolate with a narrowed apex, c. 1 cm,
slightly bifid at base. Stigma 2-lobed with oblong

Dec. 1962]

LOGANIACEAE (Leenhouts)

335

lobes, 4-5 mm 9. Fruits oblong-ellipsoid, acute,
314-4 by 134 cm, shining orange-red; calyx ap-
pressed.

Distr. Malaysia: Philippines (Sibuyan: Mt
Giting-Giting) and N. Moluccas (Talaud I.:
G. Piapi), twice collected.

Ecol. Woods on very sandy, gravelly soil,
open sunny slopes, 300-400 m. Fi. April, fr. May.

Vern. Ténggeh, Talaud.

Note. Distinctly related to F. bodenii and F.
salticola, both from New Guinea.

31. Fagraea berteriana A. GRAY ex BTH. J. Linn.
Soc. Bot. 1 (1856) 98; SEEM. FI. Vit. (1866) 164;
F. B. H. Brown, Bull. Bish. Mus. 130 (1935)
228, f. 34; YUNCKER, Bull. Bish. Mus. 220 (1959)
217; LeeNH. Bull. Jard. Bot. Brux. 32 (1962)
419.—Carissa grandis BERT. ex GUILLEMIN, Ann.
Sc. Nat. Bot. II, 7 (1837) 248, nom. nud.—F.
grandis PANCH. & SEBERT in Sébert, Not. Bois
Nouv. Cal. (1874) 184, nom. superfl—F. obovata
WALL. var. papuana F. M. Bait. Queensl. Agr. J.
3 (1898) 157.—F. peekelii GitG & BENED. Bot.
Jahrb. 54 (1916) 185, f. 10.—F. pachypoda GiLG
& BenepD. /.c. 186.—F. calophylloides GILG &
BENeD. /.c. 188, f. 11.—F. schlechteri GILG &
BENeED. ibid. 56 (1921) 550, nom. superfl.; SARLIN,
Bois et Foréts de la Nouv. Caléd. (1954) 264,
t. 127.—F. ksid GitG & BENED. Bot. Jahrb. 56
(1921) 552; KaNeH. FI. Micron. (1933) 317,
f. 161.—F. salomonensis GitG & BENED. Bot.
Jahrb. 56 (1921) 554.—F. samoensis GILG &
BENED. /.c. 551.—F. vitiensis GILG & BENED. /.c.
553, non SeeM. (1861), nom. nud.—F. galilai
Gitc & Benep. /.c. 555; KANEH. FI. Micron.
(1933) 316, f. 160.—F. sair GiLtG & BENED. Bot.
Jahrb. 56 (1921) 555, f. 3; KANEH. FI. Micron.
(1933) 318, f. 162.—F. affinis S. Moore, J. Bot.
61 (1923) Suppl. 36.—F. novae-guineae CAMMERL.
Nova Guinea 14 (1924) 117, t. 14.—F. pluvialis
S. Moore, J. Bot. 67 (1929) 49.—F. kusaiana
Hosokawa, Trans. Nat. Hist. Soc. Form. 24 (1934)
202.—F. rosenstromii C. T. Wuite, Proc. R. Soc.
Queensl. 47 (1936) 71.

Usually a small, often shrub-like tree, up to 15
(-30) m by 50 cm g, sometimes an epiphytic or
terrestrial, erect or scrambling shrub. Leaves
petioled, elliptic or oblong, sometimes slightly
ovate or obovate, 9-16 by 414-12 cm, fleshy-
coriaceous, base acute, attenuate, apex rounded;
nerves c. 10 pairs (and between every pair an in-
termediate vein which is slightly less conspicuous),
faintly visible on the upper surface, almost or
quite invisible beneath; petiole 114-4 cm, usually
slender, exauriculate; axillary scale partly adnate
to the petiole, 8-9 mm long, boat-shaped, 2-lobed
at the apex. Inflorescences cymose, often widely
branched, few- to rather many-flowered, pedicels
Y%-1'4 cm, thick, especially towards the calyx,
ebracteolate. Ca/yx rather narrowly campanulate,
144-244 cm long, usually divided for c. 14,
rarely up to halfway down. Corolla-tube tubular,
3-8 cm long when dry (up to 15 cm when alive).
Stamens inserted on a thickened ring in the tube,
anthers narrowly linear with an attenuate apex,

1-114 cm, slightly bifid at their base. Stigma
distinctly 2-lobed (the halves in bud, however,
folded upwards and appressed), lobes oblong, c.
¥%,-1 cm og. Fruits ellipsoid to globular, orange to
red when ripe, 3-514 by 214-4'4 cm; calyx-lobes
spreading.

Distr. SE. Polynesia (Marquesas: Nukuhiva,
Hiva Oa; Tubuai Is.: Tubuai, Rapa; Society Is.:

Tahiti, Tahaa; Cook Is.: Rarotonga), SW.
Polynesia (Tonga Is.: Niue, Tonga; Samoa),
Melanesia (Fiji, New Caledonia, Loyalty Is.;

New Hebrides: Vanikoro, Oba, Aneytum; So-
lomon Is.), Australia (NE. Queensland: Moss-
man R.), Micronesia (Carolines: Palau Is., Truk,
Ponape, Kusaie; Marianas: Guam, Rota), and
Malaysia: New Guinea (also in the Bismarcks
and Louisiades). Cf. LEENHOUTS, Pac. Pl. Areas
map 91.

Ecol. In open to dense, wet to dry, primary and
sometimes secondary forests, preferably along the
edges, along river-banks, creeks, efc., also in
open country, on rocks, even along the beach,
also in the mangrove; from sea-level up to 500
(-1600) m. Fl. Jan.—Dec., mainly July—Febr.,
fr. mainly Aug. The fruits are eaten by birds, main-
ly pigeons (cf. BRown, /.c.); for possible occasional
dispersal by water see RIDLEY, Disp. (1930) 219.

Uses. The timber is hard, and locally used for
house posts, other constructions, and for tools.
The very fragrant flowers are in many Pacific is-
lands highly appreciated for decoration (crowns,
leis) and for making perfumes. In New Caledo-
nia the boiled leaves are used as a medicine against
rash and skin irritation. The natives of the So-
lomon Is. use the fruit as a flytrap by first removing
the epicarp to expose the sticky interior.

Vern. Argook, kaunomori, New Britain.

Note. This species belongs to the group with
a 2-lobed stigma, linear anthers, and a thickened
ring inside the corolla-tube on which the stamens
are inserted; through these characters it is related
to F. gitingensis, F. bodenii, F. salticola, and in a
less degree to F. annulata.

Doubtful

Fagraea alteniana F. v. M. Vict. Nat. 6 (1889) 45,
nom. nud.

I saw no specimen bearing a label with this
name.

Fagraea dasyantha GitG & BENED. Bot. Jahrb. 54
(1916) 195.

Possibly related to or conspecific with F. gra-
cilipes. None of the syntypes seen, probably all
lost at Berlin.

Fagraea ledermannii GitG & BENED. Bot. Jahrb.
54 (1916) 191.

The description of the vegetative parts fits very
well with F. salticola, but the flowers are much
smaller in all parts and remind more of F. gracili-
pes or F. bodenii.

Fagraea melanochlora GiLG & BENED. Bot. Jahrb.
54 (1916) 192.

Possibly identical with F. salticola or with

F. bodenii.

336

FLORA MALESIANA

[ser. I, vol. 6?

Fagraea monticola GitG & BENED. Bot. Jahrb. 54
(1916) 189.
Apparently related to F. salticola or F. bodenii.

Excluded

Fagraea cardinalicarpa E.M. Leafl. Philip. Bot. 10
(1939) 3759, nom. illeg—Based upon ELMER
15492, a duplicate of which is-preserved in the
Herbarium of the Arnold Arboretum. This was
distributed under the manuscript name of Tern-
stroemia megacarpa Ei_m. According to Dr VAN
STEENIS it probably represents some Rubiacea,
possibly Cephaelis. It is doubtful whether the
fruit and the twig belong together. The name is
illegitimate as the description was in English only.

nom. nud.—This was the base of Ternstroemia ?
penangiana CHoisy, Mém. Soc. Phys. Hist. Nat.
Geneve 14 (1855) 108 (Guttiferae).

Fagraea pachyclados K. Scu. in Sch. & Laut.
Nachtr. (1905) 349.—This is the basionym of
Mastixiodendron pachyclados (K. SCH.) MELCH.
Bot. Jahrb. 60 (1925) 167 (Rubiaceae).

Fagraea tetragona Span. Linnaea 15 (1841)
326.—MIQUEL (FI. Ind. Bat. 2, 1857, 409) based
hereon his Alyxia spanogheana (non Alyxia te-
tragona R. Br. 1810) (Apocynaceae).

Fagraea vitiensis SEEM. Bonplandia 9 (1861) 257,
nom. nud.; ibid. 10 (1862) 37, nom. nud. = Linociera
vitiensis A. C. SmirH, Bull. Torr. Bot. Cl. 70
(1943) 549 (Oleaceae). See A. C. SmitH, J. Arn.

Fagraea dubia Watt. Cat. (1831) n. 4456, Arb. 33 (1952) 112.

2. BUDDLEJA

Houst. ex LINNE, Gen. Pl. ed. 5 (1754) 51; Sp. Pl. 1 (1753) 112; GAGNEPAIN,
Not. Syst. 2 (1912) 182; MARQUAND, Kew Bull. (1930) 177. Spelled Buddleia
by most authors.—Nicodemia TENORE, Cat. Orto Bot. Napoli (1845) 88.—
Fig. 24-25.

Shrubs, rarely trees or suffrutescent herbs; twigs, leaves (especially underneath)
and inflorescences usually covered with a dense tomentum of stellate hairs (some-
times and in some parts intermingled with capitate-glandular hairs); colleters
absent. Branches terete or quadrangular, in the latter case sometimes narrowly
winged. Leaves opposite or nearly so (in some non-Mal. spp. alternate), either
auriculate at the base or connected by a stipular line, or by leafy ‘interpetiolar
stipules’, in a few species perfoliate, entire, serrate-dentate, or rarely lobed.
Inflorescences terminal and/or axillary, thyrsoid, racemose, clustered-cymose, or
pseudo-verticillate, especially the terminal panicles often long and many-flowered.
Bracts linear. Flowers 4-merous. Calyx gamosepalous, outside usually densely
tomentose, inside glabrous. Corolla salver-shaped, sometimes campanulate, out-
side tomentose or glabrous, inside usually sparsely hairy; tube straight or curved,
lobes imbricate in bud. Stamens adnate to the corolla-tube (the only exception
being B. gynandra, in which they are described as attached to the ovary); anthers
subsessile, cleftat base, introrse, mostly included. Ovary 2-celled, placentas thickened,
with c© ovules; stigma conical or club-shaped. Fruit a septicidal 2-valved
capsule (subg. Buddleja) or a ‘berry’ (subg. Nicodemia). Seeds ~, small, often
winged; endosperm fleshy.

Distr. About 100-120 spp. in the tropics and subtropics of America, Africa, and Asia; in Malaysia one
species native, a few others naturalized or cultivated.

Ecol. Heliophilous shrubs of open terrain, shrubby vegetation, and forest borders, often invading
disturbed places, from the lowland to high up in the mountains (especially some andine species). Many
species have fragrant flowers and are frequented by many insects; the widely cultivated B. davidii is well
known to attract many butterflies. The seeds of the species with capsular fruits are wind-dispersed; those
from the species bearing berries may be dispersed by birds or other animals.

Uses. The leaves of some species are used as a wash on account of the presence of saponin; for the
same reason some species are used as a fish-poison. Moreover, some are used in medicine. A few species
are commonly planted as ornamentals.

Morph. The flowers are normally 4-merous; in B. officinalis MAxIM., however, I found in the same
inflorescence between many 4-merous flowers some S-merous ones and some with only the calyx 5-
merous; Hook. f. (Bot. Mag. t. 6323) mentions also 5-merous flowers for B. asiatica.

Dec. 1962] 337

GAGNEPAIN and MARQUAND attached much value to the place of insertion of the stamens. In many
species, e.g. B. asiatica, this character seems to be very constant indeed. In some cases, however, it seems
to represent the only differentiating character between a pair of sympatric closely related species. A
possible example is B. curviflora H. & A. and B. japonica LINDEN; furthermore, I found many specimens
which fully agreed with B. Jindleyana with the exception of the stamens which were inserted about half-
way the corolla-tube instead of at its base; according to the key given by MARQUAND these specimens
should represent B. venenifera MAKINO; a third case may be the pair B. paniculata WALL. and B. crispa
Bru. Whether these could be examples of heterostyly should be examined in the field.

The fruits in Buddleja and related genera are principally capsular; they are 2-, or (in Adenoplea) 4-
celled, with many seeds, the cavities neither completely filled up by the seeds, nor by a pulp. In the
African genera Adenoplea and Adenoplusia, and in Buddleja subg. Nicodemia they are indehiscent, in
the other genera involved and in Buddleja subg. Buddleja they are septicidal with 2 valves. The fruits of
Adenoplea and of Buddleja subg. Nicodemia have thin fleshy walls; in Adenoplusia the walls are also thin-
fleshy, but the endocarp is chartaceous; in the genera with dehiscent fruits the walls are dry and charta-
ceous or leathery. A typical character is the occurrence of large spherical glands on the inner side of the
pericarp in Adenoplea and Adenoplusia, in the first named genus all over the surface, in the last named one
only in the axils of the dissepiment. I agree with Bruce & Lewis (Fl. Trop. E. Afr. Logan. 1960, 40,
footnote) that it is misleading to describe the fruits of Adenoplea and of Buddleja subg. Nicodemia as
‘berries’, as is usually done, or even those of Adenoplusia as ‘drupes’; ‘a non-dehiscent fleshy capsule’
would be a better description, in the latter case with the addition ‘with chartaceous endocarp’.

Taxon. I have recombined the genus Nicodemia with Buddleja, as was tentatively proposed by some
authors, for instance by BRucE & Lewis, /.c. 35, 41; they only differ in the fruit characters cited above.
The merging of these two genera is supported by the karyological data published by R. J. Moore (Am. J.
Bot. 34, 1947, 527-538): the diploid number of chromosomes of Nicodemia is 38 which is also the basic
number of Buddleja, and morphologically the chromosomes are indistinguishable. The few phytochemical
data known also point to a close relationship; anatomically there seems to be no clear difference; paly-
nological and embryological data on Nicodemia are still unknown.

LOGANIACEAE (Leenhouts)

KEY TO THE SPECIES

1. Ovary and style glabrous. Fruit a dehiscent capsule.
2. Tube of the corolla 2!,-4'4 mm long, about as long as or slightly longer than the calyx; corolla
outside densely stellate-hairy ee .wbeasiatica
2. Tube of the corolla 9-12 mm long, 3-4 times as long. as ‘the calyx; corolla ‘outside either glabrous or

glandular-pubescent.

3. Corolla straight, outside glabrous. Interpetiolar stipules distinct, leafy, often divided into 2 auricles.
Inflorescences paniculate . 9 2. B. davidii
3. Corolla slightly curved, outside densely glandular- pubescent. Stipules represented by an incon-
spicuous rim. Inflorescences slender, spike-like. A native from China and Japan, in Malaysia
sometimes cultivated . 0 B. lindleyana FORTUNE
. Ovary and style pubescent. Frait fleshy, indehiscent 5 3. B. madagascariensis

1. Subgenus Buddleja

Fruit a dry capsule, dehiscent.
Distr. The area of the genus, apparently with the exception of Madagascar and the Mascarenes.

1. Buddleja asiatica Lour. Fl. Coch. (1790) 72;
ed. Willd. (1793) 90; Bru. in DC. Prod. 10 (1846)
446; Mia. Fl. Ind. Bat. 2 (1857) 363; Bru. FI.
Hongk. (1862) 231; BEppome, Flor. Sylv. 3 (1872)
163, t. 21 (IV); BRANpis, For. Fl. (1874) 318;
Kurz, Fl. Burm. 2 (1877) 250; Hook. f. Bot.
Mag. (1877) t. 6323; CLARKE in Hook. f. FI. Br.
Ind. 4 (1883) 82; K. & V. Bijdr. 9 (1903) 89,
incl. var. densiflora, salicina, & sundaica; BRANDIS,
Ind. Trees (1906) 475; KiNG, J. As. Soc. Beng.
74, ii (1908) 600; TaLsot, For. Fl. Bombay 2
(1911) 262; GaGnep. Not. Syst. 2 (1912) 189;
Koorp. Exk. FI. Java 3 (1912) 60; Dop, Fl. Gén.
I.-C. 4 (1912) 160, f. 20 (5-9); Koorp. Atlas
(1914) f. 325-326, incl. var. brevispica; MERR.
Sp. Blanc. (1918) 306; En. Philip. 3 (1923) 316;
Ript. Fl. Mal. Pen. 2 (1923) 414; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 334; BoyNTON,

Addisonia 13 (1928) 5, t. 419; MARQUAND, Kew
Bull. (1930) 195; ReHper, J. Arn. Arb. 15 (1934)
309; Merr. Comm. Lour. (1935) 310; KANEHIRA,
Formosan Trees, rev. ed. (1936) 622, f. 580;
KANJILAL & Das, Fl. Assam 3 (1939) 312 ; BAKH.
f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam.
170, p. 14; Kerr in Craib, Fl. Siam. En. 3 (1951)
53.—B. salicina LamK, Ill. 1 (1792) 291;
Porret, Encycl. Suppl. 1 (1810) 744; Bi. Bijdr.
(1826) 742.—B. neemda BucH. HAM. ex Roxs.
Fl. Ind. 1 (1820) 411; Rercu. Ic. Bot. Exot. (1824)
t. 21; CHAM. & SCHLECHT. Linnaea 2 (1827) 598,
incl. var. philippensis; Roxs. Fl. Ind. ed. Carey |
(1832) 396; Bru. in DC. Prod. 10 (1846) 446;
Mia. Fl. Ind. Bat. 2 (1857) 363; Hance, J. Linn.
Soc. Bot. 13 (1873) 112; F.-Vitt. Nov. App.
(1880) 136; VIDAL, Sinopsis Atlas (1883) t. 69 C;
MARQUAND, Kew Bull. (1930) 195.—B. acumina-

338 FLORA MALESIANA [sera evolesG?

Fig. 24. Buddleja asiatica Lour. a. Habit, x 4, b. connate leaf-sheaths, < 2, c. flower, < 5, d. opened
corolla, < 10, e. pistil, * 10, f. fruit, x 5, g. seeds, x 10 (a RAHMAT 11423, b HotstvyooGp 571, c-e
HooGLANp & PULLEN 5355, f-g KoorpDeErRS 4343).

Dec. 1962]

LOGANIACEAE (Leenhouts)

tissima BL. Bijdr. (1826) 743.—B. densiflora BL.
Bijdr. (1826) 743; Bru. in DC. Prod. 10 (1846)
447; Mia. Fl. Ind. Bat. 2 (1857) 364.—B. sundaica
BL. Bijdr. (1826) 743; Bru. in DC. Prod. 10 (1846)
446.—B. virgata (non L. f.) BLco, FI. Filip. (1837)
57; ed. 2 (1845) 38; ed. 3, 1 (1877) 70.—B. amen-
tacea KRANZL. Bull. Jard. Imp. Bot. St. Pétersb.
13 (1913) 89 & 92: MARQUAND, Kew Bull. (1930)
184.—B. arfakensis KAN. & Hart. Bot. Mag.
Tokyo 56 (1942) 157, f. 1.—Fig. 24-25.

Evergreen shrub, suffrutex, or sometimes small
tree, 1-7 m high. Branches terete or subterete,
when young densely appressed- or woolly-, white-,
grey-, or fulvous-hairy. Leaves opposite, those in
the inflorescence often more or less alternate;
connected by an, often inconspicuous, stipular
line, narrowly to oblong- or ovate-lanceolate,
4-30 by 34-7 cm, herbaceous, glabrous to rather
densely hairy above, underneath densely short to
rather long woolly-hairy; base acute, margin
remotely serrate-dentate to subentire, apex long-
acuminate, acute, nerves and veins often impressed
above; petioles 34-1 cm, tomentose. Inflorescences
terminal and/or axillary, thyrsoid, spiciform, up
to 25 by 344-2 cm, densely tomentose; flowers in
crowded or more or less remote, (sub)sessile,
1—3-(rarely more-)flowered cymes, each cyme in the
axil of a linear bract. Pedicels 14-2 mm. Flowers
4-merous, a few occasionally 5-merous. Calyx
campanulate, 2-414 mm long, cleft to 14-14,
outside more or less densely tomentose, lobes
triangular-oblong, acute. Corolla white, sometimes
light-violet or greenish, outside stellate-hairy,
inside from halfway the tube to the basal part of
the lobes woolly; tube 214-414 mm, lobes oblong-
ovate, 1-114 mm long, rounded or obtuse. Sta-
mens inserted near the throat, anthers 34 mm long,
deeply 2-lobed at the base. Ovary and style gla-
brous, style 1 mm, including the club-shaped
stigma. Capsule ovate or oblong, flattened, 3-5 by
214-3 mm, glabrous, brown. Seeds ellipsoid, laxly
enveloped by the testa which is at both ends pro-
longed into a short wing; the endosperm c. 1 mm
long.

Distr. SE. Asia from W. Pakistan and the
Deccan to South China and Formosa, the Ma-
rianas (Guam, Saipan), and throughout Malaysia.
Sometimes cultivated and naturalized in tropical
and subtropical regions.

Ecol. In open, often disturbed or secondary
vegetation, apparently more or less a pioneer of
secondary growth, in pyrogenous grassland, in
gravel-beds, on lavastreams, and landslides, from
the lowland up to about 3000 m, locally often
gregarious. Fi. fr. Jan.—Dec.

Uses. Probably mainly on account of its
saponin-content sometimes used as fish-poison
and in native medicine for several purposes (see
BuRKILL, Dict. 1935, 379; Quis. Med. Pl. Philip.
1951, 710). In Assam the flowers are eaten “‘cooked
with rats’ meat’ (KANJILAL & Das, /.c.); inthe Phi-
lippines used in making rice-wine. Sometimes used
for fuel. Cultivated as an ornamental in many
tropical and subtropical to even temperate regions.

Vern. White butterfly bush, E, kaju saludang,

Fig. 25.
Markham River (E. New Guinea)
(P. VAN RoyYEN, 1961).

Buddleja asiatica Lour., Leson Creek,

tjémara angin, Sum., (d)jugul, ki hirisan, (ki)
monjenjeén, kipirit, ki urug, sumbung luna, Java S.,
(daun)putihan, godijan, gutéhan, sanggal putung, J,
barakadjér, Sumba, hu touis, Timor, lalawal,
lawahmas, tubilit, Alor, bandau, pati-ata, Born.;
Philip.: alatin, Bag., lagindi-salasa, Bis., maligus,
Bon., dufgalau, \bn., anaiop, dumdumani, I{f.,
amuging, lg., lagien-ti-subisub, tugnang, IIk.,
ilentud ulangan, mengayan, Sub., malasambing,
salibug, sambongkala, taliknéno (talianono), Vag.,
doknam, Ving.; kukuru-in-talun, malelema, Cel.,
deraidauti, hagaiahas, hifaita, jamp, mukorrere,
taggaras, tunun, uganga, wihadzaha, wold, New
Guinea.

Notes. A variable species, especially in Java.
It is not possible, however, to subdivide it into well
circumscribed varieties.

Buddleja amentacea KRANZL. was based upon
the apical part of a flowering shoot with alternate
leaves as occurs in many species. This is one of
the fairly exceptional cases in which the leaves in
the inflorescence are strongly developed and
conspicuous. The two leaves under the inflores-
cence, however, are opposite in the type specimen;
MARQUAND, /.c., erroneously included this species
in his sect. Alternifoliae. There is further no reason
to doubt Java as its provenance; in fact, among the
Javanese material of B. asiatica | found some speci-
mens which match the type of B. amentacea
quite well.

340

FLORA MALESIANA

[ser. I voli62

2. Buddleja davidii FRANCH. Nouv. Arch. Mus.
Hist. Nat. Paris II, 10 (1887) 65; GAGNEP. Not.
Syst. 2 (1912) 188; Boynton, Addisonia 2 (1917)
9, t. 45; MARQUAND, Kew Bull. (1930) 196;
Baku. f. in Back. Bekn. FI. Java (em. ed.) 7 (1948)
fam. 170, p. 14.

Large bush up to 244 m. Branches subqua-
drangular, when young densely grey to white
tomentose. Leaves oblong-elliptic or -ovate, 8-10
by 3-4 cm, herbaceous, glabrous above except
on the base of the midrib, densely tomentose
beneath; base cuneate, attenuate, margin minutely
serrate-dentate, apex acute-acuminate; petiole c.
14 cm long, densely tomentose, glabrescent, at
the base with auriculate, usually not connate,
often relatively large stipules. Inflorescences
terminal, sometimes moreover lateral, thyrsoid,
composed of mostly short-stalked, lax, many-
flowered cymes, c. 12-20 by 3!4 cm, sparsely pu-
bescent. Bracts linear, up to c. 44 cm. Flowers
subsessile, very fragrant. Calyx slender-campanu-

late, 3 mm long, outside sparsely pubescent,
lobes c. 14-14 the total length of the calyx, acute.
Corolla lilac, the mouth orange-yellow, outside
glabrous, inside sparsely hairy above the insertion
of the stamens, tube slender, 9 mm, slightly swol-
len halfway, lobes semiorbicular, 2-214 mm long,
their margin crisped. Stamens inserted halfway the
tube, anthers 1 mm long, 2-parted at base, api-
culate. Pisti/ glabrous, style 2 mm including the
club-shaped stigma. Capsule lanceolate-ellipsoid,
8 by 114-2 mm, glabrous. Seeds thread-like, 4mm,
in the centre only slightly thickened.

Distr. Tibet and Central China (Szechuan,
Hupeh); in Malaysia and elsewhere commonly
cultivated as an ornamental, naturalized in the
Malay Peninsula (Cameroon Highlands, above
Pavit Falls).

Ecol. In a clearing in jungle. Fi. fr. Jan.—Dec.

Vern. Summer lilac, orange eye butterfly bush,
E, Japanse sering, Dutch.

2. Subgenus Nicodemia, voy. stat.

Nicodemia TENoRE, Cat. Orto Bot. Napoli (1845) 88.
Fruit indehiscent, with a thin fleshy pericarp.
Distr. Apparently restricted to Madagascar and the Mascarenes.

3. Buddleja madagascariensis LAMK, Encycl. 1
(1785) 513; Hook. Bot. Mag. (1828) t. 2824;
Bru. in DC. Prod. 10 (1846) 447; Kina, J. As.
Soc. Beng. 74, ii (1908) 601.—Nicodemia mada-
gascariensis R. N. PARKER, For. Fl. Punjab, ed. 2
(1924) 357.—Adenoplea madagascariensis EASTW.
Leafl. West. Bot. 1 (1936) 197.

Shrub, 2-3 m. Branches terete, densely silvery-
white (in the herb. rusty-)tomentose, as are the
petioles, the undersurface of the leaves, and the
inflorescences. Leaves connected by an inconspi-
cuous stipular line, ovate-oblong to ovate-lan-
ceolate, 9-11 by 3-414 cm, more or less coriaceous,
somewhat bullate, glabrous above, densely to-
mentose beneath; base rounded to acute, margin
entire, apex tapering, acute; petiole c. 1144 cm.
Inflorescences terminal, thyrsoid with some rather
long thyrsoid basal branches, c. 20 cm long, with
relatively long-stalked (c. 1 cm), rather dense, c.
7-flowered, patent lateral cymes. Bracts linear,
up to c. 114 cm. Flowers subsessile, sweet-scented.
Calyx campanulate, 3 mm long, outside densely
tomentose, lobes c. 1 mm, broadly triangular.
Corolla orange, outside densely tomentose, inside
glabrous, tube slender, c. 6 mm, lobes 2 mm long,
rounded. Stamens inserted slightly below the
mouth; anthers 114 mm long, shortly split at the
base. Ovary (except at base) and style densely
hairy, style slender, 414 mm including the club-
shaped stigma. Fruit globular, c. 21 mm, pubes-

cent, at first white, when ripe purple-blue. Seeds
ellipsoid, c. 1 mm long, smooth, brown.

Distr. A native of Madagascar, cultivated and
naturalized in tropical and subtropical regions;
in Malaysia naturalized in the Malay Peninsula
(Penang, Perak).

Ecol. Open jungle, 700-800 m. F/. Oct.—Febr.;
I saw no fruiting specimens from Malaysia.

Uses. According to STANDLEY (Contr. U.S.
Nat. Herb. 23, 1924, 1143) in Madagascar “the
leaves were formerly employed ...as a soap
substitute and the plant is used as a remedy for
asthma, coughs, and bronchitis.”

Excluded

Buddleja indica LAMK, Encycl. 1 (1785) 513.—
B. diversifolia VAHL, Symb. 3 (1794) 15, nom.
illeg.; Bru. in DC. Prod. 10 (1846) 445.—Nico-
demia diversifolia TENoRE, Cat. Orto Bot. Napoli
(1845) 88, nom. illeg.

A native from the Mascarenes and Madagascar,
twice wrongly described from Java, as already
mentioned by BENTHAM, /.c. The name B. diver-
sifolia was illegitimate as B. indica LAMK was
cited as a synonym.

Buddleja otophylla Hassk. Versl. Med. Kon. Ak.
Wet. Afd. Nat.5 (1857) 97 = B. brasiliensis Jaca. f.

Apparently based on a specimen cultivated in
the Botanical Garden, Bogor.

Dec. 1962] LOGANIACEAE (Leenhouts) 341

3. NORRISIA

GARDN. in Hook. J. Bot. Kew Misc. 1 (1849) 326.—Fig. 26.

Trees. Twigs gradually thickening from the nodes upwards, rusty pubescent
when young. Leaves petioled; connected by very broadly triangular, about | mm
high, blunt stipules. Inflorescences terminal and axillary, dichasially branched to
thyrsoid, dense and many-flowered, densely rusty tomentose; bracts narrowly
triangular. Flowers 5-merous, subsessile, with two pairs of decussate, minute
bracteoles. Ca/yx cupular, 34 (—1) mm high, tube very short, lobes broadly rounded,
densely tomentose outside, inside glabrous, without colleters. Corolla salver-
shaped, thin-fleshy, outside densely tomentose, creamy to yellowish; lobes valvate
in bud. Stamens inserted between the corolla-lobes, glabrous, exserted filaments
filiform, c. 6 mm; anthers latrorse, cells free in the lower half. Ovary obovoid,
for the greater part densely and shortly erect-hairy; 2-celled, with elliptic peltate
placentas; ovules - ; style terete, slender, glabrous, caducous, stigma faintly
capitate and slightly bilobed. Capsule septicidal, 2-valved, valves outside densely
fulvous-tomentose except the margins. Seeds few to many, small, slenderly
spindle-shaped, smooth and glabrous; endosperm fleshy.

Distr. Two closely related species, restricted to Malaysia: Sumatra, Malay Peninsula, Borneo, and
the Philippines.

Ecol. Small to medium trees of lowland forests. Insect-pollination seems probable; the tiny seeds are
in all probability wind-dispersed.

KEY TO THE SPECIES

1. Anthers oblong, 4/; mm long. Mouth of the corolla glabrous or with a few long hairs only. Most of
the nerves distinctly looped and joined at some distance from the margin. Twigs usually conspicuously
lenticellate tT er Poteet st Whig te Eee biel RagcPeRs. SiMen a Rte ah. cee WED PIN Mala CCeNSIS

1. Anthers orbicular, 1. mm g. Mouth of the corolla densely tomentose. Nervation for the greater part
open, only the upper two or three pairs of nerves looped and joined close to the margin. Twigs rarely

distinctly lenticellate .

1. Norrisia malaccensis GARDN. in Hook. J. Bot.
Kew Misc. 1 (1849) 327; Mia. Fl. Ind. Bat. 2
(1857) 359; CLARKE in Hook. f. Fl. Br. Ind. 4
(1883) 85; Kinc, J. As. Soc. Beng. 74, ii (1908)
602, incl. var. pubescens; RipL. Fl. Mal. Pen. 2
(1923) 414, var. malaccense only; Burk. Dict.
(1935) 1561, p.p.—Antonia griffithii WiGuT, Ill.
Ind. Bot. 2 (1850) 172, t. 156 b.—N. philippinensis
Evo. Leafl. Philip. Bot. 4 (1912) 1482; Merr. En.
Philip. 3 (1923) 311.—Fig. 26g, 1.

Tree 10-30 m by 32-40 cm go. Twigs terete,
nearly always densely set with minute, warty,
white lenticels. Leaves faintly pubescent to gla-
brous, elliptic to oblong, 614-10 by 234-4 cm,
thin-coriaceous, base cuneate, attenuate, apex
usually shortly acute-acuminate, rarely blunt;
midrib prominent and keeled beneath, nerves
6-8 pairs, usually distinctly ascending, fairly
strongly curved, usually distinctly looped and
joined at some distance from the margin, flat
above, prominulous beneath; petiole 14-1 cm,
slender. Inflorescences terminal, often thyrsoid,
414-84 cm long incl. the 214-414 cm long pe-
duncle. Largest bracts 114-2 mm long. Tube of the
corolla 414 mm, inside sparsely hairy, the base
excepted. Anthers oblong, c. 4/, mm. Ovary 1 mm;
style 6-744 mm. Fruits apparently scarce, 2-314

2. N. maior

by 134 mm. Seeds few, c. 2 mm long, very delicate.

Distr. Malaysia: Sumatra (West Coast Res.),
Malay Peninsula (southern half only), and the
Philippines (Sibuyan, Mindanao).

Ecol. Inand along primary forests, up to 920 m.
Fl. March-Nov., fr. Nov.

Vern. Balang, bareh bareh, Sum., kayu kar-
karas, kélat, sérapoh bukit, Mal. Pen.

Notes. Specimens from the Philippines differ
slightly from those of the Malay Peninsula and
Sumatra in the following points: petioles 5-6 mm
versus 7-10 mm, nerves 7-8 pairs, more or less
spreading versus 6—7 pairs and distinctly ascending.

Antonia griffithii WiGHT was based upon ma-
terial collected by GriFFITH in Malacca. This was
probably GriFFITH K.D. 3731. The specimens,
distributed under this number belong, however,
to both species of Norrisia. I did not see any
specimen clearly bearing the name in WIGHT’s
handwriting; the figure can refer to both species;
the same holds for the description, the only
point in favour of N. malaccensis being “leaves

. glabrous on both sides”.

2. Norrisia maior SOLER. in E. & P. Nat. Pfl. Fam.
4, 2 (1892) 37; Kina, J. As. Soc. Beng. 74, ii
(1908) 602.—N. malaccensis (non GARDN.) MERR.

342

FLORA MALESIANA

[ser. I, vol. 6?

Fig. 26. Norrisia maior SOLER. a. Habit, * 24, 6. connate leaf-sheaths, »

opened corolla, showing the bearded mouth, »
x 4, 7. placenta with seeds, * 4, k. seed, x 6.

8, e. pistil, »
N. malaccensis GARDN. g. Anther, < 8, /. leaf, x 24

2, c. flower, x 4, d. detail of

4, f. stigma, * 20, A. anther, = 8, i. fruit,

(a-f, h RIDLEY 6828, g, | SF. 40586, i-k Kep 15224).

En. Born. (1921) 490; CAmMMeRL. Bull. Jard.
Bot. Btz III, 5 (1923) 302; HeyNne, Nutt. Pl.
(1927) 1267; Burk. Dict. (1935) 1561, p.p.—wN.
malaccensis var. major Ripi. Fl. Mal. Pen. 2
(1923) 414.—Fig. 26a-f, h-k.

Tree 10-45 m by 10-75 cm g, heavily buttressed
up to 4 m high; twigs terete, lenticels inconspic-
uous to nearly invisible. Leaves elliptic, 514-10
by 214-5 cm, thin-coriaceous, usually minutely
tomentose on the midrib above, sparsely hairy on
midrib, nerves, and sometimes veins beneath;
base cuneate, attenuate, apex blunt or rounded to
fairly abruptly to tapering acuminate, acumen
short and broad, blunt to acute; midrib prominent

and rounded beneath, nerves 6—10 pairs, spreading,
faintly curved, not looped and joined with the
exception of the upper 2 or 3 pairs, petiole (4—)
6 mm. /nflorescences terminal and usually in the
axils of the upper pair of reduced leaves, usually
compoundly cymose, 3—6!4 cm long, including the
peduncle, which is up to 314 cm; bracts up to 2'4
mm. Corolla-tube 5—6 mm long, inside about the
mouth densely tomentose, further glabrous.
Anthers suborbicular, c. mm ¢. Ovary 144 mm;
style 7-8 mm. Fruits always many, 4-10 by 2-4
mm. Seeds c. 5—20 per cell, 4 mm long, delicate.

Distr. Malaysia: Sumatra (Palembang), Banka,
Riouw, Malay Peninsula (southern half only), and

Dec. 1962]

LOGANIACEAE (Leenhouts)

343

Borneo.

Ecol. Primary and secondary forests, appar-
ently often along river-banks, in swamp forests or
in temporarily inundated places, up to 450 m.
Fl. Jan., July—Oct., fr. Febr.—April, Aug., Oct.—
Nov.

Uses. In the Malay Peninsula the timber is
locally used in house building. In Sumatra the

light and soft wood is used for floors of houses
and indoor constructions, but even then it lasts
for only 3—4 years; from the buttresses, handles of
tools and oars are made.

Vern. (B)éngkaras, meérkaras, Sum., kayu
tankat, lémak hutam (or kétam ?), otak hudang,
sérapak paya, Mal. Pen., bandung, kélait, masilan
putih, simpopoit, tabak, Born.

4. GELSEMIUM

Jussieu, Gen. Pl. (1789) 150.—Medicia GARDN. & CHAmpP. in Hook. J. Bot.
Kew Misc. | (1849) 324.—Leptopteris BL. Mus. Bot. 1 (1850) 240.—Fig. 27.

Straggling shrubs. Twigs slender, terete. Leaves petioled; stipules reduced to
an interpetiolar line. /nflorescences terminal and axillary, thyrsoid to 1-flowered.
Flowers 5-merous (in G. sempervirens heterostylous). Calyx: sepals free, glabrous
or ciliate. Corolla funnel-shaped, lobes imbricate. Stamens glabrous, exserted;
filaments strap-shaped; anthers sagittate, cells free, latrorse. Pisti] glabrous;
ovary 2-celled, ovules several per cell; style filiform, stigma twice forked. Capsule
septicidal, 2-valved. Seeds several, winged.

Distr. Three species, one in SE. Asia, the others in SE. North America, Mexico, and Guatemala.

Cf. LEENH. Pac. Pl. Areas map 30.

Ecol. Shrubby vegetation at low to medium altitudes.

1. Gelsemium elegans (GARDN. & CHAMP.) BTH.
J. Linn. Soc. Bot. 1 (1856) 90; Mia. Fl. Ind. Bat. 2
(1857) 359; Bru. Fl. Hongk. (1861) 229; Kurz,
Fl. Burma 2 (1877) 249; Fors. & HeEms_. J.
Linn. Soc. Bot. 26 (1889) 117; BRANDis, Ind. Trees
(1906) 476; Dorp, Fl. Gén. I.-C. 4 (1914) 162,
f. 211-4; CAMMERL. Bull. Jard. Bot. Btzg III, 5
(1923) 295; Hu & Cuun, Ic. Pl. Sinic. (1929)
t. 97; Merr. & CHUN, Sunyatsenia 2 (1935) 305;
KANJILAL & Das, Fl. Assam 3 (1939) 314; HERK-
Lots, Hongk. Natur. 10 (1940) 28, f. 1-2.—Medicia
elegans GARDN. & CHAMP. in Hook. J. Bot. Kew
Misc. 1 (1849) 324; Bru. in ibid. 5 (1853) 56.—
Leptopteris sumatrana Bu. Mus. Bot. 1 (1850) 240,
f. 34.—G. sumatrana BoeRL. Handl. 2 (1899)
448, 457; Gipss, J. Linn. Soc. Bot. 42 (1914) 111.
—Fig. 27.

Usually a straggling half-shrub, sometimes a
large, woody climber; twigs glabrous. Leaves
ovate-lanceolate (to ovate), (7-)10-12 by 2-54
cm, thin-chartaceous to pergamentaceous, base
rounded to cuneate, apex long and slenderly
acute-acuminate; midrib sunken above; nerves
c. 5-8 pairs; petioles 34-1 cm. Inflorescences
terminal and in the upper leaf-axils, together
making a pyramidal leafy panicle; lower axillary
branches up to 6 cm, the terminal inflorescence
c. 3 cm, all sparsely pubescent. Bracts acutely
triangular to subulate, up to 4 mm long, ciliate
and pubescent on the prominent midrib beneath.

Pedicels c. 34 cm, bibracteolate. Sepals oblong-
ovate, 214-314 by 114-134 mm, acute, erect, out-
side glabrous or slightly pubescent on the thicken-
ed to keeled midrib, inside glabrous; no colleters.
Corolla 1-14 cm long, bright yellow to orange,
the tube outside brownish, mouth spotted with
red, the tube 7-9 mm, thin, fully glabrous, lobes
blunt to acute. Stamens inserted about the middle
of the tube, filaments 314-4 mm long, anthers
oblong-ovate to lanceolate, 114-2 mm _ long,
acute. Ovary oblong to obovoid-lanceolate, 2-2!
mm; style 8-12144 mm. Fruits ellipsoid, 4-10 by
3-4 mm, smooth and glabrous, papyraceous, the
valves split at the top. Seeds c. 8 per cell, elliptic
to bean-shaped, 314 mm go, warty, the central
part hairy, winged all around.

Distr. Assam, N. Burma, N. Siam, Indo-
China, S. and SE. China, Hainan, and Malaysia:
Sumatra, Borneo.

Ecol. In scrubby forests and thickets, 250-2000
m. Fl. June-Aug., Nov., Dec., fr. June-July,
Nov.

Uses. Reputed to be very poisonous, especially
the leaves and roots; mentioned to be used for
murder and suicide. The poisonous element is the
alkaloid gelsemidin.

Vern. Liman, Borneo.

Note. In specimens from N. Burma the fruits
are usually slightly larger (12 by 6 mm).

5. STRYCHNOS
LINNE, Gen. Pl. ed. 5 (1754) 86; Sp. Pl. 1 (1753) 189; A. W. Hitt, Kew Bull.

(1917) 121.—Fig. 28, 30-32.

Usually lianas, sometimes shrubs or treelets; usually provided with axillary,

344 FLORA MALESIANA [ser. I, vol. 6?

Fig. 27. Gelsemium elegans (GARDN. & CHAMP.) BTH. a. Habit, * 24, b. flower, * 2, c. opened corolla,
x 2,d. pistil, x 4,e. anther, x 6,f. fruit, x 2, g.seed, x 4 (a-e BALANSA 1041, f-g BeccaRI herb. 6642).

Dec. 1962] LOGANIACEAE (Leenhouts) 345

simple or double tendrils and sometimes with axillary thorns; stems and older
branches in some species spiny. Stipules reduced to a mostly ciliate and straight
rim connecting the leaf-bases. Leaves mostly inserted upon distinct leaf-cushions,
3-5(- ie plinerved, i.e. apart from the midrib nearly always provided with one or a
few pairs of nearly equally strongly developed basal nerves which do not fully
reach the leaf apex; penninerved in a few African species. Some pairs of scale-
like cataphylls are present at the base of new shoots, of inflorescences, and of the
branches of the latter. Inflorescences terminal or axillary, thyrsoid. Bracts scale-
like. Flowers (4-)5-merous. Ca/yx nearly completely divided, lobes in Mal. spp.
always broad, scale-like, and brown (in African and American spp. sometimes
lanceolate and green), outside usually very sparsely hairy, ciliate along the margin,
inside at the base provided with colleters. Corolla rotate to salver-shaped, white to
yellowish or greenish, thin-fleshy, always more or less thickened towards the lobes,
the basal part included by the calyx much thinner, outside usually distinctly dense-
ly papillose, mostly glabrous, inside variously hairy except the thin basal part;
lobes valvate in bud, spreading to reflexed when open. Stamens exserted; anthers
mostly slightly bifid at the base, introrse. Ovary 2-(in some African spp. 1-)celled,
with many ovules; style cylindric, stigma faintly 2-lobed. Berry usually globose or
ellipsoid, the thin to thick shell in Mal. spp. always hard, outside smooth or mi-
nutely warty, glabrous, orange to red when ripe; pulp fleshy, usually orange.
Seeds ~, or 2-1, either lenticular, orbicular to elliptic and usually convex on one
and concave on the other side with a silky or felty testa (fig. 28g, k), or irregular
castorbean-shaped and glabrous (fig. 28j); endosperm bony.

Distr. About 150-200 ssp. in the tropics and subtropics.

Ecol. Large lianas in the forests or scrambling or erect shrubs or treelets in more open vegetations.

The dispersal of the seeds is still incompletely known; in all probability this is largely due to animals,
both mammals and birds which digest the fruit pulp; according to KRuKoFF (Brittonia 4, 1942, 258)
toucans may be mainly responsible for it in America.

Uses. Many species are poisonous as they contain a sometimes high amount of alkaloids in their
seeds, roots, bark, and leaves. Most important among these alkaloids are strychnine and brucine (in
American spp. these are replaced by curare), which are used in medicine; the main source of these al-
kaloids are the seeds of S. nux-vomica, and locally those of S. ignatii. The natives, especially those in Bor-
neo and the Malay Peninsula, make use of the poison from roots and bark in composing arrow-poison.
The lianas are used as ropes. For further details see BURKILL, Dict. (1935) 2091-2100.

Taxon. It is not yet possible to give a satisfactory subdivision of the whole genus. Best known and
still in use is that by PRoGEL (in Mart. Fl. Bras. 6, 1, 1868, 271), modified and made suitable for the whole
genus by SOLEREDER (in E. & P. Nat. Pfl. Fam. 4, 2, 1892, 37). They distinguished 3 sections, viz:

Sect. Longiflorae: corolla-tube distinctly longer than the lobes.

Sect. Intermediae: corolla-tube about as long as the lobes.

Sect. Breviflorae: corolla-tube distinctly shorter than the lobes.

This division is obviously rather artificial because lines of alliance based on a complex of other
characters sometimes go across it.

In his revision of the Asiatic species HILL, /.c., accepted SOLEREDER’S classification but divided sect.
Intermediae into two sections by using as an additional character the kind of indument on the inner side
of the corolla. In this way he came to the following subdivision:

Sect. Brevitubae: identical with SOLEREDER’s Breviflorae (fig. 30b-c).

Sect. Lanigerae: Intermediae sensu SOLEREDER in part, with long woolly hairs at the mouth (fig. 32).

Sect. Penicillatae: Intermediae sensu SOLEREDER in part, with a line of erect bristle-like hairs either
across the base or across the middle of the inner face of the corolla lobes (fig. 30g).

Sect. Tubiflorae: identical with SOLEREDER’s Longiflorae (fig. 28).

As far as the Asiatic species are concerned this subdivision is more satisfactory. Especially the Pe-
nicillatae and the Tubiflorae (the latter section with the exception of S. angustiflora BrH.) seem to be
natural taxa. Among the Lanigerae a few species are somewhat anomalous (S. polytrichantha, S. curtisii,
and S. /anata), the Asiatic Brevitubae showing the greatest diversity.

Though this is a fairly good subdivision of the Indo-Malayan species, it proved to be of very little use
for the African ones. In Africa the genus shows its greatest development, and any subdivision of the genus

346 FLORA MALESIANA _ [ser. L- VolenG2

as a whole will primarily have to be framed on the African species. DUVIGNEAUD (Bull. Soc. R. Bot.

Belg. 85, 1952, 9) attempted this and distinguished 17 sections and some subsections, based upon as many

characters as possible. I feel incapable of judging the full merits of his system; in trying to fit the Asiatic

species into it I got the impression that the number of subdivisions may be somewhat too large.
In the present account I have refrained from giving a sectional subdivision, confining myself to the
mentioning of six groups of distinctly related species:

( i) S. ignatii, S. nux-vomica, and S. lucida (Tubiflorae HILL; nearest to DuvIGNEAUD’s Floribundae).

(ii) S. maingayi, S. ovata, and S. villosa (Brevitubae HILL, close to DUVIGNEAUD’s sect. Sambae).

(iii) S. curtisii, S. polytrichantha, S. oleifolia, and S. lanata (different from HILL’s Lanigerae by the rel-
atively short tube; according to the description almost identical with DuvIGNEAUD’s sect. Ligustroi-
des).

(iv) S. colubrina, S. quadrangularis, S. borneensis, S. ledermannii, S. lanceolaris, and S. flavescens (Lani-
gerae HILL, about identical with DUvIGNEAUD’s Booneae according to the description).

(vy) S. luzoniensis (close to the next group, but with a short corolla-tube).

(vi) S. axillaris and S. ridleyi (Penicillatae HILL).

Nomenclaturally the sectional and subsectional names must by typified and corrected, starting with
those of PROGEL.
Notes. A fairly complete account of the morphology and of the variability of many characters, based

upon extensive field-studies, was given by KRUKOFF & MONACHINO, Brittonia 4 (1942) 253-259.

Though after some experience it may be possible to recognize the leaves of many species, flowers are
absolutely necessary to come to a trustworthy identification. In many cases the fruits are possibly also of
value for specific distinction, but up till now the available material of undamaged fruits is insufficient as
they were usually dried and pressed, instead of preserved in liquor. According to Ducke, Bol. Tecn.

Inst. Agr. Norte n. 30 (1955) the seeds show also distinct specific characters but they should also be

preserved in liquor: seeds prepared out of dried fruits are nearly worthless, as the typical testa-characters
usually have been lost by then.

KEY TO THE SPECIES

1. Tube of the corolla distinctly longer than the limb.
2. Inflorescences axillary. Leaves Oe slender-(to caudate-)acuminate. Fruits 4-12 cm @, many-
seeded ..... oo » 2 Si Saienatn
2. Inflorescences terminal, eventually on axillary twigs with at least. one pair ‘of leaves. Leaves either
not acuminate, or with a short and broad, triangular acumen. ues 2-4(-6) cm 9, 2-4-seeded.
3. Petiole 6-11 mm. Branchlets smooth. Leaves large, 41-15 by 314-11 cm, apex blunt to shortly and
broadly triangular-acuminate, mucronulate. Inflorescences is -flowered. Style sparsely woolly
hairy. Seeds c. 2 cm g, sericeous . . . . 2. S. nux-vomica
3. Petiole 2-4 mm. Branchlets scabrous by many "small warty lenticels. Leaves small, 214-10 by 114-6
cm, apex blunt to emarginate, not mucronulate. Inflorescences few-flowered. Style glabrous. Seeds
c. 1-144 cm @, tomentose. . . A Stee ase) >: Se OL
. Tube of the corolla about as long as or sr shorter than the limb.
a Corolla inside partly or nearly completely woolly, very rarely fully glabrous, never with a whorl of
erect, bristle-like hairs (fig. 30c, 31c, 32c).
5. Tube of the corolla distinctly shorter than the limb.
6. Corolla 2-4 mm long.

7. Calyx inside thinly pubescent. Inflorescences terminal and in the upper leaf-axils, 5-20 cm long,

lax, long-peduncled . . . . « © ab OSes ayillosai
7. Calyx inside glabrous. Inflorescences axillary ‘and ‘terminal, up to if cm long, dense, short-pe-
duncled.

8. Corolla outside glabrous. Filaments 114-2 mm; anthers oblong to lanceolate.
4. S. maingayi
8. Corolla outside sparsely hairy. Filaments short, up to 134 mm; anthers ovate.

5. S. ovata
6. Corolla 5'14-10 mm long.

9. Corolla 5144-7 mm long.
10. Anthers | mm long, glabrous; filaments 2 mm. Indument of the pistil appressed and short.

Inflorescences dense, 2-444 cm... . . 7. 8. curtisii
10. Anthers 114—2 mm long, bearded; filaments ile 114, mm. . Indument ‘of the ‘pistil woolly. Inflo-
rescences Bee oo a ern) An ome en operon > FKL Sp Hama

ae eee 8-10 mm long.
. Corolla outside glabrous, inside woolly with the exception of the basal part of the tube.
Filaments 214-4 mm; anthers lanceolate. Pistil 9 mm. Nerves strongly prominent beneath,
nearly unbroken till near the apex; venation very conspicuous on both sides.
8. S. polytrichantha
11. Corolla outside with some scattered hairs about the mouth and along the sutures, inside woolly

Dec. 1962]

LOGANIACEAE (Leenhouts)

347

from slightly above the base till about halfway the lobes. Filaments 2 mm; anthers elliptic.
Pistil 6 mm. Nerves moderately prominent, broken into arches and feeble towards the apex;

venation not very conspicuous . .
5. Tube of the corolla about as long as the limb.
12. Corolla about 9 mm long . Palme
12. Corolla up to 6144 mm long.

13. Corolla inside only with a few woolly hairs near the apex of the lobes .

9. S. oleifolia
13. S. borneensis

15. S. lanceolaris

13. Corolla inside woolly especially in the mouth, sometimes fully glabrous.

14. Ovary pubescent. Filaments 1-114 mm long, anthers often | mm or more.

11. S. colubrina

14. Ovary glabrous, style often with some hairs. Anthers subsessile, c. 34 mm long (compare also
12. S. quadrangularis, the flowers of which are still unknown).

15. Corolla glabrous outside. Leaves 5—7-plinerved .
15. Corolla pubescent outside. Leaves 3-plinerved .

14. S. ledermannii
16. S. flavescens

4. Corolla on the inner surface provided with a whorl of erect, stiff, bristle- like hairs, either at the
base of the lobes or up to about halfway on these, very rarely moreover pubescent (fig. 30g).
16. Calyx inside pubescent. Corolla-lobes inside fairly densely patently short-hairy above the bristles.

19. S. ridleyi

16. Calyx inside glabrous. Apart from the bristles corolla-lobes inside glabrous.

17. Corolla 214 mm long, tube shorter than the limb. Pistil pubescent .
17. Corolla 3-4 mm long, tube about as long as the limb. Pistil glabrous .

1. Strychnos ignatii BERG. Mat. Med. 1 (1778)
146; FruckiGeER & Meyer, Pharm. J. III, 12
(1881) 1-6, cum fig.; VIDAL, Rey. Pl. Vasc. Filip.
(1886) 449, p.p., t. 2; BaILL. Hist. Pl. 9 (1888)
f. 387-388; OLIver in Hook. Ic. (1892) t. 2212;
HILL, Kew Bull. (1911) 290, t. opp. 281; ibid.
(1917) 200; Merr. Sp. Blanc. (1918) 306; BRown,
Min. Prod. Philip. For. 3 (1921) 70 & 221; Merr.
En. Philip. 3 (1923) 312; Burk. Dict. (1935) 2095;
Brown, Usef. Pl. Philip. 3 (1950) 225, f. 88.—
Ignatia amara L. f. Suppl. (1781) 149, pro fr.;
see OLIVER in Hook. Ic (1892) t. 2212.—/gnatiana
Philippinica Lour. Fl. Coch. (1790) 126, pro fr.,
nom. illeg.; see MERR. Comm. Lour. (1935) 309.—
S. tieute LescH. Ann. Mus. Hist. Nat. Paris 16
(1810) 479 & 480, t. 23; BL. Bijdr. (1826) 1019;
Rumphia 1 (1836) 66, t. 24; Mia. Fl. Ind. Bat. 2
(1857) 380; HiLL, Kew Bull. (1911) 292; Koorp.
Exk. Fl. Java 3 (1912) 58; H1LL, Kew Bull. (1917)
200; HEYNE, Nutt. Pl. (1927) 1268; non v. MALM
in Fedde, Rep. 34 (1934) 285, quae est Ervatamia
sphaerocarpa (BL.) BuRK. (Apoc.); BAKH. f. in
Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170,
p. 8.—S. ovalifolia WALL. [Cat. (1829) n. 1592,
nomen] ex G. Don, Gard. Dict. 4 (1837) 65;
Bru. J. Linn. Soc. Bot. | (1856) 103; GAMBLE,
J. As. Soc. Beng. 74, ii (1908) 616, fr. excl.; HILL,
Kew Bull. (1917) 201; Ript. Fl. Mal. Pen. 2
(1923) 425; Burk. Dict. (1935) 2097.—S. phi-
lippinensis BLco, Fl. Filip. ed. 2 (1845) 61; ed. 3,
1 (1877) 116; F.-Vitt. Nov. App. (1880) 136.
—S. maingayi CLARKE var. fructuosa CLARKE
in Hook. f. Fl. Br. Ind. 4 (1883) 88.—S. beccarii
GiLG, Notizbl. Berl.-Dahl. 1 (1897) 267; HIiLL,
Kew Bull. (1911) 291.—S. cuspidata HILL, Kew
Bull. (1909) 359; ibid. (1911) 288; ibid. (1917) 199.
—S. pseudotieuté Hiri, ibid. (1911) 287.—S.
balansae Hirt, ibid. (1917) 200.—S. krabiensis
HILL, ibid. (1940) 199; in Craib, Fl. Siam. En. 3
(1951) 58.—Fig. 28a-j.

Large liana (in Borneo sometimes a shrub or
treelet); twigs glabrous; branches greyish brown,
more or less scabrous. Leaves ovate or elliptic to
lanceolate, 4-18(-22) by 2!4-9(-12) cm, thin-

17. S. luzonensis
18. S. axillaris

to coriaceous-chartaceous, glabrous, acute to
rounded at the base, always slightly attenuate,
apex distinctly acuminate, acumen up to 134 cm
long, slender and blunt or acute; triplinerved at
or above the base; petiole 14-1 cm. Inflorescences
axillary, mostly in the axils of already fallen leaves,
laxly branched, 2-4(-7) cm long, with some
10-20 flowers, minutely pubescent. Calyx 1-114
mm high, sepals ovate, acute, outside densely
minutely tomentose, inside glabrous. Corolla
salver-shaped, 7-17 mm long, tube 5-12 mm,
inside in the lower half with some long woolly
hairs. Stamens inserted in the mouth, filaments
short, anthers deeply cleft, oblong, 1!4—2 mm long,
apiculate, glabrous. Ovary globular, c. | mm 9,
glabrous; style c. 5-12 mm, glabrous or rarely
with a few long woolly hairs about the middle;
stigma truncate. Fruits few on strongly thickened
branches, (ellipsoid to) globular, 4-12 cm 9,
pericarp up to 5 mm thick, hard and woody,
smooth and glabrous. Seeds usually several,
either lenticular, elliptic to orbicular, c. 20-35
by 16-20 by 8-9 mm, and silky to felty (fig. 28g),
or irregular castorbean-shaped, c. 2 by 1 by 1 cm,
rough but glabrous (fig. 28j).

Distr. Tonkin and Malaysia: Malay Peninsula,
Borneo, Java, and the SE. Philippines (Biliran,
Samar, Leyte, Mindanao).

Ecol. Apparently often in Dipterocarp forests
on a sandy soil, furthermore in mixed rain-forests,
and sometimes along river-banks, from the low-
land up to c. 1500 m. Fil. Jan.—Sept., fr. Jan.—
Dec. According to RIDLEY, Disp. (1930) 342 &
352, the fruits are eaten by monkeys and civet-
cats.

Uses. The seeds are the Saint Ignatius beans
of commerce, one of the sources of strychnine.
The bark and the seeds are used as a febrifuge in
the Philippines. In the Malay Peninsula, Java, and
Borneo the roots are used for poisoning arrows;
moreover, in the Malay Peninsula a fish-poison is
made from it.

Vern. Saint Ignatius bean, E, pepita de San
Ignacio, Sp, akar ipoh (pérah), a. tontong, bélai

348 FLORA MALESIANA

q.

Fig. 28. Strychnos ignatii BERG. a. Habit, * 24, b. flower, * 2, c. opened corolla, = 4, d. pistil, X 4,
e. twig with fruits, x 24, f. cross-section of fruit, the seeds embedded in the pulpa, = 24, g. Jenticular-
shaped seed, = 24, h. ditto in length-section, showing the embryo, * 2, i. ditto, showing the endosperm
which is split for the greater part, < 24, j. irregular shaped seeds (St Ignatius Beans’), * 7/4.—S. nux-
vomica L. k. Seed, * 74 (a after Rumphia l, t. 24, b-d Hort. Bogor X-G—46a, e & g RIDLEY s. 7., f.
Hort. Bot. Bogor X-G-46, h-i KL 1929A, j herb. L Carpol. 1194, k herb. L Carpol. 1202).

Dec. 1962]

LOGANIACEAE (Leenhouts)

349

hitam, ipoh akar (bésar), i.a. buah kéchul, pokok
ipoh, ya mu lek, Mal. Pen., pokru, S, tjétek, J,
bahan-lak, bina(h), b. kalikis, b. sangang, ipu kajo,
sireh-sireh, talinga basing, Born.; Philip.: agua-
son, dankagi, kanlara, katalonga, manandog,
pangaguason, pepita-sa-katbalogan, Bis., (i)gasud,
S. L. Bis., C. Bis., (pepita-sa-)katbalénga(n),
Tag., Pamp., yangi-ilagan, Mbo.

Notes. A fairly variable species comprising
three forms. The specimens from the Philippines
(S. ignatii sens. str.), part of those from Borneo
(S. cuspidata’), *S. krabiensis’ from the Malay
Peninsula, and ‘S. balansae’ from Tonkin are char-
acterized by large, relatively thin leaves which
are dark-green when dried; the nerves are distinct-
ly looping in the upper part, as some of the veins
are fairly strongly developed. Furthermore, these
specimens have the largest flowers. A second group
is formed by the commonest form from the Malay
Peninsula (‘S. ovalifolia’) and the other part of the
Bornean ones (‘S. beccarii’): these have small
leaves with a less conspicuous nervation, especi-
ally those of the Bornean ones are slightly more
coriaceous and are brown when dry, while the
flowers are smaller; the leaves are distinctly acu-
minate like in the large-leaved group. The third
form, ‘S. tieute’ from Java, is more or less inter-
mediate; it has small thin leaves, which show a
fairly coarse nervation, they are not conspicuously
acuminate, and greyish green when dried; the
flowers are small.

As to the seeds, from the Philippines I saw only
irregular-shaped ones, from Borneo and Java only
lenticular ones, and from the Malay Peninsula I
had both types of seeds from plants which other-
wise showed no difference. Therefore I conclude
that this character cannot be used for specific
distinction; KRUKOFF & MONACHINO (Brittonia
4, 1942, 258) reached the same conclusion for the
American species. The value, formerly laid upon
this character, will have been caused by the
pharmaceutical use of the lenticular-shaped seeds
of S. nux-vomica and the irregular shaped ones of
S. ignatii from the Philippines.

The type of S. krabiensis is KERR 18582 (in K),
not 18532 as cited in the original publication.

S. spireana Dor (Bull. Soc. Bot. Fr. 57, Mém.
n. 19, 1910, 19), described from Laos, is either
closely related to or conspecific with S. ignatii;
as already remarked by Hitt (Kew Bull. 1917,
199) the corolla-lobes were erroneously described
as being hairy inside.

Ignatia amara L. f. was apparently based upon
flowering material of the American Rubiacea
Posoqueria longiflora AuBL. and a fruit of S.
ignatii. Ignatiana philippinica Lour. was no
n.ore than a superfluous new name, fully based
upon the diagnosis of LINNE f.. BLANCO originally
(Fl. Filip. ed. 1, 1837, 82) also adopted the name
Ignatia amara and copied the original diagnosis,
in the later editions he changed the name into
S. philippinensis, and gave a new description of
flowering material of the true S. ignatii, correctly
including only the fruit from the original mixtum.
In doing thus he restricted the name Jgnatia amara

to the flowering, Rubiaceous part.

2. Strychnos nux-yomica LINNE, Sp. Pl. | (1753)
189; Mia. Fl. Ind. Bat. 2 (1857) 378, excl. specim.
Mal. et f. depauperata quoad est S. lucida; non
F. v. M. Fragm. 4 (1863) 44 (= S. lucida);
CLARKE in Hook. f. FI. Br. Ind. 4 (1883) 90;
SOLER. in E. & P. Nat. Pfl. Fam. 4, 2 (1892)
37, f. 20 E-J; TrImMEN, Fl. Ceyl. 3 (1895) 175;
non BoeERL. Handl. 2 (1899) 460 (= S. lucida);
HILL, Kew Bull. (1917) 183, cum fig., 341; J. M.
H[ILLieR], Kew Bull. (1919) 238; GAMBLE, FI.
Madras 5 (1923) 868; Burk. Dict. (1935) 2096;
KANJILAL & Das, Fl. Assam 3 (1939) 315; Quis.
Philip. J. Sc. 77 (1947) 141; Baku. f. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 8;
HILt in Craib, Fl. Siam. En. 3 (1951) 59.—Fig. 28k.

Liana or tree up to 20 m high. Twigs slightly
pubescent, glabrescent; branches yellowish grey,
not rough, sometimes with axillary thorns. Leaves
broadly ovate to elliptic, (414-)714-15 by (314-)
614-11 cm, pergamentaceous to chartaceous, gla-
brous, faintly cordate to rounded at the base,
apex blunt or gradually acuminate, in the latter
case with a broadly triangular acumen, often
mucronulate; 3—5-plinerved, the inner pair diverg-
ing slightly above the base, the outer pair mostly
faint, midrib often with two additional stronger
nerves in the upper leaf-half; petioles 6-11 mm.
Inflorescences terminal on short axillary branch-
lets with usually one pair of normal leaves,
mostly laxly pyramidal and fairly many-flowered,
4-414 cm long, thinly tomentose. Calyx 1-114 mm
high, sepals ovate, blunt to acute, outside fairly
densely pubescent, inside glabrous. Corolla
salver-shaped, 10-12 mm long, tube 6!4-9 mm,
inside in the lower half sparsely woolly, margins
of the lobes thickened and minutely tomentose.
Stamens inserted in the mouth, anthers subsessile,
cleft to slightly below the middle, elliptic, 134 mm,
blunt, glabrous. Ovary ovoid, 1 mm, glabrous;
gradually narrowed into the style, the latter 10
mm, sparsely woolly hairy about the middle;
stigma orbicular. Fruits few on strongly thickened
branches, globular, c. 3-4(-6) cm 9, somewhat
scabrous. Seeds c. 4, lenticular, orbicular to
elliptic, 2-244 by 134-2 cm by c. 4 mm, densely
sericeous.

Distr. Ceylon, India, Siam, Indo-China, and
Malaysia: N. Malay Peninsula (Puket, cf. HILL),
naturalized in the Philippines (Mindoro, Oriental
Proy., on the shore of the Puerto Galera Bay).

Ecol. In forests and along the shore, at low
altitudes.

Uses. Cultivated for its seeds, which are used
in pharmacy as the main source of strychnine.

Note. S. nux-blanda Hitt, distributed in Burma,
Siam, and Indo-China, differs only slightly from
S. nux-vomica, mostly by its larger, more-nerved
and more distinctly acuminate leaves and its c.
2 mm long, mostly narrower and more acute
sepals. Furthermore by the sweet seeds which
contain no or only a very small percentage of
strychnine. Probably this is only a variety or form
of S. nux-vomica.

350

FLORA MALESIANA

[ser. I, vol. 6?

3. Strychnos lucida R. Br. Prod. (1810) 469;
Bru. Fl. Austr. 4 (1869) 369; BariLey, Queensl.
Fl. 3 (1900) 1024; Hitt, Kew Bull. (1911) 287;
ibid. (1917) 194.—Lignum colubrinum timorense
RumpuH. Herb. Amb. 2 (1741) 121, t. 38.—S.
colubrina L.: Witi_p. Sp. Pl. 1, 2 (1797) 1052,
et auct. div. p.p., typ. excl.—S. muricata Kost.
Allg. Med.-Pharm. Fl. 3 (1834) 1072; Mia. FI.
Ind. Bat. 2 (1857) 380; Merr. Int. Rumph. (1917)
423; Meyer Drees, Comm. For. Res. Inst. 7.
33 (1951) 74.—S. ligustrina BL. Rumphia 1 (1836)
68, t. 25; Hitt, Kew Bull. (1911) 286; Koorp.
Exk. Fl. Java 3 (1912) 58; Hitt, Kew Bull. (1917)
193; HeEYNeE, Nutt. Pl. (1927) 1267; Burk. Dict.
(1935) 2095; Baku. f. in Back. Bekn. Fl. Java
(em. ed.) 7 (1948) fam. 170, p. 8.—S. nux-vomica
L.: Bru. J. Linn. Soc. Bot. 1 (1856) 103, p.p.;
Mia. FI. Ind. Bat. 2 (1857) 378, pro spec. Mal.,
incl. f. depauperata; F. v. M. Fragm. 4 (1863) 44;
BoeRL. Handl. 2 (1899) 460; K. & V. Bijdr. 9
(1903) 66.—S. roborans HitL, Kew Bull. (1925)
424; in Craib, Fl. Siam. En. 3 (1951) 60.

Small, often crooked tree or shrub, up to 12 m
by 24 cm go, spiny when young. Twigs sparsely
pubescent, soon glabrescent (Australian specimens
glabrous from the beginning); branches greyish,
rough by many small lenticels. Leaves ovate or
elliptic to suborbicular, 214-10 by 114-6 cm, thin-
chartaceous, glabrous, below granular and dull
(somewhat glaucous), base cuneate to faintly
cordate, usually attenuate, apex blunt to rounded,
sometimes slightly emarginate, rarely tapering and
acute; mostly 3-plinerved at the base (to 5-pli-
nerved); petiole 2-4 mm. Jnflorescences terminal,
sometimes with a pair of strong basal branches
in the upper leaf-axils, 244-3 cm long, with c.
9 flowers, fairly densely, minutely pubescent.
Calyx 1-14 mm long, sepals broadly cordate,
acute, outside short-tomentose, inside glabrous.
Corolla salver-shaped, 1-114 cm long, tube 7-12
mm, inside sometimes with a few woolly hairs.
Stamens inserted in the mouth, glabrous, filament
short, anther cleft to about or slightly below the
middle, oblong, 114-134 mm. Ovary globular, c.
1 mm 9, conically narrowed into the c. 1144 cm
long style, both glabrous; stigma truncate. Fruits
few, globular, 2-214 cm o, smooth, glabrous.
Seeds 2-3, nearly disk-shaped, 12-15 by 10-12 by
214-5 mm, densely short pubescent.

Distr. Siam (Nakawn Sawan, Rachasima,
Ayuthia, Rachaburi) to Australia (W. Australia:
Prince Regent River; Northern Territory: Arnhem
Land; N. Queensland: Cook Peninsula); in
Malaysia: E. Java (also Madura and Kangean),
Lesser Sunda Is. (Bali to Timor), S. Moluccas
(Tanimbar Is.). Fig. 29.

Ecol. A characteristic plant of land subject
to a dry monsoon, in S. Malaysia and Siam in
teak and other dryland forests, in secondary
forest, shrubberies, and savannahs, in Siam also
on limestone rocks, in Australia mainly in open
to dense savannah woodland, open places in
monsoon forests, scrubby forest, efc., up to c.
200(—400) m. F/. May, Sept.—Dec., fr. June—Dec.
(for both only very few exact dates are available).

We sa a

Fig. 29. Distribution of Strychnos lucida R. Br.,
a species of drier regions.

Uses. The bark, the bitter wood, and the
root are medicinally used.

Vern. Doro laut, kaju-bidara-pait, widoro
gunung, J, bidara patis, Md, dérgunung, Kangean,
kaju bidara laut, Bali, songga, Sumbawa, ai baku
moruk, bidara laut, kaju-nassi, (daon) kaju ular,
maba muti (or putih), Timor, botniawe, Tanimbar
Is.

Notes. The leaves are deciduous, they turn
yellow before falling.

HILv (Kew Bull. 1917, 193) already stressed the
close affinity of S. /ucida and S. ligustrina; in fact
he distinguished them only upon one character:
the first-named would be a shrub, the second one
a tree, but he already was in doubt as to the
reliability as well as the value of this character.
From the large range of Australian specimens in
the herbaria at Brisbane and Melbourne it is quite
clear that this only ‘difference’ does not exist.
The species is very homogeneous in spite of the
large area from Siam to N. Australia and the
considerable gap in the everwet parts of Malaysia.

4. Strychnos maingayi CLARKE in Hook. f. FI. Br.
Ind. 4 (1883) 88, excl. var. fructuosa (= S. ignatii);
GamMBLE, J. As. Soc. Beng. 74, ii (1908) 614,
p. maj. p.; HILL, Kew Bull. (1917) 141, cum fig.;
Rip. Fl. Mal. Pen. 2 (1923) 423.—S. laurina DC.:
CLARKE in Hook. f. Fl. Br. Ind. 4 (1883) 88,
pro fr.

Liana. Twigs glabrous; branches purplish-
brown with many orbicular, small, light yellow
lenticels. Leaves elliptic to oblong (to lanceolate),
5-10 by 214-514 cm, chartaceous to thin-coria-
ceous, glabrous, base broadly cuneate to rounded,
apex tapering acute-acuminate; 3-plinerved at the
base, nerves near the margin; petiole 5-7 mm.
Inflorescences axillary and terminal or pseudo-
terminal, axillary ones usually on a quadrangular
1-1'4 cm long peduncle, and often with a pair of
strong basal branches, thyrse oblong, (2—)5—6 cm
long, laxly branched and fairly many-flowered,
densely and shortly tomentose-pubescent. Calyx
0.9 mm, sepals broad-ovate, blunt to rounded,
outside subglabrous, inside glabrous. Corolla
rotate, 3-4 mm long, glabrous outside, inside

Dec. 1962]

LOGANIACEAE (Leenhouts)

351

in the mouth long woolly hairy, tube 14-1 mm,
lobes linear, acute. Stamens inserted in the mouth,
filaments 114-2 mm long, glabrous, anthers bifid
at the base, oblong to lanceolate, 1 mm long,
slightly acuminate, densely barbate to long-ciliate
all around. Ovary ellipsoid, 1 mm, for the greater
part sparsely shaggy hairy, grading into the style,
the latter 2-214 mm long and fairly thick, hairy
in the basal part; stigma semi-globular. Fruits
fairly many, globular, c. 24. cm o. Seeds 1-2.
Distr. Malaysia: Malay Peninsula (Perak and
Singapore); a specimen from Borneo (ENDERT
4838, W. Kutai) with only very young flower-
buds may also belong to this species.
Ecol. Dense jungle at low altitude. Fr. July.
Notes. The leaves look like those of S. curtisii
and of S. ignatii (‘ovalifolia’); the former has
usually the nerves more distinctly diverging above
the base, and the latter differs by the coarser
venation, the more abruptly acuminate leaf apex,
and the green colour of the dried leaves (in S.
maingayi the dried leaves are reddish brown).
The flowers of these three species cannot be
confused, as they belong to different sections.
The relationship of S. maingayi is mainly with
S. andamanensis Hitt (Andamans) and S. ovata,
but they are both sufficiently different.

5. Strychnos ovata HILL, Kew Bull. (1909) 360;
ibid. (1917) 143; Merr. En. Philip. 3 (1923) 313;
non Pl. Elm. Born. (1929) 252 (= S. axillaris).—
S. panayensis HILL, Kew Bull. (1917) 148, cum
fig.; Merr. En. Philip. 3 (1923) 313.—S¢trychnos
sp. 1, Hitt, Kew Bull. (1917) 206.—Fig. 30a-f.

Climber or shrub. Twigs glabrous; branches
yellowish-brown to yellowish-grey. Leaves ovate
to elliptic or oblong, 6-10 by 3-6 cm, chartaceous,
glabrous, base broadly cuneate to rounded,
mostly attenuate, apex blunt, acute, or tapering
acute-acuminate; 3—S-plinerved, the nerves nearer
to the margin than to the midrib; petiole 4-10 mm.
Inflorescences axillary and pseudoterminal or
terminal, 214-7 cm long, laxly to densely branched,
fairly many-flowered, sparsely pubescent. Calyx
1-114 mm, sepals ovate, blunt, outside slightly
tomentose, inside glabrous. Corolla rotate, 3-414
mm long, thin to fairly thick-fleshy, especially the
lobes, outside sparsely appressed-pubescent, main-
ly towards the tip of the lobes, rarely glabrous, in-
side in the mouth and on the bases of the lobes
(sometimes very sparsely) woolly, tube 1-134 mm
long, lobes oblong-ovate to linear. Stamens
inserted in the mouth, filament short (34-134 mm),
glabrous or pubescent, anthers broadly ovate
to elliptic, 34-1 mm long, bearded. Ovary globu-
lar to ovoid, 1-114 mm high, glabrous or shaggy
hairy in the upper part, style 1-214 mm, in the
basal half likewise hairy; stigma truncate to semi-
globular, somewhat broadened. Fruits few,
ellipsoid to globular, 1-214 cm, smooth, thin-
walled, with 1-2 seeds. Seeds orbicular, flat, 1 cm 9,
felty.

Distr. Malaysia: Borneo, Philippines (Palawan,
Calamianes, Mindoro, Babuyan Is., Luzon, Ticao,
Panay), and N. Celebes (Gorontalo); possibly

also on P. Penang near the Malay Peninsula.

Ecol. In thickets and forests, along river-
banks, from sea-level up to c. 250 m. F/. May-—
June, fr. Febr., May.

Vern. Philip.: Zgasud, Bis.

Notes. In the original diagnosis the length of
the petiole is given as 0.5 mm; this should be 0.5
cm.

The Bornean specimens are slightly different
from those of the Philippines by less conspicuous,
parallel (instead of reticulate) veins, distinctly
acuminate leaves, oblong-ovate (instead of linear)
corolla-lobes, and smaller and broader anthers.

KERR s.n. (coll. 26-10-1926) collected on P.
Penang (herb. E) comes very close to the Bornean
specimens; it mainly differs in some minute flower-
characters (pubescence inside of the corolla main-
ly towards the tips of the lobes; anthers sometimes,
ovary and style fully glabrous).

The relationships of S. ovata are with S. main-
gayi (Malaya and ? Borneo) and S. confertiflora
Merr. & CHUN (Hainan). The latter differs by
its shorter and more dense inflorescences and by
some flower details.

Apart from the flowers, S. ovata looks much
alike S. axillaris (‘impressinervis’), but differs by
the usually more ovate and dull greyish green
(instead of shining brown) leaves, in fruit moreover
by the larger and more lax infructescences with
larger fruits.

6. Strychnos villosa HiLL, Kew Bull. (1911) 296;
Koorp. Exk. Fl. Java 3 (1912) 58; Hitt, Kew
Bull. (1917) 143; Baku. f. in Back. Bekn. FI.
Java (em. ed.) 7 (1948) fam. 170, p. 7.—S. /aurina
(non DC.) K. & V. Bijdr. 9 (1903) 68.—S. hors-

fieldiana (non Mig.) Koorp.-ScHum. Syst. Verz.

1 (1913) fam. 245, 164.—Strychnos sp. A & B,
Koorp.-ScHum. /.c. 165 & 166 resp.—S. hir-
sutiflora Hitt, Kew Bull. (1917) 144, cum fig.,
Rpt. Fl. Mal. Pen. 2 (1923) 423.

Liana. Twigs thinly tomentose, glabrescent.
Leaves ovate or elliptic to oblong, 7-17 by 3-8 cm,
chartaceous, glabrous to pubescent on the nerves
beneath, base broadly cuneate to subcordate,
attenuate, apex tapering to abruptly acuminate,
acumen short, deltoid, acute; 5-plinerved, inner
nerves diverging at 14-114 cm above the base;
petiole 44-1 cm. Inflorescences terminal and in
the upper leaf-axils, 5-20(-30) cm long, laxly
branched, many-flowered, thinly tomentose, pe-
duncle up to 10 cm long. Calyx 34-1 mm, sepals
elliptic or ovate, acute to blunt, outside pubescent
to glabrous, inside thinly pubescent. Corolla
2(-4) mm, less than halfway connate, outside
fairly densely hirsute to subglabrous, inside partly
or nearly completely (the base always excepted)
woolly or hirsute. Stamens inserted in the mouth,
filaments up to 44 mm, glabrous or sparsely pu-
bescent, anthers ovate, 14-34 mm, apiculate and
slightly bearded. Pistil 114-4 mm, glabrous to
hirsute. Fruits several, ellipsoid to globular, 134—
21% cm, the wall thin and hard. Seeds 1-4, elliptic,
15 by 12'4 by 314 mm, woolly-tomentose.

Distr. Malaysia: Malay Peninsula (Perak,

352 : FLORA MALESIANA [ser. I, vol. 6?

®@ 1060

Fig. 30. Strychnos ovata Hit. a. Habit, x 7, b. flower, < 8, c. opened corolla, < 8, d. stamen, * 16,
e. pistil, x 16, f. infructescence, * 34.—S. axillaris COLEBR. g. Opened corolla to show the type of hairi-
ness of the Penicillatae (a-e SAN 15268, f KOSTERMANS 4946, g CurTIS 3494).

Dec. 1962]

LOGANIACEAE (Leenhouts)

353

Johore), Simalur (off NW. Sumatra), N. Borneo,
Java, and Lesser Sunda Is. (Bali, Sumbawa).

Ecol. Apparently indifferent to climatic sea-
sons, in and along rain-, beach-, and teak-forests;
up to 600 m. Fi. Jan., June-July, fr. Aug., Nov.

Vern. Akar tandu dotan, Simalur.

Notes. Well characterized by the large and lax
terminal inflorescences with small and mostly
hairy flowers.

This species shows, apart from the small and
short-tubed flowers, a distinct resemblance to
S. colubrina.

7. Strychnos curtisii K. & G. J. As. Soc. Beng.
74, ii (1908) 614, excl. specim. K1NG’s coll. 10281
& 10438 (= S. colubrina); HILL, Kew Bull. (1917)
164, cum fig., fr. excl. (= S. colubrina); RIDL.
Fl. Mal. Pen. 2 (1923) 424, fr. excl.

Climber. Twigs glabrous, yellowish brown, later
on slightly scabrous by scattered lenticels. Leaves
elliptic, 8-9 by 334—5 cm, subcoriaceous, glabrous,
base rounded and slightly attenuate, apex rounded,
more or less abruptly, shortly and bluntly acu-
minate; (sub)5-plinerved, the inner nerves diver-
ging at c. 14 cm above the base; petioles 7 mm.
Inflorescences axillary and (pseudo?)terminal,
2-414 cm long, rather dense and many-flowered,
minutely tomentose. Ca/yx 114 mm, sepals broad,
rounded, glabrous on both surfaces. Corolla ro-
tate, 6 mm long, outside glabrous, inside the lower
half of the linear-lanceolate lobes densely woolly,
tube 2 mm long. Stamens inserted in the mouth,
filaments 2 mm, glabrous, anthers oblong-lan-
ceolate, 1 mm long, apiculate, glabrous. Pistil
434, mm, densely appressed-short-pubescent on the
upper half of the ovary and the greater part of the
style. Fruit unknown.

Distr. Malaysia: Malay Peninsula.

Ecol. Evergreen forest, 200-650 m. Fi. April—
June.

Vern. Sémijo akar.

Notes. Very close to S. polytrichantha and S.
oleifolia. The differences between these three
species are mainly in such characters as larger or
smaller flowers, inside completely or only partly
woolly, and in the shape and nervation of the
leaves. Possibly they only represent a Malayan,
Bornean, and Philippine subspecies of one species,
but for the moment I prefer to keep them separate.

As a lectotype I have designated Curtis 2973,
from which I saw two duplicates from SING.

8. Strychnos polytrichantha GiLG, Notizbl. Berl.-
Dahl. 1 (1897) 267; Hitt, Kew Bull. (1911)
298, cum fig.; ibid. (1917) 156.

Climber. Twigs glabrous, branchlets yellowish
brown, fairly scabrous by scattered small lenticels.
Leaves ovate or elliptic to lanceolate, 614-14
by 2-714 cm, chartaceous to subcoriaceous, gla-
brous, base rounded to acute, attenuate, apex
broadly rounded, more or less abruptly acuminate,
acumen short and blunt; 5-plinerved, inner nerves
diverging slightly above the base, nearly as strong
as the midrib, veins mainly transverse, minute and
dense, prominulous on both sides; petiole 6-7 mm.

Inflorescences axillary and terminal, 2-8 cm long,
laxly branched to fairly compact, few- to many-
flowered, minutely tomentose, glabrescent. Calyx
1-2 mm, sepals ovate, blunt, inside glabrous.
Corolla 8-10 mm, tube 3 mm, inside densely wool-
ly the basal part of the tube excepted. Stamens
inserted in the mouth, filaments 2!4-4 mm,
glabrous, anthers lanceolate, 144 mm, minutely
apiculate, barbate or not. Pisti/ 9 mm long, ovary
and at least the basal half of the style shaggy hairy.
Fruits few, globular, 5 cm @, scabrous. Seeds
probably many.

Distr. Malaysia: Borneo.

Ecol. Up to 1800 m. Fi. April-July, Dec.

Notes. Closely related to S. curtisii (Mal.
Pen.) and to S. oleifolia (Philip.); see under the
first-named species. Among these three species
S. polytrichantha is the one best characterized by
its large, woolly flowers and its very typical
nervation and venation.

9. Strychnos oleifolia Hitt, Kew Bull. (1917)
156; Merr. En. Philip. 3 (1923) 313.

Liana. Twigs glabrous, branchlets yellowish
brown, more or less scabrous by many small
lenticels. Leaves ovate-oblong to -lanceolate, c.
11-12 by 4 cm, chartaceous, glabrous, base
broadly cuneate, slightly attenuate, apex gradu-
ally acuminate; 3—5-plinerved, the inner nerves
diverging at c. 2 mm above the base; petiole
6-10 mm. Inflorescences terminal, 7-8 cm long,
fairly laxly branched and many-flowered, thinly
short-hairy. Calyx 114 mm, sepals ovate, blunt,
sparsely hairy outside, inside glabrous. Corolla
8 mm, outside with some scattered short hairs
about the mouth and along the sutures, inside
woolly from slightly above the base of the tube to
about halfway the lobes; tube 314 mm long. Sta-
mens inserted in the mouth, filaments 2 mm,
glabrous, anthers elliptic, 1!4 mm long, glabrous.
Pistil 6 mm, thinly shaggy hairy on the upper
half of the ovary and all along the style. Fruit
unknown.

Distr. Malaysia: Philippines (Palawan, pos-
sibly also in Luzon).

Ecol. In forests at low altitudes. F/. May.

Notes. A specimen from Luzon (RAMOS
1388), which might also belong to this species,
shows a large infructescence with branches which
are only slightly thickened and with several small,
2-seeded fruits. Vegetatively this specimen is
about identical with the type from Palawan, but
as flowers are absent nothing can be said with
certainty. These fruits however, are one reason
more to keep this species apart from S. poly-
trichantha, with which it is distinctly related.
Other reasons are the differences in the leaves and
in some flower characters.

On the other side the whole group of three
species (S. curtisii, S. polytrichantha, and S.
oleifolia) is distinctly related to S. colubrina, the
main difference being the relatively very short
tube of the corolla. S. oleifolia especially, is quite
close to some Philippine forms of S. colubrina.

354 FLORA MALESIANA [Sera lanvolesG?

Fig. 31. Sérychnos colubrina L. a. Habit of *S. forbesii’, X 7/4, b. flower, x 4, c. opened corolla, * 6,
d. stamen, * 6, e. pistil, < 6, f. twig with infructescences, < 74, g. and h. frontal and lateral view of
seed, 1 (a BRASS 5890, b-e RAHMAT 652, f VAN ROYEN 4272, g—h ELMER 8960).

Dec. 1962]

LOGANIACEAE (Leenhouts)

395

10. Strychnos lanata HILL, Kew Bull. (1911) 299,
cum fig.; ibid. (1917) 165; Merr. En. Philip. 3
(1923) 312.

Liana. Twigs glabrous; branchlets brown, with
many minute lenticels. Leaves oblong(-ovate),
614-18 by 314-9 cm, chartaceous, glabrous, base
cuneate to rounded, sometimes slightly attenuate,
apex blunt or gradually blunt- to acute-acuminate;
5-plinerved; petioles 14-1 cm long. Inflorescences
axillary, 6-8 cm long, laxly branched, many-
flowered, tomentose. Ca/yx 114-2 mm, sepals ovate,
blunt to acute, glabrous. Corolla 512-7 mm,
inside from about halfway the tube to at least
halfway the lobes woolly; tube 2-214 mm. Stamens
inserted in the mouth, filaments 1-114 mm, gla-
brous, anthers oblong to ovate-lanceolate, 114-2
mm, apiculate, barbate. Pistil 5-6 mm, ovary
woolly, style in the lower half with a few long
woolly hairs. Fruit unknown.

Distr. Malaysia: Philippines (Mindanao).

Ecol. Rain-forest along river, at 350-700 m.
Fl. Sept.

Vern. Bebekang-ulongen, Sub., tolanan, Bo.

Note. This species is about intermediate be-
tween S. oleifolia and S. colubrina; from the for-
mer differing in some flower details, from the
latter mainly by the short corolla-tube.

11. Strychnos colubrina LINNE, Sp. Pl. 1 (1753)
189; non SPAN. in Hook. Comp. Bot. Mag. 1
(1835) 347, nec Linnaea 15 (1841) 325 (= S.
lucida); H1LL, Kew Bull. (1917) 157; GAMBLE, FI.
Madras 5 (1923) 868; Burk. Dict. (1935) 2095.—
Sirioides & Sirioides alter RuMeH. Herb. Amb.
5 (1747) 49, t. 29 f. 1.—Camotain Bico, FI. Filip.
(1837) 85; ed. 2 (1845) 62; ed. 3, 1 (1877) 116.—
S. laurina WALL. [Cat. (1829) n. 1591, nom. nud.;
G. Don, Gard. Dict. 4 (1837) 65, in syn.] ex DC.
Prod. 9 (1845) 13; non K. & V. Bijdr. 9 (1903) 68
(= S. villosa); Hitt, Kew Bull. (1911) 297;
ibid. (1917) 150, cum fig., incl. var. thorelii; BAKH.
f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam.
170, p. 7; non Herne, Pfil. Samml. Clemens Kina-
balu (1953) 90 (= S. borneensis).—S. acuminata
WALL. [Cat. (1829) n. 1593, p.p., nom. nud.) ex
DC. Prod. 9 (1845) 14; CLARKE in Hook. f. FI. Br.
Ind. 4 (1883) 86; Hitt, Kew Bull. (1911) 302.—
S. multiflora Bru. J. Linn. Soc. Bot. 1 (1856) 102;
F.-ViILL. Nov. App. (1880) 136; OLIver in Hook.
Ic. (1892) t. 2213; Hitt, Kew Bull. (1911) 300,
t. opp. p. 301; ibid. (1917) 162; Brown, Min.
Prod. Philip. For. 1 (1920) 406; ibid. 3 (1921)
221; Merr. En. Philip. 3 (1923) 312; Brown,
Usef. Pl. Philip. 3 (1950) 227.—S. potatorum
(non L. f.) F.-VILL. Nov. App. (1880) 136; VIDAL,
Sinopsis (1883) 33, t. 69 f. D, pro var. multiflora
VIDAL.—S. hypogyna CLARKE in Hook. f. Fl. Br.
Ind. 4 (1883) 86, pro specim. Born., typo excl.; cf.
HILL, Kew Bull. (1917) 146.—S. septemnervis
CLARKE in Hook. f. Fl. Br. Ind. 4 (1883) 88;
GamBLE, J. As. Soc. Beng. 74, ii (1908) 619;
HILL, Kew Bull. (1917) 149, cum fig., incl. var.
imberbis; Ript. Fl. Mal. Pen. 2 (1923) 423.—
S. ignatii (non BERG.) VIDAL, Rev. Pl. Vasc.
Filip. (1886) t. 1.—S. bancroftiana F. M. BAIL.

Rep. Exp. Bellenden-Ker (1889) 49; Syn. Queensl.
Fl. Suppl. 3 (1890) 47; Queensl. FI. 3 (1900)
1025, t. 43; Compr. Cat. Queensl. Pl. (1913) 340;
Hitt, Kew Bull. (1911) 301; ibid. (1917) 208.—
S. celebica Koorpb. Med. Lands PI. Tuin 19 (1898)
540 & 631.—S. minahassae Koorb. ex BOERL.
Handl. 2 (1899) 460, nom. nud.—S. kerstingii
Gitc & K. Scu. in K. Sch. & Laut. Fl. Schutzgeb.
(1901) 498; Gi_G & BENeED. Bot. Jahrb. 54 (1916)
164.—S. luzonensis Evm. Leafl. Philip. Bot. 1
(1908) 332, p.p., typo excl.—S. barbata Hitt,
Kew Bull. (1909) 359, non CnHiov. (1932); ibid.
(1911) 297; ibid. (1917) 153; Merr. Int. Rumph.
(1917) 423.—S. forbesii HILL, Kew Bull. (1909)
360; ibid. (1911) 295; ibid. (1917) 150, fr. excl.—
2? S. merrillii H1LL, Kew Bull. (1911) 297; ibid.
(1917) 161; Merr. En. Philip. 3 (1923) 312.—
S. dubia Hitt, Kew Bull. (1911) 298, cum fig.;
ibid. (1917) 156; Merr. En. Philip. 3 (1923) 312.
—S. similis Hitt, Kew Bull. (1912) 38; ibid.
(1917) 153; Merr. En. Philip. 3 (1923) 313.—
S. pycnoneura GitG & BENED. Bot. Jahrb. 54
(1916) 164, f. 4; Hitt, Kew Bull. (1917) 162.—
S. cinnamophylla GitG & BENED. Bot. Jahrb. 54
(1916) 166; Hitt, Kew Bull. (1917) 148.—S.
myriantha GitG & BENED. Bot. Jahrb. 54 (1916)
167, f. 5; Hitt, Kew Bull. (1917) 162; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 303.—S. leu-
coneura GILG & BENED. Bot. Jahrb. 54 (1916)
169.—S. curtisii K. & G.: HILL, Kew Bull. (1917)
164, pro fr.; Ript. Fl. Mal. Pen. 2 (1923) 424,
pro fr.—S. silvicola Hitt, Kew Bull. (1930) 156;
in Craib, Fl. Siam. En. 3 (1951) 61.—S. polytoma
(non GILG & BENED.) KAN. & Hat. Bot. Mag.
Tokyo 53 (1939) 11.—Fig. 31.

Liana, sometimes a tree (?). Twigs glabrous,
branchlets greyish and fairly smooth. Leaves
ovate or suborbicular to lanceolate, 8-25 by 3-12
cm, chartaceous to stiff-coriaceous, glabrous,
base subcordate to broadly cuneate, apex blunt to
rounded and rather abruptly shortly blunt-acu-
minate to gradually acute-acuminate, sometimes
even caudate; 3-—5(—7)-plinerved, inner nerves
usually diverging fairly high above the base (up
to c. 2 cm) and in the eastern races usually at a
different height, venation often coarse-scalariform;
petiole 3-15 mm. Inflorescences axillary and/or
terminal, usually large (up to c. 15 cm long),
widely branched and many-flowered (in some
western races only a few cm long and with some
flowers), partly or completely thinly tomentose or
subglabrous. Ca/yx 1-3 mm, outside (glabrous to)
sparsely to densely short-tomentose, inside gla-
brous, sepals broad-ovate to suborbicular, blunt to
rounded, rarely acute. Corolla 3-61, mm long,
about halfway connate, outside glabrous (except
sometimes in young buds) inside at least about the
mouth, at most from about halfway the tube to the
mouth, rarely also on the lobes, (mostly densely)
woolly, lobes slightly thickened only. Stamens in-
serted in the mouth, filament up to 1!4 mm, gla-
brous, anther ovate to ovate-lanceolate or elliptic
to oblong, 34-1142 mm long, blunt or minutely
apiculate, glabrous or barbate. Pistil 2-6 mm,
ovary completely or for the greater part hairy, style

356

FLORA MALESIANA

[ser. I} vole'62

often glabrous, sometimes in the basal half, rarely
completely hairy, indument sparse to fairly dense,
shaggy to woolly. Fruits many, globular, 21-3
cm 9, sometimes ellipsoid and 3!4 cm long, thin-
walled, glabrous and smooth to minutely warty.
Seeds 1-8, often 2, lenticular, 12-15 by 10-12 by
2-4 mm, minutely densely tomentose.

Distr. From Ceylon, India, Cochin-China,
through Malaysia: Malay Peninsula, Simalur I.
off NW. Sumatra, Borneo, Philippines, Celebes,
Moluccas, New Guinea, and New Britain, to the
Solomon Is. and Australia (NE. Queensland, mainly
the coastal region between Cairns and Innisfail).

Ecol. Primary and secondary forests, in New
Guinea also in mixed Araucaria forests, from
sea-level up to 1850 m. Fi. mainly July—Nov.,
fr. Jan.—Dec.

Uses. The wood, bark, and roots are medic-
inally used (snake-wood). A decoction of these
parts is used as a poison for arrows, the vine for
tying purposes.

Vern. Snake-wood, E, tali siri (kétjil), M, olor
sikasa ilir dotan, Simalur, léngkoyan, sémijo akar,
Mal. Pen., tao kwang du tuk, Mal. Pen. Surat,
ipu ako, i. tanah, Born.; Philip.: abukobuko,
bukuan, \bn., batlag, Pint., Sbl., batul, pamula-
klakin, tibanglan, Tag., bugahin, Bis., bukuan,
Neg., dafigilian Bag., fantandok, i\l., malaitmo-
a-balaein, Dun.; ranosandang, Cel., akar pamali,
aywda umali, hutuu, wale ammelaum, wari ammo,
Moluccas.

Notes. Very variable species, comprising a
great number of local races. Many of these races
were treated by HILL as separate species, and some
more he either did not have available or could not
identify. The differences between these races are
mostly very small and mainly concern the follow-
ing characters: larger or smaller flowers, corolla
more or less pubescent, anthers barbate or gla-
brous, style either glabrous or partly or entirely
hairy, furthermore differences in shape, size,
nervation of the leaves, and larger or smaller in-
florescences. Many of these characters proved to
be variable in one or some of the races and are
therefore useless for specific distinction. Even
only very few races can clearly be characterized:
best recognizable are ‘S. bancroftiana’ and ‘S.
forbesii’ from Australia and New Guinea, with
oblique, oblong to lanceolate leaves, the inner
nerves of which distinctly diverge at a different
height, and ‘S. multiflora’ from the Philippines
with large and many-flowered inflorescences,
large flowers, and large, ovate, coarsely reticulate
leaves. Towards the west the leaves, inflorescences,
and flowers are smaller; moreover, the leaves of
the continental Asiatic forms are usually narrow
and long-acuminate.

Though I did not see good specimens of S.
merrillii, 1 scarcely doubt its conspecificity with
the present species; the only difference, according
to HILL’s description, is apparently the pubescence
of twigs, leaves, and the outside of the corolla.

The nearest relative of S. colubrina is apparently
S. quadrangularis, different by the glabrous pistil
and seed.

12. Strychnos quadrangularis Hitt, Kew Bull.
(1917) 205; Rip. Fl. Mal. Pen. 2 (1923) 426;
Burk. Dict. (1935) 2099.

Climbing shrub or liana. Twigs slender, mostly
sharply quadrangular, glabrous; branches rounded
quadrangular. Leaves (ovate-)oblong to elliptic,
13-20 by 5!4-11 cm, thin-chartaceous to papy-
raceous, glabrous, beneath copper-coloured to
yellowish brown or green when dried, base cuneate
to rounded, apex (caudate-)acuminate; 3—5-
plinerved; petiole 4-5 mm. Jnflorescences axillary,
4-6 cm long, laxly branched, fairly many-flowered,
minutely pubescent. Sepals broadly ovate, acute,
glabrous on either side. Corolla small; stamens
unknown. Pistil glabrous. Fruits globular, c. 2 cm
©, said to be white. Seed 1, semiglobular, c. 1 cm
@ and 8 mm thick, glabrous.

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

Ecol. Dense jungle, from the lowland to above
1000 m. Fr. Sept.

Uses. The bark of the roots is used for arrow-
poison. See BurRKILL, /.c.

Vern. Akar ipoh chelak, a.i. padi, a.i. siat
gunong, a.i. tontong, ipoh aker.

Note. Inadequately known, doubtless closely
related to S. colubrina, mainly different by the
glabrous pistil and seed, both characters being
usually constant.

13. Strychnos borneensis LEENH. Bull. Jard. Bot.
Brux. 32 (1962) 458.—S. laurina (non DC.)
HEINE, Pfl. Samml. Clemens Kinabalu (1953) 90.—
Fig. 32.

Liana. Twigs glabrous; branchlets (yellowish)
brown, smooth or scabrous by many small lenti-
cels. Leaves oblong or elliptic-oblong, 714-914
by 3-4 cm, chartaceous, glabrous, base cuneate,
slightly attenuate, apex gradually to caudate
acute-acuminate, acumen up to 14 cm; 3-—5-
plinerved, nerves diverging at or slightly above the
base, distinct, not far from the margin, venation
dense, transverse; petiole 6-7 mm. Inflorescences
axillary, 2-7 cm long, larger ones long-peduncled
and with a pair of strong branches at the base,
laxly branched, relatively few-flowered, at least the
terminal parts fairly densely rusty tomentose.
Calyx 114-134 mm, inside sometimes at the base
with a line of erect hairs, sepals ovate, rounded to
nearly truncate. Corolla c. 9 mm long, halfway
connate, very thin, outside glabrous, inside dense-
ly woolly from slightly above the base to about
halfway the lobes, most densely so in the mouth.
Stamens inserted in the mouth, filament 1144-214
mm long, glabrous, anthers elliptic to lanceolate,
1144-134 mm long, minutely apiculate, glabrous
or with a few hairs at the base of the cells. Pistil
8-9 mm, fairly densely shaggy short-hairy from
about halfway the ovary upwards. Fruit (pos-
sibly not quite normal and galled!) on a strongly
thickened branch, globular, c. 2 cm 9, slightly
scabrous. Seeds unknown.

Distr. Malaysia: North Borneo.

Ecol. Forests, from sea-level to c. 1600 m. Fi.
Novy., Jan.

Dec.

LOGANIACEAE (Leenhouts)

357

Fig. 32. Strychnos borneensis LEENH. a. Habit,

x %, b. flower,
d. stamen,

« 4, c. opened corolla, x 4,
< 8, e. pistil, < 4 (all from Puasa
4190).

Vern. Bina, kabuh.

Note. A characteristic species, possibly related
to S. polytrichantha, but differing by the corolla
of which the tube is about half the total length.
The leaves are very similar to those of S. vanprukii
Craip (Siam).

14. Strychnos ledermannii GitG & BENED. Bot.
Jahrb. 54 (1916) 169; Hitt, Kew Bull. (1917) 179.

Liana. Twigs subglabrous; branchlets grey,
smooth. Leaves ovate to oblong-elliptic, 7-13 by
3-514 cm, coriaceous to chartaceous, glabrous,
base cuneate to broadly rounded, apex gradually
acute-acuminate; 5—7-plinerved above the base,
nerves irregular, venation coarse; petiole 5-7 mm.
Inflorescences axillary and terminal, laxly branch-
ed, up to c. 5 cm long, few-flowered, subgla-
brous. Flowers seen in bud only. Calyx 1 mm,
sepals elliptic to broadly ovate, blunt, glabrous
on both surfaces. Corolla 3 mm, halfway up con-
nate, outside glabrous, inside woolly about the
mouth. Stamens inserted in or slightly below the
mouth, anthers subsessile, elliptic, 34-1 mm,
glabrous. Pistil 244 mm, with some patent long
hairs on the style. Fruit unknown.

Distr. Malaysia: New Guinea (NE. New Gui-
nea, Etappenberg).

Ecol. Dense forest, 850 m. Fi. Oct.

Notes. Hitt, who did not see any specimen,
placed this incorrectly in his section Penicillatae.

The only one of the syntypes apparently still
preserved, LEDERMANN 9435 (dupl. in K, L),
I have indicated as lectotype.

15. Strychnos lanceolaris M1Q. Sum. (1861) 551 &
227; Hitt, Kew Bull. (1911) 295.

Probably a liana. Twigs glabrous; branchlets
yellowish, scabrous. Leaves lanceolate, 614-9 by
2-214 cm, coriaceous, glabrous, base acute, apex
caudate-acuminate, acute; 3-plinerved above
the base, nerves usually not far from the margin,
veins transverse, minute, and dense; petiole 4-6
mm. Inflorescences axillary, lax, c. 24 cm long,
narrow and with few flowers, thinly tomentose.
Calyx 1% mm, outside sparsely pubescent, inside
glabrous. Corolla 3 mm, halfway up connate,
outside very sparsely pubescent, inside with some
short woolly hairs near the apex. Stamens inserted
in the mouth, filaments short, glabrous, anthers
broad-ovate, 44 mm long, apiculate, slightly
barbate. Pistil 2 mm, sparsely pubescent. Infructes-
cences with strongly thickened branches, torus |
cm thick. Fruits probably large and many-seeded.

Distr. Malaysia: Sumatra.

Vern. Téméras akar.

Note. A species of unclear relationships.

16. Strychnos flavescens K. & G. J. As. Soc. Beng.
74, ii (1908) 617; Hitt, Kew Bull. (1917) 155;
Ript. Fl. Mal. Pen. 2 (1923) 423.

Liana. Twigs usually glabrous, sometimes.
sparsely minutely pubescent; branchlets whitish
yellow-brown, fairly smooth. Leaves elliptic-
lanceolate to ovate-oblong, 5-12 by 3-614 cm,
thin-chartaceous to coriaceous, glabrous, very
glossy light yellow-green when dried, base cuneate
to subcordate, slightly attenuate, apex gradually
acuminate, more or less caudat:, acute; 3-plinerved
at the base, nerves near the margin, distinctly
more faint than the midrib; petiole 14-1 cm.
Inflorescences terminal and axillary, up to 10 cm
long, laxly branched, many-flowered, sparsely
minutely pubescent. Flowers (4—)5-merous. Calyx
1 mm, outside with some scattered hairs, inside

358

FLORA MALESIANA

[ser. I, vol. 6?

glabrous, sepals ovate, blunt. Corolla 2-3 mm,
halfway connate or somewhat less, outside thinly
tomentose, inside about the mouth woolly or
fully glabrous; lobes often strongly thickened.
Stamens inserted in the mouth, anthers subsessile,
ovate, 14-24 mm, minutely apiculate, glabrous or
barbate. Pistil 1144-2 mm, glabrous or the style
slightly hairy. Fruits many, subglobular, c. 114-2
cm 9, smooth or slightly warty. Seeds 1 or 2, len-
ticular, 14-16 by 1114 mm, thin.

Distr. Malaysia: Malay Peninsula, Banka.

Ecol. Swampy forests, along streams, at low
altitudes. F/. April-May, Nov.

Vern. Akar kalikut, kélikuh, Banka.

17. Strychnos luzonensis Erm. Leafl. Philip. Bot.
1 (1908) 332, p.p.; em. Hitt, Kew Bull. (1911)
297; ibid. (1917) 180, cum fig.; MERR. En. Philip.
3 (1923) 312, p.p.

Scandent shrub. Twigs thinly, minutely pu-
bescent. Leaves oblong-ovate, 714-10 by 314-4
cm, thin-chartaceous, midrib on both sides slightly
pubescent near the base or glabrous beneath;
base rounded, attenuate, apex tapering acute-
acuminate; 3—5-plinerved; petiole 744 mm. In-
florescences terminal, with a pair of strong branches
in the upper leaf-axils, c. 5-7 cm long, lax,
many-flowered, thinly short-pubescent. Calyx 1
mm, outside subglabrous, inside glabrous, sepals
ovate, blunt to acute. Corolla 21 mm, tube 1 mm,
lobes thickened towards the apex, inside with a
line of erect, bristle-like hairs about halfway the
free lobes (these hairs sometimes missing in older
flowers!). Stamens inserted in the mouth, fila-
ments 14 mm, glabrous, anther broad-ovate
(subquadrangular), 4% mm, apiculate, slightly
barbate. Pistil 14, mm, thinly pubescent. Fruit
unknown.

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

Ecol. Humid forest, 500 m. F/. May.

Notes. The combination of a short corolla-
tube and a line of erect bristle-hairs is unique
among the Malaysian species.

As mentioned already by HILL, ELMer included
in his original description two different elements;
the above description has been based exclusively
upon ELMER 8251, in accordance with HILL’s
emendation. The other specimen, ELMER 7885, is
here referred to S. colubrina.

Vegetatively, S. /Juzonensis closely resembles
S. ovata; the latter species is inter alia different by
the glabrous twigs and the often blunt leaves with
the nerves closer to the margin.

18. Strychnos axillaris CoLesr. Trans. Linn. Soc.
12 (1819) 356, t. 15; WALL. in Roxb. FI. Ind. 2
(1824) 266; BL. Bijdr. (1826) 1019; CLARKE in
Hook. f. Fl. Br. Ind. 4 (1883) 89; HiLL, Kew Bull.
(1917) 169.—S. malaccensis BtTH. J. Linn. Soc.
Bot. 1 (1856) 101; GAmB LE, J. As. Soc. Beng. 74,
ii (1908) 618; Hitt, Kew Bull. (1917) 177; Rip.
Fl. Mal. Pen. 2 (1923) 425.—S. horsfieldiana Mia.
Fl. Ind. Bat. 2 (1857) 379; HiLL, Kew Bull. (1911)

294, cum fig.; ibid. (1917) 179; BAKH. f. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 7.—
S. monosperma Mia. Fl. Ind. Bat. 2 (1857) 381;
Hitt, Kew Bull. (1917) 207.—S. palembanica
Mig. Sum. (1861) 551 & 227; Hitt, Kew Bull.
(1911) 293; ibid. (1917) 179; CAMMERL. Bull.
Jard. Bot. Btzg III, 5 (1923) 304, excl. spec.
Simalur.—S. psilosperma F. vy. M. Fragm. 4
(1863) 44; Bru. Fl. Austr. 4 (1869) 369; F. M.
BAIL. Queensl. Fl. 3 (1900) 1024; Compr. Cat.
Queensl. Pl. (1913) 337, f. 316; HiLL, Kew Bull.
(1917) 171 & 341.—S. pubescens CLARKE in Hook.
f. Fl. Br. Ind. 4 (1883) 89; Gamste, J. As. Soc.
Beng. 74, ii (1908) 620, incl. var. scortechinii
K. & G.; Hitt, Kew Bull. (1911) 292; Dop, FI.
Gén. I.-C. 4 (1914) 165; Hitt, Kew Bull. (1917)
166; Ript. Fl. Mal. Pen. 2 (1923) 424; Burk.
Dict. (1935) 2098.—S. pilgeriana GiLG, Notizbl.
Berl.-Dahl. 1 (1897) 268.—S. rufa CLARKE: GAM-
BLE, J. As. Soc. Beng. 74, ii (1908) 617, p.p.—
S. schmidtii G1LG, Bot. Tidsskr. 32 (1915) 312;
HILL, Kew Bull. (1917) 170; in Craib, Fl. Siam. En.
3 (1951) 61.—? S. oophylla GitG & BENED. Bot.
Jahrb. 54 (1916) 170, f. 6; Hitt, Kew Bull. (1917)
181.—S. polytoma GitG & BENED. Bot. Jahrb.
54 (1916) 173, f. 7; HmLL, Kew Bull. (1917) 182;
non KAN. & Hat. Bot. Mag. Tokyo 53 (1939)
11 (= S. colubrina).—S. wenzelii MERR. Philip.
J. Sc. 11 (1916) Bot. 202; HiL_, Kew Bull. (1917)
178; Merr. En. Philip. 3 (1923) 313.—S. quintu-
plinervis Hitt, Kew Bull. (1917) 166; Ripv. FI.
Mal. Pen. 2 (1923) 424.—S. robinsonii HILL, Kew
Bull. (1917) 168.—S. scortechinii Hitt, l.c., cum
fig.; Ripe. Fl. Mal. Pen. 2 (1923) 425; Burk. &
Hot. Gard. Bull. S.S. 3 (1923) 61.—S. armata
Hitt, Kew Bull. (1917) 170.—S. plumosa HILL,
Lc. 171, cum fig.; in Craib, Fl. Siam. En. 3 (1951)
60.—S. arborea Hitt, Kew Bull. (1917) 172,
cum. fig.—S. penicillata Hix, lc. 178; Ripe. Fl.
Mal. Pen. 2 (1923) 425; Burk. Dict. (1935)
2098.—S. impressinervis H1LL, Kew Bull. (1917)
180; Merr. En. Philip. 3 (1923) 312.—S. mucrona-
ta HILL, Kew Bull. (1917) 181; in Craib, Fl. Siam.
En. 3 (1951) 58.—S. tesseroidea HILL, Kew Bull.
(1917) 206; Merr. En. Philip. 3 (1923) 313.—
S. cenabrei MerR. Philip. J. Sc. 20 (1922) 433;
En. Philip. 3 (1923) 312.—S. viridiflora HILL,
Kew Bull. (1925) 424, nom. illeg., non DE WILD.
(1923).—S. kawbet HILL, Kew Bull. /.c. 425; in
Craib, Fl. Siam. En. 3 (1951) 58.—S. ovata (non
HILL) Merr. Pl. Elm. Born. (1929) 252.—S.
chloropetala HILL, Kew Bull. (1930) 175; HENDERS.
J. Mal. Br. R. As. Soc. 17 (1939) 58; HiLu in
Craib, Fl. Siam. En. 3 (1951) 57.—Strychnos sp.
HILx in Craib, Fl. Siam. En. 3 (1951) 62.—Fig.
30g.

Usually a liana, sometimes a shrub or small tree.
Twigs densely to sparsely minutely pubescent and
glabrescent or glabrous; axillary spines sometimes
present. Leaves rhomboid to suborbicular and
elliptic to lanceolate, 134-18 by 1-9 cm, charta-
ceous (rarely + herbaceous) to coriaceous, smooth
to densely minutely warty on the lower surface,
above glabrous to tomentose on the midrib and
the bases of the nerves, beneath mainly on mid-

Dec. 1962]

LOGANIACEAE (Leenhouts)

359

rib, nerves, and veins, sometimes sparsely all over
the leaf; base cuneate to subcordate, often attenu-
ate (in Australian specimens sometimes decurrent
to the base of the petiole), apex tapering acute-
acuminate to slightly emarginate, occasionally
mucronate; 3—S-plinerved at the base; petiole 1 '4—
74% mm. Inflorescences axillary and/or terminal,
1—5 cm long, subsessile or distinctly peduncled,
dense to lax, always fairly many-flowered, densely
minutely tomentose to glabrous. Ca/yx 1-2 mm,
sepals ovate to suborbicular, acute to rounded,
outside glabrous to densely tomentose, inside
glabrous. Corolla 3-4 mm long, about halfway
connate, lobes strongly thickened in their upper
part, outside glabrous or very rarely sparsely
tomentose, inside with a row of erect bristle-like
hairs at the base of or up to halfway on the lobes.
Stamens inserted at c. 14 the length of the corolla
below the hair ring, filaments short, glabrous, an-
thers (broadly) ovate, 24-1 mm long, usually
minutely apiculate, barbate, rarely moreover
ciliate all around. Pisti/ 114-2 mm, glabrous.
Fruits some to many, either oblique-ovoid, 11-2
by 34-114 cm, and slightly scabrous, or globular,
34-114 cm 9 and smooth, pericarp thin, corneous.
Seeds 1-2, either elliptic, 714-15 by 6-10 by 3-5
mm, or orbicular, 6-714 mm g.

Distr. Continental SE. Asia from Assam
through Siam to Cochin-China, throughout
Malaysia (except the Lesser Sunda Is.) to Austra-
lia (E. Queensland from about Innisfail to the Mac-
Pherson Range just over the NSW frontier).

Ecol. In and along primary and secondary
forests, in dry as well as in swampy localities,
sometimes on coastal rocks, up to 2000 m. Fi.
mainly March—Aug,, fr. Jan.—Dec.

Uses. In the Malay Peninsula apparently some-
times used for the composition of an arrow-
poison.

Vern. Akar kuling, Sum., akar bidara utan,
a. ipoh, a. lada lada, bédara utara, bélai bésar,
ipoh batang, i. burong, kio ngu, lep rawk, opoh
batang, Mal. Pen., tjantélan, J, tangang, Born.;
Philip.: malaigasud, S. L. Bis., (mara) igasud;
aputu ladwan, Talaud, kebrai pakrik, New
Guinea.

Notes. This is the most variable species of the
genus in Indo-Malaysia; it comprises a fairly
large number of local races, most of them dis-
tinguished by Hitt (1917 and later) as different,
though related, species. In my opinion they can
be treated only as local races as most of the dif-
ferences between these forms are either gradual or
concern vegetative characters only; the flowers are
surprisingly uniform. Only the fruits— and accord-
ingly the seeds — show two distinctly different
forms which are geographically almost replacing
one another: the oblique-ellipsoid, large fruits
with large elliptic seeds are found in Asia and
West Malaysia, reaching as far east as the Phi-
lippines (‘S. tesseroidea’) in the north and the
Tanimbar Is. (‘S. horsfieldiana’) in the south;
the smaller globular fruits with orbicular seeds
are known from the Philippines, East Malaysia,
and Australia. I do not set much value on these

fruit and seed characters as they do not correlate
with others; besides from many forms no ripe and
undamaged fruits are available.

The races can be arranged into four groups
which possibly deserve the status of subspecies;
part of the races may be subordinated as varieties;
in the present state of our knowledge I prefer not
to go too far into details.

These four groups are:

Group A. Spines often present. Leaves usually
oblong to lanceolate, about rhomboid with the
greatest width near the base, mostly thinly char-
taceous, black when dried, smooth and usually
glabrous; base broadly cuneate, apex acute;
mostly 3-plinerved, veins inconspicuous or in-
visible. Inflorescences axillary, rarely moreover
terminal. Fruits oblique-ellipsoid. Seeds elliptic.
Throughout Siam and Cochin-China, the Malay
Peninsula, and Sumatra.

Especially the species distinguished by HILL in
this group can only be separated either with great
difficulty, or by characters which in my opinion are
of very slight importance. ‘S. armata’ is charac-
terized by the spines which are more often present
and more numerous than in the other forms;
*S. plumosa’ is the form with twigs which are most
densely tomentose, but only *S. schmidtii’ is fully
glabrous. ‘S. kawbet’? and ‘S. chloropetala’ are
nearly indistinguishable from ‘S. plumosa’.

Two forms deserve some more attention as
they are the links with two other groups. They are
the only representatives of group A in Malaysia.
The first is ‘S$. scortechinii’ from Malaya, which is
aberrant in this group by the pubescent leaves
with the nerves diverging distinctly above the base,
and which apart from the colour of the dried
leaves could as well be included in group B. The
second is *S. quintuplinervis’ from Malaya and
Sumatra, with larger and firmer leaves, more
nerves, and coarser veins; it is intermediate
between groups A and C.

Group B. Spineless. Greatest width of the
leaf usually slightly below or about the middle,
either rounded and attenuate, or very broadly
cuneate towards the base, apex usually more or
less distinctly acute-acuminate; mostly coria-
ceous and fairly stiff, the dried leaf olive- or
yellowish-brown, apparently in most forms full
of cystoliths, as the lower surface is often densely
minutely warty; usually pubescent on the nerves
beneath, often thinly pubescent all over the lower
surface; 3—5-plinerved at some distance above the
base, venation visible beneath, but minute and
not conspicuous. Inflorescences axillary and/or
terminal. Fruits oblique-ellipsoid, seeds elliptic.
Assam, Malay Peninsula, Borneo, Palawan,
Sumatra, Java, and Tanimbar Is. The main races
distinguishable are:

S. axillaris sens. str. (incl. also S. palembanica
and S. horsfieldiana), best characterized by the
rarely absent pubescence in the axils of the ner-
ves beneath; occurs in Assam, Malay Peninsula,
Sumatra, Java, and the Tanimbar Is.; it is nearly
indistinguishable from ‘S. impressinervis’ from
Palawan.

360

FLORA MALESIANA

[ser I vole?

*S. penicillata’, characterized by the distinctly
densely and minutely warty lower surface of the
leaves, with scattered appressed hairs, and by
nearly exclusively terminal inflorescences, is
restricted to the Malay Peninsula (apparently con-
fined to Perak and P. Penang).

“§. pubescens’ mainly differs from the preceding
form in the lower surface of the leaves which is
less rough, but with all the hairs inserted upon a
larger and distinct wart; inflorescences are in
Malayan specimens always axillary, in Bornean
ones usually moreover terminal; it occurs in SW.
Malay Peninsula (Singapore to Selangor) and
Borneo.

‘S. malaccensis’, differing from ‘S. penicillata’
mainly by the smooth and glabrous leaves which
are thick and shining (those of ‘*S. penicillata’
are usually dull) occurs in the Malay Peninsula
(Malacca and Singapore).

Group C. Spineless. Leaves on the average
distinctly larger than in the preceding two groups,
greatest width in or slightly below the middle,
towards the base either slightly attenuate, or very
broadly cuneate to rounded, apex acute or slightly
acute-acuminate; firmly chartaceous, dried leaves
usually shining brown to purplish-brown beneath,
grey to purple-black above, smooth and glabrous;
3—5-plinerved, nerves diverging only slightly above
the base, main veins transverse, strong, conspic-
uous beneath. Inflorescences axillary. Fruits
globular and seeds orbicular as far as known.
Borneo, Philippines, and Moluccas (Halmahera).

This group can be considered to represent one
race; in HILL’s classification it corresponds to
‘S. wenzelii’.

One specimen, PNH 12436, is aberrant by a
row of hairs at the base inside the calyx.

Group D. Spines in Australian specimens some-
times present, in Malaysian ones absent. Leaves
variable in shape, size, nervation, and colour,
always coriaceous; base and apex very variable.
Inflorescences variable, but often relatively large
and lax. Fruits globular, seeds orbicular. East
Malaysia and Australia.

This eastern group is morphologically less
clearly defined than the other ones, yet it is not
well possible to subdivide it into well defined races,
like the first two groups.

Two specimens from Celebes represent the most
western and morphologically also the most de-
viating form. They are characterized by large
ovate leaves, with a subcordate base and a blunt
apex, and by axillary, laxly branched inflores-
cences c. 5cm long, with a peduncle of about 2 cm.
These specimens show also some relationship to
“S. wenzelii’ (group C).

‘S$. robinsonii’ (Celebes and Ambon) and ‘S.
polytoma’ (New Guinea) are more or less inter-
mediate between these specimens from Celebes
and the Australian races.

The Australian races ‘S. psilosperma’ and
‘S. arborea are both characterized by the (appa-
rent) absence of tendrils, the occasional pre-
sence of spines, and furthermore by the fairly
small, mucronate leaves. ‘S. psilosperma’ is the

more variable race and is distributed all over the
area in Australia (moreover, part of the New
Guinean specimens are nearly identical); its
leaves are usually greenish when dried, they are
mostly 5-plinerved and thus relatively broad;
its inflorescences are mainly terminal, large, and
lax. ‘S. arborea’ is restricted to the vicinity of
Brisbane; the dried leaves are dark-brown, and
usually 3-plinerved, the base is cuneate and de-
current to near the base of the petiole; the in-
florescences are mainly axillary, small, and dense.

It is surprising that these Australian races,
occupying the southeastern part of the specific
range, are, apart from the fruits (those are,
however, unknown from most Asiatic forms),
nearly indistinguishable from the continental
Asiatic forms which occupy the northwestern
part of the vast area of this species.

I am not quite certain about the identity of
S. oophylla. The type is apparently lost; | saw two
specimens, which I think represent S. oophylla
(BEccARI sheet n. 6659, Fi; NGF 11008, L),
but these two specimens are different from
‘S. polytoma’, the common New Guinean race of
S. axillaris to which I have referred them with
some doubt. According to the original diagnosis
the corolla of S. oophylla would be only 144 mm
long, but from the figure it is clear that the flowers.
were in bud.

19. Strychnos ridleyi K. & G. J. As. Soc. Beng.
74, ii (1908) 621; non Dop, Bull. Soc. Bot. Fr. 57,
Mém. n. 19 (1910) 16 and Fl. Gén. 1.—-C. 4 (1914)
165 (p.p. = S. axillaris); Hitt, Kew Bull. (1917)
167; Ripv. Fl. Mal. Pen. 2 (1923) 424.

A climbing shrub. Twigs thinly pubescent.
Leaves oblong, rarely ovate-oblong, 8-10 by 4414
cm, thin-chartaceous, glabrous except for small
tufts in the nerve-axils beneath, base broadly
cuneate to rounded, apex gradually acuminate,
acumen short, broad, and acute; 3-plinerved from
the base, veins obliquely transverse; petiole 6—I1
mm. Inflorescences axillary, 144-2 cm long
(peduncle 34-1 cm), few-flowered, peduncle
fairly densely, the other parts thinly pubescent.
Calyx 1 mm, sepals suborbicular, acute, outside
glabrous, inside fairly densely appressed short-
hairy. Corolla 4 mm, up to 244 mm connate,
inside with a whorl of erect, bristle-like hairs at
the base of the lobes and fairly densely, shortly,
patently hairy above this whorl. Stamens inserted
halfway the corolla-tube, anthers subsessile,
broadly ovate, c. 0.8 mm long, bearded. Pistil
124 mm, glabrous. Fruit unknown.

Distr. Malaysia: Malay Peninsula (Singapore
only).

Ecol. Fl. May.

Note. Doubtless related to S. axillaris, but
distinctly different by the calyx and the corolla-
lobes which are pubescent inside.

Insufficiently known

Probably good species, but flowers unknown.

Dec. 1962]

LOGANIACEAE (Leenhouts)

361

20. Strychnos rufa CLARKE in Hook. f. Fl. Br.
Ind. 4 (1883) 89; GamBLE, J. As. Soc. Beng. 74,
ii (1908) 617, p.p.; Hitt, Kew Bull. (1917) 203;
RIDL. FI. Mal. Pen. 2 (1923) 426.

Large liana. Twigs patently rufous-hairy. Leaves
ovate to oblong, 9-161 by 4-614 cm, papyraceous
to chartaceous, above minutely pubescent mainly
on the midrib and the basal half of the nerves,
beneath sparsely patently pubescent all over the
surface, more densely so on nerves and veins,
moreover fairly densely minutely warty; base
rounded to faintly cordate, apex gradually acute-
acuminate; 3- to faintly 5-plinerved at the base;
petiole 3-4 mm. Flowers unknown, according to
the collector small and yellow. Infructescences ax-
illary, c. 2 cm long, with thick branches and strong-
ly broadened torus, pubescent like the twigs; |
or few fruits. Fruits globular, c. 314-4 cm 9, red.
Seeds many, elliptic, flattened, 19 by 1214 mm.

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

Eco]. Mountain forest.

Uses. Dart-poison.

Vern. Akar ipoh tanah.

Note. The pubescence and the warty lower
surface of the leaf remind of S. axillaris (‘penicil-
lata’), but the shape and size of the leaves, and
above all the large and many-seeded fruits are
quite different. The latter might point to alliance
with S. ignatii.

21. Strychnos thorelii PieERRE ex Dop, Bull. Soc.
Bot. Fr. 57, Mém. n. 19 (1910) 20; Fl. Gén. I.-C.
4 (1914) 171, t. 3 f. 11-13; H1LL, Kew Bull.
(1917) 207; in Craib, Fl. Siam. En. 3 (1951) 62.
Liana. Twigs thinly patently pubescent, gla-

414-10 by 2-4 cm, chartaceous to coriaceous,
shining above, sometimes sparsely patently hairy
on the midrib beneath, otherwise glabrous;
base broadly cuneate to subcordate, slightly
attenuate, apex gradually acute-acuminate; 3-—5-
plinerved above the base; petiole 14-1 cm,
pubescent. Jnfructescences axillary and terminal,
up to c. 10 cm long, lax, minutely pubescent,
with some fruits only, branches slender, torus
somewhat broadened. Fruits ovoid to oblong-
ellipsoid, 2144 by 114 cm, thin-shelled. Seed 1,
elliptic-lenticular, 2 by 144 cm.

Distr. Cochin-China, Siam, and Burma, in
Malaysia: northern part of the Malay Peninsula.

Ecol. In and along evergreen and bamboo
forests, up to c. 200 m. Fr. March—April.

Note. I have designated as a lectotype PIERRE
1703 (P) (erroneously cited as 1702 by Dop).

Dubious
No material seen, description insufficient.

Strychnos melanocarpa GitG & BENED. Bot.
Jahrb. 54 (1916) 172; HiLL, Kew Bull. (1917) 181.
Distr. NE. New Guinea (Sepik, Malu).
From the description it is not clear whether the
indument in the mouth consists of one row of
bristle-like hairs or is woolly. In the first-named
case this species would be closely related to S.
axillaris (different, however, by the hairy ovary),
otherwise it might be identical with S. colubrina.

Excluded
Strychnos grandis WALL. Cat. (1831) n. 4454,
nom. nud. = Anisophyllea grandis (BTH.) BURK.

brescent. Leaves oblong-ovate to lanceolate, (Rhizophoraceae).

6. GARDNERIA

WALL. in Roxb. FI. Ind. 1 (1820) 400; LeeNu. Bull. Jard. Bot. Brux. 32 (1962)
431.—Fig. 33.

Mostly climbing or creeping, glabrous shrubs. Twigs terete, sometimes with
4 faint ridges. Leaves petioled, connected by interpetiolar stipular rims; at the
base of the new shoots a number of persistent, acute-triangular bud-scales. Jn-
florescences axillary, dichasial, usually few-flowered or flowers solitary, sometimes
lax and many-flowered; bracts narrowly triangular, 1-2 mm long. Flowers long-
pedicelled, 4-5-merous. Calyx small, deeply lobed, lobes rounded, sometimes
acuminate, shortly ciliate, for the rest glabrous, inside with colleters at the base.
Corolla rotate, thin-fleshy, creamy to yellow, lobes valvate in bud. Stamens ex-
serted, filaments very short, flat; anthers bifid at the base, introrse, 4- or 2-celled
(if 4-celled the outer cells much bigger than the central ones). Pisti/ glabrous;
ovary small, 2-celled, with 1-4 ovules per cell; style mostly long and slender;
stigma capitate to 2-parted. Berry globular, red. Seeds 1 or more, elliptic to or-
bicular, concave on the hilar side, convex on the other; testa thin, endosperm
thick, cartilaginous.

Distr. Five species in SE. and E. Asia, from India to Central Japan and Java; in Malaysia | species.
Note. The relationship of this genus is clearly with Strychnos.

362 FLORA MALESIANA [ser. I, vol. 6?

Fig. 33. Gardneria ovata WALL. a. Habit, * 24, b. flower (note the cohering anthers), * 31/4, c. anther,

2-celled, x 6, d. ditto, lateral view, * 6, e. ditto, cross-section, < 6, f. calyx with pistil, * 8, g. ovary in

cross-section, showing the two ovules, * 16, /. fruit, x 114, i. seeds, x 2, /. ditto in length-section show-

ing the small embryo embedded in the endosperm, * 2, k. embryo, frontal and lateral view, * 4 (a herb.
L 908.127-599, b-g sine coll., s.n. in herb. E, h-k HENRY 10906).

Dec. 1962]

LOGANIACEAE (Leenhouts)

363

1. Gardneria ovata WALL. in Roxb. Fl. Ind. 1
(1820) 400; Pl. As. Rar. 3 (1832) 17, t. 231;
BuREAU, Fam. Logan. (1856) f. 32-34; Bru. J.
Linn. Soc. Bot. 1 (1856) 109; Kurz, Fl. Burm. 2
(1877) 227; Racrporski, Rozpr. Wydz. Mat.-
Przyrod. Ak. Umiej. Krak. 32 (1896) 315, f.
1-5; Bull. Int. Ac. Sc. Cracovie (1896) 206;
GAMBLE, FI. Madras 5 (1923) 869; KANJILAL &
Das, Fl. Assam 3 (1939) 320; BAKH. f. in Back.
Bekn. FI Java (em. ed.) 7 (1948) fam. 170, p. 9;
Kerr in Craib, Fl. Siam. En. 3 (1951) 63; LEENH.
Bull. Jard. Bot. Brux. 32 (1962) 437.—G.
wallichii WIGHT ex WALL. Pl. As. Rar. 3 (1832)
49, t. 281.—Fig. 33.

Vine or shrub. Leaves (ovate or) oblong to
lanceolate, 6—-13(—16) by 2!4-514(-8) cm, charta-
ceous to thin-coriaceous, acute and slightly de-
current at the base, gradually acute-acuminate to
caudate and sometimes mucronulate; nerves
4-10 pairs. Inflorescences erect to pendulous, 4-8
cm long, 1—3-flowered to laxly thyrsoid and more-

flowered, sometimes with a serial solitary flower
in the same axil; peduncle 34-2 cm, pedicels 14-2
cm, bibracteolate. Flowers 4-merous. Sepals
¥%4-14% by 144-2 mm. Corolla yellow to orange,
waxy, tube 14-114 mm long, lobes ovate to ellip-
tic, acute, inside papillose-pubescent, 3-414 mm.
Stamens inserted just above the base of the
corolla-tube (rarely halfway), anthers cohering
mutually, erect, broadly ovate or elliptic to ovate-
lanceolate, 114-4 mm long, truncate, 2-celled.
Ovary Ys-1144 mm, tapering into the 14-2 mm
long style, stigma faintly 2-4-lobed to 2-parted.
Ovules | per cell. Berry 34 cm g, 1- or 2-seeded.
Seeds orbicular, 44 cm @, smooth, dull grey.

Distr. NE. and SE. India, Ceylon, Yunnan,
Siam, and Malaysia: N. Sumatra (Tapanuli)
and W. Java (Preanger: Tjadas Malang), twice
collected.

Ecol. Along forest edges and climbing over
limestone rocks, at 50-2250 m. F/. March—June,
fr. May—Nov.

7. NEUBURGIA

BiumE, Mus. Bot. | (1850) 156; MARKGRAF, Bot. Jahrb. 61 (1927) 222.—Couthovia
A. GRAY, Proc. Am. Ac. Arts Sc. 4 (1859) 324.—Crateriphytum SCHEFF. ex KOORD.
[Nat. Tyd. N.I. 55 (1896) 345, nom. nud.| Med. Lands PI. Tuin 19 (1898) 540.—
Fig. 34-35.

Trees or shrubs; branches glabrous. Lateral branches mostly up to c. | cm high
coalescent with the main branch (fig. 341). Stipules interpetiolar, adnate to the
petioles, obdeltoid, in older leaves often split along a distinct suture. Leaves
petioled to subsessile; midrib often forked towards the apex, venation obscure.
Inflorescences terminal, thyrsoid, the ends cincinnate; bracts semi-amplexicaulous,
scale- to ridge-like, minute, ciliate. Flowers sessile, 5-merous. Sepals confluent at
base, suborbicular, mostly ciliate, inside at the base often with hairs and/or col-
leters. Corolla rotate to salver-shaped, white, tube thin-fleshy, the valvate lobes
thicker; with a hair ring in the mouth, inner surface of the tube either glabrous or
hairy. Stamens inserted on the upper half of the corolla-tube, usually just below
the mouth, included; filaments strap-shaped, usually very short, glabrous; anthers
introrse, 2-celled, cells up to about halfway free, the apices mostly, the bases some-
times ending in a short sterile tip, the latter often, the former rarely bearded.
Pistil mostly glabrous; ovary 2-celled with ~ ovules on a strongly thickened pla-
centa; style early caducous; stigma always about reaching the mouth, mostly +
ellipsoid, 14-44 mm high, faintly grooved, hollow (in N. sarcantha broadly trun-
cate). Fruits drupaceous, mesocarp dry, rather hard and very fibrous, these fibres
originating from the woody, rugose stone; cells slenderly spindle-shaped, slightly
curved. Seed one per cell (and usually only one per fruit), slenderly spindle-shaped.

Distr. About 10-12 spp., in East Malaysia (from the Philippines and Celebes to New Guinea),
Carolines, Solomons, New Hebrides, New Caledonia, and the Fiji Is. Cf. LEENu. Pac. Pl. Areas 38.

Ecol. Shrubs or small to medium-sized trees in the undergrowth or substage of primary, rarely
secondary forests, rarely canopy trees, often in marshy or temporarily drained localities, largely confined
to eyerwet lowland. The clustered, small, white, scented flowers probably attract insects. Once it has
been stated that the fruits were eagerly eaten by pigeons, but the rather dry, thick, corky mesocarp and
the hard pyrene make the impression that they are normally dispersed by water.

Notes. Neuburgia was assigned by BLUME to the Apocynaceae; he mentioned two species, N. tuber-

364 FLORA MALESIANA [ser. I, vol. 6?

Fig. 34. Neuburgia rumphiana LEENH. a. Habit (part of leaf showing the indumentum on the lower side),
< 24, b. flower, < 2, c. pistil, x 6, d. opened corolla, < 6, e. stamen, = 6, f. lateral and frontal view of
fruit, 1.—N. moluccana (BOERL.) LEENH. g. Longitudinal section of fruit, showing the one-seeded
fertile cell and the sterile one, 1.—N. celebica (KoorD.) LEENH. /. Cross-section of fruit (note the
corky mesocarp, the woody putamen, the two cells, one of which is fertile, and the soft tissue in the centre),

1.—N. corynocarpa (A. GRAY) LEENH. i. Detail of branching, showing the coalescence between the
lateral branches and the main branch, * 24 (a-e LAM 7787, f BW 7526, g Hort. Bot. Bog. IV-E-92,

h FRI Cel III-40, i SAUNDERS 195).

Dec. 1962]

culata and N. tubiflora. MARKGRAF, /.c., transferred the genus to the Loganiaceae; he already suggested
the identity with Crateriphytum. MERRILL (J. Arn. Arb. 23, 1942, 416) typified the genus by N. tubiflora.

Koorpers already realized the close relationship between the monotypic Crateriphytum (= N. mo-
luccana) and Couthovia. Of the two main generic differences he mentioned, the long corolla-tube provided
with a rim in the mouth, shared also by N. tubiflora, is still a conspicuous character, but I cannot attach
generic value to it; the rim is much less conspicuous than depicted by him.

CAMMERLOHER (Bull. Jard. Bot. Btzg III, 5, 1923, 306) laid special stress on a supposed third difference,
the 2-lobed stigma which is also clearly depicted by KoorDeErs. However, this appears to be an artefact
sometimes caused by the pressing of herbarium specimens; the stigma is slightly grooved and hollow
when mature, it ruptures fairly easily when pressed, and the papillae are of course found on the outside
of the ‘lobes’; in many young fruits I found an undamaged unlobed stigma!

LOGANIACEAE (Leenhouts) 365

KEY TO THE SPECIES

1. Corolla-tube c. 144 cm long.
2. Corolla-tube inside glabrous. Leaves broadly elliptic to obovate, 1714-30 by 11-22 cm. Nerves 6-8
pairs . 6. N. moluccana

2. Corolla- tube inside hairy excent at the base. Leaves oblong- ‘obovate, 16-20 by 8-10 cm. Nerves 4-6

PASM eG
1. Corolla-tube less than i cm long.

7. N. tubiflora

3. Style 414-6 mm. Calyx 234-4 mm high. Corolla 8-10 mm.
4. Leaves about orbicular. Corolla-tube inside laxly hairy from the insertion of the stamens to the

mouth. Anthers glabrous. Stigma broadly truncate .

2. N. sarcantha

4. Leaves lanceolate to elliptic. Corolla-tube inside glabrous. Anthers bearded. Stigma club-shaped to

ovoid .

3. Style up to 14 mm. _ Calyx up to 2h x mm n high. “Corolla up. to 6 mm.
5. Leaves thick and stiff, coriaceous when dry . 3 wae
5. Leaves thin, papyraceous to thin-pergamentaceous when ‘dry.

4. N. rumphiana

1. N. corynocarpa

6. Leaves elliptic to lanceolate, tapering at both ends, midrib reaching the apex. Fruit 5—6 cm long,

acute .

3. N. kochii

6. Leaves broad- -elliptic to broad- obov ate, blunt at the apex, midrib. forked near the apex. Fruit 314-4

cm long, blunt .

1. Neuburgia corynocarpa (A. GrAy) LEENH. nov.
comb.—Couthovia corynocarpa A. GRAY, Proc.
Am. Ac. Arts Sc. 4 (1859) 324; SEEMANN, FI. Vit.
(1866) 165, t. 32; GiLLespre, Bull. Bish. Mus.
n. 83 (1931) 28, f. 35; A. C. SmirH, Sargentia 1
(1942) 102.—Couthovia seemanni A. GRAY, Proc.
Am. Ac. Arts Sc. 5 (1862) 320; GILLespie, Bull.
Bish. Mus. n. 83 (1931) 29, f. 36.—Couthovia
densiflora K. Scu. Bot. Jahrb. 9 (1887) 215;
GitG & BENeED. ibid. 54 (1916) 178.—Couthovia
rhynchocarpa GiLG & BENED. /.c. 176.—Couthovia
terminalioides GILG & BENED. /.c. 177.—Couthovia
pachypoda GiLG & BENED. /.c. 178, f. 8.—Couthovia
brachyura GiLtG & BENED. l.c. 179; LANE-POOLE,
For. Res. (1925) 134.—Couthovia nymanii GILG
& BeENeED. Bot. Jahrb. 54 (1916) 180.—Couthovia
astyla GitG & BENED. /.c. 183.—Couthovia brassii
S. Moore, J. Bot. 67 (1929) 50.—Couthovia novo-
britannica Kan. & Hart. Bot. Mag. Tokyo 53
(1939) 9, f. 4; Merr. & Perry, J. Arn. Arb. 23
(1942) 410.—Couthovia collina A. C. SmiTH,
Sargentia 1 (July 1942) 101.—Couthovia leuco-
carpa MERR. & Perry, J. Arn. Arb. 23 (Oct.
1942) 412.—Couthovia yunzaingensis MERR. &
Perry, ibid. 29 (1948) 163.—Fig. 34i.

Tree, 4-40 m by up to 75 cm g, sometimes with
buttresses. Leaves oblong to broad-obovate or
orbicular, 614-17 by 3-12 cm, thin- to thick-
coriaceous, glabrous; base cuneate to subcor-
date, decurrent; apex blunt to rounded, rarely
short-acuminate; midrib distinctly forked or

5. N. celebica

not, nerves (4~-)7-9(-11) pairs; petiole 14-134
em. Inflorescences 4-10 cm long, widely branched
and many-flowered, glabrous, the terminal parts
sometimes sparsely tomentose, flowers often crowd-
ed. Calyx 1-214 mm, outside mostly glabrous, rar-
ely minutely pubescent, inside with or without
colleters and/or hairs at the base. Corolla outside
glabrous, 4-6 mm long, tube inside usually laxly
woolly at the insertion of the stamens, lobes
144-2144 mm long. Anther-cells free for c. 14-14,
hastate, 144-2 mm long, corniculate at apex,
glabrous to bearded at base. Style 1-114 mm,
stigma club-shaped, c. 4% mm. Fruits clavate,

blunt to acute, straight to curved, 134-4 by
3¥%4-14%4 cm, white.
Distr. Melanesia (Solomon Is., Fiji) and

Malaysia: New Guinea (also Kei and Aru Is.,
Los Negros, Admiralty Is., and New Britain).

Ecol. Primary (and sometimes old secondary)
rain- and swamp-forests, also on temporarily
flooded (freshwater) localities, from sea-level up
to c. 2000 m. According to SEEMANN the fruits are
much eaten by pigeons in Fiji. F/. mainly Jan.—
June, fr. Febr., July—Sept.

Uses. In the Solomon Is. the macerated bark
is applied to skin diseases.

Vern. Metan, Aru Is, masosen, Schouten L.,
kuma, rantiepi, Japen, aifim, Numfoor, badjagi,
belik, bokon, elieuw, falaka, fenam, gramgusun,
inkwam, katadi, konote, kumo, maduwi, mafu, mai-
long, male, mingo, namooi, niesebuk, pao, pegamba,

366

FLORA MALESIANA

[ser. I, vol. 6?

seruahika, situahiga, somuk, tamu, tani, teguk,
teitakka or tétaka, ufi, waffer waffer, yako(m),
New Guinea.

Notes. N. corynocarpa is a widespread species
and consists of a reticulate relationship of local
races, both in New Guinea and in the Pacific.
In the Papuan area (from the Aru Is. to the
Solomons) there is a fairly distinct clinal va-
riation as to the hairiness of the flower: the co-
rolla-tube is inside glabrous in the Aru Is., pubes-
cent in New Guinea and the Solomons, the an-
thers are bearded in the Aru Is. and New Guinea,
glabrous in part of the material from E. New
Guinea and in the Solomons. The variation in
Fiji is mainly altitudinal, “Couthovia collina
being a small-leaved form of mainly higher al-
titudes, N. corynocarpa sensu stricto with larger
leaves is mainly restricted to the lowlands; these
two forms are grading. As to the other species
described from the Pacific (from which I had not
enough material at hand to come to a definite
conclusion) Couthovia alata A. C. SmitH (Sar-
gentia 1, 1942, 104; Fiji) and Couthovia novo-
caledonica GILG & BENED. (Bot. Jahrb. 56, 1921,
547; New Caledonia) should probably, Couthovia
macroloba A. C. SmitH (/.c.; Fiji) and Couthovia
neo-ebudica GUILLAUMIN (J. Arn. Arb. 13, 1932,
23; New Hebrides: Aneityum) possibly be com-
bined with N. corynocarpa; Couthovia macrocarpa
A. C. SmitH (/.c. 105; Fiji) and Couthovia pachy-
antha A. C. SMITH (/.c. 106; Fiji) may represent
2 (or only 1?) separate, though related species.
Furthermore N. corynocarpa shows relationship
with N. sarcantha.

The type of Couthovia rhynchocarpa has wrongly
been described as lacking the hairs in the corolla-
tube.

Throughout the genus the style is rather early
caducous after fertilization, leaving a usually
somewhat broadened, flat scar; in one case GILG
& BENEDICT mistook this for a sessile stigma, and
based their Couthovia astyla on that character.

Large-leaved specimens can vegetatively be
distinguished from N. celebica not only by the
thicker leaves, but also by the more spaced, and
thus less numerous nerves.

2. Neuburgia sarcantha (GILG & BENED.) LEENH.
nov. comb.—Couthovia sarcantha GILG & BENED.
Bot. Jahrb. 54 (1916) 181, f. 9; non CAMMERL.
Nova Guinea 14 (1924) 116, quae est N. celebica.

Tree, 15-23 m by c. 30 cm; branches glabrous.
Leaves broadly obovate to nearly orbicular, 10-25
by 8-25 cm,' coriaceous, glabrous; base cuneate to
cordate, decurrent; apex rounded; midrib forked,
nerves 6—7 pairs; petiole 34-244 cm. Inflorescences
5-7 cm long, widely branched, fairly many-flower-
ed, glabrous. Ca/yx 234 mm, sepals thick-fleshy,
glabrous, inside with hairs and colleters at the
base. Corolla 8 mm long, outside glabrous, inside
laxly woolly between the mouth and the insertion
of the stamens, lobes 4 mm long. Anthers oblong,
2 mm long, the cells free for c. 144, corniculate at
apex, rounded at base, glabrous. Srfy/e thick-
cylindric, 444 mm, stigma broadly truncate, 14

mm thick, 114 mm @. Fruits ovoid, constricted at
base, 4 by 214 cm, blunt, white.

Distr. Malaysia: New Guinea (Upper Digul
River, Western Highlands, and Sepik River,
‘Felsspitze’ of the LEDERMANN-expedition, prob-
ably not far from Malu).

Ecol. In forests and Pandanus swamps, 400-—
1900 m. Fi. Sept., fr. Aug.

Vern. Bandih, mehmin.

Notes. GitG & Benepict had only material in
bud; for that reason the description given here
differs in several points from the original diagnosis.
When analyzing flower-buds of new material I
found some of the characters specially stressed
by them: a thick-fleshy corolla of which the tube
is glabrous inside.

As apparently both syntypes (LEDERMANN
13005 and 13056) are lost, I propose as a neotype
HOOGLAND & PULLEN 6207 (L, dupl. in Br,
Cans, and MEL), an excellent flowering specimen
which has mainly served for the description given
here.

N. sarcantha is best characterized by large
flowers and a truncate stigma. It seems to be
nearest related to N. corynocarpa.

3. Neuburgia kochii (VAL.) LEENH. nov. comb.—
Couthovia kochii VAu. Bull. Dép. Agr. Ind. Néerl.
10 (1907) 46.—Couthovia urophylla GILG & BENED.
Bot. Jahrb. 54 (1916) 176.—Couthovia undulati-
folia KAN. & Hart. Bot. Mag. Tokyo 56 (1942)
TSS etewee

Shrub or treelet, 2-4(—6) m; branches glabrous.
Leaves elliptic to lanceolate (sometimes oblan-
ceolate or slightly ovate), tapering at both ends,
13-1744 by 5-74 cm, thin-papyraceous, above
slightly scabrous (as are the stipules), beneath
glabrous or rarely sparsely, shortly appressed-
pubescent; base cuneate, decurrent; margin usually
slightly undulate; apex tapering acuminate, slender.
blunt to acute; midrib not forked; nerves (4—)
7-8(-9) pairs; petiole 144-114 cm. Inflorescences
114-5 cm long, densely to laxly branched, minutely
pubescent. Calyx 124-2 mm high, outside glabrous,
inside at the base usually with colleters, sometimes
with hairs. Corolla 44,-6 mm long, inside usually
glabrous or nearly so, lobes 114-3 mm long, out-
side glabrous or rarely slightly pubescent. Anthers
hastate, 1144-2 mm long, cells free for 4—/,,
corniculate at apex, not or only slightly bearded
at base. Style c. 1-144 mm, stigma slenderly
club-shaped, 34-1 mm. Fruits curved, lanceolate
to ovoid, flattened, tapering towards the acute
apex, 5—6 by 2-214 by 1 cm, red or white.

Distr. Malaysia: New Guinea.

Ecol. Primary and secondary forests, from sea-
level up to 1100 m. F/. March, May, Aug., Sept.,
fr. March, May, July.

Vern. Arora, osiwai.

Notes. Young long shoots are characterized by
4 ribs, decurrent from the leaf-bases.

The roughness of the stipules and the upper
side of the leaves is caused by numerous minute,
yellow warts, probably from cystoliths.

Ripe fruits seem to be red in Western New

Dec. 1962]

LOGANIACEAE (Leenhouts)

367

Guinea, white in the eastern half; this character
is not distinctly correlated with any other.

4. Neuburgia rumphiana LEENuH. Bull. Jard. Bot.
Brux. 32 (1962) 457.—Fig. 34a-f.

Shrub or treelet, 3-6 m high; branches glabrous.
Leaves elliptic to lanceolate, sometimes slightly
asymmetric, 16-25 by 434-14 cm, chartaceous to
thin-coriaceous, glabrous above, mainly on mid-
rib and nerves beneath sparsely and shortly ap-
pressed-pubescent, or glabrous; base acute, at-
tenuate; margin slightly undulate; apex shortly
acuminate, blunt or acute; midrib not forked,
nerves 9-15 pairs; petiole 34-3 cm. Inflorescences
c. 5 cm long, fairly dense, rather many-flowered,
densely shortly ferrugineous-pubescent, sometimes
glabrescent. Calyx 314-4 mm high, outside gla-
brous, inside at the base with hairs and colleters.
Corolla 8-10 mm long, glabrous except the mouth,
lobes 214-3 mm long. Anthers linear, 2-24, mm,
cells free for 44-14, corniculate and sometimes
with a few bristles at the apex, bearded at the
base. Style slender, 444-6 mm, stigma club-
shaped to ovoid, 34 mm. Fruits flattened-ellipsoid,
acute, 414-5 by 134-214 cm, orange(-brown).

Distr. Malaysia: West New Guinea (around
Hollandia and on Japen I.).

Ecol. Primary and secondary, sometimes
swampy forests below 100 m. FI. July, Oct., Febr.,
fr. July.

5. Neuburgia celebica (Koorp.) LEENH. nov.
comb.—Couthovia celebica Koorp. Med. Lands
Pl. Tuin 19 (1898) 537; Suppl. Minah. 1 (1918)
7, t.2; MerR. En. Philip. 3 (1923) 313.—Couthovia
calophylla GiLtG & BENED. Bot. Jahrb. 56 (1921)
548, f. 2; KANEH. FI. Micron. (1933) 313, f. 158;
J. Dep. Agr. Kyushu Imp. Univ. 4 (1935) 390.—
Couthovia sarcantha (non GiLG & BENED.) CAM-
MERL. Nova Guinea 14 (1924) 116.—Couthovia
toua Kan. Bot. Mag. Tokyo 46 (1932) 490;
Fl. Micron. (1933) 313, f. 159; J. Dep. Agr.
Kyushu Imp. Univ. 4 (1935) 390.—Couthovia
macrophylla Merr. & Perry, J. Arn. Arb. 23
(1942) 411.—Fig. 34h.

Tree or large shrub, 7-30 m by 35-80 cm g, often
with small buttresses; branches glabrous, rarely
densely tomentose. Leaves (mostly broad-)elliptic
to broad-obovate, 13-30 by 9-25 cm, herbaceous
to papyraceous, rarely chartaceous, glabrous or
sometimes partly or entirely densely short-tomen-
tose beneath; base subcordate to cuneate, de-
current; margin slightly undulate; apex round-
ed; midrib forked, nerves (7—)9—14 (—16) pairs;
petiole 1-3!4 cm. Inflorescences 15-25 cm long,
glabrous but for the minutely tomentose ultimate
branches, widely branched, many-flowered. Calyx
1-2 mm high, mostly glabrous, rarely outside more
or less densely minutely tomentose. Corolla
4-414 mm long, outside glabrous to sparsely
pubescent, tube inside pubescent near the insertion
of the stamens, lobes 114-2 mm long. Anthers
narrowly hastate, 114-114 mm, cells up to halfway
free, distinctly corniculate, glabrous. Style 1-1
mm, stigma capitate to club-shaped, 14 mm.
Fruits ellipsoid, usually stipitate, blunt or rarely

acute, 314-4 by 114-134 by 1'4 cm, orange to
red.

Distr. Micronesia (Carolines: Palau Is.) and
Malaysia: Philippines (Mindoro, Mindanao),
Celebes, Moluccas, and S. New Guinea.

Ecol. Dryland and swampy rain-forests, often
along river-banks, sometimes in Phragmites
swamps, in some localities periodically flooded by
fresh or salt water, from sea-level up to c. 400 m.
Fl. mainly Jan.—May, fr. Jan., March, July.

Uses. Timber sometimes used, mainly for
indoor-constructions.

Vern. Philip.: bali-bali, C. Bis., pagi-pagi, Mbo.,
salinuok, P. Bis., tanalak, Bag.; achéra lika,
dongkina puté, ééwé, léa léa, rondo rondo, Cel.,
a sili, Talaud I., tampel putih, Batjan, gérépiw,
porojijt, New Guinea.

Notes. Couthovia celebica was based by
KoorDERS upon two collections, viz KOORDERS
18628 and 18741 (cf. Koorp.-ScHum. Syst.
Verz. 3, 1914, 105). I have chosen as a lectotype
Koorpers 18741 (Bo; dupl. in L).

The specimens from the Carolines, described as
Couthovia toua, differ partly by slightly larger
dimensions of the flowers and fruits, and by
more coriaceous leaves. These characters, however,
are grading and some of the collections agree
entirely with Malaysian material.

6. Neuburgia moluccana (BOERL.) LEENH. nov.
comb.—? Fructus musculiformis RumpwH. Herb.
Amb. 2 (1741) 184, t. 60, pro fruct.—? Cerbera
musculiformis LAMK, Enc. 1 (1783) 62, pro fruct.
—non Banksia musculiformis GAERTN. Fruct. 1
(1788) 221.—? N. tuberculata Bu. Mus. Bot. 1
(1850) 157, nom. illeg —? N. musculiformis Mia.
FI. Ind. Bat. 2 (1856) 403, pro typo, excl. syn. N.
tubiflora; Merr. Int. Rumph. (1917) 425.—
Crateriphytum moluccanum SCHEFF. ex BOERL.
Handl. 2 (1899) 456, 460; Koorp. Bull. Inst.
Bot. Btzg 16 (1903) 10; Abb. Beschr. Crateri-
phytum moluccanum (1919) 3, t. 1; CAMMERL.
Bull. Jard. Bot. Btzg III, 5 (1923) 306.—Fig.
34g-35.

Shrub or treelet, 8-10 m high, rarely tree, up
to 30 m by 45 cm 9g, with buttresses; branches
glabrous. Leaves broad-elliptic (sometimes nearly
orbicular) to obovate, 12!4-30 by 8-22 cm,
thin-coriaceous, glabrous; base cuneate to cord-
ate, decurrent; margin slightly undulate; apex
rounded; midrib forked just below the apex,
nerves 6-8 pairs; petiole 1-4 cm. Inflorescences
5-11 cm long, widely branched, few- to many-
flowered, glabrous. Calyx 3-4 mm high, outside
glabrous, inside at the base with or without
colleters. Corolla 2 (when fresh 2.4) cm long,
slender, glabrous apart from the hair-ring in
the mouth which is inserted on a thickened,
lobed rim; lobes 14 cm long, very thick-fleshy.
Stamens inserted c. 4-7 mm below the mouth;
anthers linear, 34 mm, cells free for 14—/;,
faintly corniculate at apex and with an acute
membranous base, glabrous. Style slender, c.
114 cm, stigma ellipsoid, 4-34 mm. Fruits ovoid,
more or less flattened, slightly curved, tapering

368

FLORA MALESIANA

[sens Tee volenG2

Fig. 35. Neuburgia moluccana (BOERL.) LEENH.; note the long-tubed flowers and the stipules, especially
between the leaf pair at the left. Cult. Hort. Bot. Bogor II-I-18 (from Ambon) (JAcoss, 1958).

towards the acute apex, 414-7 by 2-234 (when
fresh —4) by 114-134 cm, orange.

Distr. Malaysia: Moluccas (Halmahera, Bat-
jan, Ceram, Ambon).

Ecol. Rain-forests, on river-banks, from sea-
level up to c. 600 m. FV. fr. Jan.—Dec.

Vern. Dodalaim, pépéké ma tuhu, Halmahera,
bandtau, Ceram.

Notes. N. celebica, the only sympatric species
which has moreover also rather large leaves, can
vegetatively easily be distinguished, as the leaves
are much thinner and the number of nerves is
larger.

Rumpuius’s Fructus musculiformis was origi-
nally based on fruit-kernels washed ashore on the
island of Ceram. I am almost certain that they
belong to a Neuburgia and, in view of the di-
mensions mentioned, probably to the present spe-
cies. In an appendix to his description, and on
his plate, he associated these fruits with a tree
found on the shore of Ceram, apparently some
Apocynacea, probably Cerbera or a related genus.
LAMARCK based his Cerbera musculiformis ex-
clusively upon Rumpuius’s description and plate
which are discordant elements. Moreover it will
hardly be possible to assign RUMPHIUs’s fruits
with certainty to a species. LAMARCK’s name can
therefore not be used as the oldest basionym.

N. tuberculata Bu. is again fully based on
Rumpuius’s description and plate, but the epithet
is illegitimate. BLUME was the first who apparently

correctly interpreted RumpuHius’s plant by in-
cluding it in his new genus Neuburgia. MIQUEL
made the legitimate combination N. musculiformis;
moreover he combined the two species distinguish-
ed by BLuME, with which I do not agree.

GAERTNER’S Banksia musculiformis was based
upon material of an Australian species of Banksia
(Proteaceae); he cited RUMPHIUS’s name in syn-
onymy with a question-mark, obviously taking
the specific epithet from it.

The name Crateriphytum moluccanum was
originally based upon material from three plants
in the Bogor Botanic Garden, viz I[-I-1, 1V—E-92,
and IV-E-92a. As a lectotype I have chosen
IV—E-92 (= Koorpers 42708) (Bo, dupl. in L,
WAG).

7. Neuburgia tubiflora BL. Mus. Bot. 1 (1850)
156; MARKGRAF, Bot. Jahrb. 61 (1927) 203, 222;
Merr. & Perry, J. Arn. Arb. 23 (1942) 416.—
N. musculiformis Mig. Fl. Ind. Bat. 2 (1856) 403,
typo excl.

Treelet; branches glabrous. Leaves oblong-
obovate, 16-20 by 8-10 cm, herbaceous to thin-
coriaceous, glabrous; base cuneate; apex blunt to
slightly blunt-acuminate; midrib forked just below
the apex, nerves 4-6 pairs; petioles 114-5 cm. Jn-
florescences c. 6 cm long, laxly branched, few-
flowered, glabrous. Calyx 3 mm high, outside
glabrous, inside at the base with colleters. Corolla
c. 2 cm long, slender, outside glabrous, the hair-

Dec. 1962]

LOGANIACEAE (Leenhouts)

369

ring in the mouth inserted on a thickened, lobed
rim, tube inside shortly patent-hairy from the
mouth to c. 1-2 mm from the base, mainly on the
main nerves; lobes c. 1 cm long. Stamens inserted
at c. 514 mm above the base of the tube; anthers
lanceolate, 3 mm long, cells free for c. 1/3, corni-
culate at apex, membranous and acute at base,
glabrous. Style c. 1 cm, stigma ellipsoid, 1—*4
mm. Fruit curved spindle-shaped, acute, 51 by
134 cm (if 2-seeded broad-ovoid, shortly stipitate,
4 by 2 by 14 cm).

Distr. Malaysia: W. New Guinea, twice col-

lected; according to BLUME also in the Moluccas,
but probably erroneous.

Ecol. Lowland forest, c. 50 m. Fi. May.

Note. The thickened rim in the throat is strong-
er developed than in N. moluccana which is
distinctly the nearest related species.

As a lectotype I have chosen ZipreLius 147a (L).

Excluded

Neuburgia ? sumatrana (MiqQ.) BoerL. Handl. 2
(1899) 392 based on Orchipeda sumatrana Mia.
Sum. (1861) 553 = Voacanga sp. (Apocynaceae).

®1962

Fig. 36. Geniostoma randianum MERR. & PERRY. a. Habit, »
opened corolla, * 4, e. stamen from in- and outside, = 8, f. pistil, x 4, g. stigma, < 8, A. fruit, x

2/;, b. stipules, < 3, c. flower, X 3, d.
1.—
G. rupestre Forst. i. Seeds, * 8 (a—b Brass 22598, c—g Brass 4522, h Brass 4995, i JUNGHUHN S./.).

8. GENIOSTOMA

Forst. Char. Gen. Pl. (1776) 12, t. 12; VALETON, Bull. Inst. Bot. Btzg 12 (1902)
1-28.—Anasser Juss. Gen. (1789) 150.—Haemospermum REINW. Syll. Pl. Ratisb.
2 (1826) 9.—Fig. 36.

Shrubs or small trees. Indument consisting of simple hairs. Twigs terete or
quadrangular, sometimes 4-winged. Leaves petioled or rarely sessile, at the base
with a short ocrea. Inflorescences axillary or sometimes (partly) ramiflorous, cy-
mose, often pseudo-umbellate or glomerulous, sometimes uniflorous. Bracteoles
present, mostly minute. Flowers 5-merous (exceptionally in some flowers 4-merous),
gynodioecious. Sepals usually only confluent at the very base, inside usually with
colleters at the base, thick, thinned out towards the almost always ciliate margin.
Corolla campanulate to rotate, (greenish-)white, tube thin-fleshy, lobes thicker,
imbricate or contorted in bud, outside glabrous or short-pubescent, inside glabrous

370 FLORA MALESIANA [ser. I, vol. 6?

or densely woolly pubescent, especially in the mouth, sometimes also near the
base. Stamens inserted in the mouth, exserted, recurved beyond anthesis; fila-
ments usually short; anthers 2-celled, often with a distinct appendage (especially
distinct in 2 flowers). Ovary 2-celled, with ~ ovules; style often very short;
stigma club-shaped or ellipsoid to globular, about as large as the ovary. Capsules
2-valved, septicidal or septifragal, black or possibly in some species green when
ripe. Seeds ~, ellipsoid to subglobular, intruded on the hilar side, minutely warty,
brown to black, embedded in a juicy yellow to red pulp; endosperm thick, charta-
ceous.

Distr. About 20-40 spp., mainly in the Pacific region as far east as the Society Is. and as far north as
Kyushu (S. Japan), absent from the Asiatic mainland, especially richly developed in New Caledonia;
in Australia, Lord Howe I., and New Zealand represented by | species each; in Malaysia at least 4 species
(3 of which are restricted to New Guinea); furthermore 2 species in the Mascarenes. Cf. LEENH. Pac. PI.
Areas 39.

Ecol. Small substage trees in the rain-forest and heliophilous shrubs in more open places, also in
mossy forests, under everwet conditions, from sea-level up to c. 2800 m.

Morph. The dehiscence of the fruit is primarily from bottom to top, secondarily also from top to
bottom, therefore the valves are coherent longest slightly below the apex; in the end the valves are shed,
and only the central part of the fruit, consisting of the dissepiment, the placentae, and the seeds, covered
by the orange-red pulpa, remains. Birds are probably mainly responsible for the dispersal of the seeds.

Specific delimitation. Both the variability of many characters and the constancy of combinations
of these differ considerably in different parts of the area of distribution. Therefore, botanists have come
to different conclusions concerning specificity of characters, hence specific delimitation, depending on the
area they studied.

In New Caledonia specific delimitation is clear: here are 12-14 well defined species, mostly endemic.
With one or two exceptions they are not closely related to those of Malaysia and other parts of the
Pacific. (See GUILLAUMIN, FI. Nouv. Cal. 1948, 85.)

In the other islands of the Pacific and in Malaysia the material is distinctly more homogeneous, but
one still meets a great variation especially in flower-characters: sepals differ in shape and size, the corolla
is sparsely or densely and short- or long-hairy in the mouth, sometimes moreover provided with a second
hair-ring near the base, stamens vary greatly in shape and size, are hairy or glabrous, and are provided
with or lack an apical appendage which also may vary considerably in shape and size, finally the pistil
may be glabrous or hairy, and the length of the style and shape of the stigma may differ considerably;
furthermore, differences may be found in the degree of branching of the cymes and in the shape and size
of the leaves.

The Pacific material comprises several small isolated island-populations, each characterized by a
combination of these differential characters, without much variation. Most of them appear closely allied
and of clearly reticulate affinity. Nevertheless, here one could gain the impression that these characters
and their combinations carry sufficient weight for specific delimitation. (See A.C. SmitH & B. STONE,
Contr. U. S. Nat. Herb. 37, 1962, 1-41, pl. 1-3.)

Application of the same standard in Malaysia (which in fact has been done by VALETON, Bull. Inst.
Bot. Btzg 12, 1902, 1—28, pl. 1) led me to the distinction of about 35 local populations which are also
mutually distinctly and reticulately allied, similarly as in the Pacific. However, they are mostly far less
clearly delimited and are often connected by transitional forms. What is worse, the characters cited above
do not appear to be constant: they show variation not only within one form, but also in otherwise almost
identical specimens from the same locality, and even between the flowers of one specimen; the sepals and
stamens may even vary within one flower. Obviously, this break-down of the constancy of characters is
primarily due to the fact that the much larger Malaysian islands offer larger continua of land through
which the effect of isolation, hence homogeneity of local populations as occurs in the Pacific, is prevented;
a second cause is probably some degree of dioecism especially in the Javanese populations.

As a result of the detail examination of a large amount of Malaysian material I have come to the con-
clusion that the above-mentioned characters can not be trusted for specific delimitation in the G. rupestre
affinity. Those which seem to me useful and constant are the size of the corolla, the presence c.g. absence
of hairs on its inner surface, and the size (and possibly the shape) of the fruit. A further character may be
found in the presence or absence of a light-coloured membranous margin along the valves of the latter,
depending on the mode of dehiscence of the fruit, septicidally or septifragally respectively.

Note. The relationship is especially with the Hawaiian genus Labordia Gaupicu., which differs by
terminal inflorescences and a corolla-tube which is distinctly longer than the lobes.

Dec. 1962]

LOGANIACEAE (Leenhouts)

Sil

KEY TO THE SPECIES

1. Corolla hairy inside, at least in the mouth.

2. Fruits up to 144 cm long. Corolla 2-414 mm long .
4-114 cm. Corolla 6-7 mm long .

2. Fruits 2 by 7
1. Corolla inside glabrous.

1. G. rupestre
2. G. randianum

3. Corolla 514 mm long. Flowers long-stalked, in Ei ag Fruits c. 10 by 714 mm. Leaves

usually 10-19 by 5-9 cm, thin .

3. Corolla 2-4 mm long. Flowers short-stalked, in fascicles or cymes. Fruits 3

more than 5 by 2 cm, stiff .

1. Geniostoma rupestre Forst. Char. Gen. Pl.
(1776) 12, t. 12; VAL. Bull. Inst. Bot. Btzg 12
(1902) 12 & 17, f. 1; A. C. SmitH & STONE, Contr.
U. S. Nat. Herb. 37, 1 (1962) 34.—Haemosper-
mum arboreum REINw. Syll. Pl. Ratisb. 2 (1826)
10; Bi. Bijdr. (1827) 1018; Hassk. Flora 28
(1845) 246 (‘Haematosperma arborescens’).—
G. haemospermum StTeub. Nomencl. (1840) 669,
nom. illeg.; BL. Mus. Bot. 1 (1850) 238, incl.var.
angustifolia, elongata, erosa, laevigata, et rugulosa;
Mia. FI. Ind. Bat. 2 (1857) 365; VAL. Bull. Inst.
Bot. Btzg 12 (1902) 21, f. 9, 10, 15 & 18; K. &
V. Bijdr. 9 (1903) 56; Koorp. Atlas 2 (1914) f.
328: Fl. Tjib. 3 (1918) 45.—G. micranthum DC.
Prod. 9 (1845) 27; F.-ViLL. Nov. App. (1880) 135.
—G. montanum Z. & M. in Mor. Syst. Verz.
(1846) 58; Bru. J. Linn. Soc. Bot. 1 (1856) 97;
Mia. FI. Ind. Bat. 2 (1857) 366, t. 33.—G. lasio-
stemon BL. Mus. Bot. 1 (1850) 239, f. 35, incl. var.
moluccanum; Mia. Fl. Ind. Bat. 2 (1857) 365.—
G. reticulata BL. Mus. Bot. 1 (1850) 239.—G.
lanceolatum Z. & M. ex Mia. FI. Ind. Bat. 2 (1856)
t. 33, nom. illeg., non BovER ex DC. 1845; Hocnr.
Candollea 6 (1936) 471.—G. cumingianum BTH.
J. Linn. Soc. Bot. 1 (1856) 97; Merr. En. Philip.
3 (1923) 310.—G. australianum F. vy. M. Fragm.
5 (1865) 19; Bru. Fl. Austr. 4 (1869) 367; F. M.
BAIL. Queensl. Fl. 3 (1900) 1022; VAL. Bull. Inst.
Bot. Btzg 12 (1902) 17, f. 2.—G. arboreum O.K.
Rey. Gen. Pl. (1891) 425, incl. var. laevigatum;
Baku. f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 170, p. 3.—? G. avene VAL. Bull. Inst. Bot.
Btzg 12 (1902) 23 & 16.—G. celebicum VAL.
Le. 19 & 15, f. 7 & 13.—G. miquelianum K. & V.
CxaVAba Ics 22& 14,.f: 11,12 & 16; K. & V.
Bijdr. 9 (1903) 58; Koorp. Exk. Fl. Java 3 (1912)
56 & 57; Atlas 2 (1914) t. 327; CAMMERL. Bull.
Jard. Bot. Btzg III, 5 (1923) 297; Baku. f. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 3.—
G. moluccanum VAv. Bull. Inst. Bot. Btzg 12
(1902) 19 & 15, f. 4; Ic. Bog. 2 (1904) 143, t. 130;
CAMMERL. Bull. Jard. Bot. Btzg III, 5 (1923)
297.—G. oblongifolium K. & V. ex VAL. Bull.
Inst. Bot. Btzg 12 (1902) 20 & 15, f. 5, 6 & 14;
K. & V. Bijdr. 9 (1903) 60; Koorp. Atlas 2 (1914)
f. 329; BAKH. f. in Back. Bekn. FI. Java (em. ed.)
7 (1948) fam. 170, p. 3; Meyer Drees, Comm.
For. Res. Inst. n. 33 (1951) 74.—G. caulocarpum
K. Scu. in K. Sch. & Laut. Nachtr. (1905) 348;
GitG & BENED. Bot. Jahrb. 54 (1916) 161.—G.
philippinense Merr. Philip. J. Sc. 3 (1908) Bot.
259.—G. batanense MeRR. ibid. 3 (1909) Bot.
427; En. Philip. 3 (1923) 309.—G. stenophyllum

. . 3. G. weinlandii
—5 mm g. Leaves rarely
4. G. arfakense

Merk. Philip. J. Sc. 7 (1912) Bot. 329, non GILG
& BeENeED. 1916; En. Philip. 3 (1923) 310.—? G.
pulgarense ELM. Leafl. Philip. Bot. 5 (1913) 1808;
Mere. En. Philip. 3 (1923) 310.—G. brevipes
Merk. Philip. J. Sc. 9 (1914) Bot. 384; En. Philip.
3 (1923) 309.—G. laxa ELM. Leafl. Philip. Bot. 8
(1915) 2746; Merr. En. Philip. 3 (1923) 310.—
G. mindanaense Em. Leafl. Philip. Bot. 8 (1915)
2747.—G. acuminatissimum GitG & BENED. Bot.
Jahrb. 54 (1916) 159, f. 3, non Merr. 1922.—G.
antherotrichum GILG & BENED. /.c. 158, f. 2.—G.
psychotrioides GILG & BENED. l.c. 160.—G.
dasyneurum GILG & BENED. /.c. 161.—G. schlech-
teri GILG & BENED. /.c. 162.—G. longipes MERR.
Philip. J. Sc. 12 (1917) Bot. 296; En. Philip. 3
(1923) 310.—G. nigrescens (BLCO) MEeErRR. Sp.
Blane. (1918) 305, pro specim., typo excl.; En.
Philip. 3 (1923) 310.—G. pachyphyllum MEeERR.
Philip. J. Sc. 14 (1919) 448; En. Philip. 3 (1923)
310.—G. lancilimbum Mer. Philip. J.Sc. 17 (1921)
304; En. Philip. 3 (1923) 310.—G. ramosii MERR.
Philip. J. Sc. 17 (1921) 304; En. Philip. 3 (1923)
310.—G. pullei CAMMERL. Nova Guinea 14 (1924)
115, t. 12.—G. fasciculata Quis. & MEeERR. Philip.
J. Sc. 37 (1928) 190.—G. aff. rupestris Forst.:
CHRISTOPHERSEN, Bull. Bish. Mus. n. 128 (1935)
176.—G. dallmannense Kan. & Hart. Bot. Mag.
Tokyo 56 (1942) 164, f. 8.—Fig. 36i.

Shrub or treelet, up to 6(—-10) m high. Branches
mostly terete, rarely quadrangular, glabrous or the
young twigs shortly brown- to fulvous-tomentose.
Leaves ovate to lanceolate, 114-17 by 14-614 cm,
herbaceous to chartaceous, rarely coriaceous,
glabrous or (mainly on the midrib beneath) short-
ly tomentose; base rounded to acute, rarely
emarginate; margin entire or rarely faintly cre-
nulate; apex blunt or acute to long-acuminate,
aristulate or not; nerves (3—)6-10(—-11) pairs;
petiole '4%4-1(-134) cm. Inflorescences mostly
axillary, sometimes partly ramiflorous, often
glomerulous, not rarely laxly paniculate, sometimes
umbelliform, with (1-)several flowers, 14-114
(-414) cm, sparsely (to densely) shortly appressed-
hairy, sometimes glabrous. Pedicels 1-5 mm.
Calyx (24-)1-2(-2'4) mm high, sepals elliptic or
ovate to broad-triangular, blunt to acute, outside
glabrous to densely short-tomentose. Corolla
(1144-)244-444 mm long, outside glabrous or
rarely fairly densely papillose-hairy, inside the
throat densely to sparsely hairy, sometimes a
second ring of hairs near the base of the tube;
lobes (34—-)1-2'%4 mm long, usually blunt, rarely
acute. Stamens variable, especially as to hairiness;

372

FLORA MALESIANA

[ser. I, vol. 6?

filaments (#/,—)'4-34(-1) mm, glabrous to rather
densely hairy; anthers 34-114 mm long, glabrous
or hairy, appendage of the connective absent to as
long as the cells, in the latter case (mainly in 2
flowers) triangular above the adnate cells. Ovary
glabrous to densely shortly tomentose; style usual-
ly very short (less than 1/; mm), in some forms
(mainly in Sumatra, W. Java, and the Philippines)
about 1(-2) mm, glabrous or rarely slightly pu-
bescent. Fruits mostly slightly flattened-globose,
rarely ellipsoid, ovoid, or obovoid, 4-714(-12)
by 4-6(-9) mm, black when ripe (sec. MERRILL
whitish in G. philippinense, red in G. stenophyllum),
septifragal (but see note).

Distr. Widely distributed, throughout Malaysia
(except in the Malay Peninsula), E. Queensland
(around Innisfail, Ravenshoe, and Rockingham
Bay), and the West Pacific (at least as far north as
the Marianas and east as far as Samoa).

Ecol. Very variable, in the substage of the rain-
forest, but apparently preferring more open
places, often in exposed sites, along forest edges
and river-banks, in young secondary forests and
shrubberies, on steep slopes and in grass fields,
under everwet conditions, in regions subject to a
dry season only, in the montane rain-forest, for
example in the Lesser Sunda Is. only between
700-2200 m; from sea-level up to 2800 m.

Wood anat. Jutrre, Nova Guinea n.s. 9 (1958)
360, f. 8 (sub Geniostoma sp.).

Vern. Itit biru, kaju tain, kitelong, S, maniran
(or meniran)-idju (or -idjo), tumon telong, SJ;
Philip.: bitig-bitig, Neg., gagadang, lv., san-
guadan, Mbo., tagobinlod, C. Bis., umum, Ig.;
bochogan, kaju-im-bulut, kaju-tai, kopopojoan-
wewene, mawotai, Cel., gandarusa-utan, Ambon.

Notes. As has been alluded to under the genus,
G. rupestre is extremely variable, both in vegeta-
tive and in floral characters; fruit characters are
less variable. There are a number of geographically
more or less restricted races. Part of these are very
local and often known from a few specimens only;
others are locally sometimes sharply delimited but
in other regions connected by intermediates. I
deem it premature to venture on a subdivision of
this species and it is doubtful that this will ever be
possible on more than a local scale.

Some of these local forms, and some single
specimens, have only with some doubt been in-
cluded in G. rupestre. G. pulgarense is known only
from the type-collection (ELMER 13197, Mt
Pulgar, Palawan, Philippines; fruiting specimen),
and deviates from the mountain form described
from Luzon as ‘G. cumingianum’; it seems to be
more closely related to the form of Mt Kinabalu,
Borneo. The type specimen of G. avene (TEYSMANN
s.n., P. Gebeh, E. Moluccas; fruiting specimen)
also comes close to G. pulgarense, with the ex-
ception only of the fruits which are septicidal with
light-coloured, membranous margins.

Two specimens from New Guinea (DocTers
VAN LEEUWEN 10345 & 10415) are characterized
by small, narrowly lanceolate leaves (2144-5 by
144-114 cm) and small flowers (calyx 0.8 mm,
corolla 2 mm); these specimens were collected

near the Rouffaer River, on the river-bank be-
tween stones, at 250-300 m.

Among the Celebes specimens, KJELLBERG 2775
from Matano (at 400 m) is mainly aberrant by the
relatively large (11-12 by 7 mm), ellipsoid to
ovoid fruits. KJELLBERG 2681 (Central Celebes
at 1200 m) and Erma 470 & 2051 (both from
Ceram, at 1550-2600 m) are deviating by very
small flowers (calyx 34—*/; mm, corolla 114-2
mm, anthers subsessile, 0.4 mm long).

Nomencl. The name G. haemospermum
STEuD. is illegitimate as it had been based upon
Haemospermum arboreum Reinw., and the epi-
theton arboreum should have been used.

The name G. lanceolatum Z. & M. ex Mia.
is illegitimate primarily as being a later homonym
of G. lanceolatum Boser ex DC. (1845), and
secondarily as this name, apparently erroneously
printed under the plate, was corrected in the text
by MIQUEL to montanum, hence not accepted by
him.

2. Geniostoma randianum Merr. & Perry, J.
Arn. Arb. 23 (1942) 409.—G. obtusum MERR. &
Perry, /.c. 410.—Fig. 36a-h.

Shrub or treelet, 2-6 m high, glabrous. Leaves
often inserted on distinct leaf-cushions, obovate
or elliptic to lanceolate, 314-434 by 114-214 cm,
stiff-coriaceous; base cuneate; apex blunt or acute
to shortly, broadly, and acutely acuminate, aristu-
late; nerves 5—7 pairs; petiole 6-8 mm. Jnflores-
cences axillary, 1—5-flowered, dichasial, 1-2 cm.
Pedicels c. 14 cm. Calyx 114-3 mm high, up to
about halfway connate, glabrous, lobes acute.
Corolla 6-7 mm long, outside glabrous, inside
more or less densely hairy in and sometimes above
the mouth; lobes acute, 2!4-4 mm. Filaments /-
2/, mm, glabrous; anthers 114-114 mm long, entire-
ly or partly lax-hairy, appendage of the connective
c. 0.1-4 mm. Pistil glabrous; style 4-3 mm.
Fruits usually 1 per axil, flattened-ovoid to obo-
void, 2 by 34-114 cm, septifragal.

Distr. Malaysia: New Guinea (Central Div.;
Milne Bay Distr.).

Ecol. On forest fringes in mossy forest and
shrubberies, 2230-2840 m.

Notes. In describing G. obtusum, MERRILL and
Perry observed already that it is closely related to
G. randianum, and though there are some differ-
ences in fruit, flowers, and leaves, I think they
should be combined.

Well characterized by the large-sized flowers
and fruit.

3. Geniostoma weinlandii K. ScH. in K. Sch. &
Laut. Nachtr. (1905) 349; GitG & BENED. Bot.
Jahrb. 54 (1916) 157, f. 1.—G. acutifolium HIERN,
Nova Guinea 8 (1909) 201.

Shrub or treelet, 2-5 m, glabrous or the young
twigs sometimes thinly tomentose. Leaves ovate
to oblong (to lanceolate), (614-)10-21 by (314-)
5-9 cm, tough-papyraceous to thin-chartaceous,
base rounded to cuneate, apex tapering to rather
abruptly acuminate, acumen short and triangular
to long and slender, acute; nerves 5—8(—10) pairs;

Dec. 1962]

LOGANIACEAE (Leenhouts)

373

petiole 14-114 cm. Inflorescences axillary and
partly ramiflorous, umbelliform, short-stalked,
3-5(—-L15)-flowered, 1-2 cm long. Pedicels slen-
der, (0.6-)0.8-1.3 cm. Calyx 2-21 mm high,
outside rather densely shortly tomentose, lobes
semi-orbicular. Corolla 54. mm long, glabrous
on both sides, lobes 3!4 mm, slightly acuminate,
broadly imbricate in bud. Stamens glabrous,
filaments up to 1 mm, anthers 1-11 mm long, the
appendage of the connective small. Pisri/ glabrous,
style 114-134 mm. Fruits obovoid to ellipsoid,
c. 1 by 34 cm, sutures distinct.

Distr. Malaysia: New Guinea and ? Moluccas
(Aru Is.).

Ecol. Mainly along forested river-banks, also
on Saccharum-covered shoals in river-beds, 50—
1750 m. Fl. fr. probably Jan.—Dec.

Vern. Djangdjangalie, Aru Is., marau, New
Guinea.

Notes. The species had been based upon two
specimens, SCHLECHTER 14495 (still represented in
BM, Brst, K) and WEINLAND s.n. (BO, BRI,
Brsit, SING), both from Kaiser Wilhelmsland,
“am mittleren Bumifluss’. As the WEINLAND
specimen bears both flowers and fruits (especially
the Wroclaw-sheet is a good one) I choose this
as lectotype; the holotype is lost in Berlin.

A sterile specimen from the Aru Is. (bb. 25404)
probably belongs to this species, though it is not
quite impossible that it represents G. rupestre
which is vegetatively not well distinguishable.

4. Geniostoma arfakense KAN. & Hat. Bot. Mag.
Tokyo 56 (Apr. 20, 1942) 163, f. 7—G. archbol-
dianum Merr. & Perry, J. Arn. Arb. 23 (Oct.
1942) 408.

Shrub or small tree, up to 10(—13?) m high.
Branches glabrous or the young parts sometimes
sparsely pubescent. Leaves glabrous or rarely
ciliate when very young, (ovate-)elliptic to lan-
ceolate, (34—)114-5(-10) by (144-)1-2(-414) cm,
chartaceous to coriaceous, base broadly to nar-
rowly cuneate, sometimes slightly rounded, apex
blunt to gradually long and slender acute-acu-
minate, sometimes aristulate; nerves 4-8 pairs;
petiole (114—)2'14-5(-7) mm. Inflorescences axil-
lary, glomerulous to laxly cymose, 34—2 cm long,
up to c. 10-flowered; pedicels slender, 144-6 mm.
Sepals mostly acute, 1-114 mm long, outside gla-
brous or minutely pubescent. Corolla 24 mm
long, lobes blunt to acute, 1144-214 mm, glabrous
(rarely outside papillose-tomentose). Stamens
mostly shortly patent-hairy, filaments very short
to ¥4 mm, anthers 974-1274 mm, appendage of the
connective up to about half as long. Ovary gla-
brous or sometimes hairy around the style base;
style almost absent or up to c. 44 mm, glabrous
or pubescent. Fruits globular to slightly ellipsoid,
3-5 mm 9, septifragal; suture inconspicuous.

Distr. Malaysia: New Guinea.

Ecol. In and mainly along primary, rarely
secondary forests, along river-banks, on grassy
deforested slopes, and in low regrowth on peaty
soil, mainly 1700-2600 m. Fi. (March, April)
June—Dec., fr. June-Nov.

Vern. Kegopa, kiaugupa, mongemandimand,
raurau, tuoparimoh, tuwapalimo.

Notes. There is some variation in details,
mainly in the shape and size of the leaves and the
number of flowers per inflorescence.

The alliance is with G. oleifolium S. Moore
from New Caledonia, but it comes also very close
to some E. Malaysian forms of G. rupestre (‘G.
pullei’ and ‘G. pulgarense’ especially) from which
it only differs by the glabrous mouth of the corolla.

Dubious

Anassera rumphii SPANOGHE, Linnaea 15 (1841)
325, nom. nud.

Neither the specimen, nor the drawing (‘‘Icon.
n. 27”) could be localized.

Geniostoma gilgii MERR. & PERRY, J. Arn. Arb. 23
(1942) 409.—G. stenophyllum GitG & BENED.
Bot. Jahrb. 54 (1916) 162, nom. illeg., non MERR.
1912.

Described from New Guinea. Possibly a sy-
nonym of G. rupestre, but the description shows
some discrepancies with that species and authentic
material could not be localized.

Excluded

Anasser moluccana LAMK, Illustr. 2 (1793) 40, is
according to BAKKER, FI. Mal. I, 5 (1957) 356-359
= Pittosporum moluccanum (LAMK) Mia. (Pitto-
Sporaceae).

Anasser laniti BLco, Fl. Filip. (1837) 112, is
according to MERRILL, Sp. Blanc. (1918) 313 =
Wrightia laniti (BLCO) MERR. (Apocynaceae).

Geniostoma acuminatissima MerRR. Philip. J. Sc.
20 (1922) 432, nom. illeg., non GiLG & BENED.
1916, is according to BREMEKAMP, Candollea 16
(1957) 93 = Lasianthus furcatus (M1Q.) BREMEK.
(Rubiaceae).

Geniostoma acuminatum WALL. in Roxb. FI. Ind.
2 (1824) 316, is according to BENTHAM, J. Linn.
Soc. Bot. 1 (1856) 69 = Urophyllum streptopodium
WALL. (Rubiaceae).

Geniostoma crassifolium BrH. J. Linn. Soc. Bot.
1 (1856) 96. A species described from the Isle of
Pines near New Caledonia, in Index Kewensis
erroneously cited as from the Philippines.

Geniostoma nigrescens (BLCO) MerR. Sp. Blanc.
(1918) 305.—Tayotum nigrescens Buco, Fl. Filip.
(1837) 105; ed. 2 (1845) 76; ed. 3, 1 (1877) 141.
MERRILL wrongly referred BLANCo’s description
to G. philippinense; the ‘illustrative specimen’
distributed by him (MERRILL, Sp. Blanc. 878) I
reckon to belong to G. rupestre. FERN.-VILLAR
(Nov. App. 1880, 136) was also mistaken by re-
ducing BLANCo’s species to Norrisia malaccensis
GarDN. In fact BLANCO’s monotypic genus cannot
represent any Loganiacea; the fruit which is said
to be enveloped by the calyx, excludes it from that
family.

374 FLORA MALESIANA [ser. I, vol. 6?

Fig. 37. Cynoctonum sphaerocarpum LEENH. a. Habit, x 24, b. flower, * 8, c. ditto, longitudinal section,
x 8, d. fruit, x 4, e. seed, x 20.—C. mitreola (L.) Britt. f. Habit, * 24, g. fruit, * 4, A. seed, x 20
(a combined from CLEMENS 29469 and JAHERI 286, b—e CLEMENS 29469, f-h HOOGLAND 4365).

Dec. 1962] LOGANIACEAE (Leenhouts) _ 375

9. CYNOCTONUM

GMEL. Syst. (1791) 443.—Mitreola LINNE, Gen. Pl. ed. 1 (1737) 377.—Ophiorhiza
LINNE, ibid. ed. 5 (1754) 74, p.p.—Mitreola BOEHMER in Ludwig, Def. Gen. PI.
(1760) 31, nom. illeg., p.p— Mitreola SCHAEFF. Bot. Exped. (1760) 11, nom.
illeg —Selenocera Zipp. ex SPAN. Linnaea 15 (1841) 316.—Parophiorrhiza CLARKE
in Hook. f. Fl. Br. Ind. 3 (1880) 85.—Fig. 37.

Herbs, annual and erect or perennial and creeping at the base. Leaves petioled
or sessile; interpetiolar stipules well developed or reduced to a transverse ridge.
Inflorescences terminal and/or axillary, usually long-peduncled, dichasial with
long cincinnate branches. Flowers on short pedicels to subsessile, 5-merous.
Calyx almost parted to the base. Corolla urceolate, lobes short, imbricate in bud,
mouth closed by a dense ring of hairs. Stamens short, included; anthers introrse,
cells free. Pisti] semi-inferior; ovary 2-celled, each cell with many, small, anatro-
pous ovules; styles 2, short, sometimes absent; stigmas more or less adnate to
each other. Capsules 2-horned, rarely globular, almost free from the calyx, both
carpels dehiscent along the ventral suture. Seeds co, small, with fleshy endosperm.

Distr. Six spp. in tropical and subtropical America (1 endemic), Madagascar (2 endemic), SE. Asia
(1 endemic), Malaysia (1 endemic), and North Australia (Arnhem Land). Cf. LEENH. Pac. Pl. Areas 33.

Ecol. Preferably on wet spots under seasonal conditions, on heavy clay soils or limestone, not or
slightly shaded.

Notes. In defining the genus Ophior(r)hiza, LINNAEUS included two species, viz O. mungos and O.
mitreola (Sp. Pl. 1753, 150). The former species is an Ophiorhiza (Rubiaceae) in the current sense, and
the type of the genus; the latter was based upon Mitreola L. 1737, which is the present Cynoctonum
mitreola. LupwiG based his genus Mitreola upon LINNAEUS Gen. Pl. 1737; as he cited, however,
Ophiorhiza L. as a synonym, his name is superfluous, hence illegitimate.

In the same year as LUDWIG, SCHAEFFER described his (new) genus Mitreola, without a reference to
LinnE, but clearly distinguishing it from Ophiorhiza L. The preface of SCHAEFFER’s publication is dated
October 1760; though the month of LuDwiG’s publication is unknown, it is at least probable that it
appeared earlier in the year and antedated SCHAEFFER’s name.

The genus is especially closely related to Mitrasacme, the main differences being the 5-merous flowers
and the semi-inferior ovary.

KEY TO THE SPECIES
1. Annual erect herb. Pistil with two distinct styles. Fruits 2-horned . = cee Gs mitreola
1. Perennial herb, creeping in the basal part and rooting at the nodes. Pistil with sessile stigmas. Fruits
globular, only slightly bilobed at the top . 5 2. C. sphaerocarpum

1. Cynoctonum mitreola (L.) Britt. Mem. Torr. Bru. Fl. Austr. 4 (1869) 349; CLARKE in Hook.

Bot. Cl. 5 (1894) 258; Hocur. Bull. N.Y. Bot.
Gard. 6 (1910) 284, incl. var. orthocarpa; MERR.
& Perry, J. Arn. Arb. 29 (1948) 163; BAKH. f. in
B ck. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170,
p. 4, incl. var. lilacina; Blumea 6 (1950) 382;
G.LEASON in Britton & Brown, Ill. Fl. U.S. ed. 2, 3
(1952) 54, fig. p. 55.—Ophiorhiza mitreola LINNE,
Sp. Pl. (1753) 150.—C. petiolatum Ge . Syst.
2 (1791) 443.—Mitreola paniculata WALL. ex
G. Don, Gard. Dict. 4 (1837) 171; DC. Prod. 9
(1 845) 9; WiGurt, Ic. 4 (1850) 14, t. 1600; Illustr. 2
(1850) t. 156 b fig. 1 (15-21); Hook. Ic. Pl. (1852)
t. 828; Mia. Fl. Ind. Bat. 2 (1857) 361; PROGEL
in Mart. Fl. Bras. 6, 1 (1865) 266, t. 71; Koorp.
Exk. Fl. Java 3 (1912) 57; CAMMeRL. Bull. Jard.
Bot. Btzg III, 5 (1923) 298, incl. var. lilacina
Back.; Back. Onkr. Suiker. (1931) 485.—Mi-
treola oldenlandioides G. Don, Gard. Dict. 4
(1837) 172, nom. illeg.; DC. Prod. 9 (1845) 9;
Hook. Ic. Pl. (1852) t. 827; Bru. J. Linn. Soc.
Bot. 1 (1856) 91; Mia. Fl. Ind. Bat. 2 (1857) 360;

f. Fl. Br. Ind. 4 (1883) 79; BoerL. Handl. 2 (1899)
449 & 458; Kina, J. As. Soc. Beng. 74, 11 (1908)
598; Dop, Fl. Gén. I.-C. 4 (1912) 155, f. 20 (1-2);
Gi_G & BenebD. Bot. Jahrb. 54 (1916) 163; ibid.
56 (1921) 547; Rip. FI. Mal. Pen. 2 (1923) 413;
GAMBLE, Fl. Madras 5 (1923) 863; KANJILAL &
Das, Fl. Assam 3 (1939) 310; HENDERS. Mal. Nat.
J. 6 (1950) 303, f. 287; Kerr in Craib, Fl. Siam.
En. 3 (1951) 52.—Selenocera secundiflora Zipp.
ex SpAN. Linnaea 15 (1841) 316.—Mitreola
petiolata ToRR. & GRAY, Fl. N. Am. 2 (1841) 45;
DC. Prod. 9 (1845) 8; Bru. J. Linn. Soc. Bot. 1
(1856) 91; ProGeEL in Mart. Fl. Bras. 6, 1 (1865)
266, t. 82 (1); Merr. Philip. J. Sc. 11 (1917)
Bot. 307; SpecuT, Rec. Am. Austr. Exp. Arnhem
Land 3 (1958) 468.—Mitreola inconspicua Z. & M.
in Mor. Syst. Verz. (1846) 55.—C. paniculatum
B. L. Ros. Proc. Am. Ac. Arts Sc. 45 (1910)
396; Merr. Pap. Mich. Ac. Sc. 23 (1938) 188.—
Fig. 37f-h.

Erect, annual herb, simple or branched from

376 FLORA MALESIANA [ser Ly volaG?

Fig. 38. Spigelia anthelmia L. a. Habit, * 14, b. stipules, x 3, c. flower, = 3, d. longitudinal section of
corolla with pistil, * 414, e. calyx with persistent basal part of fruit, x 414, f. fruit, x 3, g. seeds, x 6
(a—b, e CoeRT 624, c-d HoL_stvooGp 78, f-g RAHMAT 8401).

Dec. 1962]

LOGANIACEAE (Leenhouts)

377

the base, 5-80 cm high. Stem quadrangular to
narrowly 4-winged, glabrous. Leaves with broadly
deltoid, 1-114 mm high, interpetiolar stipules,
ovate to ovate-oblong, 1144-7 by 1-344 cm,
herbaceous, very sparsely appressed-pubescent or
glabrous apart from the appressed-ciliate entire
margin, base obtuse or acute, usually decurrent,
apex obtuse to acute; nerves 4-10 pairs, ascending;
petiole grooved, 44-3 cm long. Inflorescences
partly terminating the stem and few-leaved
branches, partly in the highest leaf-axils, widely
branched, the ultimate branches 2-15 cm, slender,
glabrous or shortly ciliate under the nodes. Bracts
narrowly deltoid, 11 —2 mm long. Flowers shortly
pedicelled to subsessile. Ca/yx obconical, 114 mm,
sepals ovate, acute, with broad, membranous
margins, glabrous. Corolla 1-2 mm, halfway
connate, lobes oblong-ovate, acute; corolla out-
side and lobes inside papillose. Stamens inserted
just above the base of the corolla-tube, glabrous;
filaments 44—!4 mm. Pistil glabrous, with 2 free
styles 14-14 mm long; stigmas globular, connate
or not. Capsules 2-horned, 2-314 mm long,
ciliate on midrib and along the margins or fully
appressed-pubescent, the horns either straight and
divaricating or curved inwards. Seeds ellipsoid,
4-34 mm long, concave on one side, smooth.

Distr. SE. United States, Central and tropical
South America, SE. Asia from Bombay to Tonkin,
in Malaysia: Malay Peninsula (peninsular Siam
and Langkawi Is.), N. Sumatra, Java (especially
common in the teak-forests of the eastern half),
Bawean, Kangean, Bali, Timor, Tanimbar Is.,
S. Celebes (also Buton I.), Philippines (Luzon,
Mindoro, Bohol), New Guinea, and N. Australia
(Arnhem Land). Introduced in the Carolines
(Yap and Palau).

Ecol. Forest edges, along roads, on grassy
plains, sometimes as a weed on fields, on sawah-
dikes efc.; on slightly shaded, badly drained,
preferably marly soils, up to 600 m alt.; in Java
especially along fire brakes in teak-forest, on
Langkawi I. on limestone rocks near the sea;
obviously bound to areas subject to a dry season.
Fl. at the end of the rainy season.

Vern. Ki fatu, Timor; Philip.:
magtindug, Mang.

Notes. The flowers are usually white. BACKER
described a var. lilacina based upon specimens
from Central Java which were characterized by
violet flowers and straight fruit lobes; the speci-
mens from Langkawi I. are also said to have pale

tagarinuk

mauve flowers, and here the fruit lobes are also
straight. HOCHREUTINER distinguished three va-
rieties based upon the shape of the fruit. As his
varieties are neither sharply delimited (var. in-
termedia comprises all specimens which do not
show one of the extreme fruit forms) nor geo-
graphically restricted (only the frequency may be
different) they seem to have no taxonomic value.

Jover (Bull. Mus. Nat. Hist. Nat. II, 6, 1934,
291-301) described two species from Madagascar:
Mitreola perrieri and Mitreola turgida, which are
closely related to the present species; both are espe-
cially different in the shape and size of the fruit.

The name Mitreola oldenlandioides WALL. was
not validly published by G. Don, as it was not
accepted by him as a species.

2. Cynoctonum sphaerocarpum LeeNnu. Bull. Jard.
Bot. Brux. 32 (1962) 417.—Fig. 37a-—e.

Perennial herb, in the basal part creeping and
rooting at the nodes. Stem terete to faintly qua-
drangular, at the top with two narrow wings most
distinct below the nodes; glabrous. Leaves gla-
brous, with narrowly triangular, 144 mm long,
acute, soon shrivelling interpetiolar stipules,
(elliptic-)oblong, 6-14 by 3-6 cm, thin-papyrace-
ous, base cuneate and decurrent, apex tapering
acute-acuminate; nerves c. 8 pairs; petiole flat
above, 1-144 cm. Inflorescences terminal and
axillary, glabrous, rather dense, 2-3 cm long,
7-8 cm peduncled, ultimate branches short.
Bracts narrowly deltoid, up to 114 mm. Pedicels
slender, 1-114 mm. Calyx obconical, 1-144 mm
long, sepals oblong-ovate, acute, with broad mem-
branous margins, glabrous. Corolla 2 mm long,
white, lobes c. 24 mm, elliptic, rounded, papillose
inside. Stamens inserted about halfway the corolla-
tube, glabrous, filaments '%4-—!4 mm, anthers
elliptic-ovate, 1 mm long. Pisti/ glabrous, stigmas
subsessile, connate. Fruits broad-ovoid, bilobed,
not or only very slightly split at the apex, c. 3 mm
9. Seeds about hemispherical to tetraedrical, c.
34 mm, minutely warty.

Distr. Malaysia: Borneo (twice collected: Mt
Kinabalu, at about 1800 m, ff. fr. May; Bt
Liangbatu).

Note. Apparently closely related to C. pedi-
cellatum (BrH.) B. L. Ros., a species described
from the Sikkim and also cited from S. China
(Yunnan, Kweichow); it differs from the Bornean
one mainly by distinct styles and mitre-shaped
fruits.

10. SPIGELIA
LInngE, Gen. Pl. ed. 5 (1754) 74; Sp. Pl. 1 (1753) 149; L. B. SmirH, Wrightia 2

(1960) 90 (Am. spp.).—Fig. 38.

Annual or perennial herbs or undershrubs. Leaves often partly in (pseudo)
whorls at the base of the inflorescence, short-petioled or sessile, the bases connected
by interpetiolar stipules or sheaths. Inflorescences terminal and/or in the upper
leaf-axils, cincinnous, sometimes reduced to a few flowers. Flowers sessile or al-
most so, 5-merous. Calyx: sepals free or connate at the base, inside at the base

378 FLORA MALESIANA [ser. I, vol. 6?

with some colleters. Corolla: tube tubular, lobes valvate in bud, shorter than the
tube. Stamens included, anthers dorsifixed, introrse, lanceolate or ovate, 2-celled.
Ovary superior, 2-celled, with many ovules. Capsule 2-lobed, 2-celled, 4-valved,
valves caducous with the exception of a cupular basal part (fig. 38e). Seeds
some per cell, globose to angular, verrucose; endosperm fleshy or cartilaginous.

Distr. About 50 spp. in tropical and subtropical America, one naturalized in W. Africa andin Malaysia.

1. Spigelia anthelmia LINNE, Sp. Pl. 1 (1753) 149;
Sims, Bot. Mag. (1822) t. 2359; DC. Prod. 9 (1845)
7; BisscHop GreV. Pl. N.I. (1883) 611; BACK. Trop
Natuur 1 (1913) 65, f. 1-4; HeyNne, Nutt. PI.
(1927) 1267; Back. Onkr. Suiker. (1931) 484;
BAKH. f. in Back. Bekn. Fl. Java (em. ed.) 7 (1948)
fam. 170, p. 4; L. B. SmitH, Wrightia 2 (1960) 98;
LEEUWENB. Acta Bot. Neerl. 10 (1961) 460, f. 1,
map |.—Fig. 38.

Annual herb, 2—25(—70) cm high, unbranched or
with some pairs of strong branches near the base;
stems erect, terete, glabrous, with a few remote
pairs of rather small leaves and an apical pseudo-
whorl of 4 larger ones. Leaves connected by
interpetiolar, broadly triangular, blunt, glabrous
stipules, ovate-oblong to ovate-lanceolate, 3-10
by 34-5 cm, herbaceous, scabrous above, glabrous
beneath, cuneate and often decurrent at the base,
attenuate at the apex; nerves 4—5 pairs, strongly
ascending; petiole 0-1 cm. Inflorescences terminal
and usually in the axils of the whorled upper
leaves, up to 15 cm long, peduncle very short,
glabrous or nearly so; bracts lanceolate, 2-3 mm
long. Flowers spaced, (sub)sessile. Sepals free,
quincuncial in bud, somewhat unequal in length,
ovate-linear-lanceolate, 2-314 mm, acute, glabrous
or outside sparsely puberulous, pale green. Corolla
salver-shaped, glabrous, white to red or purplish;

tube 614-15 mm, lobes exduplicative-valvate in
bud, triangular, 2-214 mm long.Stamens glabrous,
inserted slightly below the middle of the tube,
filaments filiform, + 1 mm, anthers attached
slightly above the base, lanceolate, 144-134 mm,
obtuse. Ovary glabrous, subglobose, 14-34 mm 9,
style cylindrical, 34 mm; stigma ovate-lanceolate,
2 mm, pubescent near the tip, caducous. Capsule
4-5 by 5-6 mm, squamulate-tuberculate mainly
in the upper half. Seeds obliquely ellipsoid or
ovoid, 2-3 by 114-2 mm, dull brown, tuberculate.

Distr. Native in America from Mexico and
Florida to Peru and Brazil, naturalized in tropical
West Africa and in Malaysia: Sumatra, Java
(mainly West and Central), and Lesser Sunda Is.
(Sumba).

Ecol. A weed of sandy seashores, river-banks,
fields, gardens, roadsides, waste places, from the
beach up to 600 m; locally often common, but on
the whole rather rare. A self-pollinating plant;
the flowers are open from 2-5 in the afternoon.
Fl. fr. Jan.—Dec.

Uses. A decoction of the roots is well known
to be a very effective vermifuge.

Vern. Wormgrass, E, sammondjo,
djukut puntir, taih manuk, 8, platikan, J.

Note. Reported to be very poisonous.

Sum.,

11. MITRASACME

LaBILL. Nov. Holl. Pl. Spec. 1 (1804) 35, t. 49; LeENH. Bull. Jard. Bot. Brux. 32
(1962) 440.—Fig. 39-47.

Annual or perennial herbs. Leaves in pairs along the stem or rosulate, (sub)-
sessile, connected at base by a membrane, stipules not separately distinguishable;
one- or curvi-nerved. Flowers either solitary in the leaf-axils, rarely terminal, or
in terminal and/or axillary cymose inflorescences, 4-merous. Calyx 4-, rarely
2-lobed, sepals about halfway connate; colleters few or absent. Corolla urceolate,
campanulate, or salver-shaped, membranous to thin-fleshy, often bearded in the
mouth; lobes exduplicate-valvate in bud. Stamens inserted on the lower half of
the corolla-tube, mostly included, filaments usually filiform; anthers 2-celled,
introrse, extrorse, or rarely latrorse. Pistil superior or slightly inferior, glabrous,
ovary 2-celled with many ovules per cell; styles 2, usually free in their basal and
connate in the upper part, rarely entirely connate or completely free; stigma
truncate to 2-lobed; anthers and stigma always at about the same height. Capsule
opening by apical loculicidal slits, 2-horned, the horns terminated by the styles,
which may be torn loose (e.g. fig. 391) or remain partly connate (e.g. fig. 42f).
Seeds ~>, minute, ellipsoid or angular; testa thin, endosperm fleshy.

Dec. 1962] LOGANIACEAE (Leenhouts) 379

Distr. About 40 spp., mainly Australian, furthermore in Tasmania, New Zealand, New Caledonia,
Malaysia, the Carolines, and in Asia from the Deccan and Ceylon to Central Japan and Korea.

Ecol. Heliophilous plants from dry to wet open places, locally often gregarious, from sea-level to
above 3000 m.

Notes. This largely Australian genus is in urgent need of revision. In the past the habit of the plant
played a great part in specific delimitation. For that reason some species, for example M. polymorpha
R. Br. and M. alsinoides R. BR., were supposed to have an extensive area of distribution, reaching from
New South Wales (both were originally described from Sydney) to continental Asia and Japan. During
the revision of the Malaysian species I found very useful and constant characters in the flower mor-
phology and the seed-coat, coinciding with sometimes small but constant characters in the stem, leaves,
inflorescences, and pubescence, but not with habit. In my opinion M. polymorpha and M. alsinoides are
restricted to subtropical eastern Australia. What, in Malaysian literature, was called ‘“‘M. polymorpha”
is the lowland form of M. pygmaea, though the habit is quite different from that of Australian speci-
mens. The Indo-Malaysian material formerly included in “‘M. alsinoides’ comprises at least 4 well de-
fined species which possess nearly the same habit.

It fell outside the scope of this work to verify the subdivision of the genus as proposed by R. BROWN
and later authors, the more so because for distinction I have used a number of characters formerly
neglected.

Many species look very much alike some of the smaller species of Hedyotis (Rubiaceae). Mitrasacme
is easily distinguished by the superior fruit (in Hedyotis distinctly adnate to the calyx for the greater part)
which is typically mitre-shaped, the superior ovary, and the short corolla-lobes (in Hedyotis about as
long as the tube) and included stamens; furthermore by the absence of raphides in the leaves (in Hedyotis
and related genera these are very conspicuous).

Morph. Crarke (in Hook. f. Fl. Br. Ind. 4, 1883, 80) mentioned of M. ‘polymorpha’: “‘stems .. .
glandular hairy below”. KLetr (in Mez, Bot. Arch. 5, 1924, 327) mentioned the presence of capitate-
glandular hairs in M. tenuiflora Btu. Neither in these nor in any other species could I find them.

The inflorescence, if many-flowered, can be best described as irregularly repeatedly umbellate. At every
node of the rachis there are 2 lateral branches; each of these bears usually 1 or more collateral much
feebler or often uniflorous ones. Both these branches or only one of them may represent the peduncle of
the next umbel; the continuation of the rachis is mostly much feebler or fully suppressed (fig. 42a).

The corolla is very thin in most species and usually much shrunk in herbarium specimens; the absolute
dimensions (especially the lower values) are therefore not very reliable.

The degree to which the styles are connate is different in anthesis and fruit, usually less in the latter
state.

KEY TO THE SPECIES

1. Flowers in distinct terminal inflorescences. Styles during anthesis free for c. 14-144; stigma distinctly
2-lobed (fig. 39j, 42e).
2. Corolla salver-shaped, 1-134 cmlong. Fruits 5-714 by 2-314 mm; styles only connected by the stigma

or free. Peduncle with some pairs of appressed, bract-like leaves .. . . . 1. M. elata
2. Corolla urceolate to campanulate, up to 14 cm long. Fruits c. 114 mm 9; styles still about halfway
connate. Peduncle without appressed, bract-like leaves . . . . .. 2. M. pygmaea

1. Flowers solitary, axillary, sometimes terminal, not in distinct peduncled inflorescences. Styles during
anthesis at least halfway free; stigma truncate or rarely slightly 2-lobed (e.g. fig. 45e).
3. Stem distinctly 4-ribbed to narrowly 4-winged, usually glabrous. Testa coarsely reticulate.
3. M. indica
3. Stem terete to somewhat flattened, nearly always pubescent. Testa minutely reticulate (fig. 47g—h)
or more often densely warty (fig. 46g).
4. Leaves on the lower surface with distinctly thickened midrib and margins, c. 244 by 114 mm;
internodes about as long as the leaves. Sepals distinctly more than halfway connate.
4. M. saxatilis
4. Midrib and margins not distinctly thickened, leaves longer and narrower, at least 4 by 1 mm;
internodes at least in the upper part of the plant much longer than the leaves. Sepals about halfway

connate.
5. Leaves and sepals with a narrow white membranous margin. spa 314 mm, glabrous. Anthers
sagittate, | mm long, introrsely dehiscent . . . . 5. M. albomarginata

5. Leaves and sepals without a white margin. Calyx up to 2, mm, ciliate. Anthers about elliptic,
up to 14 mm long, extrorsely dehiscent.
6. Flowers terminal and in the upper leaf-axils. At least the lower leaves elliptic, 4-6 mm long,
squamulate-ciliate along the margin . . . . 6. M. neglecta
6. Flowers exclusively in the upper leaf-axils. All jeaves feneeo late: 5 9 mm ions: glabrous.
7. M. bogoriensis

380 FLORA MALESIANA [ser. I, vol. 6?

1. Mitrasacme elata R. Br. Prod. (1810) 453;
DC. Prod. 9 (1845) 11; F. v. M. Fragm. 1 (1859)
132; Bru. Fl. Austr. 4 (1869) 354; F. M. Bat.
Queensl. Fl. 3 (1900) 1018; Compr. Cat. Queens].
Pl. (1913) f. 313; Ewart & Davies, Fl. North.
Terr. (1917) 219.—M. nudicaulis REINW. ex BL.
Bijdr. (1826) 849; DC. Prod. 9 (1845) 12; Mia.
Fl. Ind. Bat. 2 (1857) 361; BoerL. Handl. 2 (1899)
458; non Bru. in Hook. J. Bot. Kew Misc. 5 (1853)
56, et Aucr. Div. (= mostly M. pygmaea var.
Pygmaea, sometimes var. malaccensis, or M.
erophila LEENH.).—M. trinervis SPAN. Linnaea 15
(1841) 335; DC. Prod. 9 (1845) 560; Mia. FI. Ind.
Bat. 2 (1857) 361.—M. longiflora F. v. M. ex Bru.
Fl. Austr. 4 (1869) 354; F. M. Bart. Queensl. FI.
3 (1900) 1018; Ewart & Davies, Fl. North. Terr.
(1917) 219.—Fig. 39-40.

Erect annual, 35—85 cm. Leaves radical, rosulate
(often already withering during anthesis), sessile,
oblong to lanceolate, 45-414 by 4-114 cm,
fleshy, glabrous, narrowed at the base, apex blunt,
3(—5)-nerved. Inflorescences terminal and some-
times axillary, laxly paniculate with up to 4 lateral
branches per node (usually irregular and few-
flowered); peduncle long, c. 114 mm 9, terete,
smooth to minutely fluted, glabrous to minutely
tomentose especially near the base, with some
distant pairs of appressed bract-like leaves, acute-
lanceolate, c. 5 mm long, connate at the base,
minutely ciliate. Bracts similar but smaller and
patent. Pedicels 2-314 cm. Calyx campanulate,
3-8 mm long, for 14-34 connate, glabrous, tube
membranous, lobes oblong and acute or semi-
orbicular and apiculate, thickened. Corolla salver-
shaped, inside sparsely pubescent in the lower
half of the tube, and with a thickened fimbriate
rim in the mouth, outside (creamy or) salmon-
pink to brownish, inside creamy to white; tube
slender, 1-134 cm, lobes elliptic to oblong, (2—)
4-12 mm, shortly acuminate. Filaments c. 6 mm
long, anthers basifixed, ovate-lanceolate to linear
or sagittate, 1-2 mm, blunt, latero-introrse. Ovary
1-2 mm, tapering into the styles, which are con-
nate for the far greater part, 5-9 mm long, slender;
stigma 2-lobed, 1 mm long. Capsule for the greater
part enveloped by the persistent calyx, slender- to
globular-mitre-shaped, 5-714 by 2-314 mm, styles
either torn loose or only connected by the stigma
when mature. Seeds angular, minutely warty.

Distr. Tropical Australia and SE. Malaysia:
Lesser Sunda Is. (Sumba, Solor, Timor), and New
Guinea. Fig. 41.

Ecol. Savannahs and parklands, up to 2000 m.

Fig. 39. Mitrasacme elata R. Br. var. elata.
a. Habit, x 2/,, b. flower, X 1, d. calyx, X 2,

f. opened corolla-tube, x 2, g. anther, X 7,
h. pistil, < 3144, i. ovary, X 7, j. stigma, X 7,
k. cross-section of ovary, X 7, /. fruit, xX 2, m.

seeds, X 4, m. embryo, Xx 48.—var. brevicalyx

LEENH. c. flower, X 1, e. calyx, X 2 (a-b, d,

f-m VAN RoyeEN 5027, c and e SPANOGHE in herb.
L 908.127-526).

Dec.

LOGANIACEAE (Leenhouts)

1962]

TN

Fig. 40. Mitrasacme elata R. Br., Andjai, Kebar

Valley, W. New Guinea (vAN ROYEN & SLEUMER
6756) (SLEUMER, 1961).

var. elata.—M. elata R. Br.

Calyx 4-8 mm long, connate for 14-24, lobes
oblong, acute (fig. 39d). Fruit oblong, 6-714 mm
long.

Distr. Australia (Arnhem Land) and Malay-
sia: New Guinea (also in Fergusson I.).

Ecol. Dry savannahs and parklands, from sea-
level up to 2000 m. FI. fr. Jan.—Dec.

var. brevicalyx LEENH. Bull. Jard. Bot. Brux. 32
(1962) 446 .—M. nudicaulis REINW. ex BL.—
M. trinervis SPAN.—M. longiflora F.v. M. ex BTH.

Calyx c. 3 mm long, connate for 24—34, lobes
mostly semi-orbicular and apiculate (fig. 39e).
Fruits subglobular, c. 5 mm g.

This var. brevicalyx is primarily based upon, and
thus typified by M. trinervis SPAN. (holotype
SPANOGHE 5.7. in L).

Distr. Australia (Northern Territory and
Queensland), in Malaysia: Lesser Sunda _ Is.
(Sumba, Solor, Timor), and some dubious spe-
cimens from E. New Guinea.

Ecol. Grassland and bare slopes, at 450-1000
m. Fi. fr. March—June.

Notes. ‘M. elata’ and ‘M. longiflora’ are
clearly distinguishable in Australia, as there, apart
from the varietal characters cited above, the for-
mer is a distinctly more robust plant with much
larger flowers. The differences fade away in New
Guinea, however, where many of the specimens
are very slender and have small flowers as ‘M.
longiflora’, but share the long calyx and oblong
fruit with typical M. elata. These proved to be the
only “either ...or...’’ characters.

Fig. 41. Distribution of Mitrasacme elata R. BrR.,
representative for those species restricted to trop-
ical Australia and adjacent parts of Malaysia.

The structure of the inflorescence is somewhat
complicated. In a well developed specimen the
lower one or two nodes have four lateral branches,
two collateral ones in every axil. In many speci-
mens the central axis is less well developed or
completely suppressed, the number of lateral
branches may be less, and their size may differ
considerably.

The nearest related species is M. connata R. Br.
from tropical Australia which differs by the
following characters: leaves not rosulate but some

382

FLORA MALESIANA

[ser. I, vol. 6?

pairs close to each other at the base, linear-
lanceolate, cauline leaves 34—1 cm long, inflores-
‘cence more-flowered, styles fully connate, stigma
truncate.

Another closely allied species is the Australian
M. exserta F. v. M., which is at first sight different
by its dense, umbellate inflorescences.

2. Mitrasacme pygmaea R. Br. Prod. (1810) 453;
DC. Prod. 9 (1845) 11; Bru. J. Linn. Soc. Bot. 1
(1856) 92; Fl. Austr. 4 (1869) 357; F. M. Barn.
Queensl. FI. 3 (1900) 1019; BANKs & Sov. Bot.
Cook’s Voy. 2 (1901) t. 208; Baku. f. in Back.
Bekn. Fl. Java (em. ed.) 7 (1948) fam. 170, p. 6;
Hara, J. Jap. Bot. 30 (1955) 23, incl. var. malac-
censis, excl. syn. M. nudicaulis et trinervis; LEENH.
Bull. Jard. Bot. Brux. 32 (1962) 452.—M.
polymorpha (non R. Br.) CLARKE in Hook. f.
Fl. Br. Ind. 4 (1883) 80, incl. var. parishii, excl.
syn. M. trinervis; Forpes & HEmMsL. J. Linn. Soc.
Bot. 26 (1889) 118, incl. var. grandiflora; BOERL.
Handl. 2 (1899) 450 & 458; KiNG, J. As. Soc.
Beng. 74, ii (1908) 599; Dop, Bull. Soc. Bot. Fr.
57, Mém. n. 19 (1910) 5; Fl. Gén. I.-C. 4 (1914)
158, f. 20 (3-4); GampsLe, FI. Madras 5 (1923)
864; Merr. En. Philip. 3 (1923) 311; Rip. FI.
Mal. Pen. 2 (1923) 413; Merr. & CHUN, Sunyat-
senia 5 (1940) 171; GurmLLAuMiIN, FI. Nouv. Cal.
(1948) 285; HENpDERsS. Mal. Nat. J. 6 (1950) 304,
f. 288; Kerr in Craib, Fl. Siam. En. 3 (1951) 53;
MAKINO, Ill. Fl. Japan, rev. ed. (1954) 217, f.
650.—M. capillaris WALL. in Roxb. Fl. Ind. 1
(1820) 420; D. Don, Prod. Nepal. (1825) 129;
DC. Prod. 9 (1845) 11; Bru. J. Linn. Soc. Bot. 1
(1856) 92, excl. syn. M. trinervis; Mig. Fl. Ind.
Bat. 2 (1857) 362; Bru. Fl. Hongk. (1861) 230;
NAKAI, Fl. Korea 2 (1911) 96.—M. nudicaulis
(non Bu.) Bru. in Hook. J. Bot. Kew Misc. 5
(1853) 56, p.p., excl. syn. M. chinensis GRIsEB.; FI.
Hongk. (1861) 230; Fl. Austr. 4 (1869) 355;
Fors. & Hems.i. J. Linn. Soc. Bot. 26 (1889)
117; K. Scu. & Laur. Nachtr. (1905) 348; MerRR.
& Rotre, Philip. J. Sc. 3 (1908) Bot. 120; Dop,
Fl. Gén. I.-C. 4 (1914) 157; GitG & BENED. Bot.
Jahrb. 54 (1916) 163; Ewart & Davies, FI.
North. Terr. (1917) 219; GitG & BENED. Bot.
Jahrb. 56 (1921) 547; Merr. En. Born. (1921)
490; En. Philip. 3 (1923) 311; Rov. Fl. Mal. Pen.
2 (1923) 413; CAmMMeRL. Bull. Jard. Bot. Btzg III,
5 (1923) 301; Nova Guinea 14 (1924) 116; STEEN.
Trop. Natuur 25 (1936) 41-42, 159 f. 3; Hara,
J. Jap. Bot. 16 (1940) 155; En. Sperm. Jap. 1
(1948) 130; GuimLLAuMmIN, FI. Nouv. Cal. (1948)
285; Kerr in Craib, Fl. Siam. En. 3 (1951) 52.—
M. malaccensis WiGur, Ic. 4, 4 (1850) 15, t. 1601.
—M. alsinoides (non R. Br.) MerRR. Philip. J. Se.
1 (1906) Suppl. 116.—Androsace tonkinensis
Bonatl, Bull. Soc. Bot. Genéve II, 5 (1914) 298.—
M. micrantha Domin, Bibl. Bot. 89 (1929) 1071.—
M. setosa (non HANCE) MASAMUNE, J. Geobot.
10 (1961) 1, f. 43.—Fig. 42-43.

Erect, annual herb, up to c. 35 cm, branched
mainly at the base. Stem terete, slightly fluted,
fairly densely white hirsute in the lower part,
sometimes glabrescent or subglabrous. Leaf pairs

Fig. 42. Mitrasacme pygmaea R. BR. var. pygmaea.

a’. Habit, < 34, f. fruit, x 7.—var. malaccensis
(WIGHT) HARA. a. Habit, = 34, b. flower, X 7,
c. opened corolla (note the extrorse anthers),
X 7, d. anther, X 221A, e. pistil, < 12, g. seeds,
x 221%, (a MERRILL 3093, a’ RAHMAT 6059,
b-e BS 19208, f vAN STEENIS 6006, g ForBES 3884).

Dec. 1962]

LOGANIACEAE (Leenhouts)

383

spaced, the upper 2 pairs sometimes rosulate at
the base of the inflorescence, in small specimens
(nearly) all the leaves radical rosulate; ovate or
elliptic to linear-lanceolate, 214-19 by 1-6 mm,
herbaceous, subglabrous to rather densely hirsute
above, hirsute along the margin and beneath on
the midrib and sometimes on the nerves, base
cuneate, apex blunt to acute, 1—3-nerved. Jn-
florescences terminal, umbellate-paniculate, up
to c. 25 cm, long-peduncled, lax,!~ —1-flowered,
usually glabrous; peduncle terete to distinctly
grooved. Bracts oblong-ovate to -lanceolate, up
to 3 mm long, ciliate along the margin and some-
times along the midrib beneath. Pedicels slender,
angular, up to 4 cm (in fruit up to 144 cm).
Calyx conical to conical-campanulate, 11-24%
mm, about halfway connate or somewhat less,
glabrous except some bristles along the margin
or at the top of the acute-triangular lobes. Corolla
urceolate to campanulate, 3-5 mm long, inside
often with a few scattered hairs in the mouth,
yellow, outside white to pink with brownish or
pale orange dots or streaks, lobes 34-144 mm,
rounded. Filaments 11/,-3 mm, anthers basifixed,
extrorse, oblong-ovate to sagittate, c. 1 mm long,
acute. Pistil 2-414 mm, ovary 34-1 mm, styles free
in their basal 14—14 part, stigma 2-lobed. Capsule
globular-mitre-shaped, c. 114 mm 49, the styles
remaining connate for the greater part. Seeds
angular-ellipsoid, densely minutely warty.

Distr. Tropical Australia, New Caledonia,
through Malaysia to the Carolines (Jap, Palau),
and SE. Asia from Nepal to Korea and Japan.

var. pygmaea.— M. pygmaea R. BrR.—M. nudicaulis
AUCT.

Up to 12(—20) cm. Leaves radical rosulate, up
to 8 by 3 mm, I-nerved (fig. 42a’).

Distr. NW. India (Garkhwal, Kumaon),
Siam (Rachasima, Prachinburi), Cambodia, South
China (Yunnan), Formosa, Korea, Japan (Hon-
shu to Ryukyus), Caroline Is. (Jap), Australia
(Northern Territory, E. Queensland), New Ca-
ledonia, in Malaysia: Malay Peninsula (Malacca:
Bukit Sialang near Sampang Ampat), North and
Central Sumatra, Java (Preanger, Mt Merapi,
Mt Idjen), Lesser Sunda Is. (Bali, Lombok,
Alor, Timor), Philippines (Luzon), S. Moluccas
(Aru Is.), and New Guinea (also Goodenough I.).

Ecol. On rocks, in grasslands, open spots in
savannahs, in Eucalypt forest, along roadsides,
near solfataras and fumaroles, at (0—)1000—1800
(-3150) m. Fi. fr. Jan—Dec., mainly May-July.
The abnormal high altitude of 3150 m is near the
fumaroles on summit of Mt Agung, Bali; VAN
STEENIS (1936) discussed this peculiar habitat
and considers the seed to have been dispersed
epizoically to this spot by the pilgrimage of man,
or by monkeys, deer or other animals. See fig. 43.

Vern. Kdadnaquila, lassak, watakiki, Alor Is.

var. malaccensis (WIGHT) HARA (1955).—M.
polymorpha Auct.—M. capillaris WaAtLi.—M.
malaccensis WiGHTt.—Androsace tonkinensis Bo-
NATI.—M. micrantha DomMIN.

Up to 35 cm. Leaves not rosulate (or only the

upper 2 pairs), usually 8-19 by 2-6 mm, often
3-nerved (fig. 42a).

Distr. Nepal, Burma, Siam, Indo-China, S.
China, Hainan, Formosa, Japan (Okinawa,
Kyushu, Honshu), Caroline Is. (Palau), to Aus-
tralia (Northern Territory, Queensland), in
Malaysia: Malay Peninsula, Sumatra (also Riouw
Arch. and Billiton), E. Java (Besuki: Situbondo),
Lesser Sunda Is. (Bali, Alor, Timor), Borneo
(also Anambas Is.), Philippines (Luzon), SE.
Celebes, Moluccas (Sula Is., Ambon, Tanimbar
Is.), and New Guinea (also Sudest I.).

Ecol. In rocky localities, on bare soil, open
spots in savannahs, alang fields, lawns, road-
sides, Melaleuca forest, both in very dry spots and
in marshy places, from sea-level to 1000(—2000) m.
The flowers are only open in the middle of the
day. Fi. fr. Jan.—Dec., mainly Sept.—April.

Vern. Bingbis-lupa, Philip. Tag.,
gomme, gommoh gumiseh, New Guinea.

gomme

Fig. 43. Mitrasacme pygmaea R. BR. var. pygmaea
between mosses near fumaroles at 3150 m, Mt
Agung, Bali (VAN STEENIS, 1936).

Notes. Apart from the two varieties cited
above, there are two more: var. parishii (CLARKE)
LEENH. from Burma and var. grandiflora (HEMSL.)
LeeNH. from SE. China and Tonkin. Both are
characterized by distinctly larger flowers; the
former is connected with var. malaccensis by
intergrades, mainly in Indo-China.

As the varieties pygmaea and malaccensis are
mainly altitudinal vicariants, not every specimen
can be included in one of them. Moreover, espe-
cially towards the north (Japan) the differences
become vague also in the lowland specimens.

The only difference I found between the Aus-
tralian specimens of var. pygmaea and the
Malaysian ones is that the former have a sparsely
hirsute inflorescence.

The synonymy cited above clearly shows that
there has been considerable confusion about the
correct name of this species. This was mainly
caused by BENTHAM’s misinterpretation of M.
nudicaulis (and of M. trinervis which he put into

384

FLORA MALESIANA

[ser. I, vol. 6?

the synonymy of M. capillaris) and by CLARKE’s
concept of the Australian M. polymorpha in which
he included with some doubt the Nepalese M.
capillaris. The interpretations of these prominent
botanists were commonly followed. Only recently
Hara (1955), in his careful studies on the typifi-
cation of Japanese plants, showed the correct
application of these names.

Indeed, var. malaccensis is in habit closely alike
M. polymorpha R. Br., a species restricted to sub-
tropical E. Australia. The latter species differs
mainly from M. pygmaea var. malaccensis in the
following characters: pedicels long (2-3 cm in
flower, up to 314 cm in fruit), calyx long (5 mm),
corolla large (7 mm, the lobes 3 mm), connective
ending in a liguliform incurved appendix, stigma
truncate, fruit exceeded by the calyx, larger (c.
214 mm @Q), and with free styles, seeds ellipsoid,
rounded at both ends, rather large (0.6 mm
instead of 0.35—0.4 mm), shining black instead of
brown, sparsely warty.

Var. pygmaea is since BENTHAM (1853) confused
with an overlooked species from Assam, M.
erophila LEENH. This species differs from var.
pygmaea by the following characters: the presence
of sterile bracts along the peduncle (these are very
exceptional in var. pygmaea), the slightly zygo-
morphic corolla with relatively long lobes and a
more densely pubescent mouth, stamens which are
inserted above the middle of the corolla-tube (in
M. pygmaea distinctly below the middle), short
styles which are nearly fully connate during an-
thesis, but free in fruit, and the fairly coarsely
reticulated seeds.

In his description of M. micrantha, DOMIN
mentioned by error a 5-lobed calyx; in all three
type-specimens it is 4-merous.

In herbaria, Mollugo nudicaulis LAMK (Aizoaceae)
is sometimes mixed up with this species on account
of a superficial resemblance.

3. Mitrasacme indica WiGuT, Ic. 4, 4 (May 1850)
15, t. 1601; Illustr. Ind. Bot. 2 (1850) t. 156
f. 1-13; Bru. in Hook. J. Bot. Kew Misc. 5 (1853)
56; J. Linn. Soc. Bot. 1 (1856) 92; Fl. Austr. 4
(1869) 356; F. M. Bart. Queensl. Fl. 3 (1900)
1019; Ewart & Davies, Fl. North. Terr. (1917)
219.—M. pusilla Daz. in Hook. J. Bot. Kew
Misc. 2 (May 1850) 136.—M. crystallina GRIFF.
Not. 4 (1854) 87; Icon. (1854) t. 383 f. 2.—M.
alsinoides (non R. BR.) CLARKE in Hook. f. Fl. Br.
Ind. 4 (1883) 80; Fors. & Hems-. J. Linn. Soc.
Bot. 26 (1889) 117; TRIMEN, FI. Ceyl. 3 (1895) 170;
BoerRL. Handl. 2 (1899) 450 & 458; Dop, Bull.
Soc. Bot. Fr. 57, Mém. n. 19 (1910) 4; Fl. Gén.
I.-C. 4 (1912) 157; CAMMeERL. Bull. Jard. Bot.
Btzg III, 5 (1923) 300; GAMBLE, FI. Madras 5
(1923) 864; Merr. En. Philip. 3 (1923) 311;
Rip. Fl. Mal. Pen. 2 (1923) 413; BACKER, Onkr.
Suiker. (1931) 487; Merr. & Perry, J. Arn.
Arb. 23 (1942) 410; BAKH. f. in Back. Bekn. FI.
Java (em. ed.) 7 (1948) fam. 170, p. 5; MASAMUNE
& Syozi, Act. Phytotax. Geobot. 12 (1950) 202;
KerR in Craib, Fl. Siam. En. 3 (1951) 52; MAKINO,
Ill. Fl. Japan, rev. ed. (1954) 217, f. 649; Hara,

J. Jap. Bot. 30 (1955) 24, incl. var. indica —
Oldenlandia brachyphylla Mrrr. Fl. Manila (1912)
448.

Annual, up to 15 cm. Stems usually branched,
slender, distinctly 4-ribbed to narrowly 4-winged,
often slightly compressed, glabrous or nearly so;
internodes 2-4 times as long as the leaves. Leaves
lanceolate to ovate, 4-7 by 114-214 mm, herba-
ceous, glabrous, apex acute, l-nerved. Flowers
solitary in the upper leaf-axils. Pedicels 3-7 (in
fruit 5-25) mm, glabrous to scabrous. Calyx
conical, 134-2 mm long, glabrous, about halfway
connate, lobes lanceolate, acute. Corolla about
campanulate, 3-4 mm long, shortly and thinly
bearded in the mouth, white, lobes 1-114 mm long,
rounded. Filaments 1 mm; anthers _basifixed,
introrse, elliptic, rarely sagittate, c. 274 mm long,
sometimes with an apical appendix. Piszil 14 mm,
ovary nearly 14 mm, styles far apart at the base,
connate in the upper half, stigma broadly truncate
(to 2-lobed). Capsules subglobular, c. 2 mm @,
styles connate only at the apex. Seeds angular-
ovoid, 0.2 mm, brown, coarsely reticulate.

= —_ > =;

=e) y

— . — Fes VA ie Ll

Fig. 44. Distribution of Mitrasacme indica WIGHT,
as an example of a wide-spread species.

Distr. Ceylon, Deccan Peninsula, Upper
Burma, Indo-China (Tonkin), coastal regions of
China as far north as Shantung (Chefoo Mts),
Korea (Chemulpo), Japan (Honshu to Ryukyus),
Formosa, Hainan, Australia (Northern Territory,
Queensland), in Malaysia: Sumatra (West Coast),
Malay Peninsula (Surat, Puket, Kelantan), Java
(also Madura and Kangean Is.), Lesser Sunda Is.
(Sumba), SW. Celebes (Malino), Philippines
(Luzon), and Southern New Guinea. All citations
from Borneo seem to go back to CLARKE (1883,
1.c.); | did not see any specimen from that island.
Fig. 44.

Ecol. Open places of all kinds, on poor to
heavy soils, dry to marshy, sandy or on limestone,
from sea-level to c. 500 m. Fi. fr. Jan.—Dec.,
mainly March—June.

Notes. One of the species usually referred to
as M. alsinoides R. Br. Well distinguishable by its

Dec. 1962]

LOGANIACEAE (Leenhouts)

385

glabrous, ribbed to winged stems and its coarsely
reticulated seeds.

The names M. indica WicHtT and M. pusilla
DALz. were apparently both published in May
1850. BENTHAM (1853, /.c.) was the first who
reduced M. pusilla to M. indica. DALZIELL & GIB-
SON (Bombay FI. 1861, 155) stated under M.
pusilla: “This was afterwards named M. indica
by Wight Ic. 1601.” This remark does not neces-
sarily refer to the dates of actual publication.

4. Mitrasacme saxatilis BACK. in Cammerl. Bull.
Jard. Bot. Btzg III, 5 (1923) 301, f. 1; Onkr. Suiker.
(1931) 487.

Erect, annual (or perennial?) herb, up to 12 cm,
branched at the base. Stems terete, canaliculate,
scabrous, internodes about as long as the leaves.
Leaves sessile, ovate, 214 by 114 mm, stiff, glabrous,
broadly cuneate at base, apex acute to aristulate,
margin revolute, distinctly thickened as is the
midrib on the lower surface. Flowers solitary in the
upper leaf-axils. Pedicels 114-244 mm, slender,
scabrous. Calyx campanulate, 1144 mm long,
glabrous, connate for 2/4, lobes triangular, acute,
midrib strongly thickened as are the margins near
the apex. Corolla 2mm long, bearded in the mouth,
white, lobes 24 mm long, nearly acute. Anthers
subsessile, introrse, 14-14 mm long. Pistil small,
styles 1/4; mm long, far apart at the base, connate
in the upper half. Capsules broadly mitre-shaped,
c. 114 mm high, styles torn loose. Seeds ellipsoid,
brown, minutely warty.

Distr. Malaysia: E. Java (near Banjuwangi)
and Madura I.

Ecol. In open vegetation, on cliffs and dry
limestone hills, locally gregarious; from sea-level
up to 400 m. Fi. fr. May-July.

Note. A very characteristic species: stiff and
erect, dense, with sessile, small, ovate leaves which
have conspicuously thickened margins and mid-
rib, calyx with relatively short lobes which also
have strongly thickened margins and midrib, fruit
nearly completely enveloped by the calyx and with
very short styles.

5. Mitrasacme albomarginata LeEENH. Bull. Jard.
Bot. Brux. 32 (1962) 442.—Fig. 45.

Annual herb, up to c. 10-15 cm. Stems flattened
(also in vyivo?), minutely appressed-pubescent,
internodes up to about twice as long as the leaves.
Leaves lanceolate (rarely ovate-lanceolate) 5—6 by
114 mm, herbaceous with a narrow, white, membra-
nous margin, glabrous, contracted at base, aris-
tulate at apex, l-nerved. Flowers solitary in the
upper leaf-axils. Pedicel 14-1 cm, minutely pu-
bescent. Calyx conical-campanulate, 314 mm long,
glabrous, about halfway connate, lobes acute-
triangular, with a narrow white membranous
margin and aristulate like the leaves. Corolla
white, campanulate, 444 mm, inside with long
bristles in the mouth and some short hairs at
the apex of the lobes; lobes elliptic, acute, 2 mm
long. Stamens short, filaments 1.2 mm, strap-
shaped; anthers basifixed, introrse, sagittate, 1 mm
long, acute and with some short bristles at the

Fig. 45. Mitrasacme albomarginata LEENH. a.

Habit, = 1, b. leaf (note the membranous margin),
x 5, ec. connate leaf-sheaths, < 5, d. flower,
x 5, e. opened corolla, x 5, f. anther, X 10,
g. pistil, X 10, #. fruit, x 5, i. seed, X 20 (all

from VAN ROYEN 4847).

apex. Pistil 2 mm, ovary globular, 34 mm, styles
inserted far apart, connate in their upper half,
stigma truncate. Fruits globular, 14, mm go,
completely enveloped by the calyx-tube, styles
soon separating from each other. Seeds angular,
0.2 mm, minutely warty.

Distr. SW. New Guinea, once collected.

Ecol. In grasslands, open places with very low
vegetation, c. 8 m alt., locally common. Fi. fr.
Sept.

Note. Similar to the East Australian M.
paludosa R. Br. which is distinctly different in
several floral characters, amongst others a fleshy
corolla and extrorse anthers with a robust apical
hook.

386

FLORA MALESIANA

[ser Ie svoleioe

6. Mitrasacme neglecta LEENH. Bull. Jard. Bot.
Brux. 32 (1962) 449.—Fig. 46.

Annual herb, up to 15 cm, simple or branched,
mainly near the base. Stems terete, especially near
the base, minutely fluted, fairly densely squamu-
late-hairy to subglabrous, internodes several times
as long as the leaves. Leaves ovate or elliptic, upper
ones often (ovate-)lanceolate, 6 by 3 to 4 by 1 mm,
herbaceous, with squamulate hairs on the margin,
otherwise glabrous, apex acute; |-nerved. Flowers
terminal and in the upper leaf-axils; as the leaves
gradually become smaller and narrower higher up,
and the internodes become longer, the upper part
may assume the appearance of a lax raceme.
Pedicel 6-7 mm (in fruit up to 1 cm) long, sca-
brous. Calyx widely conical, 2 mm high, about
halfway connate, lobes lanceolate, acute, ciliate,
especially towards the apex. Corolla subcampanu-
late, white, 3 mm long, inside with some bristles
in the mouth, lobes rounded, | mm long. Filaments
c. 74 mm; anthers basifixed, extrorse, ovate to
elliptic, 4% mm long, with a minute apical appen-
dix. Pistil 1% mm, ovary broad-cylindrical, 1mm
high, styles inserted far apart, connate in the upper
4-4 part, stigma truncate. Fruits globular-
mitre-shaped, 114 mm @, styles connected only
by the stigma. Seeds elliptic, slightly flattened,
minutely papillose.

Fig. 46. Mitrasacme neglecta LEENH. a.
24, b. flower, < 4, c. opened corolla (note the

Habit,

extrorse anthers), < 4, d. anther, = 16, e. pistil,
x 8, f. fruit, x 4, g. seed, x 30 (a-e sine coll.,
s.n. in herb. L 909.67-695, f-g CoerT 917).

x 4,
d. anther, 16, e. pistil, x 8, f. fruit, xX 4,
g. and h. seeds, = 42 (all from BACKER 6218).

* 4, c. opened corolla,

Distr. Australia (Arnhem Land) and Ma-
laysia: E. Java, Madura, Lesser Sunda Is. (Bali,
Sumba, Timor), and SW. Celebes (Bt Parangpeda
near Banto Parang).

Ecol. On beach walls, in fields, in rocky lo-
calities, on limestone hills, on bare rocks, between
grass and low scrub; from sea-level up to 925 m.
Fl. fr. March-April (July).

Note. From the closely related M. indica
immediately distinguished by the terete, pubescent
stem and the slight though distinct differentiation
between the non-flowering and the flower-bearing
part of the plant.

7. Mitrasacme bogoriensis LEENH. Bull. Jard. Bot.
Brux. 32 (1962) 444.—Fig. 47.

Erect annual herb up to c. 18 cm, simple or
branched in the upper half. Stems terete, rather

Dec. 1962]

LOGANIACEAE (Leenhouts)

387

densely squamulate-hairy, internodes about as
long as the leaves, the upper ones much longer.
Leaves (ovate-)lanceolate, decreasing in size up-
wards, 5-9 by 1-114(-2'4) mm, herbaceous,
glabrous, margin revolute, apex acute, aristulate;
l-nerved. Flowers solitary in the upper leaf-axils,
by the long internodes, small leaves, and long
spreading pedicels seemingly forming a lax raceme.
Pedicel c. 34-244 cm long, pubescent. Calyx
conical-campanulate, 114-214 mm, about halfway
connate, lobes acute-triangular, sparsely ciliate a-
long the margin and on the prominent midrib
towards the apex. Corolla white, urceolate-cam-

lobes 34-114 mm, rounded. Filaments 2/4; mm;
anthers basifixed, extrorse, oblong-ovoid, 0.3—14
mm long, truncate. Pistil 1/2 mm, ovary 0.4 mm,
styles inserted far apart, connate in their upper
half, stigma broadened truncate, slightly 2-lobed.
Fruits globular-mitre-shaped, 2 mm 9, styles still
connate in their upper half. Seeds angular, rather
minutely reticulate, brown.

Distr. W. Java (Bogor), three collections.

Ecol. In grass fields, along grassy roadsides,
100-400 m. Fi. fr. April, Dec.

Note. Well characterized by its slender habit
and narrow leaves.

panulate, 3-414 mm long, bearded in the mouth,

Excluded

Cleisocratera Kortu. Kruidk. (1844) 256 was originally as a monotypic genus assigned to the Rubiaceae,
but Miquet (FI. Ind. Bat. 2, 1857, 384) included it tentatively in the Loganiaceae. MERRILL (En. Born.
1921, 576) remarked that it had been referred incidentally to Psychotria but believed this to be wrong.
KortTHALs himself already remarked its close proximity to Saprosma and VALETON (Ic. Bog. 3, 1909,
209) reduced it to the latter genus, an opinion which, after examination of the type specimen, is also
shared by Dr BAKHUIZEN VAN DEN BrINK f. and Dr VAN STEENIS (Rubiaceae).

Dolianthus C. H. Wricut, Kew Bull. (1899) 106 was originally assigned to Loganiaceae, but according
to BREMEKAMP (Kew Bull. 1936, 103) it is Rubiaceous and is allied to Amaracarpus. MERRILL & PERRY
(J. Arn. Arb. 27, 1946, 221) accept it as belonging to a set of Papuan, high altitude species of that genus
(Rubiaceae).

Gaertnera LAMK, Tabl. Enc. Bot. Ill. 2 (1791) 273, t. 167 was originally placed in Loganiaceae and has
up to the present often been treated as belonging to this family. On account of the resemblance in flower
structure (apart from the more or less superior ovary), in stipules, habit, and anatomy to the Rubiaceae-
Psychotrieae, with which it also shares the presence of raphides, recent authors on Rubiaceae include it in
that family and tribe. See Petit, Bull. Jard. Bot. Brux. 29 (1959) 377 and Verpcourt, ibid. 28 (1958)
238 (Rubiaceae).

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FLORA MALESIANA

SCIENTIFIC COMMUNICATIONS
concerning Flora Malesiana should be addressed to the General Editor,
Dr C. G. G. J. VAN STEENIS, Rijksherbarium, Leyden, Holland

Volume 1.
Volume 2.
Volume 3.
Volume 4.
Volume 5.

*

SUBSCRIPTIONS
should exclusively be directed to N.V. DE ERVEN P. NooRDHOFF,
P.O. Box 39, Groningen, Holland

Cyclopaedia of collectors & collections. 1950. pp. clii. + 639. Dfl. 55.50, £ 5.10.-, US$ 14.65.
Malaysian vegetation. Sty Jai tact oceee ts elem eds Fie

Malaysian plant geography .
General chapters and revisions. 1948 - 1954, pp. ccix + 631.
Bibliography, specific delimitation, & revisions.

In preparation.
: In preparation.
‘DA. 69. , £6.16.-, US$ 18.15.

1955-1958. pp. coealii + 596. Dfl. 92.-,

£ 9.1.-, US$ 24.20.

For sale only , Not for exchange

INDEX TO REVISED FAMILIES

Aceraceae . 4: 3, 592
Actinidiaceae s.str. 4: 37
Aizoaceae 4: 267
Alismataceae . S52 317
Amaranthaceae 4: 69, 593
Ancistrocladaceae 4: 8
Aponogetonaceae 4: il
Basellaceae 5: 300
Batidaceae 5: 414
Betulaceae 5: 207
Bixaceae s.str. 4: 239
Burmanniaceae 4: 13, 592
Burseraceae 5: 209
Butomaceae 5: 118
Callitrichaceae 4: 251
Campanulaceae . 6: 107
Cannabinaceae 4: 223
Capparidaceae . 6: 61
Caprifoliaceae . 4: 1s 598
Celastraceae-i . 6: 227
Centrolepidaceae . 5: 421
Ceratophyllaceae . 4: 41
Chenopodiaceae . 4: 99, 594
Cochlospermaceae 4: 61
Combretaceae 4: 533
Connaraceae. . . 5: 495
Convolvulaceae . 4: 388, 599
Corynocarpaceae . 4: 262
Crassulaceae . 4: 197
Datiscaceae 4: 382
Dichapetalaceaé . 5: 305
Dilleniaceae 4: 141

Dioscoreaceae
Dipsacaceae
Droseraceae
Elatinaceae
Erythroxylaceae .
Ficoidaceae
Flacourtiaceae
Flagellariaceae
Gnetaceae.
Gonystylaceae
Goodeniaceae
Haemodoraceae .
Hamamelidaceae .
Hydrocaryaceae .
Hydrocharitaceae
Hydrophyllaceae
Juglandaceae .
Juncaceae
Juncaginaceae
Loganiaceae
Malpighiaceae
Martyniaceae
Molluginaceae
Moringaceae .
Myoporaceae
Myricaceae
Najadaceae
Nyssaceae
Papaveraceae
Pedaliaceae . .
Pentaphragmataceae .
Pentaphylacaceae
Philydraceae .

293
: 290
Be if
: 203
543
: 267
1
: 245
: 336
: 349
25935
Fal
: 363
43
: 381
: 207
: 143
4: 210

haUNEaMSHRSAS

ALNSANND

Phytolaccaceae 4: 229
Pittosporaceae 5: 345
Plumbaginaceae 4: 107

Podostemaceae . . 65
Polemoniaceae .-:.* 4: 195
Pontederiaceae” . . 4:'255

Primulaceae . . . 6: 173
Proteaceae «.. apts a
Punicaceae oe RO
Restionaceae . . . 5:416
Rhizophoraceae 5: 429
Salicaceae «2s eee
Salvadoraceae. . . 4:225

Sarcospermaceae . . $32

Saururaceae 4: 47
Scyphostegiaceae . 5: 297
Simaroubaceae. 6: 193
Sonneratiaceae 4: 280, 513
Sparganiaceae o| - ot meee.

Sphenocleaceae . . 4: 27
Stackhousiaceae . . 4: 35
Staphyleaceae ao See Gee 49

Stylidiaceae 4: 529
Styracaceae . . . 4: 49
Thymelaeaceae 4: 349, 6:1
Trigoniaceae . . . 4: 59
Turneraceae . . . 4: 235
Typhaceae . . . 4: 243
Umbelliferae . 4: 113, 595
Valerianaceae. . . 4: 253
Xyridaceae . 4: 366, 598

Zygophyllaceae . . 4: 64

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N.V. DIJKSTRA’S DRUKKERIJ V/H BOEKDRUKKERIJ GEBR. HOITSEMA - GRONINGEN