FM1S2010019001001

Flora Malesiana, Series I, Volume 19 (2010) 1– 333
CUCURBITACEAE
(W.J.J.O. de Wilde & Brigitta E.E. Duyfjes, Leiden)1
Cucurbitaceae Juss., Gen. Pl. 393 (1789), nom. cons.; Miq., Fl. Ned. Ind. 1, 1 (1856)
652; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 604; Cogn. in A.DC. & C.DC.,
Monogr. Phan. 3 (1881) 340; Pax in Engl. & Prantl, Nat. Pflanzenfam. 4 (5) (1889) 9;
Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 3; Cogn. & Harms in Engl., Pflanzenr.
88, 4.275.2 (1924) 2; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 292; Keraudren
in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 3; I.Telford,
Fl. Australia 8 (1982) 158; W.J.de Wilde & Duyfjes, Fl. Thailand 9, 4 (2008) 411;
A.M.Lu, Lu Q.Huang, S.K.Chen & C.Jeffrey, Fl. China, draft (2009); H.Schaefer &
S.S.Renner, Fam. Gen. Vasc. Pl. [Kubitzki], in press.
Type genus: Cucurbita L.
Small or large climbers; herbaceous or woody; annual or perennial; leafy stems of-
ten grooved or angular; roots fibrous or tuberous; monoecious or dioecious. Probract
present or absent (see below). Tendrils simple or branched, extra-axillary, present on
each node. Leaves dispersed, simple or foliolate; leaf base usually cordate, apex acute
(–acuminate); stipules absent. Inflorescences: flowers either solitary, or fascicled, or
in racemes or in panicles; bracts minute to large, or absent; perianth usually 5-merous,
small to large, composed of mostly free sepals and free or partly fused petals, rarely ±
zygomorphic; disc usually present. Male flowers with shallow or tubular receptacle;
stamens 3 (anthers all three 2-thecous or two 2-thecous and one 1-thecous) or stamens 5
(anthers 1-thecous), filaments free or (partly) united, anthers free, coherent or connate,
thecae straight, curved, plicate or contorted, connective narrow or broad, without or
with apical appendage. Female flowers: perianth similar to male flowers; ovary largely
or completely inferior, usually 3-locular, ovules few or numerous, (secondarily) parietal
or apical, horizontal or hanging; staminodes present or absent; style either 1 and stig-
mas 3 (lobed or not) in subfam. Cucurbitoideae, or styles 3(– 5) and stigmas 2-lobed
in subfam. Fevilleoideae. Fruit small to large, either berry-like and pulpy and usually
not opening (subfam. Cucurbitoideae), or capsular with 3 apical valves (subfam. Fevil
leoideae). Seeds variable in size and shape, often compressed, unwinged or winged;
embryo large, straight; endosperm absent.
There are 2 subfamilies, see Taxonomy.
DISTRIBUTION
A family of about 120 genera and 900 species, distributed all over the world, the major-
ity in the tropics with most genera in America, a few in temperate climatic zones, but
not occurring in cold areas. Most genera have only few species. The larger genera in
America are e.g., Cayaponia, Gurania, Cyclanthera, Melothria, and Sicyos (American
1) With contributions by P. Baas (wood anatomy) and C.B. Mennes & R.W.J.M. van der Ham (pollen
morphology). — Drawings by J.H. van Os except Fig. 52.
Flora Malesiana, Ser. I, Vol. 19 (2010)
and Pacific); in Africa e.g., Momordica, Cucumis, Pilogyne, and Neoachmandra; in

south-eastern Asia e.g., Trichosanthes (c. 100 species), Hemsleya (China, c. 20 spe-
cies), Pilogyne, Neoachmandra, and Thladiantha (mainly China, 20 species, of which
1 or 2 in Malesia).
Most genera are confined to a single continental area, only a few are found in two con-
tinents, e.g. Pilogyne and Neoachmandra, both occurring in Asia and Africa.
Species distributional areas vary from those with bi-continental distributions, such as
Mukia maderaspatana, to those with intermediate ranges, like Trichosanthes globosa,
down to restricted local-endemics like Borneosicyos, or else they are known from only
one or a few collections, or from a single locality like Pilogyne trichocarpa or Anangia
macrosepala. The genus Muellerargia contains 1 species in eastern Malesia and north-
ern Australia and 1 species in Madagascar.
In Malesia there are 29 native and 10 cultivated genera, the cultivated ones frequently
with running wild species, notably in Benincasa, Citrullus, Cucumis, Cucurbita, Cy
clanthera, Lagenaria, Luffa, and Sechium. Benincasa and Luffa contain wild as well
as cultivated taxa. Introduced wild-running species are found in the genera Cayaponia
and Melothria. The largest genera in Malesia are: Trichosanthes (43 species), Pilogyne
(14 species), and Neoachmandra (12 species). Endemism is high in Borneo and New
Guinea (particularly in Trichosanthes). In total 132 species of the Cucurbitaceae are
treated in the present flora.
HABITAT & ECOLOGY
Tendril climbing Cucurbitaceae occur worldwide in virtually all sorts of habitats, pro-
vided the climate is not too cold in winter. Bryonia species in Europe hibernate by their
deep tubers which sprout in spring. Tuberous species are also common in the seasonal
tropics. The tree-shaped, non-climbing, pachycaul endemic Dendrosicyos of dry So-
cotra Island, is unusual, although in eastern Malesia some species of Neoalsomitra have
a pachycaul stem-base.
Malesian cucurbits are all climbers, most of them of small to medium length, but some
reaching the tall forest canopy, notably Alsomitra macrocarpa, a liana with a stem up
to 15 cm diameter. In general they seem to prefer more or less disturbed (primary) for-
est or scrub in the vicinity of (running) water. Limestone species sprawl on the rocks
but usually root in the deeper soil below. Species with the highest elevation in Malesia
are found in Java (Pilogyne repanda, to 2700 m), and in Papua New Guinea (Gynos
temma papuana, to 3500 m). The incompletely known Anangia macrosepala, endemic
to Kabaena Is, northern Moluccas, apparently grows each season from a subterranean
tuber in yearly burnt vegetation.
Most species of Malesian Cucurbitaceae are perennial, without or with a woody root-
stock, or with one or several tubers. Others are annual with mostly berry-like fruits that
are dispersed by birds. Both annual and perennial species of subfamily Fevilleoideae
mostly have capsular fruits which shed winged seeds. Mukia maderaspatana is usually
De Wilde & Duyfjes — Cucurbitaceae
subannual, but plants with old, woody rootstocks are known from Java. Some species
of Trichosanthes and possibly also Bayabusua clarkei are monocarpous, dying-off after
prolific fruit production.
TAXONOMY
Jeffrey (1962, 1990) made efforts to improve the classification of Cucurbitaceae, the
latest version (Jeffrey 2005) was modified on the basis of seed coat anatomy (Singh &
Dathan 1998). Jeffrey divided the family into two subfamilies: Nhandiroboideae and
Cucurbitoideae. Subfam. Nhandiroboideae (= Fevilleoideae, syn. Zanonioideae) con-
tained one tribe with 5 subtribes, two of which included Malesian genera. The subfam.
Cucurbitoideae consisted of 10 tribes, some with 2 subtribes, most of them containing
Malesian genera.
In the most recent treatment of the whole of Cucurbitaceae which is based on molecular
data as well as on morphological cladistics (Schaefer, Heibl & Renner 2009, Schaefer
& Renner (in press (for Kubitzki)), subfamilies are not recognized, instead 104 genera
are placed within 15 tribes, 9 of which occur in Malesia. These are:
Tribe 1 — Gomphogyneae (Alsomitra, Bayabusua, Gomphogyne, Gynostemma, and
Neoalsomitra).
Tribe 3 — Zanonieae (Zanonia).
Tribe 6 — Thladiantheae (Baijiania, Thladiantha).
Tribe 7 — Siraitieae (Siraitia).
Tribe 8 — Momordiceae (Momordica).
Tribe 12 — Sicyeae (Cyclanthera, Hodgsonia, Luffa, Sechium, and Trichosanthes
(including Gymnopetalum)).
Tribe 13 — Coniandreae (Kedrostis).
Tribe 14 — Benincaseae (Benincasa, Borneosicyos, Citrullus, Coccinia, Cucumis
(including Mukia), Diplocyclos, Indomelothria, Lagenaria, Melothria, Mueller
argia, Papuasicyos (including Urceodiscus), Scopellaria, Solena, and Zehneria
(including Anangia, Neoachmandra, Pilogyne).
Tribe 15 — Cucurbiteae (Cayaponia, Cucurbita).
In the present Flora Malesiana treatment two subfamilies as circumscribed by Jeffrey
(2005) are followed but under the names Fevilleoideae and Cucurbitoideae. Various
genera to be sunk by Schaefer, Heibl & Renner (2009) and Schaefer & Renner (in
press (for Kubitzki)) are maintained on the basis of flower- and fruit-morphology. The
subfamily Fevilleoideae generally are considered as more primitive (basal) then the
Cucurbitoideae. According to Mennes & Van der Ham (see under Pollen Morphology)
the two subfamilies differ significantly in pollen characters.
FEVILLEOIDEAE
Fevilleoideae Burnett, Outl. Bot. (1835) 756, 1092, 1129.

Nhandiroboideae Kostel., Allg. Med.-Pharm. Fl. 2 (1833) 722, nom. illeg.
Zanonioideae Luerss., Handb. Syst. Bot. 2 (1882) 1080; C.Jeffrey, Kew Bull. 15 (1962) 345.
Dioecious. Tendrils distally 2-branched, spiralling below point of branching. Male

flowers in panicles or racemes. Stamens 5, inserted on narrow, flat receptacle, free or
united, disc not obvious, anthers small, 1-thecous (except Alsomitra and Bayabusua:
stamens 3). Pollen small, usually striate, prolate. Styles (2 or) 3 (or 5), free. Ovary
truncate at apex, not narrowed; ovules pendent. Fruits dehiscing, capsular, apically
opening with 3 incurving valves releasing (mostly) winged seeds (except Gynostemma,
p.p. indehiscent).
Six genera in Malesia — Alsomitra, Bayabusua, Gomphogyne, Gynostemma, Neoal
somitra, and Zanonia.
Note — The genus name Nhandiroba Kostel. is invalid because it is homotypical
with Fevillea, the older name, and Fevillea L. was mentioned as a synonym. Therefore
the subfamily name Nhandiroboideae Kostel. (1833) cannot be used.
CUCURBITOIDEAE
Monoecious or dioecious. Tendrils simple, or branched mostly at or below the middle

(in Siraitia and Sinobaijiania distally 2-branched) and then either only the branches
spiralling, or the part below the point of branching also spiralling (Sinobaijiania, not in
Malesia). Male flowers solitary, fasciculate, in racemes or in panicles. Stamens 3 (or 5),
inserted in the receptacle-tube, mostly free; anthers small or large, 1- or 2-thecous, disc
present, but may not be obvious. Pollen various, larger than in subfam. Fevilleoideae,
isodiametric, reticulate. Style 1. Ovary constricted at apex; ovules mostly horizontal.
Fruit a juicy or dry berry (pepo-fruit), mostly indehiscent (in Luffa with an opercu-
lum).
Thirty-one genera in Malesia — Anangia, Baijiania, Benincasa, Borneosicyos,
Cayaponia, Citrullus, Coccinia, Cucumis, Cucurbita, Cyclanthera, Diplocyclos, Gym
nopetalum, Hodgsonia, Indomelothria, Kedrostis, Lagenaria, Luffa, Melothria, Mo
mordica, Mukia, Muellerargia, Neoachmandra, Papuasicyos, Pilogyne, Scopellaria,
Sechium, Siraitia, Solena, Thladiantha, Trichosanthes, and Urceodiscus.
POLLINATION AND DISPERSAL
Flowers in Cucurbitaceae are unisexual, the plants either monoecious or dioecious.

Hermaphroditic flowers are rare, occurring mostly in cultivated species and are typical
in Neoachmandra hermaphrodita (endemic to Thailand). Sometimes they are parthe-
nocarpic (e.g. cultivated Cucumis).
Cross-pollination by insects seems most likely throughout the family, but little is
known for certain for most species. A variety of insects, including hoverflies (Syrphi-
dae) have been seen visiting the flowers. Pollination by oil-collecting oil bees (genus
Ctenoplectra) in the tribe Thladiantheae is remarkable (Vogel 1990; Schaefer & Renner
2008a; Renner & Schaefer 2010). Trichosanthes (most species) and some related genera
(Gymnopetalum, Hodgsonia) flower at night. Their tubular, sweetly scented, flowers
are possibly pollinated by sphingidae, or other nocturnal moths. We ourselves observed
at night the very swift visits by hawk-moths to flowers in a large sprawling stand of
Gymnopetalum scabrum.
The local endemic Borneosicyos has pollen in tetrads, presumably hinting at pollina-
tor type.
In some groups, e.g., Trichosanthes and Pilogyne, the connective of the anthers is
conspicuously set with comparatively stout, short hairs, which is also possibly related
to pollination mechanics.
SEED DISPERSAL
The conspicuously coloured, berry-like, pulpy fruits of most Cucurbitaceae suggest

that seed dispersal is by animals eating the fruit. The pendulous fruit of Momordica,
(notably M. charantia), splits open at the apex, revealing the seeds enveloped in bright
red pulpy arils.
Gynostemma pentaphylla fruit is small, berry-like, green-yellow or purple, con-
taining 1 or 2 seeds only. The single globose seeds of Neoachmandra sphaerosperma
(Thailand), have been seen carried away by ants.
In South Africa the fruit of Cucumis humifructus is geocarpous and aardvarks play
a role in fruit and seed dispersal.
In many Trichosanthes species the fruit pulp is green-black and bitter, in others it is
yellowish, orange or whitish and not bitter; in T. villosa it is sweet. The fruit of Siraitia
grosvenorii (cultivated) is known for its strong sweetness (Jeffrey 1979).
As noted above, species with capsular fruit, (mainly the subfamily Fevilleoideae,
but also the fruits of Luffa aegyptiaca (synonym L. cylindrica)) open by valves or by
an operculum at the apex, through which they shed their usually winged seed, although
sometimes the wing can be narrow or absent.
Cultivated species or feral forms of these are generally dispersed by man. Their
seeds often germinate from waste fruits, the plants either temporarily establishing or
persisting for a longer time by roadsides, in waste places, abandoned fields, secondary
scrub, or on or near rubbish dumps.
GERMINATION AND SEEDLING
Germination can be hypogeal or (mostly) epigeal, even in one genus, e.g., in Momordica
cochinchinensis it is epigeal, but in the closely related M. denticulata it is hypogeal.
The first true leaves develop generally after the leaf-like cotyledons enlarge and are
shed, and soon they produce a lateral tendril. However, in Bayabusua clarkei the first
four leaves are in one whorl.
SEXUAL CONDITION
Flowers are unisexual (with few exceptions). Whether a species is monoecious or dio-
ecious can be determined by the presence of male flowers as well as female flowers or
fruits on the same herbarium specimen, or preferably by observing living plants. Sexual
condition is relatively constant within a genus, e.g. Cucumis is monoecious, Trichosan
thes dioecious (T. cucumerina excepted), Momordica species can be either dioecious or
monoecious, the economically important M. charantia is monoecious.
INDUMENTUM, COLOUR OF PLANT ON DRYING
Trichosanthes species are variously hairy: some are conspicuously villose or strigose,
but most are subglabrous, or glabrescent when the hairs disappear with age. The hairs
may be grey or brown. Most species have a whitish (or black or brown), chalky puncta-
tion (cystoliths; cf. Zimmermann 1922) originating from the hair scars or hair bases,
especially on the upper leaf-surface, rendering the leaves scabrid. In several species
white cystoliths can be found on the stem, petiole and nerves of the lower leaf-surface.
The presence of hairs within flowers can be diagnostic (Zimmermann 1922). Siraitia
is covered with blackish gland-hairs.
With experience, the drying colour of herbarium specimens is useful for identification:
species of Neoachmandra generally dry greenish, those of Pilogyne brown or blackish.
Erkens et al. (2008) made an assessment of age and greenness of herbarium specimens
as predictors for successful extraction and amplification of DNA, including some Cu-
curbitaceae.
TENDRILS
Cucurbitaceae differ from other scandent-climbing plant families by the presence of a

tendril on each node, which is never truly axillary (conspicuously so in subfamily Cu-
curbitoidae), nor opposite the leaf, but always at some angle with the leaf-petiole. The
node, bearing a leaf, a lateral tendril and often a lateral vegetative shoot, when fertile
together with a flower or an inflorescence, is very characteristic for Cucurbitaceae, an
assemblage which is known as the nodal syndrome (Rugayah & De Wilde 1997). The
morphological origin of the tendril is unknown, but most likely it is of cauline nature
(Zimmermann 1922), and a process of concaulescence towards a higher node during its
phylogeny may explain its lateral position. Whether the tendril is simple or branched,
and whether it coils exclusively above the point of branching, or below as well, is taxo-
nomically important. The latter condition is found in the Asian genera of the subfamily
Fevilleoideae and in Sinobaijiania (Cucurbitoideae, China).
LEAVES AND LEAF GLANDS
Leaves provide many useful taxonomic characters. They may be palmately or pedately
compound (with petiolulate leaflets) or simple, with the blade either entire or variously
(deeply) lobed. Lobing may be very variable within a species. The presence of small
glands on the probracts and sepals can be characteristic, and these are frequently vis-
ited by small black ants. Glands on the lower surface of the leaf blade are common and
their size, number and position provide good distinguishing characters for species of
Trichosanthes. Leaves of juvenile plants may be greatly different from those of adult
plants. The leaves of juvenile specimens of T. tricuspidata and related species are much
dissected and they may look very much the same mutually.
PROBRACTS
Characters of the probract have been neglected by most previous authors. Probracts
occur singly at each node and their presence and shape is usually typical for a genus.
The shape, consistency and the presence of glands are useful characters, for distinguish-
ing species of Trichosanthes. The probract can best be seen on young shoots. They
can be conspicuous, linear-lanceolate, ovate, concave or flat, or they can be small and
caducous; they are absent in some genera or species. The morphological origin of the
probract is unknown, but possibly they are derived from the bract of a solitary or lower
most flower in an inflorescence (Zimmermann 1922).
INFLORESCENCES AND FLOWERS
Flowers are solitary or (few-)fascicled in the leaf axils, or arranged in an axillary

raceme (indeterminate). By contraction and reduction of foliage leaves to bracts the
inflorescences may become (large) panicles. In dioecious species, male flowers are
either solitary, or more usually arranged in a pedunculate, bracteate raceme, often with
a single co-axillary male flower. Solitary female flowers develop at the nodes, or form
(peduncled) clusters, e.g. in Pilogyne. In monoecious plants, solitary female flowers
can be found at the nodes or (developing previously) beside the male raceme; which
develops later (usually, female flowers develop after the male flowers). In some species
of Trichosanthes the persistent pedicels of senescent or undeveloped solitary flowers
appear as an elongated or ‘straw-like appendage’ beside the male raceme.
Bracts — The bracteate racemes of male flowers bear persistent or caducous bracts,

either on the rachis, or in some species, higher-up on the pedicels. They differ in size,
shape, and consistency, varying from linear to obovate or rhomboid, with entire, dentate
or laciniate margins, are glabrous or sparsely to densely hairy, and may bear glands.
Flowers — The flowers are mostly unisexual, either white or yellow, and the perianths
of male and female flowers are generally similar. In nocturnal species of Hodgsonia,
Gymnopetalum, and Trichosanthes, the flowers are white, opening in the late afternoon
or at night and close before sunrise, when the fallen corollas can be found on the ground.
Other species e.g. T. cucumerina are largely diurnal.
Pedicel — The pedicel is mostly persistent and is articulated at or near its apex below
the flower. The pedicels of solitary male or female flowers are much longer than those
of the flowers in a raceme.
Receptacle-tube — The receptacle-tube is present in subfamily Cucurbitoideae. It is

shallow, saucer- or cup-shaped, or tubular and carries the free sepals and free or partly
connate petals on its rim, and the stamens inside.
Sepals — There are always 5 sepals, inserted on the margin of the receptacle (subfam.
Fevilleoideae) or on the rim of the receptacle-tube (subfam. Cucurbitoideae).
Sepals are typically much smaller than the petals; (narrowly) triangular or linear, but
are frequently enlarged, in Cucurbita moschata, or are larger than the petals in Anangia.
Their shape in male flowers, whether entire or lobed, is taxonomically important in

Trichosanthes.
Corolla — The colour of the flowers, ( i.e. of the petals), is usually either white or yel-
low. Some genera have a more indistinct flower colour: creamy, greenish creamy, or
whitish with green veins. In Trichosanthes the usually white petals are in some species
reddish veined, or either reddish or yellow-fringed. The petals of Bayabusua are by
exception purple red.
The petals are (almost) free, or in some genera united at base, e.g. in Cucumis, Mukia,
very distinctly so in Coccinia, Cucurbita.
Male flowers — The number of stamens is 3 or 5. In subfamily Fevilleoideae there are

five stamens of one theca, the filaments free, or partially or wholly connate in a column.
In subfamily Cucurbitoideae five (free) stamens only occur in the tribe Thladianthineae,
where 4 stamens are arranged in two pairs, and 1 solitary. In the other tribes there are
three stamens, of which either a pair of two-thecous anthers and the other one-thecous
(e.g., Cucumis, Mukia, Melothria, Indomelothria), or else all three stamens have a
two-thecous anther (e.g., Pilogyne, Neoachmandra). In many genera the thecae are
connivent into a globose whole, or connate into a short column (e.g., Trichosanthes
(mostly), Hodgsonia, Cucurbita). The thecae can be straight, curved, hooked, plicate or
3-plicate (S-shaped). The connective is often markedly dilated, with the thecae along a
wavy margin, and when connivent, resulting in a globose synandrium with the thecae
irregularly arranged into a tangle or ball.
The place of insertion of the stamens in the receptacle-tube is taxonomically impor-
tant, e.g., at the base in Pilogyne, at or near the throat in Neoachmandra. In the centre
of male flowers a conspicuous (nectary?) gland is often present, (which is mostly not a
pistillode, because its homologue is usually present in female flowers as a broad collar
around the base of the style).
Female flowers — The perianth is similar to the male flowers, but may differ slightly in
size and shape, e.g., the sepals in Trichosanthes quinquangulata: entire in the female,
are usually with few sidelobes in the male.
The ovary is inferior and 3-carpellate. It is very variable in detail in subfam. Cucur-
bitoideae, but it is always demarcated by a more or less pronounced constriction. In
subfam. Fevilleoideae the ovary is broadly attached to the receptacle, bearing 3 separate
styles, each usually furcate into a 2-lobed stigma. In subfam. Cucurbitoideae there is a
single style, and the stigma essentially 3-parted but quite variable, from subentire (e.g.
Benincasa, Cucurbita) to markedly divided (e.g. Anangia), with papillose or feather-
like stigmatic surface.
The basic construction of the ovary is described by Eichler (1875). Ovules are mostly
numerous, placed horizontally, attached to secondary-parietal placentas. The aberrant
ovary of Hodgsonia is derived from the basic type by assuming a reduction of the
ovules (De Wilde & Duyfjes 2001). In Fevilleoideae the three carpels often do not fuse
completely, leaving the ovary ± unilocular and the ovules are pendent (later on erect in
the pendulous fruit).
Nectary & staminodes — A ring-shaped nectary, as well as staminodes are often

present.
Fruit — In subfam. Cucurbitoideae it usually is a rather dry or juicy, indehiscent berry;

the pericarp can be woody, or firm and thick, to filmy. The ripe fruit exocarp can be
green, yellow, orange or red, or often paler (white) and flecked, striped, flamed or
banded. Seeds are few or many, attached to 3 parietal or ± septal placentas, each thought
to be formed from two carpel edges from adjoining carpels, best to be seen in the de-
veloping ovaries, because in the ripe fruit the content is juicy and pulpy. The seeds are
embedded in green-black, red or white pulp. The fruit in subfam. Fevilleoideae is a
woody capsule, opening with three incurving valves and shedding winged seeds.
Seeds — These are very variable and distinctive for genera or species. Seeds of the
capsular-fruited Fevilleoideae are usually winged (Gynostemma excepted). Seeds can
be compressed, sculptured, and often possess a ‘margin’, all useful characters. Seeds
of e.g. Diplocyclos and some Trichosanthes species seem 3-loculed with the embryo
in the middle compartment. The two wart-like outer compartments can be hollow, or
filled with large erect cells developed from the seed coat, either from the margin or
the face. Seed coat anatomy is highly characteristic and has been used in taxonomy,
notably for subdivision of the family into tribes (Sing & Dathan 1998; Jeffrey 2005).
See also Corner (1976) who provided detailed notes on the seeds of several genera of
Cucurbitaceae. Rugayah & De Wilde (1999: fig. 1), depict a choice of Trichosanthes
seeds according to the sections.
The use of the words margin and edge in the descriptions — For leaves, bracts and
sepals the word margin is used, which may be e.g. entire or dentate, etc. For seeds it is
different: seeds are usually more or less, or strongly compressed (subglobose to almost
flat), and seen at the faces there can be a distinct (or faint) narrow or broad border-
ing zone, called the margin, around the seed, differing from the rest of the seed-face
in thickness, colour or surface (the seed-face can be smooth or sculptured). The very
outer side of the seed showing the outline or profile is called the edge, described as e.g.
straight (smooth), undulate, crenulate.
CHROMOSOMES
There is a large diversity in chromosome number within Cucurbitaceae. Schaefer &

Renner (for Kubitzki, in press) report numbers for all genera, where known. It is note-
worthy that the cucumber (Cucumis sativus) and the much resembling melon (Cucumis
melo), are in different subgenera, and the chromosome numbers are: x = 7 (2n = 14) and
x = 12 (2n = 24) respectively (Kirkbride, 1993). Within the genus Cucumis the number
varies, 2n = 14, or 24, or sometimes 20, 22, 48 or 72.
USES
Jeffrey (2001) gives a detailed worldwide account of cultivated Cucurbitaceae, while

in PROSEA 8 (1993) all useful Cucurbitaceae occurring in Malesia are treated in detail
10 Flora Malesiana, Ser. I, Vol. 19 (2010)
a b c
d e f
g h i
j k l
Plate 1. Pollen of mainly Malesian Cucurbitaceae; scanning electron micrographs of more or less polar
views of 3-colporate monads, unless stated otherwise. Scale bar = 5 µm (a – f), 10 µm (g – l).
a – f: subfamily Fevilleoideae; g – l: subfamily Cucurbitoideae. a. Alsomitra macrocarpa (Blume)
M.Roem., perforate to indistinctly rugulate pollen grain (Java, De Wilde & Duyfjes 21886); b. Baya
busua clarkei (King) W.J.de Wilde, partly syncolporate striate pollen grain (Peninsular Malaysia, De
Wilde & Duyfjes 21886); c. Gomphogyne cirromitrata f. minor W.J.de Wilde & Duyfjes, striate pollen
grain (Thailand, Phonsena et al. 4013); d. Gynostemma pentaphyllum (Thunb.) Makino f. pentaphyllum,
striate pollen grain (Java, De Wilde & Duyfjes 21890); e. Neoalsomitra schefferiana (Cogn.) Hutch.
subsp. schefferiana, striate pollen grain (Moluccas, Kornassi 1433); f. Zanonia indica L. subsp. orien
De Wilde & Duyfjes — Cucurbitaceae 11
by several authors. Young sprouts and flowers of all cultivated Cucurbitaceae are ed-
ible. In addition to Gildemacher, Jansen & Chayamarit (1993) can be mentioned that
of New Guinean Trichosanthes species with red fruit pulp the fruits and seeds are ed-
ible, especially when cooked in a young state. Edible Cucurbits of SE Asia have been
enumerated by De Wilde & Duyfjes (2008 (‘2007’)).
WOOD ANATOMY
(Pieter Baas)
Most Cucurbitaceae are herbaceous, and the secondary xylem of woody vines and
shrubs in the family is very poorly documented or even completely unknown when
considering species from the Flora Malesiana region. We owe a classical study of
the internal phloem of the bicollateral primary vascular bundles to Worsdell (1915).
Zimmermann (1922) described the intricate stem vasculature of many taxa, including
species of Coccinia, Lagenaria, Momordica and Sechium. Carlquist’s detailed wood
anatomical account (1992) of 4 species included Coccinia grandis and Zanonia indica.
Metcalfe & Chalk (1950) and Schaefer & Renner (in press (for Kubitzki)) summarized
the general stem anatomy of both herbaceous and woody Cucurbitaceae. When woody,
the secondary xylem of Cucurbitaceae is typically characterized by wide, simply perfo-
rated vessels, sheathed by paratracheal parenchyma (that may be lignified or not) and/or
vasicentric tracheids, varying amounts of libriform fibres and unlignified apotracheal
parenchyma, and broad, unlignified rays (uniseriate rays are absent). Many of these
features are typical of the liana habit (Bamber & Ter Welle 1994; Carlquist 1992). The
fibres of Trichosanthes are nucleated (Rajput & Rao 1999). Successive cambia occur
in Luffa and Melothria (Carlquist 2001). The swollen stem bases of Neoalsomitra
podagrica are largely parenchymatous with little secondary growth in the outer ring of
collateral vascular bundles (pers. obs.).
POLLEN MORPHOLOGY
(C.B. Mennes & R.W.J.M. van der Ham)
The pollen of the Cucurbitaceae is nearly always shed as monads. Tetrads occur in a few
genera in subfamily Cucurbitoideae (tribes Benincaseae, Coniandreae, Thladiantheae).
Pollen grain size is small to very large (17– 208 μm) and pollen grain shape mostly
subspheroidal. The aperture system is mostly 3-zonocolporate, but 4 –16-aperturate,
talis W.J.de Wilde & Duyfjes var. pubescens Cogn., striate pollen grain (New Guinea, Hartley 10037);
g. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. borneensis, reticulate pollen grain (Borneo,
SAN 141929); h. Borneosicyos simplex W.J.de Wilde, brevicolporate gemmate tetrad (Borneo, SAN
144251); i. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. chlorocarpa, reticulate pollen
grain (Borneo, SAN 144096); j. Muellerargia timorensis Cogn., reticulate pollen grain (Sumba, Iboet
185); k. Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes subsp. marginata var. penangense
(C.B.Clarke) W.J.de Wilde & Duyfjes, brevicolporate reticulate-striate pollen grain (Sumatra, De
Wilde & Duyfjes 18234); l. Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes subsp.
belensis, reticulate-striate pollen grain (New Guinea, LAE 63476).
pantoaperturate, colpate and (operculate) porate systems also occur. The exine is usually
tectate; in the tribes Benincaseae, Cucurbiteae and Schizopeponeae intectate exines are
found. Pollen grains of Fevilleoideae (Plate 1a – f) are usually striate, while perforate
and reticulate are the most common ornamentation types in the Cucurbitoideae (Plate
1g – l). The Cucurbiteae, Schizopeponeae and Sicyeae show some clearly deviating pol-
len types. Below, Cucurbitaceae pollen is described per subfamily and/or tribe, based
on the literature provided by the ‘Bibliographic index to the pollen morphology of An-
giosperms’ (Thanikaimoni & Van der Ham 1999), a few more recent studies and own
observations. The taxonomy follows Schaefer et al. (2009), who recognised 15 tribes
with 128 genera, 37 of which occur in Malesia. The pollen of three genera is unknown
(see below). Pollen studies treating substantial numbers of genera include those by
Erdtman (1952), Marticorena (1963), Alyoshina (1971), Keraudren-Aymonin & Straka
(1984), Ayala-Nieto et al. (1988), Sharma (1991), Khunwasi (1998), Perveen & Qaiser
(2008) and Yang et al. (2008). Less inclusive studies are mentioned below.
Fevilleoideae
Tribes Actinostemmateae, Fevilleeae, Gomphogyneae, Zanonieae (19 genera; Malesia:

Alsomitra, Bayabusua, Gomphogyne, Gynostemma, Neoalsomitra, Zanonia).
Pollen grains monads, mostly small to medium-sized (17–52 μm) and subprolate,
sometimes subspheroidal or prolate. Aperture system 3-colporate. Ornamentation
mostly striate with long subparallel muri, sometimes perforate to indistinctly rugu-
late (Alsomitra, Gomphogyneae; Plate 1a), irregularly striate (Bolbostemma p.p.,
Actinostemmateae) or reticulate (Gerrardanthus, Zanonieae). Literature: De Wilde
et al. (2007a), Lira et al. (1998), Van der Ham (1999).
Cucurbitoideae
Tribe Benincaseae (32 genera; Malesia: Anangia, Benincasa, Borneosicyos, Citrullus,

Coccinia, Cucumis [incl. Mukia in Schaefer et al. 2009], Diplocyclos, Indomel
othria, Lagenaria, Melothria, Muellerargia, Neoachmandra, Papuasicyos, Pilogyne,
Scopellaria, Solena, Urceodiscus).
Pollen grains mostly monads, small to large (29–107 μm), subspheroidal, sometimes
oblate or prolate; pollen of Borneosicyos occurs in tetrads of c. 85 μm diam. (Plate
1h). Aperture system 3-, rarely 4- or 6-colporate, sometimes 3-brevicolporate to
-porate (Borneosicyos, Cucumis, Diplocyclos, Scopellaria, Urceodiscus; Plate 1h,
k, l). Ornamentation mostly perforate to reticulate, sometimes reticulate-striate with
short sinuous muri (Dactyliandra, Papuasicyos, Peponium, Scopellaria, Urceodis
cus; Plate 1k, l), reticulate-echinate (Diplocyclos), intectate echinate (Praecitrul
lus), gemmate (Borneosicyos; Plate 1h) or verrucate (Solena p.p.). The pollen of the
African-Madagascan Zehneria(?) peneyana deviates by being 6-porate (6-brevicol-
porate?), and that of Cephalopentandra by its operculate porate apertures and intec-
tate gemmate exine. Literature: De Wilde et al. (2004, 2006, 2007b), Duyfjes et al.
(2003), Van der Ham & Pruesapan (2006), Van der Ham & Van Heuven (2003).
Tribe Bryonieae (Austrobryonia, Bryonia, Ecballium; Malesia: absent).

Pollen grains monads, medium-sized to large (42 – 67 μm) and oblate spheroidal
to prolate. Aperture system 3-colporate. Ornamentation reticulate in Bryonia and
Ecballium, unknown in Austrobryonia.
Tribe Coniandreae (21 genera; Malesia: Kedrostis).

Pollen grains mostly monads, medium-sized to large (34 –104 μm), subspheroidal,
sometimes oblate or prolate; pollen of Gurania and Psiguria occurs in tetrads of
149 –185 μm diameter. Aperture system mostly 3-colporate, sometimes 3 – 4(5)-
colporate (Ceratosanthes), 4-colporate (Dieterlea) or 3 – 6-porate (Gurania, Psigu
ria). Ornamentation mostly perforate to reticulate, sometimes reticulate-striate with
short sinuous muri (Kedrostis p.p.) or locally intectate echinate (Gurania). Pollen
unknown: Tumamoca.
Tribe Cucurbiteae (13 genera; Malesia: Cayaponia, Cucurbita).

Pollen grains monads, large to very large (61– 208 μm), spheroidal. Aperture system
3–10-zono- or pantoporate; pores operculate. Ornamentation intectate echinate. Pol-
len unknown: Penelopeia. Literature: Barth et al. (2005), Teppner (2004).
Tribe Indofevilleeae (Indofevillea; Malesia: absent).

Pollen grains monads, mostly medium-sized (53 μm), oblate to subprolate. Aperture
system 3-colp(or)ate. Ornamentation coarsely reticulate.
Tribe Momordiceae (Momordica; Malesia: 5 species).

Pollen grains monads, medium-sized to large (65 –79 μm), oblate to prolate. Aperture
system 3-colporate. Ornamentation reticulate.
Tribe Schizopeponeae (Biswarea, Edgaria, Herpetospermum, Schizopepon; Malesia:

absent).
Pollen grains monads, medium-sized (23 – 49 μm; Schizopepon) or large to very
large (111–134 μm), subspheroidal. Aperture system 3-colporate (Schizopepon) or
3-porate with operculate pores. Ornamentation reticulate (Schizopepon) or baculate/
gemmate.
Tribe Sicyeae (24 genera; Malesia: Cyclanthera, Gymnopetalum, Hodgsonia, Luffa,

Sechium, Trichosanthes).
Pollen grains monads, medium-sized to very large (30 –168 μm), oblate to prolate,
but usually subspheroidal. Aperture system 3(4)-(brevi)colporate to 3-porate (e.g.
Gymnopetalum, Hodgsonia, Luffa, Trichosanthes) or 4 –16-colp(or)ate (e.g. Cycla
nthera, Sechium). Irregular aperture systems occur in several genera (e.g. Echinoc
ystis, Pseudocyclanthera, Rytidostylis). Ornamentation mostly perforate-rugulate to
reticulate, sometimes tectate echinate (e.g. Sechium), rarely verrucate (Trichosanthes
cucumerina). The pollen of Linnaeosicyos deviates within the Cucurbitaceae by the
presence of margos along the colpi and ridges connecting their apices. Literature:
Alvarado et al. (1992), Monro & Stafford (1998), Pruesapan & Van der Ham (2005),
Schaefer et al. (2008), Stafford & Sutton (1994).
Tribe Siraitieae (Microlagenaria, Siraitia; Malesia: Siraitia).

Pollen grains monads, mostly medium-sized (35 – 51 μm), subspheroidal, sometimes
suboblate or prolate. Aperture system 3-colporate. Ornamentation reticulate.
Tribe Telfairieae (Ampelosicyos, Cogniauxia, Odosicyos, Telfaria, Tricyclandra;
Malesia: absent)
Pollen grains monads, medium-sized to large (46 – 82 μm), spheroidal to subprolate.
Aperture system mostly 3(4)-colporate, sometimes brevicolporate (Tricyclandra).
Ornamentation mostly (coarsely) reticulate, sometimes reticulate-striate with short
sinuous muri (Ampelosicyos) or verrucate (Tricyclandra). Pollen unknown: Odosi
cyos.
Tribe Thladiantheae (Baijiania, Sinobaijiania, Thladiantha; Malesia: Baijiania, Thla
diantha)
Pollen grains mostly monads, medium-sized to large (26 –79 μm), oblate to pro-
late; the pollen of one Thladiantha species occurs in tetrads. Aperture system 3(4)-
colp(or)ate. Ornamentation (coarsely) reticulate. Literature: Zhang & Lu (1989).
Earlier accounts on the taxonomic and evolutionary significance of pollen characters in

Cucurbitaceae were given by Marticorena (1963), Jeffrey (1964) and Khunwasi (1998),
who agreed that the subdivision into two subfamilies, Fevilleoideae and Cucurbitoideae,
is well supported by pollen morphology. Kocyan et al. (2007) compared the pollen
diversity (unpublished data) in the family with a molecular phylogeny. He supported
the perception that the pollen characters were relatively conserved and he confirmed
the differences between the Fevilleoideae and Cucurbitoideae. Further, the special pol-
len types of the Cucurbiteae and Sicyeae (s.s.) and the parallel evolution of echinate
ornamentation in Diplocyclos, Praecitrullus (both Benincaseae) and the Cucurbiteae
and of tetrads in Gurania/Psiguria and Borneosicyos were noticed.
In the present study, pollen data of 123 out of the 128 genera in the family were
evaluated by plotting them on the molecular phylogeny including 127 genera (Khmerio
sicyos missing) by Schaefer et al. (2009). In this phylogeny, the genera traditionally
comprising subfamily Fevilleoideae (syn. Nhandiroboideae, Zanonioideae) occur in
four clades (tribes Actinostemmateae, Fevilleeae, Gomphogyneae, Zanonieae), which
together with subfamily Cucurbitoideae form a basal pentatomy (see also Schaefer &
Renner 2008b). The pollen characters considered are: 1) dispersal unit (monad, tetrad);
2) pollen grain size (17– 208 μm); 3) aperture number (2 –16); 4) aperture distribution
(zono-, pantoaperturate); 5) ectoaperture type (colpate, porate, operculate); 6) exine
(tectate, intectate); and 7) exine ornamentation (rugulate, perforate, reticulate, striate,
reticulate-striate, verrucate, echinate, gemmate).
Most common in Cucurbitaceae are medium-sized to large, 3-zonocolporate mon-
ads, which occur in all 15 tribes recognised by Schaefer et al. (2009). All genera in
the four basal tribes (Fevilleoideae), except Alsomitra (Plate 1a) and Gerrardanthus,
have striate pollen with long subparallel muri (Plate 1b – f), which clearly differs from
the reticulate-striate pollen with short sinuous muri (Plate 1k, l) found in eight genera
in three Cucurbitoideae tribes (Benincaseae: Dactyliandra, Papuasicyos, Peponium,
Scopellaria, Urceodiscus; Coniandreae: Kedrostis africana; Telfarieae: Ampelosicyos).
Papuasicyos, Scopellaria and Urceodiscus make up a clade within the tribe Beninca-
seae. Striate pollen with long subparallel muri is found in a few outgroup members of
the family (e.g. Anisophyllea, Begonia), but still it may represent a derived character
supporting the monophyly of the four Fevilleoideae tribes (Van der Ham et al. in press).
Most common in subfamily Cucurbitoideae is some form of perforate to reticulate orna-
mentation, which is present in 10 of the 11 tribes. The 3-zonocolporate reticulate pollen
of Khmeriosicyos, which genus is missing in the analysis by Schaefer et al. (2009),
clearly belongs to one of these 10 tribes, and would fit well in the tribe Benincaseae
(see De Wilde et al. 2004).
Derived pollen types occur especially in the more derived tribes of subfamily Cucur-
bitoideae: Benincaseae, Coniandreae, Cucurbiteae, Schizopeponeae and Sicyeae. These
types have combinations of the following characters: tetrads, more than three apertures,
pantoaperturate, brevicolpate or (operculate) porate ectoapertures, intectate exine and
reticulate-striate, verrucate, echinate or gemmate ornamentation. All these characters, ex-
cept maybe for pantoaperturate arrangement, evolved more than once in the derived Cu-
curbitoideae. Some groups show special combinations, e.g., operculate pores + intectate
exine + echinate ornamentation (Cucurbiteae) and 6 –12 colpate ectoapertures + echinate
ornamentation (Sicyeae: subtribe Sicyinae and Frantzia p.p.). Within the Sicyeae a trend
in aperture number is present. The more derived, the larger the number of apertures: 3(4)
(Nothoalsomitra, Luffa, Trichosanthes, Hodgsonia, Gymnopetalum), 4 (Linnaeosicyos),
4 – 6 (Marah, Echinocystis), 4 –16 (rest of tribe). Fossil Hexacolpites echinatus pollen
(6-colpate, echinate) from the Oligocene (34 – 23 mya) of Africa may be identified as
Sicyeae pollen (Muller 1981); it dates the split between the clade Marah-Echinocystis
and its sister clade at minimally 23 million years ago. Cephalopentandra (Benincaseae)
and the clade Biswarea-Edgaria-Herpetospermum (Schizopeponeae) have large to very
large, intectate, gemmate pollen with three operculate porate apertures, which would
represent a remarkable case of parallel evolution. Tetrads always have short colpi or pori
(without operculum). They are a synapomorphy of Gurania and Psiguria (Coniandreae),
but originated independently in Borneosicyos (Benincaseae).
CHEMOTAXONOMY
Detailed notes on the chemistry and chemotaxonomy of the Cucurbitaceae have been
provided by Hegnauer (1989) and Schaefer & Renner (in press (for Kubitzki)).
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seven returns from dioecy to monoecy and recent long-distance dispersal to Asia. Molec. Phylogen.
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KEYS TO THE GENERA
Note on the distinction of the subfamilies. — According to molecular phylogeny the

here recognized two subfamilies in Cucurbitaceae, Fevilleoideae and Cucurbitoideae
cannot be maintained because Fevilleoideae is paraphyletic (Schaefer & Renner 2008).
However, morphologically the distinction is quite clear, although a single quick and
user-friendly easily observable character in a collection is not always present. In prac-
tice one should check the presented characters in leads of both keys successively, which
readily will exclude the subfamilies from each other.
1 – GENERAL KEY FOR MALE FLOWERING MATERIAL
1a. Tendrils distally 2-branched. Inflorescences paniculate; bracts usually minute,

1– 2 mm long. Flowers small, petals less than 5 mm long (6 – 8 mm long in Al
somitra) whitish or pale off-yellow, or purple-red in Bayabusua, never yellow.
Stamens 3 (Alsomitra, Bayabusua), or 5 evenly spaced; filaments inserted on or
near the disc (if present). Pollen small, striate. (Styles 3. Fruit a capsule (except
Gynostemma)). — Subfam. Fevilleoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Tendrils simple or branched at or below the middle (except Baijiania and Siraitia
with tendrils distally 2-branched). Flowers solitary, or fascicled, or in racemes, or in
panicles but then petals usually larger; bracts small or large. Flowers small or large,
petals white, off-yellow or bright yellow. Stamens 3, or 5 but then 4 in two pairs
and 1 solitary; filaments inserted on the receptacle-tube. Pollen generally larger,
generally not striate. (Style 1. Fruit usually a berry). — Subfam. Cucurbitoideae . 7
2a. Large climber 15 – 30 m long. Stamens 3, anthers 3 or 5 . . . . . . . . . . . . . . . . . . . 3
b. Small or large climber 1–15 m long. Stamens 5, anthers 1 mm long or less, all one-
thecous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3a. Petals valvate. Anthers 3 – 4 mm long, 2 two-thecous, 1 one-thecous. Perianth

creamy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Alsomitra
b. Petals imbricate. Anthers c. 2 mm long, white, two filaments with two anthers, one
filament with one anther, all anthers one-thecous. Perianth red . . 4. Bayabusua
4a. Leaf blade simple, margin entire; petiole scar raised when dry. Inflorescences on
the older wood. Perianth fleshy, c. 6 mm diameter . . . . . . . . . . . . . . 37. Zanonia
b. Leaf blade often foliolate, margin entire or serrate; petiole scar not raised when
dry. Inflorescences terminal or among the leaves. Perianth membranous, small but
various of size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Filaments united into a solid column . . . . . . . . . . . . . . . . . . . . 16. Gynostemma
b. Filaments free (or united into a hollow column in Neoalsomitra hederifolia) . . 6
6a. Blade margin entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. Neoalsomitra
b. Blade margin serrate-dentate . . . . . . . . . . . . . . . . . . . . . . . . . 14. Gomphogyne
7a. Petals or corolla (petals free or partly united) 8 mm long or more (open corolla
more than 15 mm diam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
b. Petals or corolla less than 8(–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
8a. Petals yellow or greenish, or in Momordica, partly, near-white with dark blotch
at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
b. Petals white, sometimes green veined, or pink, rarely with some yellow . . . . . 19
9a. Flowers solitary, with conspicuous bract on flower stalk . . . . . . 23. Momordica
b. Flowers either solitary, or in clusters, racemes, or panicles, without conspicuous
bract on flower stalk (probract may be present or absent) . . . . . . . . . . . . . . . . 10
10a. Corolla pale greenish yellow. Filaments united. — [Flowers in short-stalked clus-
ters arranged in pseudo-racemes. Vigorous climber. Usually cultivated] . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31. Sechium
b. Corolla light or dark yellow. Filaments free (but anthers may be connate). — [Habit
of plant and disposition of flowers various] . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11a. Flower solitary or in (sub)sessile clusters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
b. Flowers in racemes or panicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
12a. Probract conspicuous. Petals free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
b. probract absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13a. Sepals larger, ± recurved. Leaf blade palmately veined or lobed . . 5. Benincasa
b. Sepals minute, patent or recurved. Leaf blade ± pinnately lobed . . . 8. Citrullus
14a. Tendrils unbranched. Flowers in sessile clusters; corolla c. 2 cm long or less, petals
connate for lower 1/3 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Cucumis
b. Tendrils branched. Flowers solitary, corolla larger, petals connate into a tube for
c. halfway . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. Cucurbita
15a. Tendrils (2-)3-more-branched at or below the middle. Probract present . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. Luffa
b. Tendrils unbranched or apically 2-branched. Probract absent or present . . . . . 16
16a. Receptacle and vegetative parts (especially leaves below) densely set with ap-
pressed blackish glandular hairs. Thecae conduplicate (hooked) . . . 32. Siraitia
b. Blackish glandular hairs absent. Thecae straight, curved, or convolute, (not
hooked) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a. Stamens 3, or 5, when 5 then four in two pairs, one single. Petals with basal scale
. . . . . . . . . . . . . . . . . . . . . . 34. Thladiantha (& 23. Momordica, p.p., namely
. . . . M. clarkeana, a species sometimes with a panicle-like male inflorescence)
b. Stamens 3. Petals without basal scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Flowers in a panicle. — Borneo . . . . . . . . . . . . . . . . . . . . . . . . 6. Borneosicyos
b. Flowers in a pedunculate raceme. — New Guinea . . . . . . . 36. Urceodiscus p.p.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (large-flowered species)
19a. Receptacle-tube short, cup-shaped. Petals largely fused. Filaments fused or tightly
coherent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Coccinia
b. Receptacle-tube elongate. Petals free. Filaments free (anthers coherent) . . . . . 20
20a. Apex of petiole with two glands at the transition to the blade . . . 20. Lagenaria
b. Apex of petiole without glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Stout climber. Probract thorn-like, triangular, coriaceous, with sharp apex. Petals
long fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. Hodgsonia
b. Plant of medium or small stature. Probract not thorn-like or absent . . . . . . . . . 22
22a. Petals not fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Gymnopetalum
b. Petals long-fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. Trichosanthes
23a. Anthers forming a closed continuous ring. — Introduced . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Cyclanthera (C. brachystachya)
b. Anthers not forming a closed ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24a. Sepals twice as long as petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Anangia
b. Sepals (as long as or) shorter than petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25a. Flowers axillary to large bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Baijiania
b. Bracts (minute or) absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26a. Flowers with short pedicels arranged in an elongate spike-like raceme. Petals
yellow. Filaments nodding at apex . . . . . . . . . . . . . . . . . . . . . . . . . 19. Kedrostis
b. Pedicels long or short; flowers in fascicles, or (sub)paniculate, or in pedunculate
short racemes. Petals white or yellow. Filaments straight . . . . . . . . . . . . . . . . . 27
27a. Anthers (thecae) plicate, contorted, or strongly hooked . . . . . . . . . . . . . . . . . . 28
b. Anthers (thecae) straight or ± curved, not plicate . . . . . . . . . . . . . . . . . . . . . . . 32
28a. Plant glabrous or scabrous. Filaments distinct; connective not produced . . . . . 29
b. Plant hairy. Leaf blade unlobed or lobed. Filaments short; connective produced
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
29a. Leaf blade unlobed. Flowers in peduncled raceme. — New Guinea . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. Papuasicyos
b. Leaf blade mostly deeply palmately lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Flowers fasciculate in leaf axils . . . . . . . . . . . . . . . . . . . . . . . . . 13. Diplocyclos
b. Flowers arranged in loose panicles. — East Java, naturalized . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Cayaponia (C. martiana)
31a. Thecae 2-plicate. Petals (corolla) yellow . . . . . . . . . . . . . . . . . . . . 10. Cucumis
b. Thecae hooked. Petals white . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. Muellerargia
32a. Plant usually glabrous. Leaf blade subcoriaceous, with short petiole . 33. Solena
b. Plant glabrous or hairy. Leaf blade membranous; petiole long (occasionally short
in Scopellaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33a. Plant coarsely rough-hairy. Flowers subsessile or short-pedicelled (pedicel 2–10

mm long), fascicled at the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25. Mukia
b. Plant glabrous or finely rough or soft-hairy. Flowers either long-pedicelled (pedi-
cel more than 10 mm long) or in a pedunculate raceme . . . . . . . . . . . . . . . . . . 34
34a. Flowers in a pedunculate raceme. Stamens 3, inserted in or near the throat of the
receptacle-tube; anthers two 2-thecous, one 1-thecous . . . . . . . . . . . . . . . . . . . 35
b. Flowers solitary or (few-)fascicled, directly inserted at the node, usually co-axil-
lary with a female flower, or in a (short) pedunculate raceme or fascicle. Stamens
3; all anthers 2-thecous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
35a. Petals (creamy-)white, apex obtuse . . . . . . . . . . . . . . . . . . . . 18. Indomelothria
b. Petals yellow, apex notched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. Melothria
36a. Flowers 1 or few, each directly inserted on the node. Stamens inserted near the
throat of the receptacle-tube; filaments short, as long as or shorter than the anther,
thecae straight. Probract absent . . . . . . . . . . . . . . . . . . . . . . 26. Neoachmandra
b. Flowers in a pedunculate raceme or fascicle. Filaments (usually) longer than
anthers, thecae straight or curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37a. Petals white, imbricate or valvate. Stamens inserted halfway or in the lower half
in the receptacle-tube, (largely) included. Probract (small, linear) present . . . . 38
b. Petals yellow, imbricate. Stamens inserted in the throat of receptacle-tube, (large-
ly) exserted. Probract absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
38a. Flowers mostly in a pedunculate raceme. Stamens mostly inserted at (or towards)
base of hypanthium-tube; filament longer than anther (shorter in Pilogyne viridi
folia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. Pilogyne
b. Flowers in a (sub)sessile fascicle. Stamens inserted about halfway the tube;
filament about as long as anther. — Norfolk Is., not in Malesia . . . . . Zehneria
39a. Pedicels crowded. Disc free. — SE Asia & West Malesia . . . . . 30. Scopellaria
b. Pedicels spaced. Disc urceolate, adnate to base of receptacle-tube. — New Guin-
ea . . . . . . . . . . . . . . . . . . . . . . . . 36. Urceodiscus p.p. (small-flowered species)
2 – GENERAL KEY FOR FRUITING AND FEMALE FLOWERING MATERIAL
1a. Flowers or fruit in panicles or racemes, however sometimes much reduced. Ovary
somewhat superior, at apex flat, broad, not constricted. Styles 3. Fruit becoming
dry, dehiscent with valves in truncate apex (indehiscent in Gynostemma, p.p.).
Tendrils 2-branched at apex. — Subfam. Fevilleoideae . . . . . . . . . . . . . . . . . . . 2
b. Flowers or fruit solitary or fascicled, or in racemes, or panicles. Ovary inferior,
at apex narrow, constricted. Style 1. Fruits usually berry-like (pepo-fruit), usually
indehiscent. Tendrils unbranched or branched at or below the middle (distally 2-
branched in Siraitia and Baijiania). — Subfam. Cucurbitoideae . . . . . . . . . . . . 7
2a. Climber to 30 m long. (Ovary half-globose, 5 –10 mm diam. in Alsomitra). Fruit
20 – 30 cm long. Seeds c. 2 cm diam., winged . . . . . . . . . . . . . . . . . . . . . . . . . . 3
b. Small or medium climber to 15 m long. Fruit much smaller. Seeds large (Zanonia)
or smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3a. Fruit half-globose mitriform, 20 – 30 cm diam. Seed entire, broadly butterfly-like
winged, c. 10 cm wide. Flowers creamy . . . . . . . . . . . . . . . . . . . . . 1. Alsomitra
b. Fruit claviform-cylindric, bluntly triangular, 8 –10 cm wide. Seed toothed, with

circular wing c. 5 cm wide. Flowers red (female flowers not known) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Bayabusua
4a. Ovary and fruit subglobose. Fruit 8 mm diam. or less, dehiscent or indehiscent.
Seeds few, not winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. Gynostemma
b. Ovary and fruit elongated. Fruit 1.5 cm long or longer, dehiscent. Seeds mostly
winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Sepals 3, petals 5. Fruit c. 8 cm long. Seeds large, 1.5 – 2 cm long, winged at both
ends, including the wings 4 – 8 cm long . . . . . . . . . . . . . . . . . . . . . . 37. Zanonia
b. Sepals and petals 5. Fruit c. 8 cm long or shorter. Seeds shorter . . . . . . . . . . . . 6
6a. Blade margin serrate. Fruit with 3 horn-like processes at apex. Seeds not winged
or winged all around (broadest at the ends). Pedicel often with 1 or 2 small
tendril(s). — East Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. Gomphogyne
b. Blade margin entire. Fruit without horn-like processes at apex. Seeds winged at
one side. Pedicel without tendril(s). — West & East Malesia . 27. Neoalsomitra
7a. Sepals much longer than petals. — Moluccas: Morotai Is . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Anangia (fruit not known)
b. Sepals shorter than petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Fruit (or ovary) ornamented with ridges, warts, or (few) (soft) spines, (irregular)
grooves, or fruit fenestrate (fruit of Momordica clarkeana and M. denticulata
almost smooth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
b. Fruit not ornamented (colour or fine texture excepted), surface glabrous or hairy
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
9a. Fruit ± curved. — Introduced from America, cultivated or locally naturalized . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Cyclanthera
b. Fruit not curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Tendrils 3- or more-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
b. Tendrils unbranched or 2-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
11a. Fruit longitudinally ridged . . . . . . . . . . . . . . . . . . . . . 21. Luffa (L. acutangula)
b. Fruit smooth, or ± bullate, or grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Fruit pear-shaped, 10 –15 cm long. Seed one . . . . . . . . . . . . . . . . . 31. Sechium
b. Fruit globose, ellipsoid or pyriform, small or large. Seeds numerous . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. Cucurbita
13a. Fruit (globose or) broadly ellipsoid, c. 10 cm diam. or more, warted (tubercled)
or spiny all over. Seeds c. 15 mm long or more, flat, margin cogwheel-like undu-
late . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. Momordica
b. Fruit (narrowly) ellipsoid or oblong, usually < 10 cm diam. Seeds generally small-
er . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a. Tendrils unbranched. Fruit echinate or aculeate-tubercled, or smooth, green or
yellow. Seeds oblong, not ornamented, edge smooth . . . . . . . . . . . . . . . . . . . . 15
b. Tendrils unbranched or 2-branched. Fruit tubercled, ribbed or fenestrate, ± beaked.
Seeds ornamented or with grooved margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15a. Fruit not squirting. Flowers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Cucumis
b. Fruit squirting. Flowers white. [Fruit occasionally smooth, not ornamented.] . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. Muellerargia
16a. Fruit green or yellow, fenestrate, hardly beaked . . . . . . . . . . . . 34. Thladiantha

b. Fruit orange or red, beaked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a. Tendrils (unbranched or) 2-branched. Fruit low-ribbed. Seeds ± oblong, margin
entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Gymnopetalum
b. Tendrils unbranched. Fruit ribbed or (few-)tubercled. Seeds ellipsoid-globose,
edge undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. Momordica
18a. Plant cultivated (but frequently also found running wild) . . . . . . . . . . . . . . . . . 19
b. Plant wild (some wild species occasionally cultivated; Cayaponia naturalized in
Java; Melothria widely naturalized) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
19a. Ripe dry fruit opening with apical operculum . . . . . . . . . . . . . . . . . . . 21. Luffa
b. Fruit without apical operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20a. Fruit frequently ± flask-shaped constricted. Petiole with 2 glands at apex . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20. Lagenaria
b. Fruit (depressed) globose, ellipsoid, or oblong. Petiole without glands at apex . 21
21a. Leaf blade more or less pinnately (veined and) lobed. Seeds brown or black . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. Citrullus
b. Leaf blade palmately (pedately) veined, unlobed or lobed. Seeds pale brown or
whitish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22a. Tendrils unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Cucumis
b. Tendrils 2 – 5-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23a. Probract distinct, hooded. Seeds c. 10 mm long, almost unmargined 5. Benincasa
b. Probract absent or not obvious. Seeds usually > 10 mm long, margined . . . . . 24
24a. Fruit globose, c. 6 cm diam. Seeds subcircular, margin with double wing. — Sirai
tia grosvenorii (Swingle) A.M.Lu & Zhi Y.Zhang, cultivated in China, and sold
in Malesia for its sweet, medicinal fruit . . . . . . . . . . . . . . . . . . . . . . 32. Siraitia
b. Fruit larger, globose, ellipsoid or cylindrical. Seeds elliptic in outline, not winged
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25a. Fruit globose, short-ellipsoid or pyriform. Seed edge not undulate . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. Cucurbita
b. Fruit long-cylindrical. Seed edge coarsely undulate . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . 35. Trichosanthes (T. cucumerina var. anguina)
26a. Fruit large, globose, with 6 large seeds 5 – 8(–10) cm long. Probract sharp, thorn-
like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. Hodgsonia
b. Fruit various, smaller, less than 10 cm long, seeds more than 6 (1 or 2 in Kedros
tis). Probract not sharp at apex, or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27a. Seeds tumid (barrel-shaped or (sub)fusiform), or pyriform (see Fig. 19b, 91b) . 28
b. Seeds either subglobose, or elongate, little or much compressed (ornamented or
not) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
28a. Fruit c. 2 cm diam., green, later red and white-blotched or -striped, (sub)sessile .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. Diplocyclos
b. Fruit > 2 cm diam., pedicelled . . . . . . . . . . . . . . . . . . . . . . . . 35. Trichosanthes
29a. Fruit 5 cm long or more, seeds 1 cm long or more. Flowers large, petals white,
long-fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. Trichosanthes
b. Fruit shorter or longer than 5 cm, seeds various. Flowers small or large; petals
yellow or white, not long-fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Tendrils several-branched (2- or more-branched). Probract conspicuous. Petals

large, longer than 2 cm, yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
b. Tendrils unbranched or 2-branched. Probract small or absent. Petals yellow or
greenish, smaller, less than 1.5 cm long, or petals white . . . . . . . . . . . . . . . . . 32
31a. Fruit brown, operculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. Luffa
b. Fruit green or usually with whitish bloom; not operculate . . . . . . . 5. Benincasa
32a. Flowers large, white; petals free, c. 20 mm long. Tendrils unbranched or unequally
2-branched near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Gymnopetalum
b. Flowers medium or small, white (whitish) or yellow; petals either (almost) free,
15 mm long or less, or petals largely fused and flowers large. Tendrils unbranched
or 2-branched towards the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33a. Flowers large, white, corolla c. 25 mm long, petals fused for c. 2/3. Fruit glabrous,
pulp or juice red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Coccinia
b. Flowers smaller, white or yellow, corolla c. 20 mm long or less, petals free or
short-connate at base only. Fruit glabrous or hairy, pulp or juice white or yellow
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34a. Fruit globose, c. 5 cm diam. Seeds with 2- or 3-winged margin. Leaf blade with
minute blackish appressed glandular hairs . . . . . . . . . . . . . . . . . . . . 32. Siraitia
b. Fruit globose, or ellipsoid or fusiform, small or large. Seeds not winged. Blackish
glandular hairs absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35a. Flowers medium, petals c. 10 mm long, yellow . . . . . . . . . . . . . . . . . . . . . . . . 36
b. Flowers small, petals c. 5 mm long or less, white, greenish or yellow . . . . . . . 39
36a. Female flowers or fruit solitary, in non-pedunculate raceme at the end of shoots,
or flowers 4 –7 in a short raceme. Fruit c. 5 cm long, smooth; seeds few or several,
large, c. 9 mm long, subglobose, smooth. — Borneo . . . . . . . . . 6. Borneosicyos
b. Female flowers or fruit on the leaf-bearing node. Fruits various; seeds numerous,
smaller, ornamented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37a Fruit hairy, glabrescent (see Fl. China). — Philippines . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. Thladiantha (T. nudiflora)
b. Fruit glabrous. — New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38a. Stigma 3 times parted, finely feather-like. Disc not obvious (absent) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. Papuasicyos
b. Stigma 3-lobed (papillose). Disc urceolate, largely adnate to the base of the re-
ceptacle-tube . . . . . . . . . . . . . . . . 36. Urceodiscus p.p. (larger-flowered species)
39a. Fruit globose, 3 – 4.5 cm across, hairy. — Borneo . . . . . . . . . . . . . . 3. Baijiania
b. Fruit globose but much smaller, c. 1 cm diam., or fruit ellipsoid or ovoid . . . . 40
40a. Seeds nearly globose, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
b. Seeds elongate, somewhat or much compressed. Petiole mostly long . . . . . . . 42
41a. Petiole short. Flowers creamy-white. Fruits 2–5 cm long. Seeds 5–20 33. Solena
b. Petiole long, usually at least as long as half the length of the blade. Flowers yel-
low. Fruit 1–1.5 cm long. Seeds 1 or 2 . . . . . . . . . . . . . . . . . . . . . 19. Kedrostis
42a. Fruit broad-ellipsoid, 2 – 5 cm long. Seeds smooth [Thecae plicate.] . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Cucumis [feral forms]
b. Fruit various, when more than 5 cm long then elongate-fusiform. Seeds smooth
or scrobiculate (pitted or warted). [Thecae plicate or not plicate.] . . . . . . . . . . 43
43a. Tendrils 2-branched. [Thecae plicate.] — East Java, naturalized . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Cayaponia (C. martiana)
b. Tendrils unbranched [Thecae not plicate.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
44a. Fruit (sub)sessile, (few-)fascicled; fruiting pedicel to 10 mm long. Corolla yellow.
(Male) flowers short pedicelled in (sub)sessile fascicles or clusters . . 25. Mukia
b. Fruit with short or longer pedicel. Corolla white or yellow. (Male) flowers long-
pedicelled or when shorter pedicelled then arranged in (small) pedunculate ra-
cemes or fascicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
45a. Seeds thickish, margined, faces pitted or warted. Corolla yellow . . . . . . . . . . . 46
b. Seeds compressed (or globose), faces smooth, glabrous or hairy . . . . . . . . . . . 47
46a. Fruit c. 1 cm diam. — West Malesia . . . . . . . . . . . . . . . . . . . . . . 30. Scopellaria
b. Fruit c. 2 cm diam. — New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . 36. Urceodiscus, p.p. (smaller-flowered species)
47a. Corolla yellow, petals ± notched at apex. Fruits purple-black; seeds hairy . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. Melothria pendula (naturalized)
b. Corolla white, petals not notched. Fruit green, red or purple; seeds smooth or
hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
48a. Seeds narrowly margined. Probract (small, linear) present. Fruiting pedicel short,
usually shorter than the fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
b. Seeds unmargined. Probract absent. Fruiting pedicel short or long . . . . . . . . . 50
49a. Stigma 3-parted, parts sessile or short-armed. Disc ring-shaped. Fruits solitary or
(few-)fascicled, small, to 1.5 cm long . . . . . . . . . . . . . . . . . . . . . . . 29. Pilogyne
b. Stigma parts long-armed, each 2-lobed. Disc 3-parted. Fruit 1.2 – 2 cm long. —
Norfolk Is. (not in Malesia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zehneria
50a. Seeds glabrous, sometimes short-winged at one side. Female flowers or male
flowers one or few, long-pedicelled, inserted at the node. Fruit globose, or el-
lipsoid, or fusiform, green or red . . . . . . . . . . . . . . . . . . . . . 26. Neoachmandra
b. Seeds hairy, not winged. Female flowers with long pedicel solitary on the node,
co-axillary with male inflorescence. Male flowers in a pedunculate raceme. Fruit
(long) fusiform, green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. Indomelothria
1. ALSOMITRA
Alsomitra (Blume) Spach, Hist. Nat. Vég. 6 (1838) 187; M.Roem., Fam. Nat. Syn. Monogr. 2 (1846)
117; Hutch., Ann. Bot. (Oxford) 6 (1942) 96, p.p. excl. A. clarkei; de Wit, Bull. Jard. Bot. Buiten-
zorg III, 18 (1949) 193. — Zanonia L. sect. Alsomitra Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 937.
— Zanonia L., p.p. (incl. Zanoniae sect. Alsomitra Blume; excl. Alsomitra auct. non (Blume)
Spach: Benth. & Hook.f., Gen. Pl. 1 (1867) 840 = Neoalsomitra Hutch.) Benth. & Hook.f., Gen.
Pl. 1 (1867) 839. — Zanonia L. sect. Macrozanonia Cogn. in A.DC. & C.DC., Monogr. Phan. 3
(1881) 927; Pax in Engler & Prantl, Nat. Pflanzenfam. 4 (5) (1889) 12. — Macrozanonia (Cogn.)
Cogn., Bull. Herb. Boiss. 1 (1893) 612; Pax in Engl. & Prantl, Nat. Pflanzenfam. 4 (5) (1897) 392;
Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 262. — Type species: Alsomitra macrocarpa (Blume)
M.Roem. (Zanonia macrocarpa Blume).
Tall climber, perennial; dioecious; young plant with slender stem creeping up, with
distichous leaves appressed to the substrate. Probract absent. Tendrils somewhat off-
axillary, 2-branched at apex; peltate adhesive discs present in juvenile plants, with or
without elongated pads in adult plants. Leaves: blade simple, unlobed, glands absent,
margin entire. Male inflorescences ± pendent, paniculate, in female inflorescences
reduced, raceme-like; bracts minute, caducous. Male flowers: bud ovoid, acute; flower
± campanulate, fleshy; receptacle-tube cup-shaped; calyx at first fused, apically with
narrow, minutely imbricately 5-lobed orifice, at anthesis tearing into 3 parts to about
half-way; petals almost free, ± patent, narrow, acute, valvate, cream-coloured; stamens
3, inserted towards the rim of the receptacle-tube, filaments short, free, anthers basi-
dorsifixed, either all 2-thecous or two 2-thecous and one 1-thecous, thecae oblong,
opening lengthwise lateral-introrse, connective broad; disc obscure. Female flowers
larger than male flowers; ovary not completely inferior, at apex ± convex, pseudo 3-loc-
ular; ovules numerous, pendent, inserted towards the apex on broad, fleshy placentas;
styles 3, not close together, stigmas fleshy; calyx and corolla together caducous, the scar
leaving a line on the ovary and fruit. Fruit solitary, pendent, capsular, unilocular, half-
globose, 20 – 30 cm diam., with flattened 3-valved apex. Seeds numerous, compressed,
smooth, ovate-elliptic, edge entire, with a butterfly-like wing extending laterally to both
sides 10 –12 cm wide.
Distribution — A genus with 1 species in southern Peninsula Thailand; in Malesia:
Peninsular Malaysia east to New Guinea.
1. Alsomitra macrocarpa (Blume) M.Roem.

Alsomitra macrocarpa (Blume) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 117; Hutch., Ann. Bot.
(Oxford) 6 (1942) 96; de Wit, Bull. Jard. Bot. Buitenzorg. III, 18 (1949) 193; Backer in Backer &
Bakh.f., Fl. Java 1 (1964) 299; W.J.de Wilde & Duyfjes, Proc. Fourth Int. Fl. Males. Symp. 1998
(2001) 103; Fl. Thailand 9, 4 (2008) 416. — Zanonia macrocarpa Blume, Bijdr. Fl. Ned. Ind. 15
(1826) 937; Ser. in DC., Prodr. 3 (1828) 299; Miq., Fl. Ned. Ind. 1, 1 (1856) 683; Cogn. in A.DC.
& C.DC., Monogr. Phan. 3 (1881) 927; Warb., Bot. Jahrb. Syst. 13 (1891) 444; Gibbs, Dutch NW
New Guinea (1917) 17, 51, 222. — Macrozanonia macrocarpa (Blume) Cogn., Bull. Herb. Boiss.
1 (1893) 612; K.Schum. & Lauterb., Fl. Schutzgeb. Südsee (1901) 589; Cogn. in Engl., Pflanzenr.
66, 4.275.I (1916) 262, f. 63, 64; Rolfe, Kew Bull. (1920) 197. — Type: Blume ‘1353’ (holo L,
barcode L0001604; iso L, barcodes L0001603 & L0001605), Java.
Momordica coriacea Cogn. in K.Schum. & Hollrung, Fl. Kais. Wilh. Land (1892) 82; Bot. Jahrb. Syst.
58 (1923) 240; in Engl., Pflanzenr. 66, 4.275.1 (1916) 264. — Type: Hollrung 775 (iso K), Papua
New Guinea, Kaiser Wilhelmsland.
Zanonia philippinensis Merr., Philipp. J. Sci. 1, suppl. 1 (1906) 241. — Macrozanonia philippinensis
(Merr.) Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 264. — Type: Clemens 324 (holo PNH, not
seen), Philippines.
Erythropalum triandrum Merr. & Quisumb., Philipp. J. Sci. 37 (1928) 143. — Type: Ramos & Edaño
BS 49096 (holo PNH, not seen; iso K), Philippines, Mindanao, Mati.
Climber, 30 – 50 m long, stem at base 15 cm thick. Shoots grooved, 3 – 4 mm diam.,

early glabrescent, at first with minute grey scurf; older twigs with raised leaf-scars.
Tendrils 6 –10 cm long, branches c. 2 cm long, adhesive pads 5 –10 mm long. Leaves:
petiole 2 – 4 cm by c. 2 mm; blade ovate(-elliptic), 8 –16 by 3 –13 cm, thinly coriaceous,
glabrous, base rounded or shallowly broadly cordate or sometimes narrowed and ± ob-
tuse, apex (obtuse or) acute-acuminate; 3(– 5) palmately veined, venation ± fine, promi-
nent on both surfaces. Lateral shoots and inflorescences, arising from a densely pale
brown pubescent area. Male inflorescences in short leafy lateral shoots, many-flowered,
2 mm e
2 cm
d
2 mm
2 mm
c
Fig. 1. Alsomitra macrocarpa (Blume) M.Roem. a. Twig with male inflorescences; b. detail of male
inflorescence, note finely hairy branches; c. male flower; d. ditto, opened, note disc-appendages at base;
e. stamens (a, b: Hoogland 4918 (Papua New Guinea); c – e: De Wilde & Duyfjes 21945 (West Java)).
d a
i'
2 mm
2 cm
2 cm
2 cm
i
g
2 cm e'
2 mm
3 mm
b
c
1 mm
3 cm
1 mm
c' f e
Fig. 2. Alsomitra macrocarpa (Blume) M.Roem. a. Branch with side branch with female inflorescence;
b. female flower just before anthesis; c. ditto opened; c'. detail, note one style with 2-lobed stigma
and one of the three minute staminodes; d. portion of twig with just developing fruit, note the typical
design of apex, showing the three valve seams and the 2-branched tendril; e. placenta with young seeds;
e'. detail; f. fruit, schematic; g. seed with butterfly-like wing; h. seedling; i. juvenile leafs; i'. detail
(a: De Wilde & Duyfjes 21864; b – c': Annon., cult. Hort Bot. Bog. sub. XII-A-1X 12; d – e': Kostermans
21278; f: De Wilde & Duyfjes 21814; g: De Wilde & Duyfjes 21792; h: De Wilde & Duyfjes 21978;
i, i': Korthals 2).
1 or 2 times branched, 5 –15 cm long, peduncle (1–)2 cm long, sometimes with 1 or 2

additional smaller inflorescences from the same leaf axil; bracts c. 0.5 mm long; female
inflorescences not or hardly branched, raceme-like, 5(–10)-flowered, 5 –10 cm long.
Male flowers: pedicel c. 2 mm long; perianth greenish creamy, 7–10 mm long; recep-
tacle-tube c. 2 by c. 3 mm; calyx membranous, in bud ovoid-conical, c. 5 mm long;
petals narrowly elliptic, 6 – 8 mm long, acute, papillose at apex; filaments 2 – 2.5 mm
long, anthers ellipsoid-oblong, 3 – 4 mm long, thecae papillose hairy; 3 minute subulate
appendages c. 1 mm long in the centre of the flower sometimes present. Female flowers:
pedicel c. 20 cm long; perianth including ovary 15 – 20 mm long; ovary c. 3/4 inferior,
half-globose-ellipsoid or ± conical, 5 –10 mm wide; calyx c. 5 mm long; petals c. 9 mm
long; styles not close together, short, stigmas elongated, 3 – 4 mm long, at apex deeply
2-lobed. Fruit green, ripening brown, opening with 3 incurving valves, 20 – 30 cm
diam., with woody exocarp, c. 2 mm thick, smooth, glabrous, inside with a soft spongy
layer, densely packed with winged seeds, line of perianth-scar 2 – 6 cm from the orifice;
fruiting pedicel 2 – 4 cm long. Seeds pale brown, 25 – 30 by 20 – 23 mm, faces smooth,
margin distinct, edge entire, wing butterfly-like, silvery-white, membranous, 10 –12 cm
wide. — Fig. 1, 2, 8e, h–j, 97g; Plate 2c.
Distribution — Southern Thailand; in Malesia: Sumatra, Peninsular Malaysia, Bor-
neo, West Java, Philippines, Sulawesi, Moluccas (Aru Islands), New Guinea (West
Papua, Papua New Guinea including Bismarck Archipelago).
Habitat & Ecology — Tall forest liana preferring rich soils, locally not rare on clayey
soils along rivers; 0 – 800 m; flowering in West Java in the rainy season (December –
January); fruiting, presumably set during the foregoing season, at the end of the rains
or later, March – April(– June).
Note — The juvenile plant is extraordinary. After germination the primary shoot
develops as a thread-thin stem with rather spaced small oblong leaves, c. 1 cm long,
creeping over the forest soil, and when a suitable tree trunk is reached it climbs up ver-
tically by means of small, peltate adhesive discs at the end of the small bifid tendrils.
By then the plant clings close to the substrate, with the leaves distichous, appressed,
and closely set. While climbing, the leaves which are hastate and in an early phase
also auriculate, gradually grow larger (and the stem only moderately thicker), until at a
height of about 5 m the leaves are c. 5 cm long and start approaching the adult shape.
2. ANANGIA
Anangia W.J.de Wilde & Duyfjes, Reinwardtia 12 (2006) 219. — Type species: Anangia macrosepala
W.J.de Wilde & Duyfjes.
Low suberect herb, tuberous(?); monoecious. Probract present in association with

flowers. Tendrils 2-branched from about the middle. Leaves: blade simple, not or shal-
lowly lobed. Flowers solitary (or 2), erect, long-pedicelled; perianth large, sepals much
longer than petals; petals free; disc absent; receptacle-tube small, shallow. Male flowers:
stamens 5, free, inserted at base of receptacle, erect, four in two pairs and one solitary,
anthers all 1-thecous, thecae sinuate. Female flowers: ovary elongate, ovules numerous,
horizontal, style 1, columnar, 3-armed, each with bifurcate stigma.
Distribution — 1 species in the Moluccas.
b
3 mm
3 mm
3 mm
d
c
3 mm
Fig. 3. a – c. Anangia macrosepala W.J.de Wilde & Duyfjes. a. Portion of sterile branch; b. lower node
showing 2-branched tendril; c. node with female flower. — d. Neoachmandra backeri W.J.de Wilde &
Duyfjes subsp. backeri. Node with male flower (a, b: Anang 325, type; c, d: Buwalda 4057).
1. Anangia macrosepala W.J.de Wilde & Duyfjes

Anangia macrosepala W.J.de Wilde & Duyfjes, Reinwardtia 12 (2006) 220, f. 1, 2. — Type: Anang
325 (holo BO; iso L), Moratai.
Herb, several-stemmed, c. 30 cm high, subglabrous; stems suberect, much branched,

sparsely minutely hairy, (1–)2 mm diameter. Probract at base of pedicels, oblong-
spathulate, 1(–1.5) cm long. Tendrils only seen on lower nodes. Leaves: petiole 1– 3
cm long; blade membranous, elliptic-rhomboid or subtriangular, 1.5 – 2 by 1.5 – 2.5 cm,
both surfaces minutely harshly hairy, cystoliths minute, inconspicuous; base truncate
or ± cuneate, narrowly decurrent on the petiole (and hence the basal veins branching
off from above the base of the blade), margin sparsely minutely dentate, apex acute.
Flowers subglabrous; mature buds not known, colour not known. Male flowers: pedicel
35 – 40 mm long, at apex not articulate; receptacle-tube c. 1.5 by 3 mm, papillose inside;
sepals (narrowly) elliptic, 10 –12 by 5 – 6 mm, base ± narrowed and ± clawed, apex acute,
glabrous but margin finely hairy (hairs c. 0.1 mm long), inner surface sparsely papillose;
a c
2 mm
Fig. 4. Anangia macrosepala W.J.

de Wilde & Duyfjes. a. Male flow-
er, showing large sepals and small
petals; b. male flower opened,
showing irregularly sinuate thecae;
c. female flower, showing large
sepals and small petals; d. female
flower opened (all: Anang 325,
b d type).
petals free, shape similar to sepals, 5 – 6 by 2.5 – 3 mm, glabrous, the main veins ending
in the margin with a minute dot; filaments c. 1.5 mm long, glabrous, anthers irregularly
long-ellipsoid, 1.5 – 2 mm long, thecae irregularly sinuate, at one side along the margin
of a broad and thin glabrous ‘connective’ (see Fig. 4b). Female flowers: pedicel 25 – 30
mm long, indistinctly articulate at apex; ovary oblong-linear, subfusiform, broadest
above the middle, 15 –17 by 2.5 – 3 mm, with 0.2 mm long hairs; perianth as in male
flowers but smaller; receptacle-tube c. 1 by 3 mm; sepals c. 8 by 4 mm; petals (estimated
because of incomplete material) c. 2.5 by 1.5 mm; style slender, c. 4 mm long, glabrous,
at apex 3-armed, each arm 2 – 2.5 mm long ending in a deeply forked stigma c. 1 mm
long, finely papillose. Fruits and seeds not known. — Fig. 3a – c, 4.
Distribution — Moluccas (Morotai, possibly also Yamdena), known only from the
type.
Habitat & Ecology — Only the collecting date of the flowering plant is known:
March 1938. Possibly it grows in grassy savanna, yearly burnt.
3. BAIJIANIA
Baijiania A.M.Lu & J.Q.Li in J.Q.Li, Acta Phytotax. Sin. 31 (1993) 50, p.p. for the type only, excluding
material from China; W.J.de Wilde & Duyfjes, Blumea 48 (2003) 279. — Type species: Baijiania
borneensis (Merr.) A.M.Lu & J.Q.Li (Thladiantha borneensis Merr.).
Thladiantha Bunge subg. Microlagenaria C.Jeffrey, Kew Bull. 15 (1962) 363, p.p.
Siraitia Merr. subg. Microlagenaria (C.Jeffrey) A.M.Lu & Zhi Y.Zhang, Guihaia 4 (1984) 30, p.p., for
the specimens from Borneo only.
Plants with tuberous rhizome(?); dioecious; hairs not glandular. Probract absent.
Tendrils coiling both below and above point of branching. Leaves: blade simple, un-
lobed. Male inflorescences racemose (sometimes ± branched), bracteate. Male flowers:
receptacle-tube cup-shaped, about as long as wide or longer than wide; petals free,
longer than sepals, creamy or greenish white; stamens 5 (3), inserted below the throat
of the receptacle, two pairs with filaments connate at base, one stamen solitary, thecae
somewhat curved, glabrous, filaments short; basal adaxial scales absent; disc at base of
receptacle conspicuous, 3-parted. Female flowers in a few-flowered raceme; staminodes
present. Fruit globose, c. 4 cm diam., pulp not sweet tasting. Seeds of medium size,
± compressed, smooth, margin faint.
Distribution — 1 species in Borneo with two varieties.
1. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li

Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li in J.Q.Li, Acta Phytotax. Sin. 31 (1993) 50, for the type
only; Beaman et al., Pl. Mt. Kinabalu 4 (2001) 206, pl. 15 b, c; W.J.de Wilde & Duyfjes, Blumea
48 (2003) 280. — Thladiantha borneensis Merr., Univ. Calif. Publ. Bot. 15 (1929) 298. — Siraitia
borneensis (Merr.) A.M.Lu & Zhi Y.Zhang, Guihaia 4 (1984) 31, p.p., specimens from Borneo only;
Zhi Y.Zhang & A.M.Lu, Cathaya 1 (1989) 26, f. 26, 27. — Type: Villamil 308 (holo UC, not seen;
iso BO, K, PNH lost?), Sabah.
Climber, 3 – 6 m, sparsely grey soft hairy and minutely papillose on young parts,
hairs c. 1 mm long; leafy stem 2.5 – 4 mm diameter. Tendrils 10–25 cm long. Leaves:
petiole 3 – 5 cm long; blade (narrowly) ovate, 10 – 20(– 30) by 5 –14(–17) cm, greenish
on drying, above with cystoliths, margin entire or faintly lobulate-dentate near the base;
(3 –)5-palmiveined and 2 or 3 pairs of veins from the midvein. Male inflorescences 8 –15
cm long, peduncle 4 – 8 cm long, flower-bearing part 4 –10 cm long, bracts persistent,
(narrowly) elliptic or spathulate, 3 –10(–15) mm long, rounded, entire, conspicuously
veined. Male flowers: pedicel 5 –15(– 30) mm long, faintly articulate 3 –10 mm be-
low perianth, the portion below articulation persistent; open flower 5 – 8 mm diam.;
receptacle-tube cup-shaped, 2.5 – 4.5 by 2 – 4 mm; sepals narrowly triangular-linear,
1.5 – 3 mm long, patent or recurved; petals imbricate, elliptic, rounded, 2 – 5 mm long,
3(– 5)-veined, papillose; stamens inserted 0.5(–1) mm below the rim of the receptacle,
filaments 0.5 –1 mm long, anthers (thecae) 2 – 3 mm long, half included or exserted;
disc central, consisting of 3 contiguous subconical parts, 1–1.5 mm high. Female inflo
rescences (1– 3)-flowered, peduncle 2 – 8 cm long; bracts as in male flowers, persistent.
Female flowers as male flowers; pedicel 5 –15(– 30) mm long; ovary ellipsoid, c. 5 mm
long, passing with short neck (c. 1 mm long) into shallow receptacle-tube c. 1 mm long;
style conspicuous, c. 2 mm long, stigma with 3 thick notched lobes; staminodes small,
0.5 –1 mm long, in 2 pairs and one solitary. Fruits solitary or 2 or 3 together (peduncle
2.5 –10 cm long), ripening orange, globose, 3 – 4.5 cm diam., grey-white soft hairy, hairs
1– 2 mm long; pericarp thin, pergamentaceous; endocarp pulpy-juicy; fruiting pedicel
0.5 –1.5 cm long, soft hairy. Seeds numerous, pale, (broadly) ovate, c. 5 mm across,
base truncate, apex rounded, edge entire.
KEY TO THE VARIETIES
1a. Male inflorescences simple, racemose or ± branched, flowers lax; pedicels (7–)
20(–30) mm long. Male perianth 8 –10 mm diam.; receptacle-tube about as long as
wide, c. 4 mm long; anthers (thecae) c. 3 mm long. — Dry land forest . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. var. borneensis
b. Male inflorescences a narrow raceme, densely set with flowers; pedicels 3 – 8 mm
long. Male perianth 5 – 6 mm diam.; receptacle-tube longer than wide, c. 3 mm long;
anthers (thecae) 1.5 – 2 mm long. — Peat swamp forest . . . . . . b. var. paludicola
a. var. borneensis
Male inflorescences grey, yellow or pale brown soft hairy, simple or ± (1- or) 2-
branched, each branch 10 – 20-flowered; bracts lanceolate or obovate-spathulate, 7–15
mm long; pedicels (7–)20(– 30) mm long, longer than bracts. Male perianth 8 –10 mm
diam., receptacle c. 4 by 4 mm, sepals 2.5 – 3 mm long, petals 5 – 6 by 3.5 mm; filaments
c. 1.5 mm long, anthers (thecae) c. 3 mm long. — Fig. 5a, b, 6a – h; Plate 2a, b.
Distribution — Borneo (Sabah and SE Kalimantan).
Habitat & Ecology — Forest edges, scrub in forest gaps, in (open or) shaded places;
90 –1000 m altitude; flowering and fruiting throughout the year.
Note — Female flowers and fruit are only known from specimens which belong to
var. borneensis.
2 mm
2 cm
d
a
2 mm
c
b
Fig. 5. a, b. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. borneensis. a. Twig with male inflo-
rescences; b. detail of male inflorescence. — c, d. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var.
paludicola Duyfjes. c. Twig with male inflorescence; d. male inflorescence (a, b: De Wilde & Duyfjes
SAN 139471; c, d: Kadir A2520).
1 cm
1 cm
1 mm
e
e'
2 mm
2 mm h b''
d
j'' j j'
1 mm
b' 1 mm b
1 mm 2 mm
i i' a' a
Fig. 6. a – h. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. borneensis. a. male flower; a'. ditto,
opened, showing stamens; b, b', b''. stamens: adaxial, abaxial and lateral view respectively; c. twig with
female inflorescence; d. detail of female inflorescence; e. female flower; e'. ditto, opened, showing
style with 3-lobed stigma and staminodes; f. petal with basal staminodes; g. twig with infructescence;
h. seed. — i – j''. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. paludicola Duyfjes. i. Male flower;
i'. ditto, opened, showing stamens; j, j', j''. stamens: adaxial, abaxial and lateral view respectively (a – b'':
De Wilde & Duyfjes SAN 141929; c – h: De Wilde & Duyfjes SAN 141932; i – j'': Kadir A2520).
b. var. paludicola Duyfjes

Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. paludicola W.J.de Wilde & Duyfjes, Blumea 48
(2003) 283. — Type: Kadir BNB A2520 (holo K; iso BO, KEP), Sabah, Sandakan.
Male inflorescences light brown pubescent; raceme slender, unbranched, many (10 –
40)-flowered; bracts lanceolate, 3 – 6 mm long; pedicels 3 – 8 mm long, hardly longer
than bracts. Male perianth 5 – 6 mm diam.; receptacle-tube c. 3 by 2 mm, sepals c. 2 mm
long, petals 3 – 4 by c. 3 mm; filaments (0.5 –)1 mm long, anthers (thecae) 1.5 – 2 mm
long. — Fig. 5c, d, 6i – j''.
Distribution — Sabah, known from 2 collections (Sandakan and Tawau).
Habitat & Ecology — Climber in peat swamp forest; lowlands.
Note — The inflorescences of Elmer 20472 is more lax than in the type, slightly
approaching those of var. borneensis, but its flowers are similar to those in the type,
Kadir BNB A2520.
4. BAYABUSUA
Bayabusua W.J.de Wilde, Sandakania 13 (1999) 1. — Type species: Bayabusua clarkei (King) W.J.de
Wilde (Zanonia clarkei King).
Tall liana, perennial; dioecious; young plant sprawling, with at base first foliage
leaves in one tier of 3 or 4. Probract not obvious. Tendrils slightly supra-axillary,
not distinctly off-axillary, 2-branched above the middle, spiralling below the point of
branching, with adhesive pads. Leaves: petiole scar raised; blade simple, margin entire.
Flowers: purple-red. petals nearly free. Male inflorescences erect, raceme-like panicles,
bracteate, with axillary clusters of few flowers. Male flowers rotate, receptacle shal-
low, sepals quincuncial, corolla lobes nearly free, contorted, revert at apex; stamens 3,
inserted near the centre, anthers 5, 1-thecous, basifixed, two filaments with 2 anthers
and one with 1 anther; thecae narrowly ellipsoid, opening lengthwise, with swollen
creamy-white connective with a brown wart near the apex; disc not obvious. Female
inflorescences small, 1- (or few)-flowered; female flowers not known. Fruit solitary,
cylindrical-clavate, capsular, c. 23 by 10 cm, apex flattened, opening with 3 valves
curving towards the inside. Seeds numerous, compressed, with 8 or 9 coarse marginal
spines, and a membranous circular wing, c. 5 cm diameter.
Distribution — 1 species in Peninsular Malaysia.
1. Bayabusua clarkei (King) W.J.de Wilde

Bayabusua clarkei (King) W.J.de Wilde, Sandakania 13 (1999) 2, f. 1, 2a–d. — Zanonia clarkei King,
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 67 (1898) 41; Ridl., Fl. Malay Penin. 1 (1922) 852. — Mac
rozanonia clarkei (King) Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 264. — Alsomitra clarkei
(King) Hutch., Ann. Bot. (Oxford) 6 (1942) 96. — Type: King’s Collector 7230 (holo CAL, not
seen; iso K), Peninsular Malaysia, Perak.
Tall climber, hanging from lofty trees, 20 – 30 m long; shoots slender, 1.5 – 4 mm
diam., early glabrescent, at first with minute red brown powdery hairs less than 0.1
mm long; older twigs with raised leaf-scars. Tendrils 10 –15 cm long, adhesive pads
1 cm
2 mm
1 mm
a
1 mm
1 cm f
Fig. 7. Bayabusua clarkei (King) W.J.de Wilde. a. Male flowering twig; b. upper portion of inflo-
rescence; c, d. male flowers seen from above and below; e. male flower, longitudinally dissected;
f. androecium showing two 2-thecous stamens and one 1-thecous stamen; g. detail of stamens (all: De
Wilde & Duyfjes FRI 41901).
d
2 cm
2 cm
a
1 cm
e
2 cm
5 mm 1 cm 3 cm
f g b
Fig. 8. Seeds, seedlings and fruits. — a – d. Bayabusua clarkei (King) W.J.de Wilde. a. Seedling, note
cotyledons; b. fruit; c. portion of tendril of older plant with adhesive pads; d. seed with circular wing. —
e. Alsomitra macrocarpa (Blume) M.Roem. Seed with butterfly-shaped wing. — f. Neoalsomitra clav
igera (Wall.) Hutch. Seed with wing. — g. Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes
var. pubescens Cogn. Seed with 2-sided wing (a, c: De Wilde & Duyfjes 21961; b, d: De Wilde & Duyfjes
21781; e: De Wilde & Duyfjes 21792; f: Kerenga et al. LAE 73815: g: De Wilde & Duyfjes 21855).
2 – 6 mm diameter. Leaves: petiole 0.5 – 3 cm long, 1– 2 mm wide; blade faintly laterally

lobed towards the base, membranous or slightly succulent, broadly ovate, 3 –15 by 2 –12
cm, membranous and brittle on drying, glabrous, base (broadly) rounded or shallowly
cordate with broad sinus, margin (sub)entire; 5(–7) palmately veined, venation lax.
Male inflorescences spike-like, indeterminate panicles, bracteate; peduncle 0.5 –1 cm
long, up-curved, rachis 5 – 20 cm long, c. 2 mm thick, purple-red; bracts, subcaducous,
5 –15 mm long, each with axillary a cluster of 2 – 5 flowers. Female inflorescences
(according to infructescences) small, less than 5 mm long, 1 (or few)-flowered. Male
flowers somewhat fleshy, pedicel c. 4 mm long; receptacle-tube shallowly cup-shaped,
c. 5 mm diam., c. 0.5 mm thick, firm-fleshy (disc); sepals broadly obtuse-triangular,
c. 2 mm long, together with the receptacle-tube showing up from beneath as a pale green
pentagon c. 9 mm diam.; perianth circular in outline, c. 12 mm diam.; petals broadly
obovate, 3.5 – 4.5 by 4 – 5 mm, inside papillose; filaments somewhat tapering, 1.2 –1.5
mm long, whitish, anthers c. 2 mm long. Female flowers not known. Fruit pendent,
green, ripening brown, smooth, glabrous, opening at the flat apex by 3 bluntly triangular
inward curving valves, 20 – 25 by 8 –10 cm, base ± narrowed, subacute, apex flattened,
pericarp woody, c. 0.5 mm thick, inside with a spongy-fibrous layer c. 10 mm thick,
line of perianth-scar 0.5 –1(–1.5) cm below the orifice; fruiting pedicel 1–1.2 cm long,
2 – 3 mm tick. Seeds densely packed in 3 rows throughout the fruit, dull brown, elliptic,
15 – 20 by 10 mm, faces finely warty, the edge with 8 (or 9) coarse, blunt spines 3 – 9
mm long, with a conspicuous membranous whitish subcircular wing around the seed,
c. 5 cm diameter. — Fig. 7, 8a – d; Plate 3, 4a, b.
Distribution — Peninsular Malaysia (Kedah, Perak, and Selangor).
Habitat & Ecology — Foothill and lower montane forest, on good soil; 100 – 800
m altitude. Flowering recorded in February, June, August, and November; fruiting in
December to February, rarely in August.
Notes — 1. Young plants have 4 verticillate, petioled pseudocotyledon-like first
leaves, then sprawling, with leaves alternate, with long internodes; the margin of juve-
nile leaves is coarsely sinuate, the apex apiculate. Fresh leaves are somewhat fleshy,
and glossy.
2. In male flowers, below each sinus between the corolla lobes, on the inner side of
the receptacle-tube, are hard-fleshy, palish, glossy, verrucose, long-triangular markings,
with at the apex, in each sinus, a minute globose creamy-white appendage, contrast-
ing with the purple-red corolla. The meaning and morphological explanation of these
structures is unknown.
3. Female plants are possibly monocarpous.
5. Benincasa
Benincasa Savi, Mem. sopra una pianta Cucurbitaceae (1818) 158; Backer in Backer & Bakh.f., Fl.
Java 1 (1964) 301; C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1530; W.J.de
Wilde & Duyfjes, Reinwardtia 12 (2007) 267; Sandakania 17 (2008 ‘2007’) 45; Fl. Thailand 9, 4
(2008) 419. — Type species: Benincasa cerifera Savi.
Stout herbaceous annual climber; plant hairy; monoecious (sometimes flowers

hermaphroditic). Probract hooded or concave. Tendrils branched, halfway or below.
Leaves: blade simple. Flowers solitary, long-pedicelled, large; sepals recurved; petals
free, yellow. Male flowers: receptacle-tube shallow; sepals entire or serrate; petals
large, obovate, margin entire; stamens 3, inserted towards the base of the receptacle-
tube; filaments free, short, anthers all 2-thecous, exserted, thecae flexuous, connective
broad, thin, 3-lobed; disc absent or low. Female flowers: perianth as in male flowers;
ovary ovoid or narrowly ellipsoid, densely hairy, style short, thick, inserted on the
disc, stigma robust; ovules numerous, horizontal; staminodes present, sometimes with
reduced anthers. Fruit a pepo, mostly waxy-whitish, smallish or large, indehiscent.
Seeds numerous, pale, compressed, margin only slightly thickened.
Distribution — One species with two forms, the cultivated form with several culti-
vars. The precise area of the wild form is not known, but it is speculated to include East
Malesia and the Pacific. The species is mainly known by the widely cultivated form
with edible fruits.
Note — Young developing leaves have small glands on the blade, and their lobe-
apices develop precociously and are conspicuously darker on drying. This phenomenon
is also obvious in not closely related Mukia, Cucumis sativus, and some other genera.
1. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes

Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes, Reinwardtia 12 (2008) 268; Sandakania 17
(2008 ‘2007’) 45, Fl. Thailand 9, 4 (2008) 421. — Cucurbita pruriens Parkinson, J. Voy. South
Seas (1773) 44.
Annual herb, shoots 2 –7 m long, stem 3 – 5 mm diam., sparsely or densely grey or

rusty hairy, hairs 2 – 5 mm long. Probract narrow- or broad-elliptic, (5 –)10 – 20 mm
long, acute or blunt. Tendrils 2- or 3-branched. Leaves: petiole 4 –12(– 20) cm long;
blade 5 –7-angular or shallowly (sometimes deeply) lobed, 7– 20 cm diam., cystoliths
not apparent, margin coarsely dentate, apex of lobes acute-acuminate. Male flowers:
pedicel 40 –150 mm long, hairy; receptacle-tube broadly cup-shaped, 4 – 6 by 10 –15
mm; sepals narrowly elliptic or linear, 10 – 20 mm long, acute, entire or (deeply) 2- or
3-lobed or serrate; petals obovate to narrowly elliptic, 25 – 40(–70) mm long, subacute
or obtuse, with minute mucro, hairy on veins outside; stamens inserted at c. 1/3 from the
base in the receptacle-tube, very densely pilose-hairy where the stamens are inserted,
the hairs screening off a small basal glabrous chamber (i.e. the basal portion of the re-
ceptacle-tube) without or with an inconspicuous disc at base, filaments glabrous, erect,
broad, 2 – 3 mm long, anthers somewhat connivent, 4 – 5 by 5 –7(–10) mm, somewhat
exserted, thecae plicate along a broad, flat 3-lobed connective. Female flowers: peri-
anth as in male flowers but somewhat smaller; pedicel 20 – 40 mm long; ovary densely
hairy, (narrowly) ovoid or (narrowly) ellipsoid (cylindrical), 15 – 20(– 40) by 5 –10(–15)
mm, style stout, 2 – 3 mm long, stigma stout, 5 –10 mm diam., undulating by 3 horse-
shoe-shaped lobes; disc a low ± undulating ring; staminodes inserted close to the disc,
mostly inconspicuous, 1– 2 mm long, flat. Fruit solitary, globose, ellipsoid or narrowly
ellipsoid, 4 –120 cm long, glabrescent; exocarp thin, leathery, when old woody, 1–1.5
mm thick; mesocarp ± absent or thick and fleshy; fruiting pedicel 3 – 5 cm long, usually
curved, leaving a concave scar after breaking off. Seeds ovate-elliptic, 7–10 by 4 –7
mm, faces not ornamented, margin narrow, finely 2-grooved, edge entire.
Distribution — The cultivated form widespread in SE Asia and elsewhere.

Uses — The young and ripe fruits and the young shoots are used as a vegetable.
KEY TO THE FORMS
1a. Fruit subglobose to narrowly ellipsoid (10 –)20 – 60(–120) by 10 – 25 cm. Seeds
8 –10 mm long. — Cultivated . . . . . . . . . . . . . . . . . . . . . . . . . . . b. forma hispida
b. Fruit globose or ellipsoid, 4 – 8 cm long. Seeds 7– 8 mm long. — Growing wild . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. forma pruriens
a. forma pruriens
Cucurbita vacua F.Muell., Fragm. 6 (1868) 186. — Benincasa vacua (F.Muell.) F.Muell., Syst. Census
Austral. Pl. 1 (1882) 76. — Type (syntypes, Telford 1982): Dallachy s.n. 28 Sept. 1867 & 20 Nov.
1867 (MEL, not seen), Queensland, Rockingham Bay.
Benincasa hispida (Thunb.) Cogn., for the wild form: I.Telford, Fl. Austr. 8 (1982) 170; Peekel, Fl.
Bismarck Archip., translated by E.E.Henty (1984) 547, f. 874.
Plants generally of a more delicate stature compared to cultivated form hispida; hairs
grey(-brown) coloured. Flowers smaller, in male 50 –70 mm diameter. Fruit globose
or ellipsoid (subacute at both ends), 4 – 8 cm long; exocarp woody, 1–1.5 mm thick;
mesocarp ± absent. Seeds 7– 8 mm long, embedded in rather juicy whitish pulp. —
Fig. 9; Plate 31c.
Distribution — Indochina, south-east through Malesia to Australia (Queensland),
Solomon Islands, and islands in the Pacific; in Malesia: Borneo, East Java, Lesser Sunda
Islands, Sulawesi, New Guinea (Papua New Guinea (New Ireland, fruit ellipsoid)).
Habitat & Ecology — Disturbed areas in lowland rainforest, forest edges, open for-
est, river banks, scrub-land and beach-vegetation; up to 700 m altitude.
Note — Forms similar to the truly wild form of Benincasa pruriens possibly are feral
forms, developed from escaped populations of the cultivated form hispida in natural
places. As the oldest collections of the wild form, with very small fruits, come from
East Malesia, Queensland, and The Pacific, this area is the most plausible original place
of occurrence of Benincasa.
b. forma hispida (Thunb.) W.J.de Wilde & Duyfjes

Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes forma hispida (Thunb.) W.J.de Wilde &
Duyfjes, Sandakania 17 (2008 ‘2007’) 47. — Cucurbita hispida Thunb., Nov. Acta Regiae Soc.
Sci. Upsal. 4 (1783) 38; Benincasa hispida (Thunb.) Cogn. in A.DC. & C.DC, Monogr. Phan. 3
(1881) 513; Craib, Fl. Siam. 1 (1931) 757; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 302; Rifai
& M.E.C.Reyes, PROSEA 8 (1993) 95. — Type: Thunberg 22775 (holo UPS, IDC microfiche),
Japan.
Benincasa cerifera Savi, Mem. sopra una pianta Cucurbitaceae (1818) 58; Ridl., Fl. Malay Penin.
1 (1922) 852. — Type: not known, described from a cultivated plant, possibly originating from
eastern Asia.
Cucurbita villosa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 931. — Type: Blume 1079 (holo L), Indonesia,
Java.
Cucurbita farinosa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 931. — Type: not found, Indonesia, Java.
2 mm
1 cm
d
2 cm
2 cm
3 mm
Fig. 9. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes forma pruriens. a. Habit; b. detail
of male flower showing stamens; c. node with female flower; d. fruit and node with fruiting pedicel;
e. seed (a, b: Brass 24290, a short-haired plant; c: Postar et al. SAN 144094, a long-haired plant;
d, e: Postar et al. SAN 144142).
Cucurbita littoralis Hassk., Cat. Hort. Bot. Alt. (1844) 190. — Type: not found, Indonesia, Java.
Gymnopetalum septemlobum Miq., Fl. Ned. Ind. 1, 1 (1856) 679. — Type: Junghuhn s.n. (holo L,
barcode L0587810), Indonesia, Weltevreden.
Stout plant with grey or rusty coloured hairs. Flowers large, in male 6 –10 cm dia
meter. Fruit subglobose or (long-)ellipsoid, (10 –)20 – 60(–120) cm long; exocarp hard-
leathery, c. 0.5 mm thick; mesocarp fleshy, 20 –100 mm thick. Seeds 8 –10 mm long,
embedded in rather firm pulp. — Plate 4c, d, 32a, c.
Distribution — Malesia: Widely cultivated.
Uses — The young and ripe fruits and young shoots are used as a vegetable.
Notes — 1. The form hispida, or Wax Gourd, comprises all cultivated or escaped
varieties or cultivars.
2. A more conspicuous disc in the (male) flower can be found in cultivated varieties,
especially those with hermaphroditic flowers.
6. BORNEOSICYOS
Borneosicyos W.J.de Wilde, Reinwardtia 11 (1998) 224; W.J.de Wilde & Duyfjes, Fl. Males. Bull. 14
(2007) 35. — Type species: Borneosicyos simplex W.J.de Wilde.
Slender liana, perennial; dioecious. Probract present. Tendrils unbranched. Leaves:

blade simple, unlobed. Flowers (pale) yellow. Male inflorescences racemose-panicu-
late, flowers short-pedicellate. Male flowers: receptacle-tube short cup-shaped, lobes
minute; petals 5, entire, free, quincuncial; stamens 3, free, inserted towards base of
tube, anthers free but connivent into a depressed globose head, two 2-thecous, one 1- or
1.5-thecous, thecae sigmoid, connectives broad, glabrous; disc absent or not obvious.
Female flowers solitary or in a short raceme co-axillary with single female flower;
ovary narrowly ellipsoid, ovules rather few, horizontal; receptacle-tube and perianth
as in the male flowers; style conspicuous, stigma 3-parted; staminodes 4 or 5, subulate,
inserted near the throat of the tube; disc present (obscure in the male ones). Fruit solitary
(always?), somewhat baccate. Seeds rather large, near-globose, faintly margined; testa
not ornamented.
Distribution — 1 (or 2?) species in Sarawak and Sabah.
1. Borneosicyos simplex W.J.de Wilde

Borneosicyos simplex W.J.de Wilde, Reinwardtia 11 (1998) 224; W.J.de Wilde & Duyfjes, Fl. Males.
Bull. 14 (2007) 35. — Type: Chow & Aban SAN 65050 (holo KEP; iso K, SAN), Sabah, Crocker
Range.
Liana 6 –12 m high, main stem to 10 mm diam., leafy stem terete, c. 2 mm diam.,
green, glabrous except for minute hairs on young parts. Probract narrowly elliptic, c. 3
mm long, acute, subentire, with minute scattered glands. Tendrils 6 –16 cm long. Leaves:
petiole 10 –15 mm long, 1.5 – 2 mm thick; blade membranous or slightly succulent,
ovate-oblong, 8 –13 by 3 – 6 cm, glabrous, glands on lower surface numerous, scattered,
less than 0.5 mm diam., cystoliths absent, base rounded or shallowly cordate, margin
entire or more usually with some minute sparse teeth up to 1 mm long, sometimes dis-
tinctly dentate in sterile shoots, apex acute-acuminate; basal veins 3(– 5)-palmate, arch-
2 cm
g
2 cm
2 mm
i
a
h
i'
1 mm 1 mm
f
c
2 mm
1 mm
c'
b e d
Fig. 10. Borneosicyos simplex W.J.de Wilde. a. Sterile leafy twig; b. portion of twig with male inflo-
rescence; c, c'. apex of partial male raceme with buds; d. male bud opened, showing one immature
stamen ± adaxially; e. immature stamen, abaxially; f. male flower, lower portion, with receptacle-tube
and sepals; g. twig with fruiting pedicels and basal portions of fruits; h. fruit; i, i' seed (a: De Wilde &
Duyfjes 21959; b – f: De Wilde & Duyfjes SAN 139474; c': Postar et al. SAN 14425; g, i, i': J. & M.S.
Clemens 26751; h: Chow & Aban SAN 65050).
c
1 mm
2 cm
2 mm
1 mm
g
2 mm 3 mm
b d e f
Fig. 11. Borneosicyos simplex W.J.de Wilde. a. Female flowering twig; b. node with developing fe-
male inflorescence, co-axillary with pedicel of fallen female flower, also showing probract; c. detail
of leaf margin, abaxial side, showing small marginal tooth and two leaf glands; d. racemose female
inflorescence; e, f. female flowers; g. ovary in cross section (a, b, d – g: De Wilde, Postar & Ubaldus
SAN 144018; c: De Wilde & Duyfjes 21959).
2 mm
a d b c
Fig. 12. Borneosicyos simplex W.J.de Wilde. a. Male flower; b. ditto, opened, showing anther head;
c. separate stamen with anther 2-thecous; d. stamen with anther 1.5-thecous (all: Postar, De Wilde &
Ameng SAN 144251).
ing-up to halfway the blade, lateral veins 2 – 4 per side. Male inflorescences ± peduncled
lateral panicles 10 –15 cm long, loose, composed of few few-flowered short racemes
2 – 4 cm long; bracts small, 1 mm or less, tardily caducous. Male flowers: pedicel c. 4
mm long, articulate c. ± halfway; expanded perianth 10(–15) mm wide; receptacle-tube
c. 4 by 5 mm; sepals green, later brown, triangular, c. 0.5 mm long; petals (narrowly)
elliptic, c. 8 by 4 mm, obtuse, 5 – 8-veined, inside and outside minutely sparsely grey-
brown pubescent (hairs c. 0.2 mm long), margin finely fimbriate; filaments subulate, c.
3 mm long, with few hairs in the lower portion; anther head c. 4 mm wide, anthers c. 3
mm diam., connective broad, glabrous. Female flowers solitary or 4 –7 in a short raceme
(peduncle 3 – 4 cm long); bracts minute; pedicel in solitary flower 10 –15 mm long, in
raceme c. 5 mm long; ovary cylindrical-oblong, 9 –10 by c. 3 mm, at apex somewhat
narrowed into a c. 2 mm long neck, minutely brownish pubescent, green with paler
irregular lengthwise flecks and faintly warty bands; receptacle-tube cup-shaped, c. 3.5
by 4 mm, inside thickened and with 5 faint disc pads; sepals shortly connate forming at
anthesis a tube 1(– 2) mm long with the lobes somewhat reflexed, acute, 1–1.5 by c. 0.5
mm; petals subacute, c. 10 by 4 – 5 mm, connate for up to 0.5 mm only, 3(– 5)-veined,
minutely papillose-hairy, short-woolly at base, apices in bud slightly contorted; style
cylindrical, at base somewhat broadened, 9 –10 by 1(–1.5) mm, stigma of 3 sessile ±
recurved parts, each elongated heart-shaped, 3.5 – 4 by c. 2 mm, papillose; staminodes
subulate, c. 6 mm long, soft-haired. Fruit at first green, darker flecked, ripening red,
smooth, glabrous, narrowly ellipsoid, (4 –)8 –10 by (2 –)4 – 4.5 cm, base rounded, apex
acute, 2 – 3 mm beaked; exocarp thin on drying; endocarp juicy, on drying leaving
thin membranes around the seeds; fruiting pedicel slender, 0.5 –1 cm long (in raceme)
or 3.5 – 5 cm long (when fruit solitary), c. 5 mm thick. Seeds 20 – 30, somewhat com-
pressed globose, 9 –10 by 8 – 9 by c. 4 mm, with a faint margin, smooth, edge entire. —
Fig. 10 –12; Plate 5.
Distribution — Sabah: Crocker Range, Kinabalu Park (Tenompok area and Langa-
nan Waterfall), but see note 2.
Habitat & Ecology — Primary montane Lithocarpus evergreen forest, (1000 –)1500 –

1800 m altitude; flowering in July and December; fruiting in July and October. Full-
grown Borneosicyos is a slender but tall liana, flowering in the tree crowns 5 –10 m
high. The immature sterile plants thrive in the moist atmosphere of the underscrub in
the montane forest, preferably not far from streams.
Note — The collection, Yii S 51389 with one open male flower only, from West
Sarawak, Gn Merubong, possibly represents an undescribed species of Borneosicyos.
It is distinct by a slightly stouter habit, and leaves with a rather deeply cordate base; its
pollen is in tetrads as in B. simplex.
7. CAYAPONIA
Cayaponia Silva Manso, Enum. Subst. Braz. 31 (1836) 31, nom. cons.; Cogn. in A.DC. & C.DC.,
Monogr. Phan. 3 (1881) 738; Pax in Engl. & Prantl, Nat. Pflanzenfam. 4 (5) (1889) 34; C.Jeffrey,
Kew Bull. 25 (1971) 201. — Type species: Cayaponia diffusa Silva Manso, typ. cons.
Climbers, perennial; monoecious (or dioecious, not in Malesia). Probract absent.

Tendrils 2- or 3-branched or unbranched. Leaves: blade simple, unlobed or lobed, or
3 – 8-foliolate, usually with scattered glands near the insertion to the petiole. Flowers
whitish or yellowish green, solitary or fascicled, arranged in racemes, or subpaniculate;
petals free or short-connate. Male flowers: receptacle-tube campanulate or subcylindri-
cal; petal lobes (ovate-)elliptic or oblong, valvate or (sub)imbricate; stamens 3, free,
inserted near the throat of the receptacle-tube (or lower down, not in Malesia), anthers
basifixed, often connivent, two 2-thecous, one 1-thecous, thecae plicate, connective not
produced; disc (pistillode) absent or small. Female flowers solitary or few-fascicled, ar-
ranged in panicles; perianth as in male flowers; ovary ovoid, ovules 1– many per locule,
erect, style long, with nectary-like annular disc at base, stigmas 3, unlobed, recurved;
staminodes small. Fruit ovoid or subglobose, berry-like, with 3 –12 seeds. Seeds little
compressed, mostly smooth, margin absent, edge entire.
Distribution — About 50 species in the Neotropics, few species in Africa, including
Madagascar; in Malesia: 1 introduced and naturalized species.
1. Cayaponia martiana (Cogn.) Cogn.

Cayaponia martiana (Cogn.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 777, including var.
acutiloba & var. tomentosa; Backer in Backer & Bakh.f., Fl. Java 3 (1968) 645; C.Jeffrey, Kew Bull.
25 (1971) 233. — Trianosperma martiana Cogn. in Mart., Fl. Bras. 6, 4 (1878) 87, t. 25. — For
typification see Cogn. (1878).
Climber or creeper to 5 m long; stem subglabrous. Leaves: petiole 4 – 8 cm long; blade

(deeply) 3 –7-lobate (upper leaves smaller, unlobed), 8 –15 by 10 – 20 cm, scabrous-
hairy, base narrowed into the petiole, margin crenate-dentate. Inflorescences elongate,
variable, grouped into large leafy panicles, bracts minute. Male flowers fasciculate;
pedicel 2 – 5 mm long, hairy, mostly with a bracteole; receptacle-tube narrowly cam-
panulate, 3–4 by c. 1.5 mm; sepals c. 1 mm long; petals whitish, 5 – 6 by 2 mm, papil-
lose-villose; filaments c. 2.5 mm long, anthers ± coherent into a head, c. 1.5 by 1 mm;
disc elongate. Female flowers as in male flowers; ovary narrowly ellipsoid, constricted
2 cm
3 mm
k
2 cm 1 mm
1 mm g
c
h 1 mm
d
0.5 mm
1 mm
b
1 mm
i j
Fig. 13. Cayaponia martiana (Cogn.) Cogn. a. Habit of node with male inflorescence; b, c. male flower,
from outside and opened respectively (thecae plicate); d. detail of portion of filament; e. infructescence
with immature fruits; f. ditto, with mature fruits; g, h. seeds; i, j. detail of lower and upper leaf blade
respectively; k. base of leaf blade with glands on lower surface (a – d, i – k: Neubauer 594; e: Koster
mans s.n., November 1938; f – h, k: herb. Martius s.n., barcode L 0587282, Brasil).
at apex, receptacle-tube villose-papillose at the throat; staminodes inconspicuous. Fruit

ripening green or yellow, yellowish striped, subglobose, c. 1 cm long; fruiting pedicel
c. 0.5 cm long. Seeds few, grey, ovoid, 5 – 6 by 3.5 – 4 by 2 – 2.5 mm. — Fig. 13.
Distribution — South America (Brazil, Paraguay); in Malaysia: East Java, Idjen Pla-
teau, where introduced and running wild; 800 –1000 m altitude; flowering and fruiting
probably throughout the year.
8. citrullus
Citrullus Schrad. Enum. Pl. Afric. Austral. 2 (1836) 279, nom. cons.; Cogn. in A.DC. & C.DC.,
Monogr. Phan. 3 (1881) 507; I.Telford, Fl. Australia 8 (1982) 173; C.Jeffrey in Hanelt, Mansfeld’s
encycl. agric. hort. crops 3 (2001) 1533. — Type species: Citrullus vulgaris Schrad., typ. cons.
Herbs, annual (or perennial with root-stock), trailing or climbing, plant scabrous or
soft-hairy; monoecious. Probract present. Tendrils branched (unbranched or spinescent,
not in Malesia). Leaves: blade simple, ± pinnately lobed. Flowers solitary, pedicelled,
medium-sized, petals (nearly) free, yellow. Male flowers: receptacle-tube short, broadly
campanulate; sepals narrow; petals ovate-oblong, obtuse, margin entire; stamens 3, in-
serted near the base of the receptacle-tube, filaments free, short, anthers coherent, two
2-thecous, one 1-thecous, thecae plicate, connective broad, flat; disc inconspicuous.
Female flowers: perianth as in male flowers; ovary ovoid or subglobose, hairy, style
short, stigma deeply 3-lobed, ovules numerous, horizontal; disc not apparent; stami-
nodes small. Fruit a pepo, solitary, (small or) large, firm-walled, indehiscent. Seeds
numerous, compressed, (narrowly) ovate in outline, (nearly) smooth, margined or not,
edge entire.
Distribution — A genus of 4 species in Africa, east to Pakistan; in Malesia: 1 species,
C. lanatus, widely cultivated.
1. Citrullus lanatus (Thunb.) Matsum. & Nakai

Citrullus lanatus (Thunb.) Matsum. & Nakai, Cat. Sem. & Spor. Hort. Bot. Univ. Imp. Tokyo
(1916) 30; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 300; H.Hara, Taxon 18 (1969) 346,
f. 5; I.Telford, Fl. Australia 8 (1982) 173; Paje & Van der Vossen, PROSEA 8 (1994) 144; W.J.de
Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 49; Fl. Thailand 9, 4 (2008) 422. — Momordica
lanata Thunb., Prodr. Pl. Cap. (1794) 13. — Type: Thunberg s.n. (UPS, photo seen), Cape Province,
Republic of South Africa.
Citrullus vulgaris Schrad., Enum. Pl. Afric. Austral. 2 (1836) 279; Craib, Fl. Siam. (1931) 760.
— Type: not known.
Citrullus edulis Spach, Hist. Nat. Vég. 6 (1838) 214; Miq., Fl. Ned. Ind. 1, 1 (1856) 662. — Type:
not known.
Annual herb, shoots to 3 m long, stem 2 – 4 mm diam., soft grey-hairy, hairs 2 – 3(– 5)
mm long. Probract narrowly elliptic, (sub)obtuse, narrowed at base, 5 –15(– 20) mm
long, glands not obvious. Tendrils 2- or 3-branched. Leaves: petiole 3 –12 cm long, soft
hairy; blade pinnately (deeply) 3 – 5-lobed, ovate to narrowly elliptic in outline, 5 – 20 by
3 –15 cm, glabrescent above, scabrid beneath, cystoliths and glands not obvious, base
shallowly cordate, margin irregularly dentate, apex and apices of lobes rounded, obtuse
or acute. Male flowers: pedicel 10 – 80 mm long, hairy; receptacle-tube 3 – 4 mm long,
3 mm
3 mm
b
h
2 mm
e 2 cm
f
1 mm
3 mm
a d
Fig. 14. Citrullus lanatus (Thunb.) Matsum. & Nakai. a. Growing branch with male and female flow-
ers; b, c, d. male flowers, from outside and opened, in d stamens segregated by hand; e. hair of stem;
f. petal; g female flower, from outside and opened; h. seed (a – g: De Wilde & Duyfjes 21803; h: De
Wilde & Duyfjes 22328).
villous-hairy; sepals linear, 3 – 5 mm long; petals 8 –15 mm long; filaments c. 2 mm

long, glabrous, anthers connivent into a globose mass c. 3 mm diameter. Female flow
ers: pedicel 5 – 40 mm long; ovary 6 –12 mm long, style 4 – 5 mm long. Fruit solitary,
ripening (pale) green or greyish green without or with stripes or blotches, or yellow,
(sub)globose or ellipsoid, 20 – 30(– 60) cm long, glabrescent, smooth; exocarp hard
but not woody; pulp (including mesocarp) white, yellow or pink(-red); fruiting pedicel
2 –7 cm long. Seeds white, brown or nearly black, 6 –12 mm long, smooth or ± rough,
margin absent or present. — Fig. 14; Plate 6c.
Distribution — Malesia: widely cultivated.
Vernacular names — Semangka (Indonesia).
Note — Citrullus lanatus, the watermelon, is popular and cultivated everywhere in
various cultivars for its fruits. Discarded seeds easily give rise to temporary spontane-
ous occurrence.
9. COCCINIA
Coccinia Wight & Arn., Prodr. Fl. Ind. Orient. 1 (1834) 347; Cogn. in A.DC. & C.DC., Monogr. Phan.
3 (1881) 528; C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1529; W.J.de Wilde
& Duyfjes, Fl. Thailand 9, 4 (2008) 423. — Type species: Coccinia indica Wight & Arn., nom. illeg.
(Bryonia grandis L. = Coccinia grandis (L.) Voigt).
Herbaceous or (sub)woody perennial climbers, early glabrescent; dioecious. Pro

bract small, caducous. Tendrils unbranched (or elsewhere 2-branched). Leaves: blade
simple, variable of shape. Flowers white (or creamy or orange, not in Malesia), petals
fused into corolla-tube. Male flowers solitary (or 2 or 3), white (elsewhere yellowish);
receptacle-tube cup-shaped; sepals 5, small; corolla campanulate, 5-lobed, lobes fused
for c. 2/3, margin entire; stamens 3; filaments fused or free but tightly appressed form-
ing a column, inserted at or near the base of the receptacle-tube; anthers two 2-thecous
and one 1-thecous (elsewhere all 2-thecous), free but mostly coherent into a globular
synandrium; thecae plicate; connectives broad; base of receptacle with a nectariferous
disc (all species?). Female flowers mostly solitary; ovary ovoid or oblong, smooth or
faintly ribbed, glabrous; ovules many, horizontal; stigmas 3, each 2-lobed. Fruit ripen-
ing red, berry-like, ellipsoid (elsewhere globose), moderate of size, pericarp thin. Seeds
numerous, (ovate-)elliptic, compressed, margin distinct, edge entire.
Distribution — A genus of c. 30 species mainly in Africa; in Malesia 1 species.
1. Coccinia grandis (L.) Voigt

Coccinia grandis (L.) Voigt, Hort. Suburb. Calcutt. (1845) 59; Backer in Backer & Bakh.f., Fl. Java 1
(1964) 305; I.Telford, Fl. Australia 8 (1982)176; Boonkerd, Na Songkhla & Thephuttee, PROSEA
8 (1993) 150; W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 50; Fl. Thailand 9, 4 (2008)
423. — Bryonia grandis L., Mant. Pl. (1767) 126. — Coccinia indica Wight & Arn., Prodr. Fl. Ind.
Orient. 1 (1834) 347, nom. illeg.; Craib, Fl. Siam. 1 (1931) 761. — Cephalandra indica (Wight &
Arn.) Naudin, Ann. Sci. Nat. Bot., Sér. 5 (1866) 16, nom. illeg.; Ridl., Fl. Malay Penin. 1 (1922)
849. — Type: Herb. Linn. No. 1153/2 (lecto LINN, designated by Jeffrey, Fl. Trop. E Africa (1967)
68 (see C.E.Jarvis, Order out of chaos (2007) 363)), India.
Momordica bicolor Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 928, including vars. a and b. — Type: Blume
1012 (holo L), Java.
2 cm
2 cm
2 cm
a l
2 mm
c 3 mm n
3 mm
2 mm 2 mm
b 2 mm
j
2 mm
d e h 1 mm g m
Fig. 15. Coccinia grandis (L.) Voigt. a. Node with male flowers; b, c. male flowers; d, e. stamens;
f. node with female flower; g, h. female flowers; i. ditto, basal portion opened; j. staminode; k. portion of
twig with immature fruits; l. node with fruit; m. seed; n. blade base, lower surface, showing glands; note:
corollas lobed to about halfway deep (a: De Wilde & Duyfjes 21734; b – e: De Wilde & Duyfjes 22270 ;
f, k: De Wilde & Duyfjes 21717; g – j: De Wilde & Duyfjes 22271A; l – n: De Wilde & Duyfjes 21739).
Coccinia wightiana M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 93. — Type: Wight 1123 (not seen),
but Bryonia grandis L. cited by Wight & Arnott (1834, l.c.) in the synonymy.
Herbaceous or soft-woody climbers to 8 m long, sparsely puberulous, early glabres-

cent; older stem to 20 mm across, with grey bark. Probract fleshy, elliptic to oblong,
2 – 3 mm long, caducous. Tendrils unbranched. Leaves: petiole 1– 2.5 cm long; blade
slightly succulent, unlobed, (3- or) 5-angular or (deeply) 3 – 5(–7)-lobed, or dissected
to 4/5 deep, 3 –10 cm across, glands small, several, near the insertion of the petiole
and along lateral veins, cystoliths in older leaves present, margin minutely (blackish)
dentate-sinuate, apex acute or rounded. Male flowers solitary (or 2 or 3); pedicel 20 – 50
mm long; receptacle-tube greenish white, cup-shaped, narrowed to the base, 6 – 8 by
c. 6 mm at throat; sepals oblong-subulate, c. 3 mm long, ± up-curved; corolla white, pale
green veined, broad campanulate, (15 –)25 – 30 mm long, inside (sparsely) pubescent,
lobes ovate-oblong, acute(-acuminate); stamens inserted c. 5 mm below the throat of
the receptacle-tube, filaments either connate or free but coherent, forming a hollow
column (4 –)5 – 6 mm long (sometimes leaving 3 openings at base), synandrium c. 5
mm diam.; the receptacle-tube below the insertion of the stamens densely white-hairy,
hiding a basal cavity, c. 3 mm deep, comprising a conspicuous green-yellow cup-shaped
disc adnate to the base of the tube. Female flowers: pedicel 1–3 cm long; ovary oblong,
10(–13) by 2(– 3) mm, narrowed at both ends, style 3 – 5 mm long, at base surrounded
by a cup-shaped disc, stigmas 3, usually 2-lobed, 5 –7 mm long, papillose; staminodes 3,
minute. Fruit green-white blotched, ripening red (starting at the apex), (narrowly) ellip-
soid, 2.5 – 6 by 1.5 – 3 cm, apex subacute, pulp red, juicy; fruiting pedicel 1– 4 cm long.
Seeds 6 –7 by 2.5 – 3 by c. 1.5 mm, ± smooth, margin narrow. — Fig. 15; Plate 7.
Distribution — Widely distributed in the Old World, northern tropical Africa, east
to Arabia through India to tropical North Australia; in Malesia: Peninsular Malaysia,
Singapore, Borneo (Sabah, cultivated), eastern Java, Philippines (Cebu), Lesser Sunda
Islands (Bali, Lombok, Flores Timor), Moluccas (Weri). Currently pantropical.
Ecology — In open forest, scrub, along waste land, roadsides, and near the coast;
from sea level to 400 m altitude; flowering and fruiting throughout the year.
Uses — Young shoots are eaten as a vegetable.
Note — Coccinia grandis prefers a (strong to) light monsoon climate. Its fruits
are (glaucous) green with contrasting white blotches in rows and turn into bright red
when ripe, except for the very base. The pulp is frequently eaten by animals, mainly by
birds.
10. Cucumis
Cucumis L., Sp. Pl. ed. 1 (1753) 1011; Gen. Pl. 5 (1754) 442; Miq., Fl. Ned. Ind. 1, 1 (1856) 670;
C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 619; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
479; J.H.Kirkbr., Biosyst. Monogr. Gen. Cucumis (1993) 19; H.Schaef., Blumea 52 (2007) 166;
Ghebret. et al., Novon 17 (2007) 176; W.J.de Wilde & Duyfjes, Adansonia, sér 3, 29 (2) (2007)
240. — Lectotype (Britton & Wilson, Scientific Survey of Porto Rico and the Virgin Islands 6, 2
(1925) 264): Cucumis sativus L.
Melo Mill., Gard. Dict. Abr. ed. 4 (1754) without pagination. — Lectotype (Swart, Index Nom. Gen.
(Pl.) Card (1960)): Cucumis melo L.
Small or medium-sized annual or subperennial climbers (rarely suberect), leafy

stem 1– 4 mm diam., plant scabrous or setose; monoecious (rarely dioecious); usually
cultivated. Probract absent. Tendrils unbranched. Leaves simple. Flowers yellow, soli-
tary or few-fascicled; pedicels short; corolla composed of partly fused petals, small- or
medium-sized. Male flowers: receptacle-tube campanulate or turbinate, small; sepals
mostly linear; petals (corolla) with margin entire; stamens 3, free, inserted about half-
way the receptacle-tube, filaments short, anthers two 2-thecous, one 1-thecous, thecae
sinuate, 3-plicate or S-shaped, connective considerably produced; disc large, gland-like,
free from the tube. Female flowers usually solitary; ovary hairy, hairs sometimes api-
cal on small protuberances, ovules numerous, horizontal; perianth as in male flowers
but somewhat larger, stigma 3-lobed, lobes lobulate; staminodes often present, small;
disc surrounding base of style, free from the tube. Fruit a (large) pepo (fleshy berry),
indehiscent, pubescent or glabrous, or with fleshy spines or tubercles, green, yellow or
orange (rarely maturing underground in C. humifructus, southern Africa). Seeds numer-
ous, pale, compressed, (narrowly) elliptic, not sculptured, unmargined with entire, acute
edge, not winged (or rarely winged).
Distribution — About 30 species in the Old World, mostly Africa; in Malesia 2 spe-
cies, widely cultivated.
Taxonomy — According to molecular research (Ghebret. et al., Novon 17 (2007)
176; H.Schaef., Blumea 52 (2007) 165) the genus Cucumis should include a number of
smaller genera, including Mukia, but for the present purpose we treat Cucumis in the
original sense.
KEY TO THE SPECIES
1a. Leaf lobes mostly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . 1. C. melo (2 forms)

b. Leaf lobes acute-acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. sativus
1. Cucumis melo L.
Cucumis melo L., Sp. Pl. ed. 1 (1753) 1011; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 482;
Craib, Fl. Siam. (1931) 760; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 301; I.Telford, Fl. Aus-
tralia 8 (1982) 189; J.H.Kirkbr., Biosyst. Monogr. Gen. Cucumis (1993) 79; C.Jeffrey in Hanelt,
Mansfeld’s encycl. agric. hort. crops 3 (2001) 1512; Paje & Van der Vossen, PROSEA 8 (1994) 153;
W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 54; Fl. Thailand 9, 4 (2008) 428. — Type:
Herb. LINN NO. 1152/8 (lecto LINN, designated by Meeuse, Bothalia 8 (1962) 61 (see C.E.Jarvis,
Order out of chaos (2007) 465)), a plant cultivated at Uppsala.
Annual or subperennial (climber or) trailer to 6 m long, plant (woolly) hairy, hirsute
or hispid. Leaves: petiole 3 –10 cm long; blade ovate or subcircular in outline, 3 –15
cm diam., (deeply) lobed or unlobed, mostly hispid, lobes mostly rounded, margin
shallowly sinuate-toothed. Male flowers few-fascicled; pedicel 3 – 25 mm long, hairy;
receptacle-tube 4 – 6 mm long; sepals 1– 6 mm long; corolla united in lower third,
lobes (narrowly) elliptic, 5 – 20 by 2.5 –15 mm, apex obtuse or occasionally subacute,
mostly finely hairy; filaments c. 0.5 mm long, anthers ± included, 1– 2(– 2.5) mm long,
connective with (entire or) 2-lobed apical extension. Female flowers solitary; pedicel
5 – 30(– 50) mm long; ovary ellipsoid, 4 –10(–14) mm long, densely fine-hairy, hairs
spreading or appressed; sepals 1.5 – 3(–7) mm long; corolla 8 –15(– 25) mm long; style
1– 2 mm long, stigma 1.5 – 2.5 mm diameter. Fruit ripening green, yellow, white or
brown, plain-coloured, or striped or mottled, globose, (narrowly) ellipsoid or (ob)ovoid,
2 – 20(–100) by 2 – 5(– 20) cm, smooth; exocarp leathery; mesocarp juicy or carnose;
fruiting pedicel 2 – 4 cm long. Seeds 4 –15 by 1– 2 mm, usually not winged.
Distribution — Widely cultivated (and with feral forms) all over the world, including
Malesia.
Note — Cucumis melo, the honey melon, known all over the world in numerous
cultivars as a vegetable or table fruit, can be found seemingly wild when germinated
from seeds from the waste and then sometimes producing forms approaching the truly
wild forma agrestis (Naudin) W.J.de Wilde & Duyfjes.
KEY TO THE FORMS
1a. Plant generally more robust, leaf blade 4 –15 cm diameter. Ovary usually with ±
spreading hairs; corolla 8 – 20 mm long. Fruits (5 –)10 – 20(–100) cm long. Seeds
8 –15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. forma melo
b. Plant more slender, leaf blade 2 – 5 cm diameter. Ovary with appressed hairs; corolla
5 – 8 mm long. Fruits 2 – 5 cm long. Seeds 4 – 8 mm long . . . . . . b. forma agrestis
a. forma melo
Plant rather robust. Leaf blade 4 –15 cm diameter. Ovary usually with ± spreading
hairs; corolla 8 – 20 mm long. Fruit (5 –)10 – 20(–100) cm long. Seeds 8 –15 mm long.
Distribution — As the species.
b. forma agrestis (Naudin) W.J.de Wilde & Duyfjes

Cucumis melo L. forma agrestis (Naudin) W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 55.
— Cucumis melo L. var. agrestis Naudin, Ann. Sci. Nat., Bot., Sér. 4, 6 (1859) 73. — C. melo L.
subsp. agrestis (Naudin) Pangalo in Zhuk., La Turquie Agricole (1933) 534; I.Telford, Fl. Australia
8 (1982) 189; J.H.Kirkbr., Biosyst. Monogr. Gen. Cucumis (1993) 81. — Type: Naudin s.n. (lecto
P, designated by Kirkbride (1993) 105), cultivated from seeds from India.
Plant slender. Leaf blade 2 – 5 cm diameter. Ovary with appressed hairs; corolla 5 – 8
mm long. Fruit 2 – 5 cm long. Seeds 4 – 8 mm long. — Fig. 16; Plate 6b.
Distribution — Africa, Asia, Australia and the Pacific; in Malesia: Borneo (Kaliman-
tan), Java, Philippines, Lesser Sunda Islands (Lombok, Sumba, Flores), New Guinea
(West Papua, Papua New Guinea); often on barren fields and waste places, at low alti-
tudes.
2. Cucumis sativus L.
Cucumis sativus L., Sp. Pl. ed. 1 (1753) 1012; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 620; Cogn.
in A.DC. & C.DC., Monogr. Phan. 3 (1881) 498; Backer in Backer & Bakh.f., Fl. Java 1 (1964)
301; I.Telford, Fl. Australia 8 (1982) 189; Gildemacher & G.J.Jansen, PROSEA 8 (1993) 157;
J.H.Kirkbr., Biosyst. Monogr. Gen. Cucumis (1993) 84; C.Jeffrey in Hanelt, Mansfeld’s encycl.
agric. hort. crops 3 (2001) 1520; W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 56; Fl.
0.5 mm
f
2 cm
a
e
1 mm 1 cm
d
1 mm
b c
Fig. 16. Cucumis melo L. forma agrestis (Naudin) W.J.de Wilde & Duyfjes. a. Habit; b, c. male flower,
from outside and opened respectively; d. fruit; e. seed; f. detail of lower surface of leaf blade (a: Brass
24287; b – f: De Wilde & Duyfjes 21859).
Thailand 9, 4 (2008) 428. — Type: Burser vol. 17, no. 97 (lecto UPS, designated by Ten Pas et al.,
Taxon 34 (1985) 291), from cultivation.
Cucurbita vittata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 932. — Type: not indicated.
Cucumis? rumphii Hassk., Abh. Naturf. Ges. Halle 9 (1866) 280; Merr., Interpr. Rumph. Herb. Am-
boin. (1917) 47, 492. — Type: Rumphius, Herb. Amboin. 5 (1747) 404, t. 146, f. 2. Possibly a
feral form.
Annual climber or trailer, with fibrous roots, shoots to 5 m long, plant scabrid by
bristly hairs. Leaves: petiole 5 –10 cm long; blade broadly ovate in outline, 10 –15(– 20)
cm diam., (shallowly or) deeply 5-lobed, the middle lobe largest, both surfaces sparsely
hairy, margin finely dentate, lobes at apex acute-acuminate. Male flowers solitary or
few-fascicled; pedicel 5 – 20 mm long, hairy; receptacle-tube 4 – 5(–10) mm long, hairy;
sepals 4 – 5(–7) mm long; petals (corolla) (10 –)20(– 25) mm long, at base fused for
1/3; filaments short, anthers 2.5 – 3 mm long, included or hardly protruding, connective
extension slender, entire or 2-lobed; disc 1– 2 mm diameter. Female flowers solitary (or
few-fascicled); pedicel 2 – 20 mm long; ovary 10(– 20 in certain cultivars) mm long,
glabrous or densely hairy; perianth as in male flowers; style short, stigma 2 – 3 mm
diameter. Fruit ripening whitish or green, or variously two-coloured (striped), white,
green or yellow(-brown), ellipsoid to cylindrical, 5 – 20(– 50) cm long, blunt at apex,
smooth or with low prickles; exocarp thin; mesocarp fleshy or pulpy, whitish or yel-
lowish; fruiting pedicel 1.5 – 4 cm long. Seeds elliptic, 7–12 mm long, not winged.
Distribution — Originally occurring in SE Asia (the wild form, forma hardwickii
(Royle) W.J.de Wilde & Duyfjes in northern India, Myanmar and Thailand), at present
cultivated worldwide for its fruits.
Uses — Cucumis sativus, the cucumber, is probably the most commonly used source
of raw or cooked vegetable.
11. Cucurbita
Cucurbita L., Sp. Pl. ed. 1 (1753) 1010; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 542; Backer
in Backer & Bakh.f., Fl. Java 1 (1964) 305; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge,
Laos & Vietnam 15 (1975) 102; Widjaja & Sukprakarn, PROSEA 8 (1993) 160; C.Jeffrey in Hanelt,
Mansfeld’s encycl. agric. hort. crops 3 (2001) 1541; W.J.de Wilde & Duyfjes, Sandakania 17 (2008
‘2007’) 58; Fl. Thailand 9, 4 (2008) 430. — Pepo Mill., Gard. Dict. Abr., ed. 4 (1754) without
pagination. — Type species: Cucurbita pepo L.
Medium-sized stout annual (or perennial not in our area) trailing or climbing herbs,
leafy stem 3 – 5(–10) mm diam., plant hairy or scabrous; monoecious; usually cultivated.
Probract absent. Tendrils (2 –)3 – 6-branched. Leaves: petiole long; blade simple, large,
± unlobed or lobed, sometimes with whitish blotches, margin finely or coarsely dentate,
apex of lobes rounded or acute. Flowers orange(-yellow), solitary (in male sometimes
2); receptacle-tube (shallowly) campanulate; sepals 5; corolla large, 6 –10 cm long in
cultivated species, petals fused for the lower half, lobes acute, patent or ± out-curved at
apex. Male flowers: pedicel long; sepals oblong or linear, or obovate and long-clawed;
corolla-lobes ovate(-oblong), (sub)entire; stamens 3, filaments free, swollen at base (in
cultivated Asian species), inserted towards the bottom of the receptacle-tube, anthers
two 2-thecous, one 1-thecous, united (fused) into an elongated whole, thecae plicate-
sinuate, connective narrow, not produced, connate, hidden by the thecae; disc (pistil-
lode) absent. Female flowers: pedicel shorter than in male flowers; ovary globose or
ellipsoid, 1(– 2) cm diam., placentas 3(– 5), ovules numerous, horizontal; perianth as in
male flowers; style short, stout, stigma-lobes 3(– 5), thickly fleshy, each lobe shallowly
lobed; disc absent or inconspicuous; staminodes short, at base of receptacle-tube. Fruit
a pepo, often with hard rind, small or (very) large, variable in shape and colour; meso-
carp firm or pulpy. Seeds numerous, compressed, medium or large, (narrowly) elliptic
in outline, little or not ornamented, margin narrow (rarely broad), edge entire.
Distribution — About 25 species. All Cucurbita species are originally indigenous
in the New World; four species are cultivated all over the world, three extensively so
in tropical as well as (in summer) in subtropical or extra-tropical regions; in Malesia:
3 species commonly cultivated, but Cucurbita ficifolia Bouché occasionally cultivated
in the highlands of the Philippines.
Uses — Flowers, cooked or fried, are eaten. Fruits and (oily) seeds are widely used
as food. Shoots are used as a vegetable. Also medicinal.
Note — Marked wild or feral forms do not occur in our area, and these have not been
further accounted for. The three (five) cultivated species are each very variable and
look-alike and can often only be recognized when completely known (habit, flowers and
fruits); especially, well-developed fruit can be characteristic. Hybridization between the
species seems difficult or absent. A brief description of each current species should suffice
for distinction and naming a plant found in our area. The species of Malesia are annual.
KEY TO THE SPECIES
1a. Sepals mostly leaf-like broadened at apex (long-clawed) or ± linear. Fruiting pedi-
cel much broadened at the transition to the fruit . . . . . . . . . . . . . . 2. C. moschata
b. Sepals linear. Fruiting pedicel not or hardly broadened at the transition to the
fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Fruiting pedicel stout, terete, spongy . . . . . . . . . . . . . . . . . . . . . . . 1. C. maxima
b. Fruiting pedicel slender, angular, not spongy . . . . . . . . . . . . . . . . . . . . 3. C. pepo
1. Cucurbita maxima Duchesne

Cucurbita maxima Duchesne, Essai Hist. Nat. Courges (1786) 7; Lam., encycl. (1786) 151; Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 544; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 622;
Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 103; Paris, Taxon 49
(2000) 316; C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1545; W.J.de Wilde
& Duyfjes, Sandakania 17 (2008 ‘2007’) 60. — Type: Lamarck s.n. (holo P).
Plant, especially the leaves, rigid. Leave blades lobed or not, ± reniform in outline.
Flowers: pedicel terete; receptacle-tube campanulate; sepals linear in male and female
flowers. Fruit ripening (blue-)green or orange, smooth or warty; fruiting pedicel stout,
short, ± terete, spongy, often with a ± fissured surface, not widened at the transition to
the fruit. Seeds c. 20 mm long.
Distribution — Cultivated.
2. Cucurbita moschata Duchesne

Cucurbita moschata Duchesne, Essai Hist. Nat. Courges (1786) 7; Backer in Backer & Bakh.f., Fl. Java
1 (1964) 305; Paris, Taxon 49 (2000) 316; W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’)
61. — Cucurbita pepo L. var. moschata (Duchesne) Lam., encycl. (1786) 152; Keraudren in Aubrév.
& J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 104; C.Jeffrey in Hanelt, Mansfeld’s encycl.
agric. hort. crops 3 (2001) 1545. — Type (Keraudren, 1975): original material not known.
Gymnopetalum calyculatum Miq., Fl. Ned. Ind., Eerste Bijv. 2 (1861) 332. — Type: Amann (‘Amand’)
(= Kurz) s.n. (holo U, barcode U0001457), Bangka.
Plants including leaves pubescent, not scabrous. Leave blades lobed, lobes acute
or obtuse. Male flowers: pedicel subterete; receptacle-tube short-campanulate; sepals
mostly linear. Female flowers pedicel ± angular; receptacle-tube as in male flowers;
sepals ± linear or mostly leaf-like broadened at apex (long clawed). Fruit (depressed)
globose, elongated or flask-shaped, often shallowly furrowed (sulcate) from apex to
base of fruiting pedicel; fruiting pedicel angled, distinctly broadened at the transition
to the fruit. Seeds variable in size, 10 –15(– 20) mm long. — Plate 32c.
Distribution — Widely cultivated.
Note — Cucurbita moschata is the most heat-tolerating species and found most
frequently in the lowlands.
3. Cucurbita pepo L.
Cucurbita pepo L., Sp. Pl. ed. 1 (1753) 1010; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 622; Cogn.
in A.DC. & C.DC., Monogr. Phan. 3 (1881) 545; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge,
Laos & Vietnam 15 (1975) 105; C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001)
1547; A.Goldman, Complete Squash (2004) 103; W.J.de Wilde & Duyfjes, Sandakania 17 (2008
‘2007’) 61. — Type: Herb. LINN NO. 1151.4 (lecto LINN, designated by Keraudren (1975) (see
C.E.Jarvis, Order out of chaos (2007) 465)).
Plant, including leaves, rigid or scabrous, in certain cultivated varieties bushy and
not trailing. Leave blades often deeply acutely lobed, lobes often lobulate. Flowers:
pedicel subangular; receptacle-tube campanulate; sepals linear in male and female
flowers; corolla lobes slightly out-curved. Fruit small, medium or very large, smooth,
sometimes costate or verrucate; fruit pulp sometimes fibrous; fruiting pedicel rather
slender, woody, angular (sulcate), not or only little thickened at the transition to the
fruit. Seeds variable in size, 7– 25 mm long.
Distribution — Cultivated.
Note — Here belong the non-trailing suberect ‘bushy’ growing (Italian) ‘courgette’
or ‘zucchini’, nowadays a popular vegetable in Europe, and also the flat ‘scallop’ or
‘patison’. See further Jeffrey (2001), and Goldman (2004).
12. Cyclanthera
Cyclanthera Schrad., Index Sem. (Gottingen) (1831) 2; Cogn. in A.DC. & C.DC., Monogr. Phan. 3
(1881) 822; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 306; Keraudren in Aubrév. & J.-F.Leroy,
Fl. Cambodge, Laos & Vietnam 15 (1967) 19. — Type species: Cyclanthera pedata Schrad.
5 mm
g' g
2 cm
c
1 mm
h
d
0.5 mm
3 mm
3 mm 2 mm f
e
b
Fig. 17. Cyclanthera brachystachya (Ser.) Cogn. a. Portion of flowering shoot; b. node with female
flower with co-axillary a male inflorescence; c. male flower, the perianth consisting of the broad
hypanthium and broadly triangular petals, sepals absent; d. circular synandrium, opening by circum-
ferencial slit; e. female flower; f. female perianth, showing cushion-shaped sessile stigma; g, g'. fruit
from outside and opened respectively; h. seed (all: De Wilde & Duyfjes 21654).
Annual or perennial herbaceous climbers, glabrescent; monoecious; cultivated or

running wild. Probract absent. Tendrils unbranched or 2- or more-branched. Leaves
simple, unlobed or deeply lobed to almost foliolate. Flowers small, pale greenish yellow,
petals free or hardly connate at base. Male inflorescences racemose or paniculate. Male
flowers: receptacle-tube small, shallow; sepals small, subulate or linear; stamens united
a short central column, anthers connate into a head or into a horizontal (peltate) ring,
opening with a horizontal slit; disc (pistillode) absent. Female flowers solitary or one
co-axillary with male inflorescence; perianth as in male flowers; ovary narrowly ovoid
or ellipsoid, somewhat oblique, rostrate, style short, stigma hemi-spherical, ovules 1 to
many per locule; staminodes absent. Fruit oblique, ovoid(-oblong), various of shape,
not juicy, glabrous or mostly with soft spines, 1- or more-locular, 5- to many-seeded,
when ripe elastically opening and jetting the seeds away, leaving central column and
placenta. Seeds few or several, compressed, angular, often toothed at the end(s), faces
smooth or rough, margin distinct (wing-like), edge irregularly coarsely toothed.
Distribution — An American genus with some 40 species; in Asia 2 species intro-
duced as vegetable; in Malesia 1 species.
1. Cyclanthera brachystachya (Ser.) Cogn.

Cyclanthera brachystachya (Ser.) Cogn., Diag. Cucurb. Nouv. 2 (1877) 64; in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 842; C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1555; W.J.de
Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’) 62. — Elaterium brachystachium Ser. in DC, Prodr.
3 (1828) 310; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 842. — Type: based on a figure
in Moçino & Sessé, Fl. Mex. Icon. t. 38, f. F, Mexico (unpubl.).
Cyclanthera explodens Naudin, Ann. Sci. Nat., Bot., Sér. 4 (1859) 160; Backer in Backer & Bakh.f.,
Fl. Java 1 (1964) 306. — Type: plant cultivated at P, originating from New Grenade, Mexico.
Vigorous but slender climber to 5 m long, lanate or (finely) hairy on the nodes, later
scabrous or glabrescent. Tendrils (1- or) 2-branched. Leaves: petiole 1– 5 cm long; blade
triangular or ovate in outline, 6 – 9 cm long, 3- or 5-lobed up to halfway deep, lobes
triangular, acute, margin finely dentate. Male inflorescences condensed 10 – 20-flowered
racemes or few-branched panicles, 1– 2.5 cm long, shorter than the leaves. Male flowers:
pedicel 1– 3 mm long; receptacle-tube c. 1.5 mm diam.; sepals less than 1 mm long, or
± absent; petals green-yellow, triangular, c. 1 by 1 mm, finely papillate; anther circular,
horizontal, c. 0.5 mm diam., opening with a circular slit. Female flowers: pedicel 3 –7
mm long; ovary obliquely ovoid, style short, stigma broadly rounded. Fruit ripening
green, 2 – 3 cm long, ovoid-reniform, with soft spines; fruiting pedicel 1– 3 cm long.
Seeds c. 8 per fruit, blackish brown, 9 –14 by 6 – 8 mm, ± 5-angled, 3- (or 5-)toothed at
base, 3-toothed at apex. — Fig. 17; Plate 6a.
Distribution — Tropical America; in Malesia: Java, locally cultivated for the young
shoots used as a vegetable; in Cibodas Botanical Garden running wild.
Note — A second species, C. pedata Schrad., with deeply pedately lobed leaves,
usually appearing as pedately 5 –7-foliolate, is widely distributed in Central and South
America, and is introduced and locally commonly cultivated as a vegetable in mountain-
ous areas in the southern Himalayas of India and in China. The species can be expected
in the future in the Malesian area. It is mentioned (PROSEA 8 (1994) 288) as occasion-
ally cultivated in the Old World tropics, including Malaysia.
13. DIPLOCYCLOS
Diplocyclos (Endl.) T.Post & Kuntze, Lex. Gen. Phan. (1903) 178; corr. C.Jeffrey, Kew Bull. 15
(1962) 354. — Bryonia L. sect. Diplocyclos Endl., Prodr. Fl. Norfolk. (1833) 68. — Type species:
Bryonia affinis Endl.
Bryonia L. sect. Bryonopsis Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 922, p.p. — Bryonopsis (Blume)
Hassk., Cat. Hort. Bogor. (1844) 188, p.p.; Naudin, Ann. Sci. Nat., Bot., Sér. 4, 18 (1862) 193, nom.
inval., (not Bryonopsis Arn. (1841) = Kedrostis).
Ilocania Merr., Philipp. J. Sci., C 13 (1918) 65. — Type species: Ilocania pedata Merr. (= Diplocyclos
palmatus (L.) C.Jeffrey).
Subannual or perennial climbers; monoecious. Probract present. Tendrils 2-branched.

Leaves: blade simple, (deeply) palmately lobed. Inflorescences (from D. palmatus):
male flowers few to many in fascicle-like short racemes, female flowers 1– 5 co-axillary
with male flowers, or in fascicle-like short racemes, or solitary; flowers pale green-
ish (creamy-)yellow. Male flowers: receptacle-tube cup-shaped, narrowed to the base;
sepals small, linear; corolla of partly fused petals, lobes entire; stamens 3, inserted in
upper part of the receptacle-tube, anthers two 2-thecous, one 1-thecous, anthers free
but coherent, thecae plicate, connective broad; disc low, annular or indistinct. Female
flowers: perianth as male flowers; ovary (broadly) ovoid or subglobose, ovules few,
horizontal, style 3-armed, stigmas ± 2-lobed, papillose; staminodes 3, elongate; disc
inconspicuous. Fruit berry-like, solitary or few-fascicled, broadly ovoid or ellipsoid or
subglobose, shortly rostrate or not, exocarp thin. Seeds tumid, pyriform with strongly
convex faces, margin thick, edge grooved.
Distribution — A genus of c. 5 species in the Old World, all in Africa, of which 1
species also widely distributed in Austral-Asia.
1. Diplocyclos palmatus (L.) C.Jeffrey

Diplocyclos palmatus (L.) C.Jeffrey, Kew Bull. 15 (1962) 352; Keraudren in Aubrév. & J.-F.Leroy, Fl.
Camb., Laos, Viêt-Nam 15 (1975) 109, pl. 11: 4–7; I.Telford, Fl. Australia 8 (1982) 178; W.J.de
Wilde & Duyfjes, Fl. Thailand 9, 4 (2008) 432. — Bryonia palmata L., Sp. Pl. ed. 1 (1753) 1012.
— Type: Herb. Hermann 2: 58, No. 353 (lecto BM, barcode BM000621700, designated by Jeffrey
(1962)), Sri Lanka.
Bryonia affinis Endl., Prodr. Fl. Norfolk. (1833) 68 (no. 125). — Bryonopsis affinis (Endl.) Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 479. — Type: Bauer s.n. (holo W), Norfolk Is.
Bryonia pedata Hassk., Flora, Beibl. (1842) 32. — Bryonopsis pedata (Hassk.) Hassk., Cat. Hort. Bot.
Bogor. (1844) 189. — Type: Java, not further indicated.
Zehneria erythrocarpa F.Muell., Hooker’s J. Bot. Kew Gard. Misc. 8 (1856) 51. — Bryonopsis lac
iniosa (L.) Naudin forma erythrocarpa (F.Muell.) M.Mizush., J. Jap. Bot. 41, 9 (1966) 259. — Type:
Mueller s.n. (K), Australia.
Bryonopsis erythrocarpa Naudin, Ann. Sci. Nat., Bot., Sér. 4, 18 (1862) 194. — Bryonopsis laciniosa
(L.) Naudin var. erythrocarpa (Naudin) Naudin, Ill. Hort. 12 (1865) pl. 431; Cogn. in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 478. — Type: a plant cultivated at P from seeds imported from
India.
Bryonia laciniosa auct. non L.: Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 927; Naudin, Ann. Sci. Nat., Bot.,
Sér. 4, 12 (1859) 139. — Bryonopsis laciniosa auct. non (L.) Naudin: Naudin, Ann. Sci. Nat., Bot.,
Sér. 4, 18 (1862) 195, p.p., excl. type; Ann. Sci. Nat., Bot., Sér. 5, 6 (1866) 30; Cogn. in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 477; Cogn. & Harms in Engl., Pflanzenr. 88, 4.275.2 (1924) 160;
Craib, Fl. Siam. (1931) 762; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 301. (Bryonia laciniosa
L. = Cayaponia laciniosa (L.) C.Jeffrey.)
Bryonia quinquefolia Noronha, Verh. Batav. Genootsch. Kunsten 5 (1790) 48, nom. nud.
Climber 2 –7 m tall, perennial, old plants with tuberous rootstock, leafy stem 1.5 – 3
(– 4) mm diam., glabrous except few hairs in young parts. Probract membranous or
fleshy, obovate, 2 – 5 mm long, ± concave, wrinkled when dry. Tendrils 10(– 20) cm
long, branched at or below the middle. Leaves: petiole 2 –7 cm long, sometimes towards
apex with course stiff upward directed hairs; blade membranous, circular in outline,
deeply (3 –)5(–7) palmately lobed nearly to the base, 4 –16 cm diam., glabrous, except
for few hairs on veins on lower surface, glands small, usually a few close to the inser-
tion of the petiole, cystoliths generally not apparent, lobes (narrowly) elliptic, to 11 cm
long, margin ± dentate-undulate or entire, minutely dentate-mucronate. Inflorescences
up to 10 male flowers of various stage of development, mostly accompanied by up to
5 female flowers, or a few female flowers without male flowers. Male flowers: pedi-
cel 2 – 20 mm long; receptacle-tube 2 – 4 mm long and wide, glabrous; sepals linear,
0.5 –1(– 2) mm long; corolla 5 –7(– 9) mm long, tube c. 2 mm long, lobes ± ovate, 3 – 5
mm long, obtuse, apex ± mucronate; filaments inserted towards the throat of the recep-
tacle-tube, (0.5 –)1–1.5 mm long, glabrous or slightly hairy at base, anthers 2 – 3 mm
diameter. Female flowers: pedicel 1– 5 mm long; ovary ovoid, 4 – 5 by 2.5 – 3.5 mm,
often whitish striped; corolla lobes c. 5 mm long; style 1.5(– 2) mm long, style arms
1– 2(– 3) mm long, stigmas stout; staminodes 1(– 3) mm long. Fruit 1– 5 in a cluster at
the nodes, ripening bright red with (usually) pure white bands or rows of white blotches,
1.5 – 2(– 2.5) cm diam.; fruiting pedicel 0.1– 0.5 cm long. Seeds c. 12 or less, grey or
pale brown, obovoid, 5(– 8) by 2.5 – 3 by c. 4 mm, narrowed to the base, faces much
protruding, smooth, margin broad.
Distribution — Widely distributed in Africa and SE Asia, from India and South
China (not recorded for Yunnan), Japan (Ryukyu Is.) through Indochina south-east to
North and NE Australia and western Pacific, incl. Norfolk Is.; in Malesia: Sumatra,
Java, Philippines, Sulawesi, Lesser Sunda Islands, Moluccas (Buru), and New Guinea
(Papua New Guinea); not yet recorded from West Papua.
Habitat & Ecology — Open places and fringes of primary and secondary forests,
along stream banks, in old gardens, thickets, and grassland; on limestone as well as on
granite bedrock; from sea level to 1800 m altitude.
Notes — 1. Telford (1982) remarks that plants from eastern Malesia, the Pacific and
Australia, with larger ellipsoid fruits (Bryonia affinis) may be subspecifically distinct.
2. Cooked shoots are eaten in Papua New Guinea and cooked young fruits in Su-
lawesi (PROSEA 8 (1994) 290).
1a. Leaf blade deeply lobed, ± to 1/5 deep; middle leaf lobe 20 mm wide or more . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. var. palmatus
b. Leaf blade deeply lobed, nearly to the base; middle leaf lobe 2 –7 mm wide . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. pedata
b
2 cm 3 mm
e
c
2 mm
2 mm
g f d
Fig. 18. Diplocyclos palmatus (L.) C.Jeffrey var. palmatus. a. Portion of twig with male and female
flower buds on the nodes; b. node with male and female flower buds; c. male flower; d. ditto, opened;
e. stamens; f. (immature) female flower; g. ditto, opened (a, b, f, g: De Wilde & Duyfjes 21653; c – e:
De Wilde & Duyfjes 21707).
a. var. palmatus
Leaf blade deeply lobed, ± to 1/5 deep; middle leaf lobe 20 mm wide or more. —
Fig. 18, 19; Plate 8.
Distribution — As the species
c
2 cm
2 cm 1 mm
a b
Fig. 19. Diplocyclos palmatus (L.) C. Jeffrey var. palmatus. a. Node with infructescence; b. seed;
c. seedling; d. leaf, showing small glands near the blade base (a, b: De Wilde & Duyfjes 21653; c, d:
De Wilde & Duyfjes 21707).
Fig. 20. Diplocyclos palmatus (L.) C. Jeffrey var. pedata (Merr.)

W.J.de Wilde & Duyfjes. Node with deeply incised leaf blade
(Ramos BS 27490).
2 cm
b. var. pedata (Merr.) W.J.de Wilde & Duyfjes

Diplocyclos palmatus (L.) C.Jeffrey var. pedata (Merr.) W.J.de Wilde & Duyfjes, Reinwardtia 12
(2009) 405, f. 1. — Ilocania pedata Merr., Philipp. J. Sci., C 13 (1918) 65. — Type: Ramos BS
27552 (lecto US, designated by De Wilde & Duyfjes, Reinwardtia 12 (2008) 268), Ilocos Norte,
Bangui.
Leaf blade deeply lobed, nearly to the base; middle leaf lobe 2 –7 mm wide. —
Fig. 20.
Distribution — Philippines (Ilocos Norte).
14. GOMPHOGYNE
Gomphogyne Griff., Account Bot. Coll. Cantor (1845) 26; Hook.f. in Benth. & Hook.f., Gen. Pl. 1
(1867) 838; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 632; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 923; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15
(1975) 22; W.J.de Wilde & Duyfjes, Thai Forest Bull., Bot. 35 (2007) 50; Fl. Thailand 9, 4 (2008)
435. — Type species: Gomphogyne cissiformis Griff.
Delicate or slender herbaceous climbers, 0.5 – 5 m long, annual or biennial, dioecious;

roots fibrous, without tuberous rootstock. Probract absent. Tendrils 2-branched at apex.
Leaves: blade pedately foliolate (or simple, not in Malesia), ovate or subcircular in out-
line, membranous, leaflets up to 9, petiolulate, lateral ones usually smaller, elliptic or
narrowly elliptic, base acute, margin finely or coarsely serrate-dentate, teeth mucronu-
late, apex acute or acute-acuminate; cystoliths absent or inconspicuous. Flowers white,

small; sepals and petals free. Male inflorescences usually lateral, few- or many-flow-
ered, paniculate with ultimate branches fine, raceme-like; minute linear bracts and lower
portion of pedicels persistent. Female inflorescences few-branched or unbranched, with
1– 3 flowers on a slender peduncle; pedicel slender, usually with 1 or 2 small, simple
and subopposite tendrils close to the flower. Male flowers: pedicel articulate at or below
halfway; buds with subacute apex; corolla rotate; receptacle narrow, flat or shallow; pet-
als imbricate in bud; stamens 5, inserted in or near the centre of the receptacle, subpatent,
anthers small, 1-theous, opening extrorse; disc inconspicuous or absent. Female flowers:
ovary cylindrical-clavate, 1- or 3-locular, ovules 1 to several per placenta, pendulous,
styles 3, free, stigma 2-lobed; staminodes absent. Fruit few or solitary, ripening green,
small or medium-sized, capsular, cylindrical-clavate, longitudinally striate or ribbed, or
irregular-veined, apex truncate, 3-valvate, with three horn-like processes partly formed
by the styles at the angles. Seeds up to 12, imbricately arranged in 3 rows, (narrowly)
elliptic in outline, compressed, tubercled or scrobiculate, not winged or the margin with
a membranous or corky wing either only at the end(s) or all around.
Distribution — A genus of six species in SE Asia; in Malesia 1 species.
1. Gomphogyne peekelii W.J.de Wilde & Duyfjes

Gomphogyne peekelii W.J.de Wilde & Duyfjes, Thai Forest Bull., Bot. 35 (2007) 62, f. 6, map 1. —
Type: Peekel 118 (holo BO), Papua New Guinea, New Ireland, Neu-Mecklenburg, Ugana Distr.
Gynostemma pedatum auct. non Blume: Peekel, Fl. Bismarck Archip., translated by E.E.Henty (1984)
551, f. 878.
Climber 1– 4 m long; leafy stem 1– 2 mm diam., subglabrous. Leaves: petiole 2.5 –7

cm long; petiolules (0.2 –)0.3 – 0.6 cm long; blade 3 – 5-foliolate, i.e. the lateral leaflets
of 3-foliolate blade either deeply lobed or 2-foliolate, 5 –14 cm diam.; leaflets narrowly
or broadly elliptic, 3 – 9 cm long, pinnately veined, glabrous (hairy veins in Bismarck
Archipelago). Male inflorescences 3 – 9 cm long, loose, once or twice branched with
2 – 5 lateral branches from the rachis, glabrous or finely hairy, few-flowered; peduncle
1.5 – 3 cm long; rachis straight or somewhat zigzag, 3 – 5 cm long, lateral branches 1(– 2)
cm long, ultimate spike-like racemes 3 – 5 mm long, set with bracts and c. 0.5 mm long
basal portions of pedicels; bracts 0.5 – 3 mm long. Male flowers: pedicel 2.5 – 3.5 mm
long, sparsely minutely hairy; perianth 3 – 4 mm diam.; receptacle shallow, 0.1– 0.2 by
0.5 –1 mm, sparsely hairy outside; sepals 1–1.5 mm long; petals ovate, elliptic or ob-
long, 1.5 – 2 mm long, subacute; filaments 0.5 – 0.6 mm long, glabrous; anthers ovoid or
ellipsoid, c. 0.3 mm diameter. Female inflorescences: few-flowered; peduncle c. 3 cm
long. Female flowers not seen. Fruit 1 or 2, ripening green, 5 – 5.5 cm long, with (6 –)9
rib-like veins, sharply raised, in-between with coarse reticulate venation, base tapered,
at apex c. 1.5 cm wide, apical horns 2 – 3 mm long, somewhat out-curved; fruiting
pedicel c. 1 cm long, with 1 or 2 tendrils c. 2 cm long. Seeds c. 12(?), pale, compressed,
elliptic, c. 6 –7 by 3.5 mm, very finely tubercled, margin absent, edge entire, with a
corky wing all around the seed, 2 – 4 mm broad. — Fig. 21.
Distribution — Moluccas (Seram), New Guinea (West Papua (Vogelkop), Papua
New Guinea (East New Britain (Gazelle Peninsula), New Ireland)).
3 mm
2 mm 1 mm
g
2 mm
1 mm
c 2 cm g
1 mm
d e
Fig. 21. Gomphogyne peekelii W.J.de Wilde & Duyfjes. a. Portion of branch with male inflorescences;
b. detail of male inflorescence; c – e. male flowers; f. fruit; g. seeds (a, b: Vink BW 11487; c: F. & M.
Panoff 361; d, e: Van Royen & Sleumer 6818; f, g: Peekel 118, type).
Ecology — Primary and secondary forests, along streams, in disturbed places and

abandoned gardens; sandy and damp loamy soil; from sea level to c. 700 m altitude;
fruiting in May.
15. GYMNOPETALUM
Gymnopetalum Arn., Madras J. Lit. Sci. 12 (1840) 52; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879)
611; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 387; Ridl., Fl. Malay Penin. 1 (1922) 846;
Backer in Backer & Bakh.f., Fl. Java 1 (1964) 302; W.J.de Wilde & Duyfjes, Blumea 51 (2006)
282, Fl. Thailand 9, 4 (2008) 442; Kocyan et al., Mol. Phyl. Evol. 44 (2007) 553. — Lectotype
(Pfeiffer, Nom. 1 (1873) 1526): Gymnopetalum tubiflorum (Wight & Arn.) Cogn. (Bryonia tubi
florum Wight & Arn.).
Annual or perennial, climbing or widely-creeping herbs, frequently rooting at the

nodes, subglabrous or (densely) hairy, leafy stem (1–)2 mm diam.; monoecious. Pro
bract present or absent. Tendrils unbranched or 2-branched. Leaves: blade simple,
unlobed or lobed, margin sparsely finely dentate; glands not obvious. Flowers: petals
white, free, margin entire or short-fimbriate. Inflorescences in male: flowers either soli-
tary (or 2) on the nodes, long-pedicelled, or (co-axillary) in an erect several-flowered
bracteate raceme, with flowers sessile or short-pedicelled; in female: flowers solitary.
Male flowers: in bud folded into an elongate body, receptacle elongate, broadened in
upper half and slightly constricted at the throat; stamens 3, filaments inserted about
halfway the receptacle tube, free, short; anthers two 2-thecous, one 1-thecous, thecae
plicate, united into an elongate synandrium, included; disc consisting of linear ribs ad-
nate to the base of the tube, or of 3 lobes. Female flowers: ovary with many horizontal
ovules; perianth as in male; style long, stigmas 3 (or single, deeply 3-lobed), included;
disc not obvious; staminodes minute or absent. Fruit ripening orange or red, globose or
ovoid-oblong, (2 –)3 – 6 cm long, smooth or (low) ribbed, mesocarp yellow, soft-canose,
pulp greenish black. Seeds numerous, ± compressed, not or little ornamented, margin
present or absent, edge entire.
Distribution — A genus of 4 species in South India, Sri Lanka, North India, South
China, east to East Malesia; in Malesia 3 species.
Note — The type species of Gymnopetalum is G. tubiflorum from Sri Lanka. Cogni-
aux, 1881, divided the genus into two sections: sect. Tripodanthera (Roem.) Cogn. to
which the three Malesian species belong and sect. Gymnopetalum. Based on molecular
data Gymnopetalum is nested within Trichosanthes (Kocyan et al, 2007) with the Male-
sian species wide apart from the single species G. tubiflorum.
KEY TO THE SPECIES
1a. Leaves finely bullate, densely hairy beneath, at least on the veins. Fruit ellipsoid
or globose, rounded at apex, not ribbed. — Widespread . . . . . . . . . 3. G. scabrum
b. Leaves not bullate, thinly hairy or subglabrous beneath. Fruit ellipsoid-fusiform,
narrowed at apex, distinctly or faintly ribbed, or not ribbed . . . . . . . . . . . . . . . . . 2
2a. Male bracts cuneate or rounded at base, sessile, without glands. Sepals entire
or rarely ± lobed. Male flowers in raceme pedicellate, pedicels persistent. Fruit
ribbed. — Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. G. chinense
2 cm
f
g
3 mm 1 mm
3 mm 3 mm
3 mm
d
b
c
Fig. 22. Gymnopetalum chinense (Lour.) Merr. a. Node with male inflorescence, note persistent pedicel
of fallen co-axillary male flower; b. apex of male raceme; c. male flower, opened, showing 3-lobed
disc at the base of the receptacle-tube; d. unfolded petal; e. female flower bud, opened, showing disc
at the base of the receptacle-tube; f. node with mature fruit; g. seed (a, f, g: De Wilde & Duyfjes 21719;
b – e: De Wilde & Duyfjes 21722).
b. Male bracts narrowly cuneate at base, sub sessile or short-stipitate, with scat-
tered glands below. Sepals deeply lobed. Male flowers in raceme sessile. Fruit not
ribbed. — East Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. G. orientale
1. Gymnopetalum chinense (Lour.) Merr.

Gymnopetalum chinense (Lour.) Merr., Philipp. J. Sci. (1919) 256; W.J.de Wilde & Duyfjes, Blumea
51 (2006) 283; Reinwardtia 12 (2008) 268, Fl. Thailand 9, 4 (2008) 443. — Evonymus chinensis
Lour., Fl. Cochinch. (1790) 156. — Type: Untraced. Neotype: Levine 1705 (holo A, designated by
De Wilde & Duyfjes, Reinwardtia (2008)), South China.
Bryonia cochinchinensis Lour., Fl. Cochinch. (1790) 595. — Gymnopetalum cochinchinense (Lour.)
Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40 (1871) 57; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2
(1879) 611; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 391; Ridl., Fl. Malay Penin. 1 (1922)
846; Craib, Fl. Siam. 1 (1931) 755; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 302. — Type:
Loureiro s.n. (BM), Vietnam.
Momordica surculata Noronha, Verh. Batav. Genootsch. Kunsten 4 (1790) 21, nom. inval.
Gymnopetalum quinquelobum Miq., Fl. Ned. Ind. 1, 1 (1856) 681. — Type: Horsfield s.n. (holo U,
barcode U0001464), Java, Soerakarta.
Trichosanthes laciniata Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 107. — Type: Curtis in
Ridley 8350 (holo K, not seen), Peninsular Malaysia, Langkawi.
Climbing or creeping herbs to 6 m long, finely scabrous and sparsely hairy or sub-
glabrous. Probract (minute, caducous or) absent. Tendrils unbranched. Leaves: petiole
2 – 5 cm long; blade membranous, ovate or triangular in outline, or ± 3- or 5-angled or
(deeply) lobed, 4 –12 cm diam., ± sparsely scabrous-pubescent on both surfaces, upper
surface not bullate, cystoliths usually faint. Male inflorescences: flowers solitary or in
erect racemes 10 – 25 cm long; peduncle 4 –12 cm long; bracts sessile, oblong, 15 – 20
mm long, (deeply) 2 – 5-lobed, without glands, base cuneate or rounded. Male flowers:
pedicel persistent, 30 –70 mm long in solitary flowers, 5 –15 mm long for flowers in the
raceme; receptacle tube narrow, widened in upper 1/3 where containing the synandrium,
25 – 35 by 3 – 4(– 5) mm at throat, outside and inside ± grey pubescent, throat yellow in-
side; sepals linear, rarely ± lobed, 6 – 9 mm long, spreading to recurved; petals obovate,
short-clawed, 20 – 30 by 12 –15 mm, indistinctly veined, somewhat hairy, yellow at very
base; stamens inserted 12 –15 mm below receptacle throat, filaments short, synandrium
9 –10 by c. 3.5 mm; disc short, 3-lobed. Female flowers: pedicel 5 – 20(– 40) mm long;
ovary narrowly ellipsoid, 10 –12 by 2.5 – 4 mm, pubescent, (faintly) 10-ribbed; style
18 – 20 mm long, stigmas 4 – 6 mm long, included; staminodes minute. Fruit broadly
fusiform, 2.5 – 5 by 1.5 – 3 cm, beaked mainly by receptacle remnant, glabrescent, sharp-
ly 10-ribbed; fruiting pedicel 1– 4 cm long. Seeds narrowly elliptic, 7– 8 by 2 – 3 by 1.5
mm, faces not ornamented, with marginal groove. — Fig. 22, 26a, a'; Plate 9.
Distribution — Widespread; north-eastern part of India, China, through Indochina;
in Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Philippines, Su-
lawesi, Lesser Sunda Islands east to Flores.
Ecology — Forest edges, clearings, scrub, in hedges and in open fields, on sandy
soil, to 500 m altitude; flowering and fruiting throughout the year.
Note — The flowers of G. chinense open at night and are wilting the following
morning.
2 cm
5 mm
b f d c
Fig. 23. Gymnopetalum orientale W.J.de Wilde & Duyfjes. a. Node with male inflorescence; b. apex
of male inflorescence; c. male flower, opened, note disc composed of line-shaped thickenings adnate
to the receptacle-tube; d. unfolded petal; e. node with female flower, and probract; f. female flower,
opened (a – d: Wieringa 1811; e, f: Schmutz 4301).
2. Gymnopetalum orientale W.J.de Wilde & Duyfjes

Gymnopetalum orientale W.J.de Wilde & Duyfjes, Blumea 51 (2006) 290, f. 5. — Type: De Wilde &
Duyfjes 21937 (holo L), Lombok.
Climber to 5 m long, minutely grey or brown hairy, subglabrescent; leafy stem c.

2 mm diameter. Probract absent or various in size, (ob)ovate, up to 1 cm long, ir-
regularly sharp-dentate, glandular. Tendrils (simple or) unequally 2-branched in lower
half. Leaves: petiole 2.5 –7(– 9) cm long; blade membranous, ovate or subcircular in
outline, 6 –14 cm diam., usually (3 –)5 sharp-angular or (deeply) lobed, petiole and veins
minutely scabrid-hairy on both surfaces, blade scabrous by cystoliths above. Inflores
cences: male flowers either solitary with pedicel 20 – 60 mm long, or usually a solitary
flower co-axillary with a later developing long-peduncled few- to 20-flowered raceme,
10 – 30 cm long, flowers (sub)sessile; bracts subpersistent or (late) caducous, rhomboid
or ovate in outline, 10 –15 by c. 10 mm, sharply 5(–7)-angular or -lobed, up to half its
length, glandular beneath, subsessile or to 5 mm stipitate; rachis 5 – 20 cm long, pedun-
cle 5 –13 cm long, c. 2 mm diam.; female flowers solitary. Male flowers (sub)glabrous
but petals minutely hairy especially on the veins; receptacle-tube 35 –70 mm long,
tapering, at throat 5 – 8 mm wide; sepals ± recurved, lanceolate or long-triangular, 5 –15
mm long, with few narrow lobes to 9 mm long; petals (narrowly) obovate, 20 – 30 mm
long, at base with some fringes to 5 mm long, margin crenulate-lobate, apex 1– 2 mm
mucronate; stamens inserted c. 15 mm below the throat in the receptacle-tube, filaments
2 – 3 mm long, synandrium 10 –12 by 2 – 4 mm; disc consisting of 3 elongate, acute,
carnose bodies 3 –15(– 20) mm long, adnate to the basal portion of the receptacle-tube;
pistillode absent. Female flowers: pedicel 10 – 20 mm long; ovary narrowly ellipsoid,
7–16 by 2 – 3 mm, harshly minutely hairy (hairs c. 0.1 mm long), faintly 6 – 8-ribbed;
receptacle-tube c. 30 mm long; sepals (5 –)10 mm long, with a few sidelobes; petals c.
20 mm long; style 15 – 20 mm long, stigma deeply 3-lobed, c. 10 mm long; disc absent;
staminodes minute, inserted c. 5 mm above base of the tube. Fruit ellipsoid-fusiform,
5 – 6 by 3 – 4 cm, shortly beaked, harshly fine-hairy, glabrescent, sometimes with scat-
tered wartlets; fruiting pedicel 2 –7 cm long. Seeds numerous, long pear-shaped, rather
acute at one end, 7– 8 by 3.5 by 2.5 – 3 mm, faces shallowly verrucose-rugose, not
margined. — Fig. 23, 26d, d'.
Field-notes — Flowers opening between 23.30 h and 24.00 h, still expanded at 12.00
h, with a spicy odour, visited by midges. Fruits eaten by crows.
Distribution — Sulawesi, Lesser Sunda Islands (Bali, Lombok, Flores), Moluccas
(Seram, Babar Is.).
Habitat & Ecology — Climber in scrub and forest edges; damp sites; 25 –1500 m
altitude; flowering mainly June to January; fruiting in February.
Note — Gymnopetalum orientale resembles G. chinense, but the latter is less robust
in all parts. The disc in the male flowers, consisting of carnose elongate bodies adnate
to the receptacle-tube is similar to those in most of Trichosanthes. Gymnopetalum
orientale is reminiscent of Trichosanthes quinquangulata because of its leaf size and
shape, its lobed sepals and sometimes ± fringed petals.
3. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes

Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes, Reinwardtia 12 (2008) 268; Fl. Thailand
9, 4 (2008) 445. — Trichosanthes scabra Lour., Fl. Cochinch. (1790) 589. — Type: Untraced.
Neotype: Poilane 11322 (holo P, designated by De Wilde & Duyfjes, Reinwardtia (2008); iso L),
Vietnam, Annam.
Trichosanthes lucioniana Náves ex F.Villar, Fl. Filip., ed. 3 [F.M. Blanco] (1880) Nov. App.: 95, pl.
460 (see note); Merr., Sp. Blancoan. (1918) 13.
Gymnopetalum integrifolium (Roxb.) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40 (1871) 58;
C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 612; King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 67
(1898) 31; Ridl., Fl. Malay Penin. 1 (1922) 846; Cogn. & Harms in Engl., Pflanzenr. 88, 4.275.2
(1924) 179; W.J.de Wilde & Duyfjes, Blumea 51 (2006) 286. — Cucumis integrifolius (‘integri
folia’) Roxb., Fl. Ind. (1832) 724. — Type: Wallich Cat. 6730 (KW, IDC microfiche), Myanmar,
Ponlong.
Gymnopetalum leucostictum Miq., Fl. Ned. Ind. 1, 1 (1856) 680; Backer in Backer & Bakh.f., Fl. Java 1
(1964) 302. — Type: Junghuhn s.n. (lecto L, barcode L0589693, designated by De Wilde & Duyfjes
(2006); iso U, barcode U0001465), Java, Weltevreden.
Climbing or creeping herbs to 5 m long, stem (densely) long grey or brownish hairy.
Probract green-yellow, lanceolate, unlobed or (deeply) 2- or 3-lobed, (1–)1.5 – 2.5 cm
long, acute, sometimes caducous. Tendrils unbranched or unequally 2-branched near
the base. Leaves: petiole 1– 5 cm long; blade subcoriaceous, circular, or reniform, or
broadly ovate in outline, or 5-angular, 2 –11 cm diam., subglabrous above, densely
coarse-hairy beneath, at least on the veins, when fresh bullate above, cystoliths in older
leaves present, margin entire or finely dentate-mucronate or ± coarsely lobulate or
wavy-dentate, apex rounded or subacute, reticulation of smaller veins distinct beneath.
Male inflorescences: flowers solitary or in bracteate racemes; bracts sessile, 10 – 20 mm
long, with upper margin regularly, finely, densely laciniate or few-lobed, base cuneate.
Male flowers: pedicel persistent, 20 –120 mm long in solitary flowers, 10(– 20) mm
long for flowers in the raceme; receptacle tube (strongly) narrowed below insertion
of stamens, 15 – 20(– 30) by 6 –7 mm at the throat, outside and inside grey pubescent,
throat inside yellow; sepals narrowly triangular, lanceolate, entire or ± lobed, recurved,
(4 –)5 – 8 mm long; petals obovate, ± clawed, c. 20 by 15 mm, distinctly veined; stamens
inserted c. 10 mm below throat of the hypanthium-tube; filaments 2 – 2.5 mm long, ±
glabrous, synandrium 8 –12 mm long, 2 – 2.5 mm wide, apex of synandrium narrow,
flat, hairy, bright yellow when fresh; disc consisting of 3 short linear bodies. Female
flowers solitary; pedicel 10 – 30 mm long; ovary ellipsoid, 8 –10 by 6 –7 mm, long-pu-
bescent; receptacle tube cylindrical, c. 10 by 5 mm; style 7–10 mm long, stigma lobes
erect, c. 2 mm long; disc at base of the tube, minute or absent. Fruit short ellipsoid
or globose, (2 –)3 – 4 cm long, finely (sparsely) hairy, glabrescent, not ribbed; fruiting
pedicel 1– 3(– 5) cm long. Seeds narrowly elliptic, 6 – 9 by 2.5 – 3 by 1.5 – 2 mm, faces
small, almost smooth, demarcated by groove from broad, rounded margin.
Distribution — India, Sri Lanka, Myanmar, Thailand, South China, Laos, Cambo-
dia, Vietnam; in Malesia: Peninsular Malaysia (Penang, Perak, Selangor), Singapore,
Java, Philippines (Cebu), Sulawesi, and Lesser Sunda Islands (Bali)-; not known from
Sumatra and Borneo.
Ecology — Roadsides and disturbed places, to 500 m; flowering and fruiting through-
out the year.
1 cm
3 mm
c 3 mm
1 cm
e b
d
Fig. 24. a, b. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. pectinatum W.J.de Wilde
& Duyfjes. a. Portion of twig with solitary male flowers; b. node with male raceme. — c – e. Gym
nopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. scabrum. c, d. Male flower from outside and
opened respectively, note the disc composed of 3 line-shaped bodies; e. unfolded petal (a, b: De Wilde
& Duyfjes 21692, type; c – e: Murata, Fukuoka & Sukasdi J1421).
3 mm
g f
e 1 mm
2 cm
3 mm
c
Fig. 25. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. scabrum. a. Twig with one male
raceme, and one solitary male flower in bud; b. twig with solitary male flowers; c. male flower; d. male
flower opened; e. female flower; f. female flower opened; g. detail of female flower showing base of
style with traces of staminodes and disc (a: Phonsena, De Wilde & Duyfjes 3515; b: d’Alleizette s.n.,
barcode L0589688; c – g: De Wilde & Duyfjes 22269).
a a' b b'
2 cm 2 mm
c c' d d'
Fig. 26. Fruits and seeds of Gymnopetalum species. — a, a'. Gymnopetalum chinense (Lour.) Merr. —
b, b'. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. pectinatum W.J.de Wilde & Duyfjes.
— c, c'. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var scabrum. — d, d'. Gymnopetalum
orientale W.J.de Wilde & Duyfjes (a, a': De Wilde & Duyfjes 21719; b, b': De Wilde & Duyfjes 21693;
c, c': Pruesapan et al. KP-74; d, d': Verheijen 3819).
Note — The plate belonging with the name Trichosanthes lucioniana depicts Gym
nopetalum scabrum, although the deeply lobed leaves and elongate fruit are sugges-
tive of G. chinense. It clearly is not Trichosanthes cucumerina L. as stated by Merrill
(l.c.).
1a. Bracts of male raceme (irregularly) shallowly or deeply few- or many-lobed or ±

irregularly narrowly laciniate. Fruit globose . . . . . . . . . . . . . . . . . a. var. scabrum
b. Bracts of male raceme regularly ± fine-laciniate in upper half. Fruit ellipsoid. —
East Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. pectinatum
a. var. scabrum
Leaf blade subcircular in outline or 3 – 5-angular, margin entire or shallowly den-
tate. Bracts of male raceme variously rather few-lobed. Sepals narrow-triangular,
lanceolate, entire or shallowly few-lobed. Fruit (sub)globose, 2 – 3(– 4) cm diam.,
glabrescent. — Fig. 24c – e, 25, 26c, c'; Plate 10a – c.
b. var. pectinatum W.J.de Wilde & Duyfjes

Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. pectinatum W.J.de Wilde & Duyfjes,
Blumea 51, 2 (2006) 287, f. 1c, 4a, b. — Type: De Wilde & Duyfjes 21692 (holo L), East Java,
west of Pasuruan.
Leaf blade subcircular or angular in outline, margin (sub)entire. Bracts of male

raceme with upper margin regularly, finely, densely laciniate. Sepals subentire or
shallowly few-lobed. Fruit ellipsoid with obtuse apex, c. 4 cm long, finely hairy, not
ridged. — Fig. 24a, b, 26b, b'; Plate 10d.
Distribution — East Java; known only from the type.
Habitat & Ecology — Among grasses on sawah-dike on clay soil; at sea level.
Note — The status of var. pectinatum is unclear. The finely incised male bracts give
the plant a very distinct aspect. The ellipsoid fruit is unique and suggests a relationship
with G. chinense; the fruit in the latter species is ridged, with a tapered apex. Possibly
var. pectinatum is of hybrid origin.
16. gynostemma
Gynostemma Blume, Bijdr. Fl. Ned. Ind. 15 (1825) 23; Hook.f. in Benth. & Hook.f., Gen. Pl. 1 (1867)
839; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 633; Cogn. in A.DC. & C.DC., Monogr. Phan. 3
(1881) 912; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 305; W.J.de Wilde & Duyfjes, Blumea
52 (2007) 264; Fl. Thailand 9, 4 (2008) 447. — Pestalozzia Zoll. & Moritzi, Syst. Verz. Zoll. (1846)
31, nom. superfl. — Type species: Gynostemma pedatum Blume.
Small or medium climbers to 2 m long, main branches subherbaceous or soft woody,

leafy stem 1– 2 mm diam., glabrous or finely hairy, roots tuberous or not; dioecious or
monoecious. Probract absent. Tendrils 2-branched at apex. Leaves: petiole long; leaf-
lets petioluled; blade simple, not lobed or subpedately 3- or 5- (or 7-) foliolate, green
on drying, margin dentate, cystoliths not obvious. Flowers (greenish )white, small,
rotate; buds globose; sepals 5, free; petals 5, free, imbricate in bud, long-triangular,
sometimes ± erose towards apex; disc inconspicuous. Male inflorescences paniculate,
2 – 4-times branched, few- or many-flowered, the ultimate branches slender with few
or several flowers ± clustered in minute short racemes; bracts small, linear, glabrous
or finely hairy. Male flowers: pedicel short, articulate, basal part persistent; receptacle
narrow, flat or saucer-shaped; petals (long-)triangular or narrowly elliptic, apex acute or
(long-)acuminate; filaments united into a staminal column, at apex with 5 (sub)sessile
or short-stipitate anthers in a ring, anthers small, 1-thecous, short-ellipsoid, opening
with a vertical slit, extrorse. Female inflorescences as in male inflorescences but less
ramified and smaller. Female flowers as male flowers but somewhat larger; pedicel short
(or long); ovary subglobose, largely inferior, the inferior part not narrowed below the
perianth, (2- or) 3- or 5-locular, each locule with 2 hanging ovules; sepals and petals
inserted along broad, slightly convex apex of ovary; styles (2 or) 3 or 5, erect, free or
± contiguous, glabrous, each at apex with a 2-branched stigma; staminodes sometimes
present. Fruit ripening green, brown or purple, (sub)globose, less than 10 mm diam.,
with subpersistent perianth and styles (the latter becoming spaced and the perianth-
scars leaving a thin line around the fruit), either a dry berry, not dehiscent or ± capsular,
opening with 3 valves (not in Malesia). Seeds 1– 5, not much compressed, ovoid or
subtriangular in outline, faces verrucose or wrinkled, margin narrow or broad, with ±
square, grooved edge.
Distribution — A genus of about 10 species, occurring in Sri Lanka and in South
and NE India, ranging through China east to Japan and to New Guinea; in Malesia: 3
species.
Note — Although the androecium is characteristic for the genus Gynostemma, two
species of the genus Neoalsomitra, namely N. angustipetala (Craib) Hutch. (from Thai-
land) and N. hederifolia (Moluccas), have a more or less similar androecium.
KEY TO THE SPECIES
1a. Monoecious. Inflorescences few-flowered. Plant delicate. — Papua New Guinea;

subalpine; 2500 – 3200 m altitude . . . . . . . . . . . . . . . . . . . . . . . 2. G. papuanum
b. Dioecious. Inflorescences many-flowered. Plant stouter . . . . . . . . . . . . . . . . . . . 2
2a. Styles 5. — Eastern Papua New Guinea; montane; 1000 – 2500 m altitude . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. G. intermedium
b. Styles (2 or) 3. — Widespread, lowland and montane areas; in New Guinea 200 –
1000(–1800) m altitude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. G. pentaphyllum
1. Gynostemma intermedium W.J.de Wilde & Duyfjes

Gynostemma intermedium W.J.de Wilde & Duyfjes, Blumea 52 (2007) 265, f. 1a, b. — Type: Pullen
422, female flowers (holo L; iso CANB), Papua New Guinea, Eastern Highlands Province.
Gynostemma pentaphyllum auct. non (Thunb.) Makino: P.Royen, Alpine Fl. New Guinea (1982) 2008,
p.p., f. 613a.
Small climber or creeper, root not known, 1– 2 m long; leafy stem 1–1.5 mm diam.,
sparsely hairy, glabrescent; dioecious. Leaves: petiole 2 – 3.5 cm long; petiolules 0.5 –
1.5 cm long; blade membranous, (3 –)5 –7-foliolate, ovate or circular in outline, 3 – 6
cm diam., leaflets up to 5 by 2(– 2.5) cm, middle leaflet largest, with 5 –7 pairs of
rather faint lateral veins, upper surface sparsely hairy mainly on veins, lower surface
subglabrous, margin (shallowly) coarsely serrate. Male inflorescences many-flowered,
paniculate, (2 or) 3 times branched, 10 –15 by 5 –10 cm, ultimately with up to 5 flow-
ers in fascicles or short racemes up to 15 mm long; peduncle c. 3 cm long, finely
pubescent; bracts minute, less than 1 mm long. Male flowers: pedicel 2 – 3 mm long,
(sub)glabrous, articulation indistinct, at c. 1/3 from the apex; perianth glabrous, 4.5(– 5)
mm diam.; receptacle shallow, 0.6 – 0.9 mm wide; sepals ovate-narrowly elliptic, c. 1
by 0.5 mm; petals triangular-ovate, c. 2 by 1(–1.5) mm, entire, with narrowed apiculate
1 cm
1 mm
2 mm 1 cm
d
Fig. 27. a, b. Gynostemma intermedium W.J.de Wilde & Duyfjes. a. Habit of shoot with female in-
florescences; b. female flower. — c, d. Gynostemma papuanum W.J.de Wilde & Duyfjes. c. Habit of
fruiting shoot; d. fruit (a, b: Pullen 422, type); c, d: Stevens LAE 55693).
apex; staminal column (0.3 –)0.5 mm long, synandrium c. 0.5 mm diam., anthers sub-
sessile, subglobose, c. 0.2 mm diameter. Female inflorescences as in male, paniculate,
rather many-flowered, 1(– 3) times branched, 5 –10 cm long; peduncle 1.5 – 3 cm long,
(sparsely) hairy. Female flowers: pedicel 3 – 4 mm long, articulation not seen; ovary
(hemi-)globose, c. 1.5 mm diam., (sparsely) hairy or glabrous; perianth c. 4 mm diam.;
styles 5, each c. 1 mm long. Fruit few or up to 20 per infructescence, shiny, ripening
green or dark purple, globose, 0.6 –1(–1.2) cm diam., with 5 style-remnants and 5 (not
opening) valve-sutures, glabrous; fruiting pedicel c. 0.5 cm long. Seeds 3 or 4 per berry,
pale brown, rounded-triangular, c. 4 by 3.5 by 2 mm, with ± flattish apex, coarsely
sparsely warted or irregularly ridged, margin narrow. — Fig. 27a, b.
Distribution — New Guinea (Papua New Guinea (Madang, Eastern Highlands, and
Morobe Provinces)).
Habitat & Ecology — Climber on tree ferns; along stream sides and gullies in moun-
tain forest; 1500 – 2500 m altitude; flowering and fruiting throughout the year.
Note — Gynostemma intermedium occupies the altitudinal montane zone between
G. pentaphyllum of the lowlands and G. papuanum of the subalpine zone.
2. Gynostemma papuanum W.J.de Wilde & Duyfjes

Gynostemma papuanum W.J.de Wilde & Duyfjes, Blumea 52 (2007) 267, f. 1c, d. — Type: Vink 17364
(holo L), Papua New Guinea, Southern Highlands Province.
Gynostemma pentaphyllum auct. non (Thunb.) Makino: P.Royen, Alpine Fl. New Guinea (1982) 2008,
p.p., f. 613b–d.
Small climber, 0.5 –1 m long, root not known; leafy stem less than 1 mm diam.,
subglabrous or sparsely hairy; monoecious. Leaves: petiole 1.5 – 2.5 cm long; petiolules
0.2 – 0.4 cm long; blade filmy or membranous, (3- or) 5-foliolate, ovate in outline, 3 – 6
by 2.5 – 4.5 cm; the middle leaflet much longer than the lateral ones, with c. 5 pairs
of faint lateral veins, both surfaces subglabrous or sparsely hairy, margin shallowly
crenate-dentate. Inflorescences few-flowered, 1– 2(– 3) cm long, consisting of 1 (or 2)
female flowers and later developing 1– 3(– 5) male flowers, inflorescence solitary or
± fascicled, sometimes only female or only male flowers present in one inflorescence,
sometimes short peduncled inflorescences aggregated in raceme-like short shoots;
branches and pedicels hairy; peduncle 0.5 – 2 cm long; bracts narrow, acute, 1 mm long
or less. Male flowers: pedicel 2 – 5 mm long, glabrous or hairy, articulate at c. 1/3 from
the apex; perianth glabrous, c. 3 mm diam.; receptacle shallow, less than 0.5 mm wide;
sepals ovate-narrowly elliptic, 0.7(–1) mm long; petals ovate-narrowly elliptic, c. 1.5
mm long, entire or somewhat incised, long-acuminate in upper 1/3; staminal column c.
0.3 mm long, synandrium 0.3(– 0.4) mm diam., anthers sessile, subglobose, c. 0.2 mm
diameter. Female flowers: pedicel 2 – 5 mm long, articulate at apex; ovary (sub)hemi-
globose, c. 0.6 mm diam., glabrous; perianth 2.5 – 3 mm diam.; sepals c. 0.5 mm long;
petals ovate-narrowly elliptic, 1(–1.5) mm long, acuminate; styles 3 (rarely 4), with
stigma each c. 1 mm long. Fruit solitary, ripening light green, globose, 0.5 – 0.7 cm
diam., with persistent sepals and styles and with 3 (not opening) valve-sutures; fruiting
pedicel 0.4 – 0.6 cm long. Seeds 2 or 3 per berry, (pale) brown, ± triangular-ovate, 2.5 – 3
by 2 – 2.5 by 1 mm, faces irregularly low-warty, margin narrow. — Fig. 27c, d.
Distribution — New Guinea (Papua New Guinea (West Sepik, East Sepik, West-
ern Highlands (Kubor Range), Southern Highlands (Mt Giluwe), Eastern Highlands
(Chimbu), and Milne Bay (Mt Suckling) Provinces)).
Habitat & Ecology — Open places in low forest, mossy forest, gullies, low scrub,
grasslands, alpine shrubbery and in Nothofagus forest; also on limestone; (1600 –)2100 –
3500 m altitude; flowering and fruiting all year round.
3. Gynostemma pentaphyllum (Thunb.) Makino

Gynostemma pentaphyllum (Thunb.) Makino, Bot. Mag. (Tokyo) 16 (1902) 179.
For literature and synonyms see under the forms.
Herbaceous or subwoody climber, 2 – 8 m tall, root perennial but not(?) tuberous,

main stem to 1 cm thick, bark fissured, grey; leafy stem slender, 1(– 2) mm diam.,
glabrous or hairy, hairs grey or rarely brown; dioecious. Leaves: petiole 3 – 6 cm long;
petiolules 0.2 – 0.7 cm long; blade simple or foliolate, subcircular (or ovate) in outline,
4 –10(–17) cm diam.; leaflets 3 – 5(–7), ovate to narrowly elliptic, 3 –10 by 1– 5 cm,
outer leaflets smallest, 3 – 6(–7)-pinniveined, veins faint or distinct, margin (remotely)
finely (or coarsely) dentate, upper surface variously (appressed) hairy or subglabrous
(except veins), lower surface (sub)glabrous. Male inflorescences many-flowered, small
or large, raceme-like or 2 – 4-branched, broadly paniculate, 5 –15(– 30) cm long, lateral
branches to 10 cm long, hairy or subglabrous, ultimately with (1 or) 2 – 5 flowers in
fascicles (or in condensed racemes), finely hairy or subglabrous; peduncle 1–7 cm
long; bracts 1– 2(– 5) mm long. Male flowers: pedicel 1– 3(–7) mm long, articulate
in the middle, the basal part persistent after flowering; perianth (2.5 –)3 – 5(– 8) mm
diam.; receptacle shallowly hollowed, c. 0.5 mm wide; sepals narrowly elliptic-linear,
(0.5 –)1 mm long; petals triangular, (narrowly) elliptic, 1– 2(– 3.5) by 0.5(–1) mm, apex
acuminate or long-acuminate, adaxially usually ± hairy or papillose; staminal column
0.5(–1) mm long, synandrium 0.5(–1) mm diam., anthers subglobose, sessile or short-
stalked, c. 0.2 mm diameter. Female inflorescences as male inflorescences, but smaller.
Female flowers: pedicel 2 – 4 mm long, subapically articulate; perianth 2 – 4 mm diam.;
ovary subglobose, c. 1 mm diam., glabrous or sparsely hairy; styles (2 or) 3, with stigma
each 0.5 –1 mm long. Fruit green, ripening pale yellow or blackish, globose, 0.5 – 0.8
cm diam., glabrous or (sparsely) hairy; fruiting pedicel 0.2 – 0.5 cm long. Seeds usu-
ally 2, subcircular, triangular or cordate in outline, 3 – 4 mm diam., little or moderately
compressed, faces irregularly verrucose or wrinkled, margin broad or narrow.
Distribution — Widespread, as for the genus; India, Sri Lanka, east through China,
to Japan and Korea, south-east through Indochina and Thailand; common throughout
Malesia.
Habitat & Ecology — In a variety of habitats: damp or not too dry places in scrub
and forest, often near water courses, also on limestone; at 200 – 2400 m altitude; pos-
sible also at sea level in the lowlands. Flowering and fruiting all through the year, but
less frequent in April and May.
Notes — 1. Densely hairy plants with 3-foliolate leaves from Myanmar, China, and
Thailand, known as G. burmanica Chakrav. were placed in the synonymy of G. penta
phyllum by De Wilde & Duyfjes (2007, 2008), but the former species can probably
better be regarded as a species of its own.
1 mm
2 cm
a
1 mm
1 mm 2 mm
g
f c
Fig. 28. Gynostemma pentaphyllum (Thunb.) Makino forma pentaphyllum. a. Habit of branch with
female inflorescences; b. detail of female inflorescence; c. female flower; d. gynoecium; e. node with
infructescence; f. fruit; g. seed (a – d: Kock 8; e – g: Soegandiredja 281).
2 cm
f 2 mm
0.5 mm
1 mm
c d e
Fig. 29. a. Gynostemma pentaphyllum (Thunb.) Makino forma simplicifolium (Blume) W.J.de Wilde
& Duyfjes. Habit of male inflorescence. — b – f. Gynostemma pentaphyllum (Thunb.) Makino forma
pentaphyllum. b. Detail of male inflorescence; c. male bud; d, e. male flowers; f. androecium seen from
above (a: Van Ooststroom 14084; b – f: De Wilde & Duyfjes 21779).
2. Gynostemma pentaphyllum is very variable, comprising various not sharply de-

marcated local forms, or ecotypes. To acknowledge some of the variation, the following
forms are recognized:
a. forma pentaphyllum
Gynostemma pentaphyllum (Thunb.) Makino, Bot. Mag. (Tokyo) 16 (1902) 179; Backer in Backer &
Bakh.f., Fl. Java 1 (1964) 306; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam
15 (1975) 25, pl. 5: 1–7; W.J.de Wilde & Duyfjes, Blumea (2007) 268, f. 2b – f, 3; Fl. Thailand 9,
4 (2008) 450. — Vitis pentaphylla Thunb., Fl. Jap. (1784) 105. — Type: Thunberg 5858 (UPS, not
seen), Japan.
Gynostemma pedatum Blume, Bijdr. Fl. Ned. Ind. 15 (1825) 23; C.B.Clarke in Hook.f., Fl. Brit. Ind.
2 (1879) 633 (‘pedata’); Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 913; Craib, Fl. Siam.
(1931) 766. — Pestalozzia pedata Zoll. & Moritzi, Syst. Verz. Zoll. (1846) 31. — Zanonia pedata
(Blume) Miq., Fl. Ned. Ind. 1, 1 (1856) 683. — Type: Blume 1429 (lecto L, barcode L0588327,
designated by De Wilde & Duyfjes (2007); iso L (6 sheets)), Java, Tjanjor & Krawang.
Zanonia laxa Wall., Pl. Asiat. Rar. 2 (1831) 29. — Alsomitra laxa (Wall.) M.Roem., Fam. Nat. Syn.
Monogr. 2 (1846) 118. — Pestalozzia laxa (Wall.) Thwaites, Enum. Pl. Zeyl. 2 (1859) 124. — Gy
nostemma laxum (Wall.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 914; Craib, Fl. Siam.
(1931) 766; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 26.
— Type: Wallich 3727 (K; K-W), India, Silhet.
Plant as the species, not including the forms dasycarpum, fasciculare, and simplici
folium. The leaves of the typical form are 3- or 5- (or 7-)foliolate. — Fig. 28, 29b – f;
Plate 11a – c.
Distribution — Widespread; in Malesia: Sumatra, Peninsular Malaysia, rare in Bor-
neo, where it is replaced by forma dasycarpum, all over Java, Philippines, Lesser
Sunda Islands, Moluccas, and New Guinea; not known from Sulawesi; from sea level
to 1500(– 2000) m altitude.
Uses — The form is cultivated for its medicinal properties. Tea prepared from the
leaves, for instance, is used for reduction of blood pressure.
b. forma dasycarpum (C.Y.Wu) W.J.de Wilde & Duyfjes

Gynostemma pentaphyllum (Thunb.) Makino forma dasycarpum (C.Y.Wu) W.J.de Wilde & Duyfjes,
Blumea (2007) 269, pl. 1c. — Gynostemma pentaphyllum (Thunb.) Makino var. dasycarpum
C.Y.Wu, Acta Phytotax. Sin. 21 (1983) 362. — Type: Li 534 (holo KUN, not seen), China, Yun-
nan.
Gynostemma winkleri Cogn. in H.Winkl., Bot. Jahrb. Syst. 48 (1912) 118. — Type: Winkler 2757 (holo
B†; iso K, L), South Kalimantan, Batu Babi.
Plant as forma pentaphyllum, but fruit hairy.

Distribution — Myanmar, southern China, Thailand; in Malesia: all over Borneo.
Habitat & Ecology — Primary forest, shaded and open places, damp places, river-
banks, and hillsides; often on limestone, also on basalt bedrock; mostly in the lowlands
to 1650 m.
c. forma fasciculare W.J.de Wilde & Duyfjes

Gynostemma pentaphyllum (Thunb.) Makino forma fasciculare W.J.de Wilde & Duyfjes, Blumea 52
(2007) 270. — Type: Johansson, Nybom & Riebe 384 (holo BO; iso L, S), Central Sulawesi.
Plant as forma pentaphyllum, but male flowers long-pedicelled, pedicels 4(–7) mm

long, fascicled.
Distribution — Endemic to Central Sulawesi.
Habitat & Ecology — Primary and disturbed primary forest on alluvial soil; from
sea level to 2200 m.
Note — In Sulawesi the typical form pentaphyllum seems absent.
d. forma simplicifolium (Blume) W.J.de Wilde & Duyfjes

Gynostemma pentaphyllum (Thunb.) Makino forma simplicifolium (Blume) W.J.de Wilde & Duyfjes,
Blumea 52 (2007) 271, f. 2a. — Gynostemma simplicifolium Blume, Bijdr. Fl. Ned. Ind. 15 (1825)
24 (‘simplicifolia’); Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 915; Backer in Backer &
Bakh.f., Fl. Java 1 (1964) 306. — Type: Blume 1493 (lecto L, barcode L0588361, designated by
De Wilde & Duyfjes (2007); iso L), Java, Mt Krawang.
Plant as forma pentaphyllum, but leaves simple, occasionally with a few 2 (or 3)
foliolate leaves among the simple leaves. — Fig. 29a.
Distribution — India, Myanmar, China; in Malesia: Peninsular Malaysia (Pahang),
Borneo (Sarawak, Sabah), West Java, Philippines (Basilan).
Habitat & Ecology — Damp and shaded places in primary and old secondary forest,
along roadsides in logged forest and river beds; yellow sandy and reddish clay-lateritic
soils, also on limestone; 100 –1600 m altitude.
Note — Simple-leaved (not foliolate) forms have been found almost all over the
area of the species. Plants from the type locality in West Java have a relatively long
petiole, almost as long as the rather narrow leaf blade. In simple-leaved forms from
other localities the petiole is generally shorter.
17. HODGSONIA
Hodgsonia Hook.f. & Thomson, Proc. Linn. Soc. London 2 (‘1853’, 1854) 257; Cogn. in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 348; Hutch., Fam. Fl. Pl., Dicot. ed. 3 (1973) 300, f. 109; Ridl.,
Fl. Malay Penin. 1 (1922) 843; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 304; W.J.de Wilde
& Duyfjes, Blumea 46 (2001) 169; Fl. Thailand 9, 4 (2008) 454. — Type species: Hodgsonia het
eroclita (Roxb.) Hook.f. & Thomson (Trichosanthes heteroclita Roxb.).
Liana, to 30 m long, leafy shoots 4 –7 mm diam., almost glabrous; dioecious. Pro

bract c. 5 mm long, thorn-like, with glands on lower surface. Tendrils 2- or 3-branched.
Leaves: blade coriaceous, subcircular in outline, 15 – 25 cm diam., palmately (deeply)
3 – 5-lobed, scattered minute glands often present, margin entire, tertiary veining on
lower surface prominent. Flowers large, puberulous; corolla rotate; petals free, white,
long-fringed, the in bud exposed portion conspicuously veined. Male inflorescences
stout bracteate racemes, hairy; bracts elliptic or oblong, entire, 5 –10 mm long, glandular
on lower surface, often inserted on the pedicel, subdeciduous. Male flowers: receptacle-
tube elongate, widened towards or near the apex; calyx lobes minute, with few glands
on lower surface sometimes also on the tube; petals cuneate; stamens 3, filaments short,
free, inserted towards the apex of the receptacle-tube, anthers one 1-thecous, two 2-
thecous, united into a head, largely included, thecae linear, plicate, connective narrow,
not produced; disc consisting of 3 elongate parts, either free or largely adnate with basal
portion of the tube. Female flowers solitary, resembling male flowers; ovary subglo-
bose, 6-locular, ovules 6 or about 12, in each locule 1 or 2 (or 3) collaterally attached
at the bottom (rarely the apex) of the locules; style filiform, stigma large, obconical,
3-lobed, partly exserted; staminodes and disc absent. Fruit a large drupe, hard-skinned,
depressed globose, 12 – 25 cm diam., filled with firm pulp and 6 large, subovoid, deeply
veined pyrenes, each containing 1 or 2 (or 3) seeds. Seeds large.
Distribution — An Asian genus of 2 species ranging from East India and South China
through Indochina to Borneo and West Java; in Malesia 1 species.
Note — Hodgsonia is related to Trichosanthes L., which has similar flowers, but
differs in fruits containing many smaller horizontal seeds. The pollen of Hodgsonia is
different from Trichosanthes.
1. Hodgsonia macrocarpa (Blume) Cogn.

Hodgsonia macrocarpa (Blume) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 349; Backer in
Backer & Bakh.f., Fl. Java 1 (1964) 305; W.J.de Wilde & Duyfjes, Blumea 46 (2001) 174, f. 2a:
3 – 4. — Trichosanthes macrocarpa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 935. — Type: Blume s.n.,
(holo P), Java, Gunung Salak.
Trichosanthes hexasperma Blume, Bijdr. 15 (1826) 935; Ser. in DC., Prod. 3 (1828) 315; M.Roem.,
Fam. Nat. Syn. Monogr. 2 (1846) 95; Hassk., Pl. Jav. Rar. (1848) 192; Miq., Fl. Ned. Ind. 1, 1
(1856) 678. — Type: Blume s.n.(holo L, barcode L0589332; iso L, barcodes L0589331, L0589330),
Java, Gunung Salak.
Trichosanthes kadam Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 331. — Hodgsonia kadam (Miq.)
Greshoff, Bull. Kol. Mus. 30 (1904) 163, with figure; Lewkowitsch, Chem. Techn. Anal. Oils 2
(1914) 515 (with references to literature on medicinal and chemical properties, mainly of the oily
seed). — Type: Diepenhorst in Teysmann s.n. (2097 HB), (holo U, barcode U0001466; iso BO),
Sumatra (Priaman).
Hodgsonia capniocarpa Ridl.; Fl. Malay Penin. 1 (1922) 843. — Type: Ridley s.n. (lecto K, designated
by De Wilde & Duyfjes (2001)), Peninsular Malaysia, Pahang.
Habit as for the genus. Leaves: petiole 3 –7 cm long; blade usually 3-lobed, lower
surface puberulous or glabrescent, small glands several, especially towards the base,
margin entire. Male inflorescences 12 – 20(– 30) cm long, peduncle (2 –)10 – 20 cm long,
3 – 5 mm thick; flowers (5 –)10 – 20. Male flowers: pedicel 2 –10 mm long, bract ovate,
c. 5 mm long, inserted up to 10 mm from the base; receptacle-tube 50 –70 mm long,
basal part 30 – 45 mm by 3 – 4 mm, the tube above the middle widening into a dilated
section 20 – 25 mm long, 10(–15) mm wide at the throat; sepals 1(–2) mm long; petals
20 – 30 mm long, white, outside at base rusty puberulous, threads c. 50 mm long, white,
pendent(?), spiralling(?). Stamens inserted at about the middle of the receptacle-tube,
i.e. where widening, filaments (5 –)7 mm long; synandrium elongate, apex truncate,
10 –12 by 3.5 – 4 mm, included; disc parts narrowly elliptic, 4 – 6 mm long, free. Female
flowers: pedicel 3 – 4 cm long; receptacle-tube c.3 cm long (including c. 6 mm long
solid basal part which remains on top of the ovary after anthesis), gradually widening
to c. 7(–10) mm wide throat; style 15 – 20 mm long, stigma 13 –14 by 6 – 8 mm, lobes
Fig. 30. Hodgsonia macrocarpa (Blume) Cogn. Habit of

twig, with male inflorescence (De Wilde & Baya Busu
FRI 41401).
2 cm
truncate; ovary 10 –12 mm diam., with soft brown hairs c.1 mm long, pustules absent
(always?), ovules 6, one in each locule. Fruit ripening greyish green, 12 –18 cm diam.,
densely grey or brown hairy, glabrescent, sometimes with large, scattered, dark-col-
oured wart-like lenticels, not grooved; fruiting pedicel 4 – 8 cm long, 10 –15 mm wide.
Pyrenes subovoid, hardly compressed, 6 – 9 cm long, containing 1 seed, faces of pyrenes
smooth, margin absent, edge entire. — Fig. 30, 31; Plate 11d.
d
b c
5 cm
2 cm 1 cm
a e
Fig. 31. Hodgsonia macrocarpa (Blume) Cogn. a. Opened submature male flower bud, showing an-
droecium and disc-lobes at base of hypanthium tube; b. submature female bud; c. ditto, longitudinal
section, showing style and stigma; d, e. ovary, longitudinal and cross section, showing position of the
ovules. (a: De Wilde & Baya Busu FRI 41401; b – e: King’s Collector 4021).
Distribution — Peninsular Thailand and South Myanmar; in Malesia: Sumatra, Pe-

ninsular Malaysia, Borneo, and West Java.
Habitat & Ecology — Moist places in primary and disturbed forests, forest fringes,
and roadsides, mostly near riversides; 100 – 250 m altitude; flowering at night, flowers
fragrant; flowering February and April, fruiting December and April.
Uses — The roasted fatty seeds are edible; Whitmore KEP FRI 576 reported: “cotyle
dons edible after seed roasted and bitter skin removed”. The empty stone-seeds (pyrenes)
can be found on the forest floor, gnawed open by rodents. The ashes of burnt leaves are
used in healing wounds.
Note — The species is easily recognisable by its characteristic thorn-like probracts,
present also in sterile shoots on each node.
18. INDOMELOTHRIA
Indomelothria W.J.de Wilde & Duyfjes, Blumea 51 (2006) 5, f. 1b, 2a; t. 1; Fl. Thailand 9, 4 (2008)
458. — Type species: Indomelothria chlorocarpa W.J.de Wilde & Duyfjes
Climbers 1– 5 m long, (sub)perennial, leafy stem 1– 2 mm diam., green on drying;

monoecious. Probract absent. Tendrils unbranched. Leaves: blade simple, entire or
shortly hastately lobed, subpinniveined to palmiveined. Flowers 5(–10) mm diam.,

sepals minute, linear or subulate, ± patent; petals white (but see note on I. blumei),
free, obovate-elliptic, valvate in bud; receptacle-tube campanulate or urceolate. Male
inflorescence a slender peduncled lax raceme, sometimes co-axillary with 1 female
flower, pedicels persistent; bracts absent. Male flowers: sepals with or without adaxi-
ally a short spur; stamens 3, inserted close to the throat of the receptacle-tube, filaments
short, thick, anthers two 2-thecous, one 1-thecous, only included at base, thecae ±
lateral, straight or slightly curved, connective broad, not produced; disc free, at apex ±
3-parted. Female flowers solitary or co-axillary with male raceme; ovary narrowly el-
lipsoid, ± with slender neck, glabrous; stigma consisting of 3 short arms, each arm with
numerous, thread-like appendages; staminodes absent; disc a faintly 3-lobed annulus,
largely free from the receptacle-tube. Fruit solitary or co-axillary with male raceme,
ripenning green, narrowly ellipsoid, fusiform, 5 –7 cm long, glabrous, pulpy; dry exo-
carp membranous-cartilaginous, smooth. Seeds numerous, compressed, ovate-elliptic,
faces little convex, not sculptured, but with dense appressed hairs, margin absent, edge
entire, not winged.
Distribution — A genus of 2 species extending from South Myanmar and South
Thailand to West Malesia; in Malesia 2 species.
KEY TO THE SPECIES
1a. Leaf blade elliptic, usually two times longer than wide or longer. Male perianth
c. 4 mm diam., sepals without(?) an appendage. Fruit 4(– 5) cm long; seeds 2.5 – 4
mm long. — Widespread; lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . 1. I. blumei
b. Leaf blade ± ovate. Male perianth c. 6 mm diam., sepals with an adaxial appendage.
Fruit 4 – 8 cm long; seeds c. 8 mm long. — Lowlands or mountains; two subspecies,
each with a restricted area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. I. chlorocarpa
1. Indomelothria blumei (Ser.) W.J.de Wilde & Duyfjes

Indomelothria blumei (Ser.) W.J.de Wilde & Duyfjes, Blumea (2006) 6; Fl. Thailand 9, 4 (2008)
459. — Bryonia blumei Ser. in DC., Prodr. 3 (1828) 305. — Bryonia heterophylla auct. non (Lour.)
Raeusch.: Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 925 (see under Solena heterophylla Lour.). — Mel
othria marginata (Blume) Cogn. var. ? var. heterophylla (Blume) Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 594, p.p.; in Engl., Pflanzenr. 66, 4.275.I (1916) 93, p.p. — Type: Blume s.n. (holo
L, barcode L0129721), Java, Batavia “inter frutices juxta Bataviam”.
Subperennial(?) climber, 1– 3 m long; stem glabrescent. Tendrils glabrous. Leaves:

petiole 1– 2 cm long, rough-hairy; blade (ovate-)elliptic to narrowly elliptic or long-
triangular, 4 – 8 by 1– 4 cm, both surfaces scabrous, but veins shortly rough-hairy, cys-
toliths often inconspicuous, base subtruncate or cordate, occasionally short-hastate,
margin entire or shallowly sparsely dentate, teeth to 4 mm long. Male inflorescences:
finely scabrous or subglabrous; peduncle slender, 1– 2 cm long, raceme 0.1– 0.5(–1.5)
cm long, with up to 10 flowers. Male flowers (incompletely known, mainly from 1
open flower in Kerr 3735, Thailand): pedicel (0.5 –)1– 2 mm long; corolla 4 – 5 mm
diam.; receptacle-tube campanulate, 1.5 – 2 mm long and wide, inside curly hairy in
upper half, hairs 0.5 mm long or less; sepals 1 mm long, with sparse minute hairs or
glabrous, adaxial appendage not apparent; petals ovate, 2 by 1.5 mm, (sub)obtuse, on
both sides (papillose) hairy; filaments 0.5 mm long, anthers subcircular in outline, 1 mm
diam., thecae curved, connective with few minute hairs; disc subglobose, 1 mm diam.,
irregularly lobed. Female flowers not seen. Fruit ripening colour not known, 4(– 5) by
1–1.5 cm; fruiting pedicel 0.3 –1 cm long. Seeds greyish brown, 2.5 – 4 by 2 – 2.5 by
c. 1 mm.
Distribution — Widespread but rarely collected: Myanmar, Thailand; in Malesia:
Sumatra, East Kalimantan, and West Java.
Habitat & Ecology — Forest edges, scrub, open marshy forest, marshland; sea level
to 300 m; flowering and fruiting May to July and November.
Notes — 1. Lörzing 3385 (Sumatra) recorded the flowers as yellow. More material
and fieldstudy is needed. The observation of the flower colour, regarded as an important
genus character, needs special attention. The petals in I. chlorocarpa are white.
2. This widespread species is apparently easily overlooked as no recent collections
are known; the latest are of 1925 (Posthumis 1046, Sumatra; Endert 2080, East Kali-
mantan). Possibly a rare species.
2. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes

Indomelothria chlorocarpa W.J.de Wilde & Duyfjes, Blumea (2006) 7, f. 3. — Type: De Wilde, Postar,
Tajuddin & Good SAN 143915 (holo L; iso SAN), Sabah, Imbak River area.
Subperennial climber, 2 – 5 m long, glabrescent (hairs c. 0.1 mm long); main stem

c. 5 mm thick. Tendrils glabrous or harshly hairy. Leaves: petiole 1.5 – 4 cm long,
(scabrous-)hairy, hairs to 1 mm long; blade unlobed or occasionally hastately lobed,
subcircular, ovate(-elliptic) or elongate-triangular in outline, 5 –12 by 4 – 8 cm, both
surfaces glabrous but scabrous, except for sparse-hairy veins, cystoliths sometimes ap-
parent, base broadly rounded or shallowly truncate-cordate, margin entire or sparsely
shallowly dentate. Male inflorescences glabrescent; peduncle slender, 1– 2 cm long, 0.2
mm thick; raceme (0.2 –)0.5 – 3 cm long, 3 – 25-flowered. Male flowers: pedicel 2 –10
mm long; perianth 5 – 8 mm diam.; receptacle-tube urceolate, 3.5 – 4.5 by 2.5 – 3.5 mm,
outside glabrous, inside with a broad band of dense, c. 1 mm long hairs, 1.5 – 2 mm
below the throat; sepals (0.5 –)1 mm long, glabrous, adaxially with a short spur (Fig.
32a, f); petals (ob)ovate-elliptic, 3 – 4 by (2 –)2.5 – 3 mm, blunt or rounded, papillose
(gland-)hairy, (creamy-)white, ± recurved at anthesis and exposing anthers; filaments
c. 0.5 mm long, rather thick, anthers broad-ellipsoid or subcircular, 1.2–1.5 mm diam.,
orange, contrasting with the white corolla, thecae short-ellipsoid, ± curved, sublateral
with broad connective (Fig. 32); disc subglobose, c. 1.5 mm diam., 3-parted in the upper
third. Female flowers: pedicel 2 –10 mm long; ovary long-fusiform, glabrous, 10(–15)
by 2.5 mm, at apex ± narrowed into a neck 2 – 3 mm long; perianth as in male flowers
but larger, 10 –12 mm diam., inside faintly hairy; style c. 4 mm long, glabrous, stigma
c. 3 mm diam., consisting of 3 thick lobes densely set with slack (pendent) hairs 1–1.5
mm long; staminodes absent; disc a depressed ring-shaped cushion, c. 1 by 2.5 mm.
Fruit ripening dark green, paler striped, (4 –)5 – 8 by 1–1.5 cm; pulp greenish white;
fruiting pedicel 0.4 –1.5 cm long. Seeds 7– 8 by c. 4 by c. 1.5 mm.
2 cm
d
j
2 cm
2 cm
b 1 mm
2 mm
2 mm
g'' k l
a
1 mm 2 mm
g'
1 mm
e
g f i
Fig. 32. a, b. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. chlorocarpa. a. Female flower,
opened; b. fruit. — c – l. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. halimunensis
W.J.de Wilde & Duyfjes. c. Portion of male branch; d. leaf; e. male inflorescence; f. male flower,
opened; g, g', g'' stamens; h. portion of female branch; i. female flower bud; j. node with fruit; k. seed;
l. embryo (a: De Wilde SAN 141909; b: Postar et al. SAN 143729; c, e – g: De Wilde 21876; d: De Wilde
21877; h – j: De Wilde & Duyfjes 21927, type; k, l: De Wilde 21875).
Distribution — Borneo (Sabah, NE Kalimantan), Java (Gn Halimun).

Field-notes — Flowers creamy-white, anthers orange. Fruits dark green, paler striped,
smelling of cucumber. The flowers sometimes develop from short shoots on the older
wood, in shady environment, close to the ground.
KEY TO THE SUBSPECIES
1a. Male pedicels 2–4 mm long. Male disc with rounded lobes. — NE Borneo . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. subsp. chlorocarpa
b. Male pedicels 5–10 mm long. Male disc with subacute lobes. — West Java (Gn
Halimun) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. halimunensis
a. subsp. chlorocarpa
Male pedicels 2 – 4 mm long; male disc with broadly rounded lobes. Fruiting pedicel
0.5(–1) cm long. — Fig. 32a, b; Plate 17a, b.
Distribution — Borneo (Sabah, NE Kalimantan).
Habitat & Ecology — Shaded in (disturbed) primary forest and forest edges; re-
corded from limestone and brown loamy soil; at altitudes from sea level to 1000 m;
flowering and fruiting mainly from July to January.
b. subsp. halimunensis W.J.de Wilde & Duyfjes

Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. halimunensis W.J.de Wilde & Duyfjes,
Blumea 51 (2006) 9, f. 3: c – l. pl. 2. — Type: De Wilde & Duyfjes 21927 (holo L; iso BO), Java,
Gn Halimun.
Male pedicels 5 –10 mm long; male disc with subacute lobes. Fruiting pedicel c. 1
cm long. — Fig. 32c – l; Plate 17c, d.
Distribution — West Java: Gn Halimun.
Habitat & Ecology — Shaded in montane forest, along rivulets and on slopes; 800 –
1500 m; flowering and fruiting throughout the year.
19. Kedrostis
Kedrostis Medik., Philos. Bot. 2 (1791) 69; W.J.de Wilde & Duyfjes, Reinwardtia 12 (2004) 129.
— Type species: Kedrostis africana (L.) Cogn. (Bryonia africana L.).
Cerasiocarpum Hook.f. in Benth. & Hook.f., Gen. Pl. 1 (1867) 832; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 728; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 298. — Type species: Cerasio
carpum zeylanicum (Thwaites) C.B.Clarke (Aechmandra zeylanica Thwaites).
Herbaceous or woody climbers, (sub)perennial, leafy stem 1– 2 mm diam., glabrous

or hairy, with or without rootstock or tubers; monoecious or dioecious (not in Asia?).
Probract absent. Tendrils unbranched (elsewhere 2-branched). Leaves: petiole long;
blade simple, subentire or (deeply) lobed. Flowers small; petals yellow (elsewhere
whitish). Male inflorescences in short or long pedunculate racemes; pedicels short, per-
sistent; bracts mostly absent. Male flowers: receptacle-tube cup-shaped; sepals small;
petals free (elsewhere shortly united), imbricate, rounded at apex, glabrescent or short
(glandular) hairy; stamens 3 (elsewhere 5 and all anthers 1-thecous), inserted in or
close to the mouth of the receptacle-tube; filaments short, anthers two 2-thecous, one
1-thecous, with or without apically produced connective; thecae straight or little (much)
curved, disc basal, minute, unlobed or not apparent and fused with the base of the re-
ceptacle-tube. Female flowers single or in a short, few-flowered raceme, sometimes
co-axillary with male raceme; ovary ovoid, glabrous or hairy, 2- or 3- (or 4)-locular;
ovules few; style distinct; stigmas 2 (or 3), each irregularly lobed; staminodes absent
or usually 3 (or 5), inserted in the mouth of the tube; disc absent or not apparent. Fruit
solitary or few-fascicled, ripening red, fleshy, glabrous or hairy, subglobose or ovoid,
indehiscent (elsewhere fusiform, rarely dehiscent). Seeds 1 or 2 (elsewhere few), sub-
globose, usually smooth, not ornamented (elsewhere ornamented), without margin,
edge entire.
Distribution — An Old World genus of c. 25 species, of which 4 (5) species in Asia;
not in Australia; in Malesia 3 species.
KEY TO THE SPECIES
1a. Leaves with sparse appressed coarse hairs . . . . . . . . . . . . . . . . . . . . . . 2. K. hirta

b. Leaves glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Male raceme to 15 cm long. Fruiting pedicel 0.5 –1 cm long. Fruit globose or ±
transversely ellipsoid, 1–1.2 by 1–1.7 cm . . . . . . . . . . . . . . . . . . . . 1. K. bennettii
b. Male raceme 1(–1.5) cm long. Fruiting pedicel c. 0.2 cm long. Fruit (narrowly)
ovoid, c. 1.5 by 1–1.2 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. K. monosperma
1. Kedrostis bennettii (Miq.) W.J.de Wilde & Duyfjes

Kedrostis bennettii (Miq.) W.J.de Wilde & Duyfjes, Reinwardtia 12, 2 (2004) 130, f. 1. — Bryonopsis
bennettii Miq., Fl. Ned. Ind. 1, 1 (1856) 657. — Cerasiocarpum bennettii (Miq.) Cogn. in A.DC.
& C.DC., Monogr. Phan. 3 (1881) 729; in Engl., Pflanzenr. 66, 4.275.I (1916) 239, p.p., excl. f. 55
and material from India and Sri Lanka; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 298. — Type:
Horsfield s.n. (holo U, barcode U0001454; iso K, U, barcode U0001455), Java.
Climber 2 – 5 m long, glabrous (in young parts with minute greyish hairs and brown
gland-hairs), cystoliths usually obvious; monoecious or seemingly dioecious. Leaves:
petiole 3 – 6 cm long; blade unlobed, or ± 5-angular or shallowly (deeply) 3(– 5)-lobed,
subcircular or (narrowly) ovate in outline, 5 –17 by 8 –15 cm, glabrous, margin minutely
remotely dentate. Male raceme: peduncle 3 –7 cm long, solitary or co-axillary with
female flower(s); raceme spike-like, up to 15 cm long, (10 –)20 – 40-flowered, flowers
rather spaced. Male flowers: pedicel persistent, 1– 2(– 5) mm long, faintly articulate
0.5 –1 mm below receptacle-tube; receptacle-tube (2 –)2.5 – 3 by (2 –)3 – 5 mm, throat
densely hairy, hairs white, 0.5 –1 mm long; sepals triangular, (0.5 –)1 by 0.7–1 mm,
glabrous (except few minute brown gland-hairs); petals ± recurved, (ob)ovate, 2.5 – 3
by 2 – 2.5 mm, apex obtuse or rounded, in apical part densely fine hairy outside, densely
gland-hairy inside; filaments 1(–1.5) mm long, with minute (gland) hairs, anthers con-
nivent, dorsifixed above halfway, giving the anthers a ‘nodding’ aspect, (1–)1.5 mm
2 cm
b
e
1 mm 3 mm
h''
c
3 mm
h' k
2 cm
h
2 mm 2 mm
i j
g f
Fig. 33. Kedrostis bennettii (Miq.) W.J.de Wilde & Duyfjes. a. twig with male inflorescences; b. twig
with female inflorescence; c. ditto, in detail; d. twig with fruits; e. twig with two male inflorescences
and two fruits at the node; f. male flower; g. male flower, opened; h, h', h''. anther, seen abaxially,
adaxially, and laterally respectively; i. female flower; j. female flower, opened; k. seed (a: Koorders &
Koorders-Schumacher 44080; b, c, e – k: De Wilde et al. SAN 144527; d: Amdjah 218).
long, c. 1(–1.5) mm diam., connective broad and swollen, split at base and apex to c. 1/4
deep, apical lobes broadly rounded, thecae narrow, c. 1 mm long, straight, ± introrsely
opening; disc (or pistillode) absent. Female flowers 1– 4 (sub)fascicled at the node;
pedicel 2 – 5 mm long; ovary ovoid(-oblong), 3 – 4 mm long, 2 – 3 mm wide, glabrous;
perianth as in male flowers; staminodes 5, minute, either c. 0.5 mm long or larger and
broader, petal-like, 1(–1.5) mm long; style 2.5 – 3 mm long; stigma c. 2.5 mm diam.,
2-lobed, lobes sessile, (± woolly) hairy, each shallowly 2- (or 3)-lobed; ovary 2- (or
4?-)locular, each locule in basal part with 1 ovule. Fruit ripening shiny yellow or red,
smooth, on drying pale brown, finely wrinkled and dull, globose or ± transversely el-
lipsoid, 1–1.2 by 1–1.7 cm; fruiting pedicel 0.5 –1 cm long. Seeds 1 or 2, greyish or
brownish, subglobose, 7– 8 mm diam., faintly low-margined, smooth. — Fig. 33; Plate
12c, d, 13a.
Distribution — Sumatra, Borneo (Sabah, East Kalimantan), West & Central Java,
Sulawesi, Lesser Sunda Islands (Bali).
Habitat & Ecology — A rare species of scattered distribution on damp, rich soil, also
known from limestone; to 1400 m.
Note — The collection van Balgooy 7551 from Bali, has in the male raceme cadu-
cous bracts, 1– 3 mm long.
2. Kedrostis hirta W.J.de Wilde & Duyfjes

Kedrostis hirta W.J.de Wilde & Duyfjes, Reinwardtia 12, 2 (2004) 132. — Type: De Wilde & Duyfjes
19257 (holo L), Sumatra, Aceh (Gn Leuser National Park, Ketambe).
Climber, 2 – 5 m; sparsely (brown-)grey hairy, leafy stem 1.5–5 mm diam., with

curved hairs 0.1(– 0.2) mm long, cystoliths not obvious; monoecious. Leaves: petiole
3 – 4 cm long, with curved hairs c. 0.5 mm long, the lower portion ± twisted on drying;
blade 3- or 5-lobed to 1/3 – 3/4, subcircular in outline, 9 –13 cm diam., lobes narrowly
triangular or narrowly elliptic, 3 – 9 cm long, both surfaces sparsely appressed hairy,
hairs c. 1 mm long, margin with sparse brown-black soft teeth to 1 mm long. Male
raceme 5 –10-flowered in a condensed spike 0.5 –1 cm long; peduncle c. 1.5 cm long,
co-axillary with female flowers. Male flowers: pedicel c. 1 mm long; flowers not known.
Female flowers solitary or 2 – 5 fascicled at the node or with (presumably) male flowers
arranged in 0.5 – 3 cm long lateral short-shoots. Fruit subglobose-ellipsoid, 1–1.3 cm
long, dull brown, coarsely wrinkled on drying; fruiting pedicel c. 0.3 cm long. Seed 1,
globose, c. 7 mm diam., dark brown with a faint paler margin, smooth.
Distribution — Sumatra, only known from the type.
Habitat & Ecology — Forest edges or open places in forest, to 400 m; fruiting in
July.
Note — Kedrostis hirta is similar to K. bennettii, the latter is glabrous and has a long
many-flowered male inflorescence, to 15 cm long.
3. Kedrostis monosperma W.J.de Wilde & Duyfjes,

Kedrostis monosperma W.J.de Wilde & Duyfjes, Gard. Bull. Singapore 61, 1 (2009) 205, f. 1, 2.
— Type: Rahim Ismail KEP 100114 (holo KEP; iso K, L, SING), Peninsular Malaysia, Pahang,
Gn Benom Game Reserve.
a
2 cm
3 mm
2 mm
Fig. 34. Kedrostis monosperma W.J.de Wilde & Duyfjes. a. Apex of branch with female inflorescences
with immature and mature fruit; b. node with male inflorescence (all flowers fallen off); c. node with
compound female inflorescence with female flower buds and immature fruits (a, c: Siti Munirah et al.
FRI 65736; b: Rahim Ismail KEP 100114, type).
Climber 5 –7 m long, glabrous (except ovary), cystoliths not obvious; monoecious.

Leaves: petiole 1.5 – 3 cm long; blade unlobed, (broadly) ovate, 9 –13 by 4 –7.5 cm,
glabrous, base ± rounded or truncate or faintly hastate, margin (sub)entire, apex 1–1.5
cm acuminate. Male raceme: peduncle 0.2 – 0.5 cm long, co-axillary with female inflo-
rescence; raceme spike-like, 1(–1.5) cm long, 10 – 20-flowered, flowers ± densely set.
Male flowers: pedicel 1–1.5 mm long, persistent; mature flowers not known. Female
inflorescence an axillary (lateral) or terminal few-branched short-shoot 10 – 30 mm
long, the branches 5 –10 mm long, each with 5(–10) flowers of different stages of de-
velopment in loose clusters; bracts subulate, c. 0.5 mm long, caducous. Female flowers:
pedicel 1– 2 mm long; ovary ovoid, c. 5.5 mm by 3 mm, base rounded, gradually nar-
rowed to the apex in upper half, (sparsely) fine-hairy, hairs 0.2 mm long of many serial
cells, 2-loculed, each locule with 1 ovule; receptacle-tube 2(– 2.5) mm across, inside
glabrous; sepals (long) triangular, 0.5 –1 mm long; petals (ob)ovate, c. 3 by (1.5 –)2
mm, apex broadly rounded, both surfaces papillose hairy, hairs 0.1 mm long, style c.
3 mm long, stigma 2-lobed, 2 – 2.5 mm wide, lobes ± lacerate, papillose; staminodes
minute, fleshy, 0.2 – 0.3 mm long, two paired and one solitary. Fruit ripening glossy
orange-red, drying glossy, (narrowly) ovoid, c. 1.5 by 1–1.2 cm, glabrescent; fruiting
pedicel c. 0.2 cm long. Seeds 1 per fruit, (pale) brown, globose, c. 7 mm across, faintly
low-margined, smooth. — Fig. 34; Plate 12a, b.
Distribution — Peninsular Malaysia, Pahang, Ulu Krau, known from 2 collections.
Habitat & Ecology — Dense primary lowland forest on hillsides with rich soil;
flowering and fruiting in April and November.
Notes — 1. Kedrostis monosperma is similar to K. bennettii, the latter with a wider
distribution in western Malesia, but not known from peninsular Malaysia. In K. bennettii
the disposition of the fruit is different, single or few subaxillary to leaves, the fruits are
dull on drying, globose or transversely ellipsoid, 1- or 2-seeded, with a longer fruiting
pedicel, 5 –10 mm long; the male racemes are much longer, to 15 cm long with longer
peduncle, 3 –7 cm long; the petioles are generally longer, 2 – 6 cm long. As far as can
be judged from the few female flowering collections of K. monosperma, the ovary is
hairy, the perianth glabrous inside, in K. bennettii the ovary is (sub)glabrous and the
perianth hairy in the throat.
2. The older stem of the liana is c. 5 mm thick, and the bark has whitish corky
patches.
20. LAGENARIA
Lagenaria Ser., Mém. Soc. Phys. Genève 3, 1 (1825) t. 2 [Mém. Fam. Cucurbitacées (1825), t. 2];
C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1531. — Type species: Lagenaria
vulgaris Ser.
Robust annual (or perennial) climbing herbs, leafy stem 2 – 5 mm diam., plant finely
hairy; monoecious (or dioecious). Probract absent (or present, not in Asia). Tendrils
(unbranched or) 2-branched. Leaves: petiole long, usually with a pair of glands at apex;
blade simple, unlobed or lobed. Flowers solitary, large; petals white, free. Male flowers:
pedicel long; receptacle-tube campanulate or elongate; sepals narrow; petals obovate,
2 mm
i
h
j
2 mm 1 mm
k
3 mm
3 mm
b e
2 cm
d
3 mm
3 mm
c
3 mm
2 cm
Fig. 35. Lagenaria siceraria (Molina) Standl. a. Habit of node with male flower; b, c. male flowers;
d. stamens; e. petal; f. habit of node with female flower; g. female flower; h, i. blade bases with glands
at transition to petiole; j. leaf margin showing glands, abaxial view; k. seed (a–e, g: De Wilde & Duyfjes
21873C, culta, Netherlands; f, i: Phonsena et al. 5683, Thailand; h, j, k: Wiriadinata 472, culta, Sumba).
margin entire or ± crisped; stamens 3, filaments free, inserted on the tube towards the
base, anthers coherent but free, two 2-thecous, one 1-thecous, thecae plicate and often
also much contorted, connective broad; disc present. Female flowers: pedicel shorter
than in male flowers; ovary hairy, ovules numerous, horizontal; perianth as in male
flowers, but receptacle-tube short; style short, stigma deeply 3-lobed or consisting of
3 subsessile parts, each ± entire or 2-lobed; disc absent; staminodes small. Fruit ripen-
ing green or yellowish, large, hard-shelled, inside fleshy, indehiscent. Seeds numerous,
compressed, margin ± distinct, edge entire.
Distribution — About 6 species in Africa, one also in South America; in Malesia: 1
species, Lagenaria siceraria, widely cultivated.
1. Lagenaria siceraria (Molina) Standl.

Lagenaria siceraria (Molina) Standl., Field Mus. Nat. Hist., Bot. Ser. 3 (1930) 435; Backer in Backer
& Bakh.f., Fl. Java 1 (1964) 302; Widjaja & M.E.C.Reyes, PROSEA 8 (1993) 190; W.J.de Wilde
& Duyfjes, Sandakania 17 (2008 ‘2007’) 64; Fl. Thailand 9, 4 (2008) 460. — Cucurbita siceraria
Molina, Sag. Stor. Nat. Chili (1782) 133. — Type: Molina (not seen), South America, Chile.
Cucurbita lagenaria L., Sp. Pl. ed. 1 (1753) 1010. — Type: Herb. LINN No. 1151/1 (lecto, LINN, des-
ignated by Jeffrey in Milne-Redhead & Polhill (ed.), Fl. Trop. E. Africa, Cucurbitaceae 17 (1967)
51 (see C.E.Jarvis, Order out of chaos (2007) 465)), a cultivated plant at Uppsala.
Lagenaria vulgaris Ser., Mém. Soc. Phys. Genève 3, 1 (1825) t. 2; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 417; Craib, Fl. Siam. 1 (1931) 756. — Type: not indicated.
Robust climber or trailer to 5 m long, glabrescent; usually cultivated. Probract ab-

sent. Tendrils 2-branched. Leaves: petiole 4 – 20 cm long, laterally at apex with 2 small
(raised) glands; blade unlobed or obscurely 3 – 9-lobed, reniform, suborbicular or ovate
in outline, 5 – 30 cm diam., cystoliths not obvious, margin (sparsely) finely dentate,
soft hairy, apex obtuse or acute. Male flowers: pedicel 40 – 300 mm long; perianth
40 – 60(–120) mm diam.; receptacle-tube (long-)campanulate, slightly bulbous at base,
10 –15 mm long, soft hairy; sepals spaced, narrowly triangular or linear, c. 5 mm long;
petals broadly obovate, 20 – 40(– 50) by (10 –)15 – 35 mm, soft hairy, margin (sub)entire;
stamens inserted below halfway in the receptacle-tube, anthers forming a ± elongate
whole, filaments free, 2 – 3 mm long, glabrous; disc gland-like, at base of tube. Female
flowers: pedicel 5 –7 mm long; ovary cylindrical, reversed flask-shaped or obovoid,
10 – 20 mm long, villose; receptacle-tube 2 – 3 mm long; sepals and petals as in male
flowers but somewhat smaller. Fruit solitary, ellipsoid or variously flask-shaped, often
with a long ‘neck’, (6 –)10 – 80 cm long, up to 20 cm wide, glabrescent; fruiting pedicel
5 –10 cm long. Seeds pale brown, subtruncate at both ends, (7–)10 – 20(– 25) mm long,
± 2-horned on the broader end, faces shallowly ornamented and with two submarginal
ridges. — Fig. 35; Plate 13b, 32b.
Distribution — Worldwide cultivated.
Notes — 1. The flowers are open during the night.
2. Lagenaria siceraria, very variable in fruit- and seed-size and shape, was already
known in pre-Columbian times in South America, but it possibly originates from Africa.
Small-fruited forms, not necessarily wild forms, are known from e.g. the Moluccas or
from Africa. Small immature fruits are offered as a vegetable on markets.
21. LUFFA
Luffa Mill., Gard. Dict. Abr., ed. 1– 4 (1754) without pagination; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 455; G.J.Jansen, Gildemacher & Phuphathanaphong, PROSEA 8 (1993) 194;
C.Jeffrey in Hanelt, Mansfeld’s encycl. agric. hort. crops 3 (2001) 1539. — Type species: Luffa
aegyptiaca Mill.
Annual or subperennial, climbing herbs to 10 m long, leafy stem 2 – 5 mm diam.,

puberulous, late glabrescent, scabrid; monoecious (one species dioecious, not in our
area); wild, cultivated or running wild. Probract fleshy, less than 5 mm long, with
glands. Tendrils branched. Leaves: blade simple, mostly lobed. Flowers medium or
large; petals folded in bud, yellow, free, margin entire. Inflorescences: male flowers in
peduncled bracteate racemes; bracts narrowly ovate, to c. 5 mm long; female flowers
solitary, often co-axillary with male inflorescence. Male flowers pedicelled; receptacle-
tube small, shallow; sepals enclosing petals in bud; stamens 3 or 5: two in pairs and one
single, the paired ones free or the filaments variously fused, then appearing as 3 stamens
of which two 2-thecous, one 1-thecous, filaments free, inserted at base of the recepta-
cle-tube, anthers free, but usually connivent into a subglobose synandrium, connective
mostly broad, thecae marginal, convoluted-plicate; disc not apparent. Female flowers:
receptacle-tube and perianth as in male flowers; ovary elongated or ovoid, smooth (or
spiny not in Malesia), ovules numerous, horizontal; style short, stigma 3-parted, each
part 2-lobed; staminodes small; disc absent. Fruit subglobose to (long-)cylindrical,
smooth or ribbed (or with short spines), when ripe dry and fibrous within, dehiscing by
an apical operculum. Seeds numerous, medium-sized, compressed, (narrowly) elliptic
in outline, faces smooth or rugate, margin mostly distinct, edge entire.
Distribution — About 7 species, of which 4 in the Old World (mainly Africa) and 3
in America; in Malesia: 2 species, both widely cultivated in all tropical regions.
KEY TO THE SPECIES
1a. Plant flowering during the night, flowers pale yellow. Fruit 10-ribbed. Leaves rather
pale green, angular or lobed up to 1/3 deep . . . . . . . . . . . . . . . . 1. L. acutangula
b. Plant flowering during daytime, flowers bright yellow. Fruit not ribbed. Leaves
dark green, variable of shape, lobed up to 4/5 deep . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. L. aegyptiaca (with two forms)
1. Luffa acutangula (L.) Roxb.

Luffa acutangula (L.) Roxb., Hort. Bengal. (1814) 70; Fl. Ind. 713. 1832; Craib, Fl. Siam. 1: 756. 1931;
Backer in Backer & Bakh.f., Fl. Java 1 (1964) 300; W.J.de Wilde & Duyfjes, Sandakania 17 (2008
‘2007’) 67; Fl. Thailand 9, 4 (2008) 461. — Cucumis acutangulus L. (1753) 1011. — Cucurbita
acutangula (L.) Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 932. — Type: not designated (see note in
C.E.Jarvis, Order out of chaos (2007) 464).
Medium-sized climber to 5 m long, stem 2 – 4 mm diam., glabrous or subscabrous.

Probract with 2 – 5 glands. Tendrils 3 – 5-branched. Leaves: pale green; petiole 5 –12 cm
long; blade unlobed or shallowly 5 –7-lobed, subcircular in outline, 5 – 20 cm diameter.
3 mm
3 mm
c
1 mm
b
f
3 mm
a
3 mm
3 mm
d
2 mm 1 cm
j h g i
Fig. 36. a – f. Luffa aegyptiaca Mill. forma aegyptiaca (cultivated form). a, b. Female buds, from out-
side and opened respectively; c, d. female flower, from outside and opened respectively; e. staminode;
f. petal. — g, h. Luffa aegyptiaca Mill. forma sylvestris (Miq.) W.J.de Wilde & Duyfjes (wild form).
g. Fruit, opened, operculum removed; h. seed. — i, j. Luffa acutangula (L.) Roxb. (cultivated form).
i. Apex of male inflorescence with buds; j. seed (anomalous) (a – f: De Wilde & Duyfjes 22305; g, h. De
Wilde & Duyfjes 21861; i. De Wilde & Duyfjes 21679; j. De Wilde & Duyfjes 21743).
Flowers opening at night; pale yellow; male and female perianth similar; male flowers
in a peduncled raceme; female solitary or usually co-axillary with male raceme. Male
flowers: pedicel 10 – 40 mm long; receptacle-tube shallowly campanulate; sepals nar-
rowly elliptic; petals obovate, c. 20 mm long, apex rounded or emarginate; stamens
3, filaments 3 – 4 mm long. Female flowers: pedicel 50 –100 mm long; ovary obovate-
ellipsoid, 10-ridged. Fruit long-ellipsoid, 10-ridged, 10 – 30 by 4 –10 cm, apex obtuse
or acute, green, glabrous; pulp fibrous, operculum small. Seeds numerous, grey-black,
ovate-elliptic in outline, 10 –12 by 6 – 8 mm, smooth or rugose, not winged, margin
narrow, ± square. — Fig. 36i, j; Plate 33b.
Distribution — Originally in South Asia (India); in Malesia: widely cultivated.
2. Luffa aegyptiaca Mill.

Luffa aegyptiaca Mill., Gard. Dict., ed. 8 (1768) without pagination, based on Momordica luffa L.,
Sp. Pl. ed. 1 (1753) 1009; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 300; W.J.de Wilde &
Duyfjes, Sandakania 17 (2008 ‘2007’) 68; Fl. Thailand 9, 4 (2008) 462. — Type: a cultivated plant,
no specimen located.
Momordica cylindrica L., Sp. Pl. ed. 1 (1753) 1009. — Luffa cylindrica (L.) M.Roem., Fam. Nat. Syn.
Monogr. 2 (1846) 63.; Craib, Fl. Siam. 1 (1931) 756; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cam-
bodge, Laos & Vietnam 15 (1975) 47; I.Telford, Fl. Australia 8 (1982) 179. — Type: Herb. LINN
No. 1150.9 (lecto, designated by Wunderlin, Ann. Missouri Bot. Gard. 65 (1978) 329 (see C.E.Jarvis,
Order out of chaos (2007) 680; however, there is still no clarity about the type).
Luffa petola Ser. in DC., Prodr. 3 (1828) 303; Miq., Fl. Ned. Ind. 1, 1 (1856) 667. — Type: “Petola”
in Rumphius, Herb. Amboin. 5 (1747) 405, t. 147.
Luffa subangulata. Miq., Fl. Ned. Ind. 1, 1 (1856) 667. — Luffa acutangula (L.) Roxb. var. subangu
lata (Miq.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 461. — Type: Horsfield s.n. (holo
U (barcode U0001467); iso BM).
Climbers 5 –15 m long, stem 2 – 5 mm diam., glabrescent, scabrous. Probract 3 – 5

mm long, acute, with (1–)4 – 8 glands. Tendrils 2 – 6-branched. Leaves: petiole 2 –15
cm long; blade dark green, palmately 3 – 5(–7)-lobed, c. 1/4 deep or much deeper, lobes
various, suborbicular in outline, 5 –15 cm diam., glands small. Flowers opening at day-
time; bright yellow. Male flowers in peduncled raceme, 5 – 35 cm long, peduncle 5 –12
cm long; bracts (narrowly) ovate, c. 5 mm long, glandular; pedicel 2 –10 mm long;
receptacle-tube c. 5 mm wide, mostly hairy inside; sepals triangular, acute(-acuminate),
5 –10 mm long, with few glands; petals broadly rounded, 20 – 45 by 15 – 30 mm; sta-
mens 3 or 5, exerting from receptacle-tube, anthers 3 – 5 mm diameter. Female flowers:
pedicel 10 – 30 mm long, ovary ellipsoid or cylindrical, 15 – 30 mm long, finely hairy,
smooth or obscurely ribbed. Fruit ripening (light and dark) yellow green, globose, short-
to long-ellipsoid or cylindrical, 3 – 20(– 50) cm long, not ribbed, glabrous; pulp fibrous,
operculum small, beaked; fruiting pedicel 1– 6 cm long. Seeds numerous, dull blackish,
elliptic in outline, 7–12 by 4 – 8 mm, with narrow membranous wing-like border.
Distribution — Old World, Australia, naturalized in tropical America; in Malesia:
widely cultivated and locally with feral or truly wild forms.
Vernacular names — Vegetable Sponge, Loofah.
2 cm
i
c
f
0.5 mm 3 mm
h
1 mm
3 mm
1 mm
d
3 mm
g
2 mm
2 cm
b a
Fig. 37. Luffa aegyptiaca Mill. forma sylvestris (Miq.) W.J.de Wilde & Duyfjes (wild form). a. Node
with male inflorescence; b. apex of male inflorescence; c. male flower, opened; d. detail of male pedi-
cel with bract, shifted up along pedicel; e. node with fruit; f. operculum of fruit; g. seed, same shape
as in cultivated form, but smaller; h, i. detail of upper leaf blade surface with coarse hairs inserted on
cistolyths (all: De Wilde & Duyfjes 21861).
KEY TO THE FORMS
1a. Plant robust. Fruit ellipsoid to cylindrical, 10 – 20(– 60) cm long . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. forma aegyptiaca
b. Plant less robust in all parts. Fruit globose, 3 – 5 cm diam. or (broadly) ellipsoid,
3 – 8(–10) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. forma sylvestris
a. forma aegyptiaca
This is the cultivated form, very variable in fruit-size; often escaped into wild (sec-
ondary) vegetations, and at present occurring in all tropical areas. — Fig. 36a – f; Plate
13c, d, 33a.
b. forma sylvestris (Miq.) W.J.de Wilde & Duyfjes

Luffa aegyptiaca Mill. forma sylvestris (Miq.) W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’)
70. — Luffa sylvestris Miq., Fl. Ned. Ind. 1, 1 (1856) 666. — Type: “Petola silvestris” in Rumph.,
Herb. Amboin. 5, p. 409, t. 150, Ambon.
Luffa aegyptiaca Mill. var. leiocarpa (Naudin) Heiser & E.E.Schill., Biotropica 20 (1988) 186; G.J.
Jansen, Gildemacher & Phuphathanaphong, PROSEA 8 (1993) 196. — Luffa cylindrica (L.)
M.Roem. var. leiocarpa Naudin, Ann. Sci. Nat. Bot., Sér. 4, 12 (1859) 121. — Luffa leiocarpa
(Naudin) F.Muell., Fragm. 3 (1862) 107. — Type: Specimen from cultivation (P, not seen).
Tendrils (unbranched or) 2- or 3-branched. Fruit globose, 3 – 5 cm diam., or ellipsoid,

5 – 8(–10) cm long; fruiting pedicel 1– 3 cm long. Seeds brown-black, c. 7 by 4 mm.
— Fig. 36g, h, 37.
Distribution — Known from Jemen, India, Australia and the Pacific, and elsewhere;
in Malesia: East Java (possibly feral), Lesser Sunda Islands (Lombok, Flores), Moluc-
cas (Kai Is.), New Guinea (Papua New Guinea (East New Britain; Morobe, Gulf (Purari
delta), and Milne Bay Provinces)).
Habitat & Ecology — Common, often coastal.
Note — Forma sylvestris comprises all wild-growing and naturalized small-fruited
feral forms. The precise area of occurrence of the wild form is unknown.
22. MELOTHRIA
Melothria L., Sp. Pl. ed. 1 (1753) 35; Naudin, Ann. Sci. Nat., Bot., Sér. 4, 6 (1859) 148; Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 572, p.p.; in Engl., Pflanzenr. 66, 4.275.I (1916) 75, p.p.;
C.Jeffrey, Kew Bull. 15 (1962) 343; Wunderlin, Ann. Missouri Bot. Gard. 65 (1978) 332; W.J.de
Wilde & Duyfjes, Blumea 51 (2006) 9, f. 1b, 2b; t. 1. — Type species: Melothria pendula L.
Low climbers, subannual, leafy stem 1–1.5(– 2) mm diam., green on drying; monoe
cious. Probract absent. Tendrils unbranched, glabrescent. Leaves simple, unlobed or
lobed, palminerved. Flowers small, 5(– 8) mm diam., yellow; sepals minute, narrow,
subpatent; petals free, ovate-elliptic, valvate in bud; receptacle-tube campanulate. Male
inflorescence a slender peduncled raceme, with congested flowers, mostly with 1 female
flower co-axillary. Bracts absent. Male flowers: receptacle-tube cup-shaped; stamens 3,
inserted at c. 1/3 below the throat of the receptacle-tube, filaments free, short, dorsifixed
1 cm
i
2 mm
2 mm
a
1 mm
j l k
2 mm
1 mm
b
h f
1 mm
1 mm
1 mm
g c e d
Fig. 38. Melothria pendula L. a. Twig with male inflorescences; b. node with one male inflorescence
and one pedicel from fallen female flower; c. male bud; d, e. male flowers; f. anthers with short, narrow
filaments; g, h. female flowers; i. node with dried fruit showing seeds; j. fruit (from spirit); k. seed;
l. seed, membrane removed (all: De Wilde & Duyfjes SAN 141901).
near apex, anthers two 2-thecous, one 1-thecous, included, thecae lateral, straight, con-
nective narrow, not produced; disc subglobose. Female flowers solitary or co-axillary
with male raceme; pedicel long; ovary (ellipsoid-)narrowly ovoid, with a slender neck;
stigma consisting of 3 erect carnose lobes, papillose; staminodes 3, minute; disc annular,
free from the receptacle-tube. Fruit solitary, with long slender fruiting pedicel, ellipsoid,
small, 1–1.5 cm long, glabrous, juicy. Seeds numerous, compressed, ovate-elliptic, not
sculptured, margin absent, not winged, edge entire.
Distribution — About 10 species in the New World; 1 weedy species introduced in
West Africa and Asia.
Note — The here presented genus description and the description of M. pendula are
largely based on Asian material of that species.
1. Melothria pendula L.
Melothria pendula L., Sp. Pl. ed. 1 (1753) 35; Naudin, Ann. Sci. Nat., Bot., Sér. 4, 6 (1859) 148; Cogn.
in A.DC. & C.DC., Monogr. Phan. 3 (1881) 586; in Engl., Pflanzenr. 66, 4.275.I (1916) 87; Wunder-
lin, Ann. Missouri Bot. Gard. 65 (1978) 333; Correll & H.B.Correll, Fl. Bahamas Arch. (1982)
1429, f. 624; Diggs, Lipscomb & O’Kennon, Sida (1999) 570; T.-W.Hsu, J.-J.Peng & H.-Y.Liu,
Taiwania (2001) 193, f. 1– 3; W.J.de Wilde & Duyfjes, Blumea 51 (2006) 10, f. 4; pl. 3c, d. — Type:
Herb. LINN No. 51.1 (lecto LINN, designated by Wunderlin (1978)), “Habitat in Canada, Virginia,
Jamaica”. Taken from ‘The Linnean Plant Name Typification Project’, with Type Image. (See also
C.E.Jarvis, Order out of chaos (2007) 664)).
Bryonia filiformis Roxb., Fl. Ind. 3 (1832) 727; Bot. Surv. India, fasc. 7 (1978) pl. 24, left-hand plant.
— Type: not seen.
Annual creeper or climber, to 4 m long, sparsely hairy, glabrescent. Leaves: petiole

1.5 – 2.5 cm long, hairy, hairs 0.5 mm long; blade subcircular or ovate in outline, 3 – 6 by
3 – 6.5 cm, both surfaces (sparsely) scabrous-hairy, cystoliths minute, margin remotely
dentate or shallowly undulate. Male inflorescences: hairy as the petiole; peduncle 1– 2
cm long, 0.2 mm thick; raceme short, 0.1(– 0.5) cm long, with 2 – 6(–10) crowded flow-
ers. Male flowers: pedicel 3 – 5(–7) mm long; perianth 5(– 8) mm diam.; receptacle-tube
c. 2.5 by c. 2 mm, outside sparsely hairy, inside minutely glandular, especially in the
throat; sepals c. 0.5 mm long; petals patent, 2.5 – 3 by (2.5 –)3 mm, obtuse or retuse,
finely glandular; filaments c. 0.5 mm long, slender, anthers ellipsoid, c. 1.5 mm long,
connivent but free; disc c. 1 mm diameter. Female flowers 1 (or 2); pedicel 10 – 20 mm
long; ovary c. 5 mm long, with a c. 1 mm long neck, glabrous; perianth as in male flower
but larger; corolla c. 8 mm diam.; style c. 2 mm long; stigma-lobes 3, erect, each 2-lobed
at apex, ± connivent, c. 2 mm long, carnose, papillose, partly exserted; staminodes c. 0.5
mm long, inserted about halfway the receptacle-tube; disc a carnose annulus, c. 1 mm
high. Fruit solitary, ripening purple-black, (narrowly) ellipsoid, 0.8 –1.5 by 0.7–1 cm;
pericarp when ripe thin, when dry leaving the seeds shining through; fruiting pedicel
2 – 4 cm long. Seeds numerous, 3.5 – 4 by 2 – 2.5 mm, silvery whitish hairy. — Fig. 38;
Plate 16a, b.
Distribution — A variable widespread species in America: from Texas south to Ar-
gentina; introduced into West Africa and tropical Asia: South China (Macau), Taiwan;
in Malesia: Peninsular Malaysia (Selangor), Borneo (Sabah), Philippines, Sulawesi,
Lesser Sunda Islands (Bali).
Habitat & Ecology — Roadsides, beach forest; in crop fields and oilpalm plantations;
at low altitudes; flowering and fruiting throughout the year.
Note — In the material studied from Malesia the female flowers have minute stami-
nodes. Naudin (1859), who knew the species well, cultivated it in the botanic garden
at Paris from seeds sent by the French consul at Macau as well as from seeds from
America. He commented on the often hermaphroditic condition of the female flow-
ers.
23. MOMORDICA
Momordica L., Sp. Pl. ed. 1 (1753) 1009; Gen. Pl. ed. 5 (1754) 440; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 427; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 299; C.Jeffrey in Hanelt, Mans-
feld’s encycl. Agric. hort. crops 3 (2001) 1521; W.J.de Wilde & Duyfjes, Bot. Zhurn. (Moscow &
St. Petersburg) 87, 3 (2002) 133; Fl. Thailand 9, 4 (2008) 464; H.Schaefer., Heibl & S.S.Renner,
Proc. Roy. Soc. London, Ser. B, Biol. Sci. 276 (2009) 843; H.Schaef. & S.S.Renner, Molec. Phylo-
gen. Evol. 54, 2 (2010) 553. — Lectotype (Britton & Millsp., Bahama Fl. (1920) 425): Momordica
balsamica L.
Small or large climbers, annual or perennial, glabrous or pubescent; monoecious

or dioecious; wild or cultivated. Probract (in Asia) absent. Tendrils unbranched (or
outside Malesia, rarely proximally 2-branched, or in Africa rarely spinose). Leaves:
blade simple, unlobed or lobed, or (sub)pedately 3-foliolate (in Africa more-foliol-
ate). Flowers medium or large, sometimes ± zygomorphic, petals imbricate, white,
cream(-white) or yellow, free, margin entire. Inflorescences: male flowers solitary or
in loose pseudo-racemes, each flower stalk with small or large (sub)persistent hooded
bract (see note); female flowers solitary, with bract lower on the stalk. Male flowers:
pedicel short or long; receptacle-tube short, broad; sepals (narrowly) ovate; petals in
1– 3 with an incurved scale inside at the base; stamens 3 (or 2, not in SE Asia), anthers
one 1-thecous, two 2-thecous, filaments short, usually free, inserted in the throat or at
base of the receptacle-tube; thecae plicate (in Africa sometimes little curved), free or
mutually coherent, connective sometimes swollen; pistillode absent but disc at bottom
of receptacle-tube may be obvious. Female flowers: perianth as in male; ovary ellipsoid
or oblong-fusiform, smooth, ribbed, warty or papillose; ovules (in Asia) mostly numer-
ous, horizontal; stigma deeply 3-lobed, lobes 2-fid or notched (also unlobed in Africa);
staminodes absent or minute. Fruit solitary, pendulous, (ovoid-)ellipsoid, fusiform or
subglobose, ± fleshy, often ornamented with soft spines, warts or ridges, glabrous,
indehiscent or ± 3-valved, with few or numerous seeds in orange or red pulp. Seeds lit-
tle or distinctly compressed, smooth or sculptured, margin distinct or faint, edge often
undulate, sometimes grooved.
Distribution — About 40 species in the Old World, most species in Africa; 10 species
in Asia; in Malesia 5 species.
Uses — Young shoots and fruits of some species are used as vegetables, or used for
medicinal purposes.
Note — Male inflorescences are axillary. The male flowers are solitary or in fascicles or
in racemes. In Asia each stalked solitary flower represents a one-flowered inflorescence, in
which the flower is subtended by a single smallish or large sessile bract. The portion below
the bract is the peduncle; the portion above the bract the pedicel. In Asia, the thence one-
flowered peduncles are solitary, or (in M. clarkeana) arranged in loose fascicles or pseudo-
racemes. Likewise this can be observed in female inflorescences and consequently the
fruit stalk consists of the peduncle and the pedicel, usually with a bract scar at base of
the pedicel.
KEY TO THE SPECIES
1a. Plant monoecious. Leaves simple, deeply lobed. Male bract conspicuous, inserted
below the middle of the flower stalk. — Often cultivated . . . . . . 1. M. charantia
b. Plant dioecious. Leaves simple, unlobed or lobed, or 3-foliolate. Male bract either
inconspicuous, or conspicuous, inserted at the apex of the flower stalk. — Not or
rarely cultivated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Petiole and/or lower blade margin (usually) with conspicuous glands. Plant vigor-
ous. Male perianth c. 50 mm diam. or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
b. Petiole (and lower blade margin) without glands. Plant more slender. Male perianth
less than 50 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3a. Leaf blade unlobed, at least c. 1.5 times longer than broad; not or hardly fetid when
crushed; green on drying; tertiary venation sharp, with fine areoles. Fruit smooth
or partly with sparse slender soft spines . . . . . . . . . . . . . . . . . 4. M. denticulata
b. Leaf blade unlobed, or lobed, or 3-foliolate, in outline about as long as wide, always
less than 1.5 times longer than broad; fetid when crushed; dull green or brown on
drying; tertiary venation coarse. Fruit variously soft-spiny or tubercled (rarely
nearly smooth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. M. cochinchinensis
4a. Leaf blade 3-foliolate. — Seram, Ambon . . . . . . . . . . . . . . . . . . . . . 6. M. trifolia
b. Leaf blade simple (unlobed or lobed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Male flowers solitary on single, axillary peduncle; bract conspicuous, 10–30 mm
long. Fruit ornamented (soft spiny or warted, or with lengthwise ribs, or both).
Seeds numerous, 10 – 20, edge shallowly undulate . . . . . . . . . . 5. M. subangulata
b. Male flowers fascicled or in short raceme, the branches (peduncles) slender; bract
minute, 1– 2 mm long. Fruit ornamented (with few sparse minute spines) or smooth.
Seeds few, c. 6, edge with c. 8 conspicuous undulations . . . . . . . 2. M. clarkeana
1. Momordica charantia L.
Momordica charantia L., Sp. Pl. ed. 1 (1753) 1009; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
436; Craib, Fl. Siam. 1 (1931) 757; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 299; Keraudren
in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 42; I.Telford, Fl. Austr. 8 (1982)
167; M.E.C.Reyes, Gildemacher & G.J.Jansen, PROSEA 8 (1994) 207; W.J.de Wilde & Duyfjes,
Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002) 135; Sandakania 17 (2008 ‘2007’) 72; Fl. Thai-
land 9, 4 (2008) 464; H.Schaefer, Heibl & S.S.Renner, Proc. Roy. Soc. London, Ser. B, Biol. Sci.
276 (2009) 843. — Type: Herb. Clifford: 451, Momordica 2, (lecto BM, barcode BM000647445,
designated by Jeffrey in Milne-Redhead & Polhill, Fl. Trop. E Africa, Cucurbitaceae 31 (1967) (see
C.E.Jarvis, Order out of chaos (2007) 680)), India.
Momordica indica L. (in Stickman), Herb. Amboin. (1754) 24. — Type: “Amara indica” in Rumph.
Herb. Amboin. 5 (1747) 410, t. 151 (lecto, designated by Merr., Interpr. Herb. Amboin. 33 (1917)
495).
Momordica muricata Willd., Sp. Pl. 4 (1805) 602. — Type: Hort. Malab. 8 (1688) t. 10 (designated by
Chakravarty, Fasc. Fl. India 11 (1982) 92).
Slender climber 2(– 4) m long, annual, sparsely to densely hairy, partly glabrescent;
monoecious. Leaves: petiole 1.5 – 5 cm long, without glands; blade usually deeply pal-
mately 5 –7(– 9)-lobed, reniform or suborbicular in outline, 2.5 –10 cm diam., glands
minute, occasionally a few towards the blade base, lobes (ob)ovate, narrowed at base,
margin sinuate-dentate, ± mucronate. Flowers solitary, ± hairy; petals yellow. Male
flowers: stalk slender, with bract below middle; peduncle 0.5 – 3 cm long; bract reni-
form-suborbicular, 5 –15 mm diam., apex ± mucronate, margin subentire; pedicel 20 – 60
mm long; receptacle-tube cup-shaped, 2 – 4 mm long and wide; sepals (narrowly) ovate-
elliptic, 4 – 6 by 2 – 3 mm, acute, pale green; petals (narrowly) obovate, 10 – 20 by 7–15
mm, apex ± mucronate, basal scales 2; filaments 1.5 – 2 mm long, inserted in the throat
of the receptacle-tube, anthers coherent; disc cup-shaped, c. 1.5 mm diameter. Female
flowers: peduncle 0.5 – 5 cm long; bract 1–10 mm diam.; pedicel 10 – 50(–100) mm
long; ovary 8 – 30 by 2 – 4 mm, narrowly rostrate, muricate-tuberculate; sepals narrow,
oblong-lanceolate, 2 – 5 mm long; petals smaller than in male, 7–12 mm long; style c.
2 mm long; staminodes whitish, c. 0.5 mm long. Fruit ripening orange, (ovoid-)ellip-
soid or narrowly ellipsoid, narrowed at the ends, at apex usually rostrate, 2 –11(– 40,
cultivated) cm long, 2 – 4(– 6) cm wide, soft, sharp or blunt tuberculate in 6 –10 ridges,
and usually soft spiny in between, splitting incompletely with 3 valves exposing orange-
red pulp; fruitstalk 3.5 –15 cm long. Seeds few (to numerous), pale brown, ± square,
compressed, 8 –11(–15) by 5 – 8 mm, sculptured, margin grooved.
Distribution — Tropical and subtropical Africa and South, East, and SE Asia, to Aus-
tralia and the Pacific; also common in America where introduced; in Malesia through-
out.
Habitat & Ecology — Scrub and secondary places, forest edges, at low and medium
altitudes; also in seasonal climate.
Note — In the treatment of De Wilde & Duyfjes (2008), M. charantia was divided
into two forms, viz. forma charantia, the cultivated form, and forma abbreviata (Ser.)
W.J.de Wilde & Duyfjes, the wild form.
KEY TO THE FORMS
1a. Plant generally stouter. Fruit (narrowly) elliptic to oblong, 5 – 40 cm long, surface
with bluntish soft thorn-like protuberances and/or rounded warts in more or less
longitudinal rows. — Cultivated or escaped near villages . . . a. forma charantia
b. Plant more delicate. Fruit ± fusiform, 2 – 5 cm long, with c. 6 rows of few, low,
broad-based soft prickles with acute apex. — Growing wild . b. forma abbreviata
a. forma charantia
The forma charantia comprises all cultivated varieties, including plants with small
as well as with larger fruits. It should be noted that therefore taxa with small densely
muricate-tubercled fruits and named Momordica muricata (Willd., 1805; based on
Rheede 1688: 19, t. 10) also belong under forma charantia. Most likely the cultivated
form originated from the wild form in India (Williams & Ng, Ann. Bogor. 6 (1976) 111,
but Schaefer et al., 2009, conclude that it is of African origin). — Plate 33b.
Distribution — Widely cultivated.
Uses — Cultivated mostly in larger-fruited cultivars. Immature fruits and young
shoots are used as vegetables (PROSEA 8 (1993) 207); the fruit is medicinal (PROSEA
12, 1 (1999) 357).
Vernacular names — Peria, Peria laut, Periok (Malay).
b. forma abbreviata (Ser.) W.J.de Wilde & Duyfjes

Momordica charantia L. forma abbreviata (Ser.) W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’)
73, f. 8. — Momordica charantia L. var. abbreviata Ser. in DC., Prodr. 3 (1828) 311; Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 437, p.p., excl. M. muricata Willd. — Type: not indicated,
presumably in G, not seen.
This is the wild form, widespread in the Old World and already long-time natural-
ized in tropical America. It generally is more delicate than the cultivated varieties under
forma charantia, the latter including, however, frequently cultivated plants with small
fruits of only c. 5 cm long or more. — Plate 15a, b.
Note — The distinction between forma abbreviata and forma charantia seems rather
sharp on the characters given in the key, but vegetatively and in the flowers both forms
are similar, though forma abbreviata is more delicate in all parts.
2. Momordica clarkeana King

Momordica clarkeana King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 67 (1898) 35; Ridl., Fl. Malay
Penin. 1 (1922) 848; Cogn. & Harms in Engl., Pflanzenr. 88, 4.275.2 (1924) 39; W.J.de Wilde &
Duyfjes, Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002) 135, f. 1. — Type: King’s collector
8340 (lecto K, designated by De Wilde & Duyfjes (2002)), Perak.
Climber 6 – 8 m long, perennial; (sub)glabrous; dioecious. Leaves: petiole 2 – 8 cm

long, at base often ± distorted on drying, without glands; blade unlobed, ovate in outline,
4 –14 by 3 –12 cm, without glands, base cordate, margin remotely dentate, apex acute-
acuminate, short-mucronate. Flowers in male (solitary or) 2 – 4 fascicled, the fascicles
subaxillary to reduced leaves, usually arranged in loose raceme-like lateral shoots to
10 cm long; petals pale yellow. Male flowers: peduncle (almost) absent; bract basal,
spathulate, minute, 1– 2 mm long, ± dentate, glandular; pedicel slender, 15 – 30 mm
long; receptacle tube cup-shaped, c. 2 by 2.5 mm, brownish; sepals broadly ovate, c.
4 by (2 –)3 mm, obtuse, brown, paler to the margin, glabrous, margin short-fimbriate;
petals obovate-elliptic, c. 10 by 6 mm, apex obtuse (or subacute?), papillose-hairy;
other details including androecium not investigated by lack of material. Female flow
ers not known. Fruit solitary (or rarely 2 per node), ripening orange or red, (broadly)
ovoid, or pear-shaped, 4.5 –7 by 3.5 – 5 cm, base and apex broadly rounded apex 2 – 3
mm beaked, smooth, glabrous or with few minute sharp tubercles; exocarp thin, hard-
leathery; indehiscent; peduncle absent; bract not seen (absent?); fruiting pedicel (fruit
stalk) slender, 3 – 5 cm long. Seeds few (c. 6), (black-)brown, elliptic or subcircular,
1 mm
2 cm
c 2 mm
Fig. 39. Momordica clarkeana King. a. Male inflorescence; b. nearly mature male flower bud; c. por-
tion of stem with mature, hardly ornamented fruit; d. seed (a, b: Scortechini 1605; c, d: Rahim Ismail
KEP 98534).
thickish, 15 –18 mm across; margin edges with c. 8 conspicuous undulations; faces

sometimes sculptured. — Fig. 39; Plate 15d.
Distribution — Endemic to Peninsular Malaysia (Perak and Kelantan).
Habitat & Ecology — Open places in primary forest, hillsides, also on limestone, at
low altitudes to c. 200 m.
Notes — 1. The cogwheel-shaped seeds of M. clarkeana resemble those of M. cochin
chinensis, but the former are smaller.
2. This is a rare species characterized by its fruit. Male flowers, however, are poorly
known and female flowers still unknown.
3. Momordica cochinchinensis (Lour.) Spreng.

Momordica cochinchinensis (Lour.) Spreng., Syst. Veg. 3 (1826) 14; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 444, incl. var. villosa; Ridl., Fl. Malay Penin. 1 (1922) 848; Craib, Fl. Siam. 1
(1931) 758; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 38,
f. 8; I.Telford, Fl. Austr. 8 (1982) 167; M.E.C.Reyes, Gildemacher & G.J.Jansen, PROSEA 8 (1994)
206; W.J.de Wilde & Duyfjes, Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002) 137; Sandaka-
nia 17 (2008 ‘2007’) 75; Fl. Thailand 9, 4 (2008) 466, f. 21. — Muricia cochinchinensis Lour., Fl.
Cochinch. 2 (1790) 596. — Type: Loureiro s.n. (holo BM), from Cochinchina (Vietnam).
Momordica cochinchinensis (Lour.) Spreng. var. minor Cogn. in A.DC. & C.DC., Monogr. Phan.
3 (1881) 446. — Type: Beccari PP 608 (lectotype FI, designated here), West Papua (Vogelkop
Peninsula, Andai).
Momordica macrophylla Gage, Rec. Bot. Surv. India 3 (1908) 61; Gagnep., Fl. Indo-Chine 2 (1921)
1072; Craib, Fl. Siam. 1 (1931) 758. — Type (syntypes): Gallatly s.n., (CAL, not seen), & Mokim
253, (CAL, not seen), both Myanmar.
Momordica sphaeroidea Blanco 1837, Fl. Filip. (1837) 771; ed. 2 (1845) 531; ed. 3, 3 (1879) 174,
t. 380; Merr., Sp. Blancoan. (1918) 371. — Type: Merrill: Species Blancoanae 818 (neotype US,
designated here; iso L).
Passiflora saponaria Blanco 1837, Fl. Filip. (1837) 650. — Modecca? saponaria Blanco, Fl. Filip.
ed. 2 (1845) 453; ed. 3, 3 (1879) 53; Merr., Sp. Blancoan. (1918) 371. — Type: Merril: Species
Blancoanae 86 (neotype US, designated here; iso L).
Momordica ovata Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 446; Cogn. & Harms in Engl.,
Pflanzenr. 88, 4.275.2 (1924) 34. — Type: Cuming 1780 (lecto P, designated by De Wilde & Duyfjes
(2002); iso L), Philippines.
Momordica suringarii Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 434; Merr., Enum. Born.
(1921) 584; Cogn. & Harms in Engl., Pflanzenr. 88, 4.275.2 (1924) 23. — Type: Korthals s.n.
(Cissus 987) (lecto L, barcode L0125166, designated by De Wilde & Duyfjes (2002); iso L),
Sumatra.
Stout perennial climber to 20 m long, all parts (sub)glabrous, older bark pale, warty
and fissured; dioecious. Leaves: petiole (3 –)5 –12 cm long, with 1– 6 conspicuous raised
glands, but glands occasionally absent; blade entire or 3(– 5)-palmately lobed, or 3-fo-
liolate (leaflets ± elliptic, with short petiolule), broadly ovate or subcircular in outline,
(3 –)7–16(– 20) cm diam., margin entire or variously dentate, commonly with sessile or
stalked glands, strongly foetid when crushed, brown on drying; venation with areoles
1– 2 mm diam., the ultimate veinlets faint. Flowers somewhat hairy, in male solitary,
or several in a bracteate raceme to 5(–10) cm long; petals creamy, the three inner ones
with a dark blotch at base. Male flowers: stalk with bract subapical; peduncle 5 –15
cm long; bract cucullate, suborbicular or reniform, 20 – 40(– 60) mm wide, scabrous
or pilose inside, base rounded or cordate, margin ± entire, apex sometimes subacute,
sometimes with few glands at apex or towards base, sometimes woolly hairy; pedicel
3 –10(–15) mm long; receptacle tube saucer-shaped, 4(– 5) by 10(–15) mm, blackish;
sepals (long)triangular or (narrowly) ovate-elliptic, (8 –)10 –16 by 4 – 8 mm, acute,
blackish, scabrid or glabrous; petals (narrowly) elliptic, (25 –)40 – 60(–70) mm long,
subacute, conspicuously veined, scales broad, ligulate, 5 –7 by 5 mm, yellow, directed to
the base of the androecium, rendering the perianth somewhat irregular; filaments erect,
fleshy, 5 –7 mm long, broadly inserted at base of receptacle tube, anthers variable in
length, connivent (but free), connective swollen, each theca with a fleshy downwards
directed appendage; disc inconspicuous. Female flowers: stalk 3 –10 cm long; bract
elliptic, 5 mm long or less, ± median; ovary ellipsoid-oblong, 12 –15 mm long, densely

soft-muricate; receptacle tube small, narrow; sepals narrowly elliptic, 4 –10 mm long;
petals as in male; style (6 –)10 mm long. Fruit ripening orange-red, irregularly burst-
ing, ovoid, (broadly) ellipsoid or subglobose, (6 –)10 –15(– 20) by (4 –)6 –10(–15) cm,
apex ± pointed, pericarp mostly densely soft tuberculate or soft-spiny (spines to 10
mm long), rarely almost smooth; fruit stalk 3 –12 cm long, with bract (scar) median or
above. Seeds numerous, brown or grey-black, circular, elliptic, or ovate, 15 – 30 mm
across, 5 – 8 mm thick, margin coarsely undulate-tubercled; faces finely sculptured in a
patchy pattern. — Fig. 42e – g; Plate 14a, b, d.
Distribution — Widespread, from NE India and South China, through Indo-China to
North Australia and into the Pacific; absent from South India and Sri Lanka; in Malesia:
Sumatra, Peninsular Malaysia, Borneo, Java (Kebun Raya, Bogor, probably an old
introduction in a waste shrubby garden corner), Philippines, Sulawesi, Moluccas, New
Guinea (West Papua, and Papua New Guinea); not known from Singapore.
Habitat & Ecology — Forest and (degraded) forest edges on a wide range of land
forms and soils, usually in the lowlands to 1000 m altitude.
Uses — Fruits (unripe) are used as vegetable, or medicinal, especially by the Viet-
namese.
Vernacular name — Teruah (Malay).
Notes — 1. The roots are tuberous.
2. Momordica cochinchinensis is accepted as a variable, widely distributed species.
The species is particularly heterophyllous. Variable characters used as criteria for de-
scribing species now considered as synonyms concern the blade (either entire, lobed
or unlobed, or foliolate), the presence or absence of glands on the petioles, the flowers
being either solitary or grouped in short raceme-like inflorescences (short-shoots), and
also the size (and shape) of the fruit and their mode of ornamentation with soft warts
or spines. The length of the fruit stalk (peduncle plus fruiting pedicel) is very variable.
How this relates to the Malesian populations needs further investigation. The synonym
M. suringarii has mainly 3-foliolate leaves.
4. Momordica denticulata Miq.

Momordica denticulata Miq., Fl. Ned. Ind. 1, 1 (1858) 1090; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 447; Merr., Enum. Born. Pl. (1921) 583; Cogn. & Harms in Engl., Pflanzenr. 88,
4.275.2 (1924) 31; W.J.de Wilde & Duyfjes, Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002)
139. — Type: Diepenhorst s.n., (lecto U, barcode U0249679, designated by De Wilde & Duyfjes
(2002); iso BO), Sumatra (Priaman).
Momordica racemiflora (Miq.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 448; Merr., Enum.
Born. Pl. (1921) 583; Cogn. & Harms in Engl., Pflanzenr. 88, 4.275.2 (1924) 39. — Momordica
denticulata var. racemiflora Miq., Fl. Ned. Ind. 1, 1 (1858) 1091. — Type: Teijsmann s.n. (holo
U) Sumatra, Bondjol.
Momordica acuminata Merr., J. Malayan Branch Roy. Asiat. Soc. 1, 87 (1923) 45; Cogn. & Harms
in Engl., Pflanzenr. 88, 4.275.2 (1924) 32. — Type: Ramos 1303 (holo PNH, not seen; iso K, L),
Sabah.
Perennial climber to 8 m long, most parts (excluding inflorescences) glabrous, older

bark whitish grey, fissured, not tuberculate; dioecious. Leaves: petiole 2 – 4 cm long,
2 mm
2 cm
5 mm 5 mm 3 mm 2 mm
d e c h
Fig. 40. Momordica denticulata Miq. a. Habit of male with single inflorescences; b. ditto, with inflo-
rescences in short racemes; c. bract subtending young male flower bud; d. male flower; e. basal part of
male flower, showing androecium (papillose hairs omitted); f. stamens; g. node with blown-over female
flower; h. female flower, petals fallen off (a: De Wilde 21955; b – f: De Wilde 18079; g, h: Lorence
Lugas 2775).
without or with few glands in the upper half; blade entire, sometimes coriaceous, hardly
foetid when crushed, greenish on drying, (narrowly) ovate-elliptic, (7–)9 –15(– 20) by
4 – 9(–12) cm, ± shiny on both sides, glands on blade surface absent, sometimes present
on the basal blade margin, base truncate or cordate, margin entire or variously (sparsely)
sharply dentate, sometimes only one sharp tooth on each basal lobe, apex subobtuse
or acute-acuminate; veins 3(– 5) palmate from base, ascending, and few pinnate from
midrib, intercostal venation sharp, forming areoles c. 1 mm diameter. Flowers very
like those of M. cochinchinensis, somewhat scabrous or hairy, in male solitary or up
to 30 grouped in bracteate racemes to 5(–15) cm long; petals creamy-white, hairy or
bearded towards the base, the three inner ones with or without a black blotch at base.
Male flowers: stalk with bract subapical; peduncle 2 – 8 cm long, glabrous or minutely
hairy; bract circular or broadly ovate, (15 –)20 – 35(– 50) mm diam., scabrid on both
surfaces, base cordate, margin finely minutely hairy, apex obtuse or acute, with or with-
out glands; pedicel 5(–15) mm long; receptacle tube saucer-shaped, 2 – 3 by 8 –12(–15)
mm, ± blackish or not; sepals subcoriaceous, (narrowly ovate or) triangular, 10 –15
by 5 – 8 mm, acute(-acuminate), blackish green or yellow-brown, somewhat scabrous;
petals (narrowly) elliptic, 30 – 50 mm long, apex rounded or acute(-acuminate), dis-
tinctly veined, scales broad, lingulate, 5 –7 by 5 mm, yellow, directed to the base of the
androecium, rendering the perianth somewhat irregular; androecium 6 –10 mm long;
thecae with or without appendage. Female flowers: stalk (peduncle and pedicel) 4 – 6
cm long; bract acute, 2 – 3 mm long, inserted near the base of the stalk; ovary fusiform,
narrowed in the apical part, 18 – 25 by 4 – 5 mm, densely minutely hairy; receptacle
tube c. 3 mm wide; sepals linear, 7– 8(–10) mm long, acute; style slender, 5 –7 mm
long. Fruit ripening green-yellow or red, (ovoid-)ellipsoid-oblong, 8 –14 by 5 –10 cm,
apex narrowed into 10(–15) mm long beak, exocarp ± leathery, without ornamentation,
sometimes minutely scabrid; fruit stalk 5 –10 cm long, with bract scar near the base.
Seeds numerous, brown blackish, compressed, subcircular, cogwheel-shaped, 20 – 30
mm across, 6 –7 mm thick; margin with a double row of c. 10 blunt wart-like bulges,
faces finely sculptured in a blotchy pattern. — Fig. 40, 42a – d; Plate 14c.
Distribution — North and Central Sumatra, Peninsular Malaysia, most of Borneo
(not known from South Kalimantan).
Habitat & Ecology — Forest edges and clearings, and along roadsides, to 1200 m
altitude.
Note — Momordica denticulata is similar to M. cochinchinensis, especially in the
male flowers, but is quite different in general appearance of the leaves, which are about
twice as long as broad, somewhat chartaceous, glossy green with finer venation, the
petioles shorter with the glands in the apical part, and in the fruit, which is ellipsoid,
almost smooth.
5. Momordica subangulata Blume

Momordica subangulata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 928, excl. ref. to Rumph. 5, f. 150 =
Luffa cylindrica; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 443, p.p.; Ridl., Fl. Malay
Penin. 1 (1922) 848; Craib, Fl. Siam. 1: 758, p.p. 1931; Backer in Backer & Bakh.f., Fl. Java 1
(1964) 299; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 41, pl.
7, f. 1, 2; W.J.de Wilde & Duyfjes, Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002) 145, f. 4;
Fl. Thailand 9, 4 (2008) 468. — Type: Blume ‘769’ (lecto L, barcode L0001618, designated by De
Wilde & Duyfjes (2002); iso L, barcodes L0001619 & L0001620), Java.
Note — De Wilde & Duyfjes (2002) distinguished in M. subangulata two subspecies,

subsp. renigera (G.Don) W.J.de Wilde (North India, South China, Myanmar, and North
Thailand) and subsp. subangulata; only subsp. subangulata occurs in Malesia.
subsp. subangulata
Climber 1– 2.5 m long, stem slender, possibly with subperennial rootstock, almost
glabrous; dioecious. Leaves: petiole 2 – 6 cm long, without glands; blade unlobed or
shallowly 3 – 5-lobed, ovate in outline, 3 – 20 by 2.5 – 8 cm, glands absent, margin var-
iously dentate. Flowers solitary, minutely hairy; petals yellow. Male flowers: stalk
with bract subapical; peduncle 4 – 8 cm long; bract subcircular or reniform, 10 – 30
by (10 –)15 – 20 mm, apex rounded or ± acute-acuminate, margin entire or crenulate,
finely hairy, inside glabrous or soft hairy; pedicel 3 – 5 mm long; receptacle tube sau-
cer-shaped, c. 2 by 4 mm, green or blackish; sepals (ovate-)oblong, (4 –)6 –10 by 3 – 5
mm, obtuse or notched, blackish, apical portion not out-curved; petals obovate-elliptic,
20 – 30 by (10 –)15 –18 mm, broadly obtuse, of which three with a dark blotch at base
(always?), scales 2, broadly rounded; filaments c. 3 mm long, inserted almost at the
base of the receptacle tube, dark-coloured outside, papillose-hairy towards base, anthers
coherent in a subglobose mass, with five coarsely papillose extensions corresponding
with the thecae; disc as minute appendages near insertion of stamens. Female flowers:
stalk 3 –10 cm long; bract minute, at or below middle; pedicel 15 – 90 mm long; ovary
fusiform, narrowed at apex, 8 –12 mm long, 2 – 3 mm wide, (finely) densely pale brown
papillose-hairy and with 8 –10 lengthwise irregular (verrucose) low ridges; sepals linear,
c. 4 mm long; petals obovate-oblong, 20 – 25 by 10(–15) mm, obtuse; style c. 5 mm
long. Fruit ripening yellow-orange, ovoid or ellipsoid, narrowed at apex, 3 – 5 cm long,
subglabrous or finely hairy, and with 8 –10 irregular crested lengthwise ridges; fruit stalk
4 –15 cm long. Seeds 10 – 20, grey-brown or blackish, somewhat compressed, 6 –7 mm
across, c. 6 mm thick, not sculptured, margin (edge) shallowly undulate. — Fig. 41;
Plate 15c.
Distribution — Myanmar?, Thailand, Laos, and South China; in Malesia: Sumatra,
Peninsular Malaysia, and West Java.
Ecology — Open scrub, forest edges, 50 –1150 m altitude.
6. Momordica trifolia L.
Momordica trifolia L. (‘Stickman’), Herb. Amboin. (1754) 24; Rugayah & W.J.de Wilde, Blumea 42
(1997) 481 (in note under Trichosanthes wawrae); C.Jeffrey & W.J.de Wilde, Taxon 48 (1999) 600
(see note); Brummitt, Taxon 53, 3 (2004) 814. — Momordica trifoliolata L., Sp. Pl. ed. 2 (1763)
1434; Willd., Sp. Pl. 4 (1805) 604. — Type: “Poppya silvestris”, Rumph., Herb. Amboin. 5 (1747)
t. 152, f. 2. Epitype: Eyma 2794 (holo BO, here designated; iso L), Seram, Wae Tuba.
Momordica rumphii W.J.de Wilde, Bot. Zhurn. (Moscow & St. Petersburg) 87, 3 (2002) 144, f. 2h–j,
nom. superfl. — Type: Eyma 2794 (holo BO; iso L).
Slender perennial climber, roots not known, glabrous; dioecious. Leaves: petiole 2 – 4
cm long, glands absent; petiolules 0.6 – 0.8 cm long, glands absent; blade 3-foliolate,
2 cm b
a h 2 mm
2 mm
f
2 mm
2 mm
g d c e
Fig. 41. Momordica subangulata Blume subsp. subangulata. a. Habit of male flowering twig; b. ditto,
female; c. bract subtending male flower; d. outer petal, at base with in-curved scale; e. male flower
opened, showing androecium bract removed; f. stamens; g. fruit; h. seed (a, c – f: De Wilde 21897;
b: Kerr 7291; g, h: Grashoff 473).
2 cm
j
h
i
a
f
b
Fig. 42. Momordica, fruits and seeds. — a – d. Momordica denticulata Miq. a. Fruit; b. ditto, opened;
c. seed; d. seedling with cotyledons remaining in the seed. — e – g. Momordica. cochinchinensis (Lour.)
Spreng. e. Fruit; f. portion of pericarp (ornamentation differing from that in e); g. seed. — h–j. Momordica
trifolia L. h. Node with male inflorescence and male flower; i. fruit; j. seed (a, b: De Wilde 21951; c: Lor
ence Lugas 2449; d: De Wilde 22055; e: Van Balgooy 4958; f, g: Van Balgooy3017; h – j: Eyma 2794).
subcircular in outline, 8 –11 cm diam., apex acuminate, leaflets (narrowly) ovate, the
two outer ones unequal-sided, 5 –7 by 2 – 3.5 cm, margin sparsely dentate (teeth c. 1
mm long). Flowers in male solitary or occasionally up to 3 per node, (sub)glabrous;
petals pale yellowish(?). Male flowers: stalk with bract subapical; peduncle 1.5 – 2 cm
long; bract rather closely subtending the flower, suborbicular, 10 –15 mm diam., base ±
cordate, apex rounded with minute acuminate tip, margin greyish puberulous; pedicel
7–10 mm long; receptacle-tube cup-shaped, tapered, 2.5 – 3 by 3 – 4 mm, blackish; se-
pals narrowly elliptic, c. 5 by 2 mm, narrowly obtuse, pale (green-)brown, puberulous;
petals ± (narrowly) elliptic, 13 –15 by 5 – 6 mm, base ± clawed, apex obtuse or subacute,
puberulous; inside of receptacle-tube and androecium not investigated. Female flowers
not known. Fruit solitary, stalk slender, c. 2 cm long (Rumphius, t. 152, f. 2); fruit ripen-
ing orange, broadly ovoid-ellipsoid or subglobose, 1– 2 mm beaked at apex, c. 4.5 by 4
cm, sparsely muricate (warts 1– 2 mm high), pulp not known. Seeds few, c. 5 per fruit,
brown-black, thick but flat, circular in outline, c. 15 mm diam., c. 8 mm thick; margin
with a double row of 8 or 9 coarse warts or undulations, edge finely corrugated. — Fig.
42h – j.
Distribution — Moluccas (West Seram and Ambon).
Habitat & Ecology — Climbing and hanging vine in roadside at low altitude.
Note — Momordica trifolia is a local endemic species in West Seram and Ambon of
which, beside Eyma 2794 (male and separate fruit), no recent collections are known.
In its area it occurs together with resembling but stouter M. cochinchinensis, a species
with 3-foliolate forms known from outside the Moluccas, readily distinct by much larger
flowers, larger fruits, and conspicuous glands on the petiole.
24. Muellerargia
Muellerargia Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 360; I.Telford, Fl. Australia 8 (1982)
188; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 298. — Type species: Muellerargia timorensis
Cogn.
Small herbaceous climber, annual, leafy stem c. 2 mm diam., green-brown on dry-

ing; monoecious. Probract conspicuous or not obvious. Tendrils unbranched. Leaves
simple, unlobed or lobed, palmiveined. Flowers 4 – 5 mm diam.; petals imbricate, white,
free. Male inflorescences few- to many-flowered long-peduncled umbelliform clusters
or short racemes; bracts absent. Male flowers: pedicel long, persistent; receptacle-tube
campanulate; sepals linear, minute; petals ovate-elliptic; stamens 3, included, inserted
at c. 1/3(–1/2) from the base in the receptacle-tube, filaments short, anthers two 2-
thecous, one 1-thecous, free but tightly connivent forming a subglobose androecium,
thecae hook-like curved in the upper part (in Madagascar straight), connective rather
broad, produced (in Madagascar not produced); disc consisting of 3 elongate parts, star-
shaped arranged, adnate to the base of the receptacle-tube (in Madagascar disc absent).
Female flowers solitary, coaxillary with male inflorescence or not; pedicel long; ovary
ovoid, tapered into a narrow neck, tuberculate, or soft-bristly (or finely hairy), ovules
many, horizontal, placentas 2 (or 3); stigma 2(or 3)-lobed (in Madagascar subglobose);
staminodes 3, minute; disc absent. Fruit solitary, squirting the seeds; exocarp mem-
branous, warted or soft-spiny; pulp juicy. Seeds numerous, compressed, ± oblique or
curved, not or hardly ornamented, margin faint with entire, square edge (in Madagascar
rounded).
Distribution — A genus of 2 species, of which 1 in Madagascar; in Malesia: 1 spe-
cies.
1. Muellerargia timorensis Cogn.

Muellerargia timorensis Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 630; in Engl., Pflanzenr.
66, 4.275.I (1916) 135; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 298; I.Telford, Fl. Australia
8 (1982) 188. — Type: Bauer 56 (lecto W, designated here), Timor.
Melothria subpellucida Cogn., Bull. Acad. Roy. Sci. Belgique, sér. 3, 14 (1887) 356. — Type: Persieh
s.n. [et comm. F. von Mueller] (BR, not seen), Australia, Endeavour River.
Zehneria ejecta F.M.Bailey, Queensl. Fl. 2 (1900) 699. — Melothria ejecta (Bailey) Cogn. in Engl.,
Pflanzenr. 66, 4.275.I (1916) 108. — Type: Cowly s.n. (BRI, not seen), Australia, Thursday Is-
land.
Herbaceous climber, 1– 3 m long, hairy, hairs short or long. Probract not obvious or
leaf-like, often obliquely curved and ± clasping the stem, up to 15 mm long. Leaves:
petiole (1–)2 – 4 cm long; blade membranous, in outline (broadly) ovate or subcircular,
(3 –)5-angular or up to halfway lobed, 2.5 –10 by 2 – 9 cm, densely or sparsely fine hairy
or long-pilose on both surfaces, often scabrous above with minute cystoliths, margin
denticulate. Male inflorescences: peduncle 3 –10 cm long, at apex with a 5 – 20(– 40)-
flowered subumbel or flowers in a dense 1– 2 cm long raceme. Male flowers: pedicel
(5 –)8 –15 mm long, finely hairy; receptacle-tube c. 2 by 2.5 mm, finely gland-hairy,
throat inside with hairs c. 0.3 mm long; sepals linear, 0.4 – 0.6 mm long; petals elliptic
(ob)ovate, 2 – 2.5 by 1.5 – 2 mm, densely 0.1 mm long gland-hairy on both surfaces;
filaments c. 0.2 mm long, anthers connivent into a synandrium c. 1.5 mm diam., connec-
tive broad, with truncate apical c. 0.5 mm long extension, hairy at apex, thecae strongly
curved, hook-like; disc consisting of three carnose ± flat lobes, each c. 1 mm high, at
base connate and adnate with the base of the receptacle tube. Female flowers solitary,
pedicel 10 – 30 mm long; perianth as in male; ovary 4 –7 by 3 – 4 mm, at apex with a
2 – 2.5 mm long neck, smooth or mostly with scattered or dense sometimes ± upward
curved protuberances to 1 mm long, all densely finely minutely hairy; style c. 1 mm
long, stigma 1.5 – 2 mm long, irregularly 2-lobed, each lobe unequally 2- or 3-lobed
again, lobes slender, finely papillose; staminodes minute, inserted at c. 1/4 from the base
in the receptacle-tube, linear, truncate. Fruits ripening green, 1.5 – 2.5 by 1.5 – 2 cm, at
apex tapering into a beak to 0.5 cm long with at apex the withered perianth persisting,
tubercles or protuberances to 1 cm long; fruiting pedicel rather stout, straight, 1.5 – 3
cm long. Seeds narrowly ovate, c. 8 by 3 mm, base ± narrowed, apex subtruncate, faces
shallowly finely scrobiculate or smooth, margin faint, with square edge. — Fig. 43.
Distribution — Australia: Christmas Is., western Australia and Queensland; in
Malesia: Java (Madura), near SW Sulawesi (Pasitaloe Is.), Lesser Sunda Islands (Lom-
bok, Sumba, Flores, Timor), Moluccas (Wetar), New Guinea (Papua New Guinea (Cen-
tral Province, Tavai Creek area)).
Habitat & Ecology — Found in the margin of low monsoon forest, in Eucalyptus
savanna, along the coast in scrub and on rocky beaches, on loamy soil or limestone;
from sea level to 60 m altitude; flowering and fruiting mostly in April.
1 cm a
1 cm
f
1 cm
1 mm
1 mm
d
1 mm
e
1 mm
c g
Fig. 43. Muellerargia timorensis Cogn. a. Habit of branch with male inflorescence and fruit; b. ditto,
with probract at the node; c. male flower, openend; d. androecium, stamens connivent; e. stamens;
f. node with female flower; g. female flower, perianth opened; h. seed (a, c – e, h: Pullen 6848; b:
Schmutz 1491; f. Loeters 1377; g. Afriastini 1598).
Note — Pullen 6848 (Tavai Creek) noted: ‘the fruits when ripe are easily dislodged
from the vine, at which time they squirt seeds from the ruptured end for a consider-
able distance’. The synonym Zehneria ejecta, refers to the squirting fruits, like in the
Mediterranean Ecballium elaterium (L.) A.Rich., the Squirting cucumber.
25. MUKIA
Mukia Arn., Madras J. Lit. Sci. 12 (1840) 50; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 623;
C.Jeffrey, Kew Bull. 15 (1962) 343; in Hooker’s Ic. Pl. 37, 3 (1969) 2, t. 3661– 3664; Keraudren
in Aubrév. & J.-F.Leroy, Fl. Camb., Laos, Viêt-Nam 15 (1975) 57; W.J.de Wilde & Duyfjes, Thai
Forest Bull., Bot. 34 (2006) 38; Fl. Thailand 9, 4 (2008) 471. — Melothria L. sect. Mukia (Arn.)
Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 622. — Type species: Mukia scabrella (L.) Arn.
(Bryonia scabrella L. = Mukia maderaspatana (L.) M.Roem.).
Small climbers, shoots herbaceous, (sub)annual or with a (thick) perennial root, leafy
stem c. 2 mm diam., whole plant scabrid-hairy; monoecious. Probract absent. Tendrils
unbranched. Leaves: blade simple, green on drying; apices of (lobes of) developing
leaves distinct, broadened, glabrous, often brown on drying; petiole long or short (leaves
subsessile). Flowers small; petals yellow, (almost) free, imbricate in bud; disc free from
the receptacle-tube. Male inflorescences in a fascicle at the node, with few to 10(– 20)
short-pedicelled flowers; bracts absent. Male flowers: pedicel 2 –10 mm long, slender;
receptacle-tube urceolate-campanulate; sepals minute, long-triangular or linear, out-
curved; petals elliptic or (ob)ovate, free or short-connate; stamens 3, inserted slightly
above halfway the receptacle-tube, filaments short, glabrous, anthers one 1-thecous and
two 2-thecous, included, thecae lateral, straight, connective narrow, ± hairy, at apex
hardly or shortly produced; disc depressed globose. Female flowers 1– 6 fascicled,
usually separate from male flowers; pedicel short; ovary globose to narrowly elliptic,
ovules few or many; perianth as in male; style thick, glabrous; stigma consisting of 3
elongate, carnose, papillose lobes, each lobe shallowly 2-lobed again; staminodes usu-
ally present; disc annular. Fruits 1– 6, clustered, subsessile or short-pedicelled, ripening
red, sometimes darker flecked, (sub)globose or ellipsoid, 0.5 – 3 cm long, juicy; exocarp
membranous or cartilaginous, smooth. Seeds few or many, whitish or pale brown, sub-
globose or compressed, ornamented or not, margin distinct, edge subentire, sometimes
grooved.
Distribution — A genus of about 9 species, including some 3 unpublished Australian
species, distributed in the tropics of the Old World: Africa (1 species); in SE Asia from
Pakistan east to China and south-east through Indo-China to Australia; in Malesia: 3
species.
Notes — 1. Molecular data (Ghebretinsae et al., Novon (2007) 176, and Schaefer,
Blumea (2007) 165) indicate that Mukia is nested within Cucumis.
2. The broadened glabrous apices in immature leaves are also seen in various other
genera, e.g. Benincasa, and Cucumis.
KEY TO THE SPECIES
1a. Fruit ellipsoid, 2– 4 cm long. — East Malesia: Moluccas & Vogelkop Peninsula . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. M. rumphiana
b. Fruit globose or ellipsoid, up to 1.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Fruit ellipsoid; exocarp thin, collapsing about the seeds, translucent. Seed faces
flat, not or hardly ornamented. Hairs of petiole spreading or curved downward. —
Sumatra, Borneo, Java, Philippines . . . . . . . . . . . . . . . . . . . . . . . . . 1. M. javanica
b. Fruit globose; exocarp thicker, wrinkling or not, not translucent. Seed faces convex,
scrobiculate. Hairs of petiole spreading or curved upward. — Widespread . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. M. maderaspatana
1. Mukia javanica (Miq.) C.Jeffrey

Mukia javanica (Miq.) C.Jeffrey in Hooker’s Ic. Pl. 37, 3 (1969) 3, t. 3661: 1–10; Keraudren in Aubrév.
& J.-F.Leroy, Fl. Camb., Laos, Viêt-Nam 15 (1975) 58, f. 10: 6 – 8; W.J.de Wilde & Duyfjes, Thai
Forest Bull., Bot. 34 (2006) 40; Fl. Thailand 9, 4 (2008) 474. — Karivia javanica Miq., Fl. Ned. Ind.
1, 1 (1856) 661. — Melothria javanica (Miq.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
625; in Engl., Pflanzenr. 66, 4.275.1 (1916) 129; Backer in Backer & Bakh.f., Fl. Java 1 (1964)
298. — Cucumis javanicus (Miq.) Ghebret. & Thulin, Novon 17 (2007) 177. — Type: Horsfield
s.n. (holo U, barcode U0226804; iso BM, K), Java.
Climber or creeper to 3 m long, stem scabrous hairy. Leaves: petiole 2 – 5(–10) cm

long, scabrid-hairy, hairs curved downward; blade 5-angular or variously (3 –)5-lobed
up to halfway deep, broadly ovate(-hastate), subcircular in outline, 2 –10 cm diam.,
upper and lower surface subglabrous or variously (scabrid-)hairy, denser on the veins
beneath, cystoliths not apparent, margin to 2 mm dentate. Male flowers 3 – 6, rarely with
few female flowers mixed; pedicel 1– 2(– 4) mm long; receptacle-tube 1.5 – 3 by 1.5 – 2
mm, scabrid hairy; sepals 0.5 –1.5 mm long; petals ovate, 1.5 – 2.5 mm long, apex su-
bacute, glabrous except for few hairs on outer surface; filaments less then 0.5 mm long,
anthers 1.5(– 2) mm long; disc 1–1.5 mm diameter. Female flowers (1–)2 – 4; pedicel
c. 1 mm long; ovary ellipsoid(-oblong), 4 – 5 mm long, (sub)glabrous or finely hairy,
hardly constricted at apex; perianth as in male glabrous; disc c. 1 mm high. Fruits 1– 3,
fascicled, ellipsoid, 1–1.5 cm long, juicy, with filmy pericarp showing the seeds when
dry, glabrous; fruiting pedicel 0.1– 0.2 cm long. Seeds 8 –18, (pale brown or) whitish,
obovate, strongly compressed, c. 5 by 3.5 – 4 by 1–1.5 mm, faces flat, ± depressed,
smooth or irregularly low-warted, with a broad 2-grooved margin. — Plate 16c.
Distribution — Northern India, east to China; in Malesia: Borneo, Java, Philippines;
not known from Sulawesi, Lesser Sunda Islands, and New Guinea.
Habitat & Ecology — Roadsides, (disturbed) forest and scrub edges; to 1200 m
altitude; flowering and fruiting throughout the year.
2. Mukia maderaspatana (L.) M.Roem.

Mukia maderaspatana (L.) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 47; C.Jeffrey in Hooker’s Ic.
Pl. 37, 3 (1969) 5, p.p., excl. t. 3662: 1– 8 (which concerns Mukia gracilis (Kurz) W.J.de Wilde
& Duyfjes); Keraudren in Aubrév. & J.-F.Leroy, Fl. Camb., Laos, Viêt-Nam 15 (1975) 60, f. 10:
9; I.Telford, Fl. Australia 8 (1982) 183, f. 40: a – g; W.J.de Wilde & Duyfjes, Thai Forest Bull.,
Bot. 34 (2006) 43; Fl. Thailand 9, 4 (2008) 475, pl. 35: 2. — Cucumis maderaspatanus L., Sp. Pl.
(1753) 1012; H.Schaef., Blumea 52 (2007) 167. — Melothria maderaspatana (L.) Cogn. in A.DC.
& C.DC., Monogr. Phan. 3 (1881) 623; in Engl., Pflanzenr. 66, 4.275.1 (1916) 126; Craib, Fl.
Siam. Enum. (1931) 764; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 298. — Type: “Cucumis
Maderaspatensis fructu minimo” in Plukenet, Phytographia (1692) t. 170, f. 2. (lecto, designated by
Meeuse, Bothalia 8 (1962) 14); typotype: Herb. Sloane 95: 201 (BM-SL, not seen (see C.E.Jarvis,
Order out of chaos (2007) 363)), “habitat in India”.
Bryonia scabrella L. in L.f., Suppl. Pl. (1781) 424; Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 926; Miq., Fl.
Ned. Ind. 1, 1 (1856) 658. — Mukia scabrella (L.) Arn. in Hook., J. Bot. 3 (1841) 276; C.B.Clarke
in Hook.f., Fl. Brit. Ind. 2 (1879) 623. — Mukia maderaspatana (L.) M.Roem. var. scabrella (L.)
Kurz, J. Asiat. Soc. Bengal 46 (1877) 104. — Type: India, without collector, cult. Uppsala (holo
LINN 1153/11, not seen).
Bryonia althaeoides Ser. in DC., Prodr. 3 (1828) 306. — Mukia althaeoides (Ser.) M.Roem., Fam. Nat. Syn.
Monogr. 2 (1846) 47. — Type: without collector (G-DC holo, not seen, microphoto in K, L), Timor.
Climber to 4 m long, stem scabrous or stiff-hairy. Leaves: petiole (0.1–)0.5 – 9 cm

long, scabrous and hispid with short or long, erect or upward curved hairs (but see note);
blade unlobed or 3 – 5-lobed, broadly ovate, subcircular or broadly hastate in outline,
2 –10 cm diam., upper and lower surface hispid or scabrous-hairy, more densely so on
the veins, cystoliths dense and minute and not apparent, margin variously up to 5 mm
dentate. Male flowers in fascicles of 2 – 20 (occasionally mixed with few female flow-
ers); pedicel 2 – 5(–10) mm long; receptacle-tube 1.5 – 4 by 1– 2 mm, with upward ap-
pressed hairs; sepals 1(–1.5) mm long; petals ovate, 1.5 – 3(– 4) mm long, apex subacute,
glabrous except for the mid-vein outside; filaments less than 0.5 mm long, anthers 1– 2
mm long, the connective usually shortly produced; disc c. 1.5 mm diameter. Female
flowers solitary or up to 8; pedicel 1– 4 mm long; ovary subglobose or broadly ovoid,
3 – 3.5 by 1.5 – 2 mm, with scattered or dense (stiff) hairs. Fruit 1– 5(– 8) in axillary
clusters, green and pale green striped, ripening red, darker striped or not, globose,
0.5 –1.5 cm diam., glabrous or with few coarse hairs; exocarp thin but not filmy, coarsely
wrinkled when dry, seeds not shining through; fruiting pedicel 0.2 – 0.5 cm long. Seeds
10 – 20, whitish or pale brown, obovoid, moderately compressed, 3 – 4 by 2 – 2.5 by
1.5 – 2 mm, margin narrow, ± rounded, faces not separated by a groove, faces convex,
variously warted, or pitted or nearly smooth. — Fig. 44, 45; Plate 16d.
Distribution — Widespread: Africa, SW and SE Asia, including Yemen, Pakistan,
India, Sri Lanka, north-east and east to China, Ryukyu Is., Indo-China to Australia
(where very variable in indumentum, including villose); in Malesia: Sumatra, Peninsu-
lar Malaysia, Borneo, Java, Philippines, Sulawesi, Lesser Sunda Islands, New Guinea
(West Papua and Papua New Guinea); no collections seen from Moluccas.
Habitat & Ecology — In a large variety of (degraded) scrub-land, among grasses,
roadsides, forest (edges), in Papua New Guinea in Eucalyptus savanna; also on lime-
stone; from sea level to 1200 m altitude; flowering and fruiting throughout the year.
Notes — 1. Specimens from savanna areas in eastern Papua New Guinea (e.g. Hey
ligers 1176, Darbyshire 696, Henty & Katik NGF 38646, and Pullen 6804) have com-
paratively large fruits, c. 1.5 cm diameter. Pullen 6804 will be accommodated into a
new species of Cucumis by Sebastian et al., PNAS (2010).
2. The discriminating character of the of curving of the hairs on the petiole, viz. up-
ward in M. maderaspatana and downward in M. javanica works well in most material
1 mm
a
1 mm
2 mm
2 cm b
g
1 mm
c
2 mm 1 mm
d e f
Fig. 44. Mukia maderaspatana (L.) M.Roem. a. Portion of twig with male inflorescences; b. detail of
male inflorescence; c. male flower, opened; d. female inflorescence; e, f. female flower, from outside
and opened respectively; g. infructescence; h. detail of lower leaf blade surface (a – f, h: De Wilde &
Duyfjes SAN 141920; g: De Wilde & Duyfjes SAN141930).
3 mm
c
d
1 mm
2 cm
1 mm
3 mm
Fig. 45. Mukia maderaspatana (L.) M.Roem. a. Male inflorescence; b. ditto; c. male flower, opened
and petal; d. female flower bud; e. fruit; f. seed, side view and face (all: Henty NGF 49703).
of SE Asia and West Malesia. However, one should be aware that in some specimens
from East Malesia (where M. javanica does not occur) the position of the hairs may be
ad variance: sometimes retrorse in Lesser Sunda Islands or often erect or curved to all
directions in New Guinea.
3. The collection Henty NGF 49703 (see Fig. 45), from savanna in East Papua New
Guinea, differs in having a delicate habit, long-pedicelled, minute, slender male flowers,
a 5 –10 mm long pedicel, the connective of anthers not produced, solitary female flow-
ers and fruits, the fruit c. 1.2 cm diam., containing c. 25 scrobiculate seeds, and in not
up-curved hairs on the petiole. Its determination as M. maderaspatana is provisional.
4. Plants may be perennial with a thick old woody rootstock or annual and soon
flowering, with fibrous roots. The specimens Raap 499 and Insani SAN10 (Java) will
be placed by Sebastian et al., l.c., into a separate species Cucumis althaeoides, but we
cannot corroborate this on morphological grounds.
3. Mukia rumphiana (Scheff.) W.J.de Wilde & Duyfjes

Mukia rumphiana (Scheff.) W.J.de Wilde & Duyfjes, Thai Forest Bull., Bot. 34 (2006) 45. — Melothria
rumphiana Scheff., Ann. Jard. Bot. Buitenzorg 1 (1876) 25. — Cucumis rumphianus (Scheff.)
H.Schaef., Blumea 52 (2007) 168. — Type: Teijsmann 7496 (lecto L, barcode: L0589472, desig-
nated by De Wilde & Duyfjes (2006)), Ternate. (For synonyms see under the subspecies).
Climber 3 – 5 m long, roots perennial, leafy stem 2(– 3) mm diam., with stiff hairs, ±
upward directed. Leaves: petiole 1– 3 cm long, scabrous by stiff upward directed hairs;
blade 3 – 5-angular or -lobed up to c. halfway (rarely deeper), broadly ovate in outline,
4 – 8 by 4 –11 cm, upper surface finely scabrous-punctate, lower surface grey scabrid-
hairy, margin sinuate-dentate. Male flowers in fascicles of 3 –15; pedicel 4 –7 mm long;
receptacle-tube long-campanulate, 3 – 4 by 1.5 – 2 mm; sepals 1–1.5 mm long, outside
and inside densely fine-hairy; petals ovate-elliptic, 2 – 3 by 1.5 – 2 mm, apex acute(-acu-
minate), wholly finely-hairy at outside; filaments c. 0.5 mm long, anthers oblong, c. 2
mm long, at apex almost without or with one or two small exsertions; disc depressed,
c. 1 mm diameter. Female flowers solitary; pedicel 1– 2 mm long; ovary ovoid, 4 – 6 by
2 – 3 mm, (densely) soft- or coarse-hairy, the hairs patent or ± upward directed; stamin-
odes minute. Fruit solitary, whitish, ripening red, ellipsoid, 2 – 4 by 1.5 – 2.5 cm, either
densely soft-hairy or sparsely hairy and later on glabrescent; exocarp thin, not or hardly
translucent; fruiting pedicel 0.4 –1 cm long. Seeds numerous, pyriform-ovoid, only little
compressed, (4 –)5 – 6 by 3 – 4 by 2 mm, margin narrow, with 2 grooves; faces flattish,
shallowly verrucose-rugose.
Distribution — Sulawesi, Moluccas (Ternate, Bacan, Sula Islands, Buru, Ambon),
New Guinea (West Papua (Vogelkop Peninsula)); 2 subspecies.
Habitat & Ecology — Forest edges, hedges, shrubberies near the coast; on limestone;
at low altitudes. Flowering and fruiting throughout the year.
1a. Fruit sparsely long-hairy, hairs 1– 2 mm long . . . . . . . . . . . . a. subsp. rumphiana

b. Fruit densely short-hairy, hairs up to 0.5 mm long . . . . . . . . b. subsp. tomentosa
a. subsp. rumphiana
Mukia celebica (Cogn.) F.M.Bailey, Queensl. Fl. 2 (1900) 700, for the type only; C.Jeffrey in Hooker’s Ic.
Pl. 37, 3 (1969) 11, t. 3664. — Melothria celebica Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
625; in Engl., Pflanzenr. 66, 4.275.1 (1916) 128. — Type: Forsten 96 (holo L), Sulawesi, Tondano.
Melothria javanica auct. non (Miq.) Cogn.: Merr., Interpr. Herb. Amboin. (1917) 491. — Cucumis
murinus ruber Rumph., Herb. Amboin. 5 (1750) 463, t. 171, f. 1 & a.
2 cm
2 mm
2 mm
c'
2 mm
b
c
d
1 mm
1 mm
g
2 mm i
h
2 mm
f 2 cm
e
Fig. 46. Mukia rumphiana (Scheff.) W.J.de Wilde & Duyfjes subsp. tomentosa W.J.de Wilde & Duyfjes.
a. Twig with male inflorescences; b. male inflorescence; c, c'. male flower, from outside and opened
respectively; d. stamens; e. node with immature female flower; f, g. female flowers, from outside and
opened respectively; h. node with fruit; i. seed (all: De Wilde & Duyfjes 21757).
Ovary (densely) hairy, hairs 1– 2 mm long. Fruits ± glossy with sparse hairs 1– 2 mm
long.
Distribution — Sulawesi (Minahassa); northern Moluccas (Bacan, Ternate, Sula Is-
lands, Buru, Ambon); New Guinea (West Papua (Vogelkop Peninsula)).
Habitat & Ecology — Sea level to 600 m altitude; flowering and fruiting throughout
the year.
b. subsp. tomentosa W.J.de Wilde & Duyfjes,

Mukia rumphiana (Scheff.) W.J.de Wilde & Duyfjes subsp. tomentosa W.J.de Wilde & Duyfjes, Thai
Forest Bull., Bot. 34 (2006) 47, f. 3, 4f. — Type: Van Balgooy 4782 (holo BO; iso K, KYO, L),
Buru.
Ovary and fruits densely velvety grey-hairy, hairs to 0.5 mm long. — Fig. 46.
Distribution — SW Sulawesi, Moluccas (Buru).
Habitat & Ecology — Limestone area; 500 –1000 m altitude; flowering and fruiting
August to January.
Note — The distribution of the subspecies rumphiana and tomentosa seem to ex-
clude each other, but both subspecies are found on Buru.
26. NEOACHMANDRA
Neoachmandra W.J.de Wilde & Duyfjes, Blumea 51 (2006) 12, f. 1c, 2c; t. 1. — Type species: Neoach
mandra japonica (Thunb.) W.J.de Wilde & Duyfjes (Melothria japonica Thunb.).
Achmandra Arn., Madras J. Lit. Sci. 12 (1840) 49, p.p., not for the lectotype which is Kedrostis; Arn.,
J. Bot. 3 (1841, ‘Aechmandra’) 274.
Small climbers, usually annual, leafy stem 0.5 – 2 mm diam., plant green on drying;
monoecious. Probract absent. Tendrils unbranched. Leaves simple (seemingly 5-fo-
liolate in N. pentaphylla, New Caledonia), palmiveined. Flowers 5(–10) mm diam.,
petals usually valvate in bud, (creamy-)white, free. Male inflorescences consisting of
1– 4(– 8) long-pedicelled flower(s) at the node, usually co-axillary with one (or more)
long-pedicelled female flower(s); bracts absent. Male flowers: pedicel (3 –)10 – 50 mm
long, persistent; receptacle-tube campanulate; sepals minute, narrowly elliptic or linear,
usually recurved; petals (narrowly) elliptic; stamens (2 or) 3, inserted in the upper half
of the receptacle-tube, filaments short, anthers all 2-thecous, included or ± exserted, the-
cae lateral, straight, ± divergent, connective broad, at apex truncate or acutely angular,
or sometimes minutely produced; disc free, (depressed-)globose. Female flowers 1 or
2, frequently co-axillary with previously developed male flower(s); pedicel (short or)
long; ovary globose or ellipsoid-fusiform, usually with slender neck at apex, glabrous;
stigmas 3 or stigma consisting of (2 or) 3 almost free lobes, lobes entire or lobed, papil-
lose-hairy; staminodes usually present; disc annular, free from receptacle-tube. Fruit
1 (or 2), usually with long fruiting pedicel, globose, ellipsoid, narrowly ellipsoid or
fusiform, 0.5 –7 cm long, apex beaked or not, glabrous, (pink-)white or red, juicy or
pulpy; dry exocarp membranous or cartilaginous, smooth or minutely pustulate. Seeds
few or numerous, pale, compressed (elsewhere also globose), ovate-elliptic, faces not
or but little convex, not sculptured, margin absent or obscure, with entire rounded edge,
base of seed without or with a short or long wing.
Distribution — A genus of c. 30 species distributed in the tropics of the Old World:

Africa, Madagascar, and in SE Asia from India, China, Japan south-east to Australia
and the Pacific; in Malesia: 12 species.
key to the species
1a. Petals more than 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b. Petals less than 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2a. Leaf blade (narrowly) triangular, at base broadest and truncate . . . . . . . . . . . . . 3
b. Leaf blade narrowly elliptic, at base narrowed. — New Guinea . 7. N. lancifolia
3a. Male flowers 2 – 4 on each node (female flowers not seen). Receptacle-tube 3.5
mm long. — Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. N. macrantha
b. Male flowers single (beside 1 female flower) on each node. Receptacle-tube c.
1.5 mm long. — New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . 3. N. clemensiae
4a. Leaf blade (3 –)5-lobed, lobes to (1/5 –)halfway deep or more . . . . . . . . . . . . . . 5
b. Leaf blade lobed less than halfway deep or leaves hastate or unlobed . . . . . . . . 6
5a. Fruit ellipsoid, 1(–1.5) cm long. — Philippines . . . . . . . . . . . . 2. N. boholensis
b. Fruit globose, 0.5(–1) cm diameter. — New Guinea . . . . . . . . . . . . 4. N. filipes
6a. Fruit fusiform, long-tapering at base and apex . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Fruit globose, ellipsoid or short-cylindrical, at base short-cuneate or more or less
rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7a. Fruit 5 – 6 cm long. — Papua New Guinea . . . . . . . . . . . . . . 10. N. morobensis
b. Fruit 2.5 – 4.5 cm long. — Java, Lesser Sunda Islands . . . . . . . . . . 1. N. backeri
8a. Fruit whitish. Seeds 5.5 –7 mm long. — Submontane; South Sumatra, Java . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. N. leucocarpa
b. Fruit whitish or green. Seeds 2 – 5 mm long. — Lowlands . . . . . . . . . . . . . . . . 9
9a. Leaf blade narrow, long-triangular, mostly conspicuously long-bristly hairy on
the veins beneath, hairs 0.5 –1.5 mm long. Ovary sparsely hairy. — Philippines .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. N. scaberrima
b. Leaf blade broader, less conspicuously hairy on veins beneath, hairs 1 mm long
or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Fruit 2 – 4 cm long, often irregularly speckled or blotched. — East Java, Lesser
Sunda Islands, Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. N. nesophila
b. Fruit 1– 2 cm long, not speckled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11a. Fruit ellipsoid, 1.5 – 2 cm long, green. Dry exocarp opaque, seeds not shining
through. — Moluccas(?), Papua . . . . . . . . . . . . . . . . . . . . . . 5. N. idenburgensis
b. Fruit globose or short-ellipsoid, 1–1.5 cm long, whitish. Dry exocarp filmy, trans-
parent, when dry seeds mostly shining through. — Widespread . . . . 6. N. indica
1. Neoachmandra backeri W.J.de Wilde & Duyfjes

Neoachmandra backeri W.J.de Wilde & Duyfjes, Blumea 51 (2006) 14. — Type: Jaag 466 (holo L),
East Moluccas, Alor Islands.
Annual (or subperennial) 1– 2 m long climber, leafy stem 0.5 –1 mm diam., sparsely
finely woolly pubescent, glabrescent. Leaves: petiole 1– 2(– 3) cm long, sparsely scab-
rous-hairy; blade 3- or 5-angular, or (deeply) 3-lobed, 2 – 5(–7) cm long and nearly

as wide, above scabrous-hairy or with cystoliths, base broadly rounded or (deeply)
cordate, margin shallowly repand-dentate. Male flowers 1 (or 2) per node, co-axillary
with later developing female flower; pedicel 8 –15 mm long; expanded perianth 5 –10
mm diam.; receptacle-tube 1.5 – 2 by 1.5 – 2.5 mm, inside at throat finely woolly-hairy;
sepals 1–1.5 mm long; petals 2.5 – 5 mm long, subacute, the margin and upper surface
minutely papillose-hairy; stamens inserted halfway in the receptacle-tube, filaments 1
mm long, anthers 1.5 mm long and wide, connective wide at apex, minutely produced in
the middle; disc 1–1.5 mm diameter. Female flowers (subsp. balinensis): single; pedicel
10(– 20) mm long; ovary narrowly ellipsoid-fusiform, 8 –12 by 1–1.5 mm, glabrous; re-
ceptacle-tube broadly campanulate; style 1.5 mm long, stigma 2 – 5 mm diam., consist-
ing of 3 deeply 2-lobed parts, densely (long-)lanose-papillose; staminodes lanceolate,
c. 1.5 mm long; disc c. 1 mm high. Fruit solitary, narrowly ellipsoid-fusiform, 2.5 – 4.5
by 0.8 –1 cm; dry pericarp (exocarp) cartilaginous, light brown, faintly or distinctly
(5- or) 6-ribbed; fruiting pedicel 2 – 4.5 cm long. Seeds numerous, (narrowly) elliptic,
3 – 3.5 by 1.5 – 2 mm, with faint paler margin.
Distribution — Java (Kangean Archipelago), Sulawesi, Lesser Sunda Islands, Moluc-
cas (see further under the subspecies).
1a. Expanded male corolla 5 –7 mm diam.; petals 2.5 – 4 mm long. — Lowlands . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. subsp. backeri
b. Expanded male corolla 8 –10 mm diam.; petals 4 – 5 mm long. — Montane . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. balinensis
a. subsp. backeri
Expanded corolla 5 –7 mm diameter. Fruit 3 – 5 cm long. — Fig. 3d.
Field-notes — Fruits said to be red when ripe and edible (Backer 28153).
Distribution — Java (Kangean Archipelago), Sulawesi (Djampea Is.), Lesser Sunda
Islands (Alor Kecil), Moluccas (Tanimbar Islands: Jamdena Is.).
Habitat & Ecology — Climber in open scrub and on beaches; sea level; flowering
and fruiting March to June.
b. subsp. balinensis W.J.de Wilde & Duyfjes

Neoachmandra backeri W.J.de Wilde & Duyfjes subsp. balinensis W.J.de Wilde & Duyfjes, Blumea
51 (2006) 15. — Type: De Wilde & Duyfjes 21732 (holo L; iso BO), Bali (Bedugul, growing wild
in forest edges in the Botanical Garden).
Expanded corolla 8 –10 mm diameter. Fruit c. 3 cm long.

Field-notes — Tiny climber. Fruits dark green, possibly red when ripe.
Distribution — Lesser Sunda Islands (Bali).
Habitat & Ecology — Climbing on tree trunks in forest edges; 1400 –1800 m.
2. Neoachmandra boholensis (Merr.) W.J.de Wilde & Duyfjes

Neoachmandra boholensis (Merr.) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 15. — Melothria bo
holensis Merr., Philipp. J. Sci. 29 (1926) 495. — Type: Ramos BS 42779 (holo UC; iso K), Philip-
pines, Bohol (Kalingohan).
Annual 1– 2 m long climber, leafy stem c. 1 mm diam., sparsely fine-hairy, glabres-

cent. Leaves: petiole 1–1.5 cm long, bristly hairy; blade subcircular in outline, 2 – 6
cm diam., (3 –)5-lobed to 1/5 deep to nearly to the base, segments narrowly elliptic or
lanceolate, upper surface scabrous, with cystoliths, lower surface bristly hairy along the
veins, base truncate or shallowly cordate, margin entire or coarsely shallowly repand-
dentate, apex acute. Male flowers 1 (or 2) per node, usually co-axillary with 1 (or 2)
longer-pedicelled female flowers; pedicel 10 –12 mm long; expanded perianth c. 5 mm
diam.; receptacle-tube 2 – 2.5 by 2.5 – 3 mm, at throat finely hairy; sepals 1 mm long (or
less); petals 2 – 2.5 mm long, blunt or rounded, inside at apex papillose-hairy; stamens
inserted close to the throat of the receptacle-tube, filaments c. 1/4 mm long, anthers
c. 1 by 1 mm, connective broad, at apex truncate, faintly produced in the middle; disc
(0.5 –)1 mm diameter. Female flowers: pedicel 15 – 25 mm long; ovary narrowly el-
lipsoid, c. 5 by 2 mm, subglabrous (with sparse hairs); style 1(–1.5) mm long, with 3
style-arms c. 0.5 mm long, each with a subellipsoid papillose stigma-lobe of c. 1 mm
diam.; staminodes less than 1 mm long. Fruit 1 (or 2), ellipsoid, 1.5 – 2 by 1–1.5 cm,
base rounded, apex narrowly rounded; dry exocarp thin, cartilaginous, almost smooth,
pale brown; fruiting pedicel slender, 2 – 2.5(– 4) cm long. Seeds numerous, elliptic, c.
3 by 1.5 – 2 mm, margin faint.
Field-notes — Corolla white. Ripe fruits green.
Distribution — Philippines (Bohol, Mindanao, Sulu Archipelago).
Habitat & Ecology — Climber in recent clearings, also in mangrove forest; sea level
to 600 m; flowering and fruiting throughout the year.
Note — Except for the deeply lobed leaves and rather large fruits, this species comes
close to the widespread N. indica of which it is possibly only a regional form.
3. Neoachmandra clemensiae (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes

Neoachmandra clemensiae (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 17.
— Melothria clemensiae Merr. & L.M.Perry, J. Arnold Arbor. 29 (1948) 168. — Type: M.S. Clemens
41287 (holo A, not seen; iso MICH), Papua New Guinea, Morobe (Sugu-Gaeng).
Annual or subperennial 2 m long climber, leafy stem c. 1 mm diam., sparsely fine-

hairy, glabrescent; ?polygamous (see under female flowers). Leaves: petiole 0.9 –1.4
cm long, densely bristly hairy; blade unlobed, narrowly triangular, 4 – 6 by 2 – 3 cm,
upper surface scabrous with cystoliths, lower surface sparsely scabrous-hairy, on veins
hairs longer and denser, base broadly truncate, margin entire, with short teeth less than
1 mm long. Male flowers single, or often co-axillary with 1 female flower, glabrous;
pedicel c. 20 mm long; expanded perianth c. 15 mm diam.; receptacle-tube c. 1.5 by
3 mm, at throat hardly hairy; sepals 0.5(–1) mm long; petals 6 –7 by 2 – 3(– 4) mm,
subacute, subglabrous; stamens inserted slightly below the throat of the receptacle-
tube, filaments less than 0.5 mm long, anthers 1.5 mm long, 1.5 mm wide, connective
broad, at apex truncate, not or slightly produced; disc depressed globose, less than
1 mm diameter. Female flowers: pedicel 30 – 40(– 45) mm long; ovary much elongated,
12(–15) by 1–1.5 mm, at apex contracted into a narrow neck 2.5 – 3 mm long, glabrous;
receptacle-tube c. 2.5 by 3.5 mm; style 1.5(– 2) mm long, stigma-lobes each c. 1.5 mm
diam.; staminodes c. 2 mm long, inserted slightly above the base of the receptacle-tube,
each consisting of a 1–1.5 mm long filament with at apex a reduced 2-thecous anther
0.5 mm long, lined with few hairs; disc nearly 1 mm high, the upper rim ± undulating.
Fruit unknown.
Distribution — New Guinea (Papua New Guinea, Sugu-Gaeng, Morobe Province);
known only from the type.
Habitat & Ecology — Along mountain trail on wet place; 1000 –1200 m; flowering
in April.
Note — The type and only known collection M.S. Clemens 41287 is a richly flower-
ing specimen, with one male and one female flower at each node. The female flowers
all seem to be hermaphroditic, as the staminodes bear well-developed anthers, though
much smaller than those of the male flowers. The staminodes are much lower inserted
in the receptacle-tube compared to the stamens in the male flowers. The thecae in the
staminodes apparently do not open, and do not produce fertile-looking pollen.
4. Neoachmandra filipes (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes

Neoachmandra filipes (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 18. — Melo
thria filipes Merr. & L.M.Perry, J. Arnold Arbor. 29 (1948) 167. — Type: M.S. Clemens 11027 (holo
A; iso MICH), Papua New Guinea, Morobe (Wantoat).
Annual or subperennial 1– 2 m long climber; leafy stem 0.5 –1 mm diam., sparsely

hairy. Leaves: petiole 1– 3 cm long, finely rigidly hairy; blade circular or broadly ovate
in outline, 4 –11 cm diam., palmately (3 –)5-lobed to c. 3/4 deep to (nearly) to the base,
lobes narrowly elliptic or lanceolate (or linear), mid-lobe usually largest, up to 8 by
1(– 2) cm, upper surface densely ± appressedly scabrous-hairy, lower surface bristly
hairy on the veins, cystoliths not obvious, base truncate or widely shallowly emarginate,
margin entire or widely minutely repand-dentate. Male flowers single, co-axillary with a
single somewhat later developing longer-pedicelled female flower; pedicel 12 – 20 mm
long; expanded perianth 6 – 8 mm diam.; receptacle-tube 1.5 by 2 mm, finely hairy in the
throat; sepals 1 mm long; petals 3 – 4 mm long, subacute, inside very finely hairy; sta-
mens inserted halfway the receptacle-tube, filaments slender, 1(–1.5) mm long, anthers
0.8 by 1 mm, thecae diverging to the apex, connective broad, at apex angularly protrud-
ing; disc depressed globose, 1 mm diameter. Female flowers: pedicel 15 – 25(– 40) mm
long, filiform; ovary nearly globose, 2 mm diam., glabrous, at apex contracted into a
narrow neck 1 mm long; receptacle-tube 1.5 by 2 mm, finely hairy in the throat; sepals
0.5(–1) mm long; petals 2 – 2.5 mm long, inside papillose-hairy; style 1.5 mm long,
stigma 3-lobed, depressed-globose, nearly 1 mm diam., papillose-hairy; staminodes 0.3
mm long; disc 0.5 mm high. Fruit solitary, subglobose, 0.5(–1) cm diam.; dry exocarp
thin, papery, smooth, pale brown; fruiting pedicel very slender, 1.5 – 6(– 8) cm long.
Seeds 4 –10 per fruit, ovate, 5 by 3 – 3.5 mm, smooth, with thickened rounded margin.
Field-notes — Locally common. Flowers creamy or white and the ripe fruits pink or
orange.
Distribution — New Guinea (Papua New Guinea (Central, Western Highlands, East-

ern Highlands, Morobe, and Madang Provinces)).
Habitat & Ecology — In tall grassland, clearings and forest edges in montane area;
1000 – 2000 m; flowering and fruiting throughout the year.
5. Neoachmandra idenburgensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes

Neoachmandra idenburgensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 21.
— Melothria idenburgensis Merr. & L.M.Perry, J. Arnold Arbor. 30 (1949) 57. — Type: Brass
14100 (holo A; iso BM, BO), West Papua, Idenburg River.
Annual c. 1 m long climber; leafy stem c. 1 mm diam., glabrescent. Leaves: petiole

1– 2.5 cm long, (sparsely) hairy; blade ovate, (long) triangular or subhastate, 3.5 –11 by
2 – 6 cm, upper surface finely scabrous with minute cystoliths, lower surface smooth,
veins sparsely finely hairy, base with broadly angular sinus up to 2 cm deep, basal lobes
subobtuse. Male flowers 1 or 2, frequently co-axillary with one equally long-pedicelled
female flower (or fruit); pedicel 7–12 mm long, (sub)glabrous; expanded perianth c. 7
mm diam.; receptacle-tube 2 by 2 mm, glabrous, minutely hairy in the throat; sepals 1
mm long; petals 2.5 – 3 mm long, inside and on margin towards apex minutely glandu-
lar-hairy; stamens inserted close to the throat of the receptacle-tube, filaments 0.3 mm
long, anthers 1.3 by 1.3 mm, connective truncate, faintly protruding in the middle; disc
depressed globose, 1 mm diameter. Female flowers not seen (ovary described in original
description as fusiform). Fruit solitary, co-axillary with pedicel of dropped male flower,
ellipsoid, 1.5 –1.7 by 1–1.2 cm, base rounded, apex (narrowly) rounded with minute
point; dry exocarp thinly pergamentaceous, opaque, not transparent; fruiting pedicel
c. 1 cm long. Seeds numerous, ovate-elliptic, 3.5 – 4 by 2 – 2.5 mm, with narrow paler
margin, basal wing c. 2 mm long.
Distribution — New Guinea (West Papua, Idenburg River, Bernhard Camp); known
only from the type.
Habitat & Ecology — The herbarium label mentions the plant as “occasional in
fringe vegetation along stream in rainforest”; at 50 m altitude; flowering and fruiting
in April.
6. Neoachmandra indica (Lour.) W.J.de Wilde & Duyfjes

Neoachmandra indica (Lour.) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 21. — Melothria indica
Lour., Fl. Cochinch. (1790) 35; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 297. — Aechman
dra indica (Lour.) Arn., J. Bot. (1841) 274. — Zehneria indica (Lour.) Keraudren in Aubrév. &
J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 52, pl. 9: 5 – 8; W.J.de Wilde & Duyfjes,
Thai Forest Bull. Bot. 32 (2004) 19. — Type (Keraudren, 1975): Loureiro s.n., Tourane, Vietnam;
untraced. Neotype: Squires 14 (BM, designated by Jeffrey, Kew Bull. 34 (1980); iso K, UC, W),
Vietnam, Hue.
Bryonia geminata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 924. — Type: Zippelius 1564 (lecto L, desig-
nated by De Wilde & Duyfjes (2006)), West Java.
Cucumis luzonicus Blanco, Fl. Filip. (1837) 861; Merr., Sp. Blancoan. (1918) 470. — Type: Merrill:
Species Blancoanae 848 (neo US, designated by De Wilde & Duyfjes (2006), not seen; iso K, L,
W), Rizal Province, Luzon.
Subannual slender 0.5 –1.5 m long creeper or climber, largely glabrescent, hairs less
than 0.5 mm long. Leaves: petiole 1– 3 cm long, finely hairy or glabrescent; blade tri-
angular or (deeply) hastately 3(– 5)-lobed, 2.5 –7(–10) by 3 – 5.5(–7) cm, base broadly
shallowly cordate or subtruncate, margin faintly dentate. Flowers: perianth 4 –7 mm
diam., male flowers solitary (or paired?), usually co-axillary with 1 or 2 previously de-
veloped, longer-pedicelled female flower(s), glabrous (but sparsely gland-dotted). Male
flowers: pedicel slender, 10 –15 mm long; receptacle-tube ± narrowly cup-shaped, 1.5 –
2 by 1.5 – 2 mm, glabrous, except for hairy fringe at throat inside; sepals (0.5 –)1 mm
long, glabrous, recurved; petals 2 – 3.5 by 1.5 – 2 mm, inner and outer surface minutely
gland-hairy and papillose; filaments 0.5 mm long, glabrous, inserted 0.5(–1) mm below
the throat of the receptacle-tube, anthers obovoid-rhomboid, 1–1.5 mm diam., thecae
straight, 1 mm long, connective broad, broadest at apex, subtruncate with short 0.2 mm
projection, (partly) finely hairy; disc elongated, ± obovate-elliptic, 1 mm long. Female
flowers: pedicel (10 –)15 – 30 mm long; ovary ellipsoid to narrowly ellipsoid, 3(– 4) by
1.5 – 2 mm, glabrous, neck 0.5 –1 mm long; perianth as in male flowers; style 1.5(– 2)
mm long, stigma 1 mm diam., composed of 3 apically deeply notched lobes, papillose;
staminodes linear, 1 mm long, inserted halfway the tube; disc thick-carnose, 1 mm high.
Fruit 1 or 2 per node, ripening whitish, subglobose or (short) ellipsoid, not apiculate,
0.8 –1.2 by 0.8 –1 cm; exocarp thinly leathery or membranous (when dry often leaving
the seeds shining through); fruiting pedicel 1.5 – 3 cm long. Seeds 15 – 25 per fruit, ±
obovate, 2 – 4 by 2.5 – 3 mm, smooth, unmargined, with entire, rounded edge.
Distribution — Widespread: from South India and South China east to West Malesia;
in Malesia: Borneo, Java, and the Philippines; not known from Sumatra and Peninsular
Malaysia. In northern India the distinction with N. odorata is not sharp, as is the distinc-
tion with N. japonica in southern China.
Habitat & Ecology — In and along waste gardens, forest edges, shaded (damp)
roadsides; 0 – 500(–1400) m; flowering and fruiting mainly July to December.
7. Neoachmandra lancifolia W.J.de Wilde & Duyfjes

Neoachmandra lancifolia W.J.de Wilde & Duyfjes, Blumea 51 (2006) 22. — Type: Paijmans 1107
(holo CANB; iso LAE), Papua New Guinea, south side of Goodenough Bay, Milne Bay Province.
Annual(?) herbaceous c. 2 m long climber; leafy stem c. 1 mm diam.; early gla-

brescent. Leaves: petiole 0.5 – 0.9 cm long, with dense, scabrous, upwards-directed
hairs; blade lanceolate, parallel-5-nerved, 5 – 8 by 0.5 –1 cm, both surfaces densely
short-scabrous hairy, hairs directed towards the apex, base narrowed but ± rounded or
subcordate, margin with sparse teeth 0.5 mm long or less, apex subacute with mucro.
Male flowers 3 – 5 per node (female flowers not present), subglabrous; pedicel 20 – 25
mm long; expanded perianth c. 15 mm diam.; receptacle-tube narrowly campanulate,
3.5(– 4) by c. 2.5 mm, inside at throat densely finely hairy; sepals 2.5 – 3 mm long, with
an odd hair; petals 6 – 8 by 3 – 4 mm, along the margin minutely papillose; stamens
inserted at c. 1/3 from the apex in the receptacle-tube, filaments slender, (1–)1.5 mm
long, anthers c. 1.5 by 1 mm, exserted, hairy along the thecae, connective broad, with
truncate apex, minutely produced in the middle; disc obovoid, c. 1.5 high, 1 mm wide.
Female flowers and fruits not seen.
Distribution — New Guinea (Papua New Guinea: Milne Bay Province, south side
of Goodenough Bay); known only from the type.
Habitat & Ecology — In grassland on plateau; at c. 250 m altitude.
Note — This species seems closely related to N. scaberrima which has much smaller
flowers.
8. Neoachmandra leucocarpa (Blume) W.J.de Wilde & Duyfjes

Neoachmandra leucocarpa (Blume) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 23. — Bryonia leu
cocarpa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 924. — Bryonopsis? leucocarpa (Blume) Miq., Fl.
Ned. Ind. 1, 1 (1856) 657. — Melothria leucocarpa (Blume) Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 601, p.p.; in Engl., Pflanzenr. 66, 4.275.I (1916) 101, p.p.; Backer in Backer &
Bakh.f., Fl. Java 1 (1964) 297. — Type: Blume s.n. (lecto L (barcode L0130099), designated by
De Wilde & Duyfjes (2006)), Java.
Zehneria deltoidea Miq., Fl. Ned. Ind. 1, 1 (1856) 655. — Melothria rauwenhoffii Cogn. in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 597 (not Melothria deltoidea (Schumach.) Benth. in Hook., Niger
Fl. (1849) 368). — Type: Junghuhn s.n. (holo U, barcode U0001468; iso L, barcode L0130089),
Java, Gunung Merapi.
Zehneria deltoidea Miq. var. subintegerrima Miq., Fl. Ned. Ind. 1, 1 (1856) 655. — Type: Horsfield
s.n. (holo U, barcode U0122933), Java.
Melothria rauwenhoffii Cogn. var. pengalenganensis Hochr., Candollea (1934) 287. — Type: Hochreu
tiner 1316 (holo G; iso L), Java, Pengalengan Plateau.
Annual or subperennial to c. 5 m long climber, widely branched, leafy stem 1– 2 mm

diam.; minutely sparsely hairy, glabrescent. Leaves: petiole 1.5 – 3.5 cm long, scabrid
short-hairy, blade broadly triangular or subhastate, 4 –11 by 3 – 9.5 cm, upper surface
glabrous except for the scabrid-hairy veins, with numerous cystoliths, lower surface
glabrous, without cystoliths, base cordate, truncate or broadly emarginate, margin entire
or shallowly dentate. Male flowers 1 or 2 per node, usually co-axillary with 1 or 2 long-
er-pedicelled female flower(s); pedicel 25 – 45 mm long, glabrous; expanded perianth
8 –10 mm diam.; receptacle-tube c. 3.5 by 3 mm, densely minutely papillose-hairy in
the throat; sepals 1 mm long; petals 3 – 5 mm long, subacute, inside minutely papillose-
hairy; stamens inserted near the throat in the receptacle-tube, filaments 0.5 mm long,
anthers c. 2 by 1.5 mm, connective broad, at apex truncate with a short mucro in the
middle; disc ± globose, 1–1.5 mm diameter. Female flowers 1 or 2; pedicel (10 –)30 – 60
mm long; ovary fusiform, c. 9 by 3 mm, glabrous, with a c. 2 mm long neck; style
c. 4 mm long, stigma depressed globose, 1.5 by 1 mm, 3-lobed, and each lobe again
2-lobed, papillose; staminodes 0.5 mm long; disc 1 mm high, shallowly 3-lobed. Fruit
1 or 2, ripening pink-white, subglobose or ellipsoid, 1– 2 by 0.8 –1.5 cm, base subobtuse,
apex (sub)obtuse, 1 mm beaked; dry exocarp thin, pale brown, smooth; fruiting pedicel
(1–)4 – 6 cm long. Seeds 5 –12 per fruit, elliptic, 6 –7 by c. 4 mm, margin faint, without
wing. — Fig. 47; Plate 18a – c.
Field-notes — Lower leaf surface very pale. Anthers dorsally with a ridge (as indi-
cated in a drawing by Kuhl & Van Hasselt s.n., barcode L0130076). Fruits at first green-
ish white, ripening pink-white. Cotyledons in the seedling measure c. 1.5 by 1 cm.
Distribution — Central and southern Sumatra (3 collections), all over Java.
Habitat & Ecology — Forest edges and damp or shady places in montane forest,
often near streams; 850–1750 m; flowering and fruiting throughout the year.
2 cm 2 cm
b
1 mm
c
1 mm f f'
2 mm
2 mm
1 mm 1 mm
i h
g e d
Fig. 47. Neoachmandra leucocarpa (Blume) W.J.de Wilde & Duyfjes. a. Twig with male and female
flowers; b. twig with fruits; c. seedling; d, e. male flowers; f, f'. stamens, showing thecae with broad
connective, ad- and abaxially respectively; g. female flower; h. fruit; i. seed (a, b, d – g: De Wilde &
Duyfjes 21843; c, h, i: De Wilde & Duyfjes 21946).
9. Neoachmandra macrantha W.J.de Wilde & Duyfjes

Neoachmandra macrantha W.J.de Wilde & Duyfjes, Blumea 51 (2006) 25. — Type: Ramos & Edaño
BS 40215 (holo P; iso K, UC), Philippines, Luzon, Mt Data.
Possibly annual 1– 2 m long climber; leafy stem 0.5 –1 mm diam.; sparsely hairy.
Leaves: petiole 0.5 –1.5 cm long, scabrous-hairy in upper midrib groove; blade entire,
(narrowly) triangular, 4 – 6.5 by 2 – 3.5 cm, upper and lower surface scabrous with short
hairs and cystoliths, veins with scabrid hairs, base truncate, margin shallowly sparsely
repand-dentate, teeth 1–1.5 mm long, apex long-acuminate with short mucro. Male
flowers: 2 – 4 (of different ages) at each node (female flowers not seen), subglabrous;
pedicel 25 – 35 mm long; expanded perianth c. 15 mm diam.; receptacle-tube c. 3.5 by
3 mm, finely hairy in the throat; sepals 1–1.5 mm long; petals (6 –)7 by (3 –)4 mm, apex
subacute, inside papillose-hairy; stamens inserted about halfway the receptacle-tube,
filaments 1–1.5 mm long, anthers 1.5 by 1 mm, connective broad, at apex truncate and
faintly produced in the middle; disc (depressed) globose, 1 mm diameter. Female flow
ers and fruit unknown.
Distribution — Philippines, Luzon, Mt Data; known only from the type; flowering
in September.
10. Neoachmandra morobensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes
Neoachmandra morobensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 25.
— Melothria morobensis Merr. & L.M.Perry, J. Arnold Arbor. 29 (1948) 167. — Type: M.S. Cle
mens 11330bis (holo MICH; iso A), Papua New Guinea, Morobe Province.
Subperennial 2 – 3 m long climber; leafy stem 1(– 2) mm diam.; minutely hairy, gla-
brescent. Leaves: petiole (1.5 –)2 – 4.5 cm long, densely harshly hairy, blade entire or
3-lobed up to halfway, broadly or narrowly triangular or subhastate, 4 –10 by 5 –14 cm,
upper surface densely (appressed) grey hairy or scabrous, with or without cystoliths,
lower surface sometimes less hairy, without cystoliths, base broadly rounded, truncate
or broadly emarginate with shallow sinus, margin entire or shallowly repand-dentate,
apex long-acuminate; 7 palmately veined. Male flowers 1– 4 per node, usually co-axil-
lary with 1 female flower; pedicel 15 – 25 mm long, sparsely hairy; expanded perianth
7– 9 mm diam.; receptacle-tube 2 – 2.5 by 2.5 – 3 mm, minutely papillose-hairy in the
throat (hairs less than 0.1 mm long); sepals 1.5 – 2.5 mm long; petals 2.5 – 3 mm long,
minutely hairy inside; stamens inserted at about 1/3 from the apex in the receptacle-
tube, filaments 0.5(– 0.7) mm long, anthers 1.5 by 1 mm, connective broad, at apex
slightly convex or straight; disc subglobose, flattened at apex, 1 by 1.5 mm. Female
flowers single; pedicel 12 –15 mm long; ovary narrowly ellipsoid; perianth immature.
Fruit solitary, narrowly ellipsoid, fusiform, 4.5 – 5.5(– 6) by (1–)1.5 – 2 cm, base acute,
apex beaked; dry exocarp coriaceous, (faintly) 6-ribbed or -lined; fruiting pedicel very
slender, c. 1.5 cm long. Seeds numerous, 3.5 – 4 by 2 – 2.5 mm, margin narrow, faint, at
base with a wing. — Fig. 48c, c'.
Field-notes — Flowers white, creamy or pale yellow; androecium orange. Fruits
glaucous, pale green, darker striped.
Distribution — New Guinea (Papua New Guinea: Central and Morobe Provinces).
Habitat & Ecology — In abandoned gardens, clearings in rainforest, disturbed forest,

along creeks; at 50 –1300 m altitude; flowering and fruiting throughout the year.
Note — The seeds in Neoachmandra have in the dried state often a wing-like ap-
pendage, possibly formed by the fruit pulp (endocarp), but in N. morobensis this is more
obvious than in other species. The pale yellow colour mentioned on a field label needs
further confirmation.
11. Neoachmandra nesophila W.J.de Wilde & Duyfjes

Neoachmandra nesophila W.J.de Wilde & Duyfjes, Blumea 51 (2006) 26. — Type: Soejatmi 10 (holo
BO; iso A, K, L, SING, US), Java.
Annual or subperennial 1– 3 m long climber; leafy stem 1–1.5 mm diam.; with sparse
minute hairs or glabrous. Leaves: petiole 1– 3(– 5) cm long, subglabrous or sparsely
or densely fine-hairy; blade ovate-triangular or (long-)triangular, 2.5 – 9 by 2.5 – 8 cm,
2 mm
a' b' c'
3 mm
b
c
a
Fig. 48. Fruits and winged seeds of Neoachmandra. — a, a', b, b'. Neoachmandra nesophila W.J.de
Wilde & Duyfjes. — c, c'. Neoachmandra morobensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes
(a, a': Soejatmi 10, Java, type; b, b': De Wilde & Duyfjes 21935, Lombok; c: Takeuchi & Ama 16328,
Papua New Guinea; c': Streimann & Kairo NGF 30905, Papua New Guinea).
sometimes broadly hastately 3-lobed, lateral lobes to 3 by 5 cm, upper and lower sur-
face (sub)glabrous, with or without (small) cystoliths and minute sparse hairs on the
veins, base broadly rounded, truncate or widely cordate, margin entire or shallowly
sparsely dentate. Male flowers 1(– 3) per node, frequently co-axillary with a single
female flower; pedicel 10 –15 mm long, (sub)glabrous; expanded perianth 6 – 9 mm
diam.; receptacle-tube 2 – 2.5 by c. 2.5 mm, outside glabrous, densely hairy in the throat
(hairs 0.5 mm long); sepals 0.5 –1 mm long; petals c. 3 mm long, outside glabrous or
with some stout hairs, inside sparsely gland-hairy; stamens inserted close to the throat
in the receptacle-tube, filaments 0.5 mm long, anthers 1–1.5 mm diam., exserted, con-
nective ± truncate and at base with a conspicuous crest-like outward protrusion of c. 1
mm long (always?, see note 3); disc depressed obovoid, 1–1.5 mm long. Female flowers
1 (frequently co-axillary with male flower) or 2; pedicel usually much longer than in
male, 25 – 45 mm long, (sub)glabrous; ovary narrowly ellipsoid, 8 –10 by 1.5 – 2(– 2.5)
mm, base acutish, narrowed into a slender neck 2.5 – 3.5 mm long, glabrous; recepta-
cle-tube 2 – 3.5 by 2.5 – 3.5 mm; style c. 2 mm long, stigma subglobose, c. 2 mm diam.,
3-lobed, each lobe deeply 2-lobed, papillose; staminodes subulate, c. 1 mm long; disc
less than 1 mm high. Fruit solitary (or 2), ripening green-red or green with darker stripes
or yellow, narrowly ellipsoid, 2 – 4 by 1.2 –1.5 cm, base rounded, apex ± narrowed into
an acute-acuminate c. 5 mm long beak; dry exocarp usually with small darker patches
and spots arranged in longitudinal rows; fruiting pedicel 2 – 6 cm long. Seeds numerous,
3.5 – 4.5 by 2 – 3 mm, margin faint, at base with a wing. — Fig. 48a, a', b, b', 49; Plate
18d, 19a, b.
Distribution — East Java; Philippines (Luzon, Mindoro); North Sulawesi; Lesser
Sunda Islands (Lombok, Sumbawa, Flores, Timor (deviating, see note 1)); Moluccas
(Halmahera, Banda).
Habitat & Ecology — Damp thickets and forest edges, near streams; at low altitudes
to 500(– 2100) m; flowering and fruiting mainly from January to July.
Notes — 1. Two collections from Timor, Forbes 3919 & 3957, from c. 700 m alti-
tude, deviate in the petiole and lower leaf surface, bearing rather dense and soft hairs,
and in the much longer male pedicels which are 20 – 60 mm long. The seeds are similar
to those of N. nesophila. More material is needed to elucidate whether we are dealing
with a separate taxon.
2. Neoachmandra nesophila, as it is conceived here, comprises specimens with an
unusual variation in fruit size: 2 – 4 cm long. A number of specimens from the same
distributional area, mostly rather incomplete, resemble N. nesophila but have smaller
fruits, 1.5 – 2 cm long, without the ornamentation of fine darker-coloured patches and
dots. These specimens also resemble the type and only collection of N. idenburgensis,
from West Papua, although the latter is so much different in other characters from
N. nesophila that it is retained here as a distinct species. The true status of the smaller-
fruited specimens alluded to above remains uncertain. A choice of these specimens is:
Clemens 18282 (Luzon); Conklin & Del Rosario PNH 72640 (Mountain Province, 2000
m); Ramos & Edaño BS 44466 (Sulu Islands); Gaerlan PPI 23101 (Mindanao, 2110
m); De Haan 1769 (Halmahera, 20 m).
3. In De Wilde & Duyfjes 21935, in spirit, from Lombok, the connective is of a firm
but watery texture and bears the described protrusion. This protrusion was not seen in
a
c 2 cm
b
3 mm
1 mm
f
1 mm
Fig. 49. Neoachmandra nesophila W.J.de Wilde & Duyfjes. a, b, c. Male flowering, female flowering,
and fruiting node, respectively; d. male flower mature bud, from outside and opened; e. female flower
from outside and opened; f. fruit (all: De Wilde & Duyfjes 21935).
boiled dried flowers in other collections. Probably the details of the connective largely
get lost during the drying process.
12. Neoachmandra scaberrima (Merr.) W.J.de Wilde & Duyfjes

Neoachmandra scaberrima (Merr.) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 29. — Melothria
scaberrima Merr., Philipp. J. Sci., C (1909) 330. — Type: Elmer 5862 (lecto L, designated by De
Wilde & Duyfjes (2006); iso BO, K, PNH†), Philippines, Luzon, Benguet.
Annual or subperennial 1– 3 m long climber; leafy stem 1–1.5 mm diam.; with

sparse hairs, glabrescent. Leaves: petiole 1– 2.5 cm long, shortly scabrid or harshly
hairy; blade narrowly triangular or sublanceolate, 4 –10 cm long, unlobed, or base short
or long hastately lobed (see also the note), basal lobes up to 5 by 1 cm, upper surface
scabrous by dense coarse appressed hairs, with or without cystoliths, lower surface
less densely hairy, but on the veins with conspicuous grey bristly patent hairs 0.5 –1.5
mm long, rarely shorter, base (sub)truncate, apex (long-)acute, margin entire, minutely
sparsely denticulate. Male flowers 1 or 2 per node, frequently co-axillary with 1 much
longer-pedicelled female flower; pedicel 8 –10 mm long, with scattered 0.2 – 0.3 mm
long hairs; expanded perianth 6 –7 mm diam.; receptacle-tube by 2 mm, outside with
few hairs, minutely hairy in the throat; sepals 0.5 mm long; petals 2 – 2.5 mm long,
outside frequently with few hairs, inside densely glandular-hairy; stamens inserted at c.
1/4 from the apex in the receptacle-tube, filaments 0.5 mm long, anthers 1.5 by 1 mm,
connective broad, truncate, with short mucro in the middle; disc subglobose, 1–1.5 mm
diameter. Female flowers 1 (and then frequently co-axillary with male flower) or 2 per
node, larger than male flowers; pedicel 20 – 40 mm long, sparsely hairy, glabrescent;
ovary narrowly ellipsoid, 5 –7(–10) by 1–1.5(– 2.5) mm, gradually narrowed into a
slender neck c. 2 mm long, sparsely hairy; receptacle-tube 3 – 3.5 by 3 – 3.5 mm; sepals
1 mm long; petals 2.5 – 4 mm long; style c. 3 mm long, with 3 short arms each bearing
a subglobose pilose-hairy stigma of c. 1.5 mm diam.; staminodes 0.5 –1 mm long; disc
1 mm high, ± 3-lobed or not. Fruit solitary (or 2), ripening yellow, (narrowly) ellipsoid,
3 – 3.5 by 1.5 – 2 cm, base shortly acute or subobtuse, apex acute or up to 0.5 cm beaked;
dry exocarp faintly 6-lined near apex; fruiting pedicel 2 – 4.5 cm long. Seeds numerous,
4 – 5 by 2.5 – 3.5 mm, margin faint, at base with a short wing.
Distribution — Philippines: Luzon (Benguet, Rizal); one deviating collection from
Mindanao (see note).
Habitat & Ecology — Thickets, shrubbery, along streams, grassy roadsides; at 300 –
1000 m altitude; flowering and fruiting throughout the year.
Note — Two deviating specimens are: Ramos BS 14608, the only collection from
Mindanao, and Loher 2136, from Central Luzon. Ramos BS 14608 deviates in having
broader triangular or 3 – 5-lobed leaves, and Loher 2136 in having narrow, hastate leaves
with the veins only shortly scabrous hairy, short female pedicels, only 5 –10 mm long,
and smaller fruits, only c. 2 cm long. The latter is likely a local variety, or both may
belong to the specimens discussed under N. nesophila (see there).
27. NEOALSOMITRA
Neoalsomitra Hutch., Ann. Bot. N-S, 6 (1942) 97; W.J.de Wilde & Duyfjes, Blumea 48 (2003) 100;
Fl. Thailand 9, 4 (2008) 484. — Type species: Neoalsomitra sarcophylla (Wall.) Hutch. (= Zanonia
sarcophylla Wall.).
Alsomitra auct. non (Blume) Spach: M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 117, p.p. (non Zanonia
sect. Alsomitra Blume); Hook.f. in Benth. & Hook.f., Gen. Pl. 1 (1867) 840; Cogn. in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 928; in Engl., Pflanzenr. 66, 4.275.I (1916) 11.
Tiny or stout climbers, annual or perennial, with or without tuberous rootstock;

dioecious or monoecious. Probract absent. Tendrils 2-branched at apex. Leaves: blade
simple or (pedately) foliolate, subcircular in outline, lateral leaflets sometimes smaller
and unequal sided, margin mostly entire. Flowers small; sepals free; petals imbricate
in bud, white or cream, free (or very short-connate at base). Inflorescences lateral
(and terminal), in male mostly many-flowered, paniculate with ultimate branches fine,
raceme-like, pedicels persistent; in female paniculate or raceme-like, fewer-flowered;
bracts present, small. Male flowers rotate or broadly bowl-shaped; receptacle small, flat
or shallow; disc absent or in N. schultzei inconspicuous; stamens 5, inserted centrally,
± out-curved, filaments free or partially or wholly fused, anthers small, 1-thecous,
extrorse, often with a minute dark dot adaxially; disc absent. Female flowers: ovary
cylindrical-clavate, imperfectly 3-locular, ovules 5(–10) per placenta, pendulous, styles
3, short, stigma 2-lobed (deeply lunate); staminodes absent (present in N. plena, Thai-
land). Fruits few or several, green, ripening brown, small- or medium-sized cylindrical-
clavate capsules, apex truncate, 3-valvate, perianth-scar 0.5 – 2(– 3) mm from orifice.
Seeds alternate in each row (wings imbricate), compressed, faces (finely) tubercled or
smooth, margin narrow or broad, sometimes double because of deep circumferential
groove (not or hardly so in N. sarcophylla), edge subentire or finely or coarsely dentate,
with apically a ± oblique membranous translucent wing.
Distribution — A genus of 11 species, distributed from NE India and South China
to Australia and into the Pacific, 1 species, N. clavigera, covering the whole area of the
genus; 6 species in Malesia.
Key to the species
1a. Leaf blade (of adult or fertile shoots) simple, unlobed or lobed . . . . . . . . . . . . . 2
b. Leaf blade (of adult or fertile shoots) foliolate . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2a. Dioecious(?). Plant tuberous. Inflorescences (infructescences) twice branched. Petals
adaxially papillose. Filaments almost completely fused. Fruit glabrous. — Kangean
Archipelago, Timor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. N. hederifolia
b. Monoecious. Plant not tuberous(?). Inflorescences not or once-branched. Petals
adaxially not papillose. Filaments connate in the lower half. Fruit glabrous or
hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Plant, including fruit, densely soft-hairy, drying (dark) brown. — SE Papua New
Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. N. pilosa
b. Plant subglabrous or sparsely hairy, drying green. Fruit glabrous . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. N. schultzei
4a. Leaf blade 3-foliolate, petiole less than 0.5 cm long; leaflets carnose, wrinkled
on drying, apex usually obtuse. Fruit 3–4 cm long. Seeds shortly 2-horned. — SE
Continental Asia, East Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . 4. N. sarcophylla
b. Leaf blade 3- or 5-foliolate, petiole 0.5 – 3(– 5) cm long; leaflets membranous, not
wrinkled on drying, apex (obtuse or) acute. Fruit 2 – 8 cm long. Seeds star-shaped
by coarsely 5 –7 (double) dentate margin (edge) . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Filaments partly united. Fruit c. 2 cm long. Stem towards base thorny. Leaflets
without marginal glands. — East Malesia . . . . . . . . . . . . . . . . 5. N. schefferiana
b. Filaments free. Fruit 4 – 8 cm long. Stem not thorny. Leaflets often with glands on
margin at base of lateral leaflets. — SE Continental Asia including South China
through East Malesia to Australia and the Pacific . . . . . . . . . . . . . . 1. N. clavigera
1. Neoalsomitra clavigera (Wall.) Hutch.

Neoalsomitra clavigera (Wall.) Hutch., Ann. Bot. (Oxford) 6 (1942) 101; C.Jeffrey, Cucurbitaceae
E. Asia (1980) 3; W.J.de Wilde & Duyfjes, Blumea, 48 (2003) 105, f. 1c; Fl. Thailand 9, 4 (2008)
487. — Zanonia clavigera Wall., Numer. List (1831) 3725, nom. nud.; Pl. Asiat. Rar. (1831) 28, pl.
133. — Alsomitra clavigera (Wall.) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 118; C.B.Clarke
in Hook.f., Fl. Brit. Ind. 2 (1879) 634; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881). — Type:
Wallich 3725A (lecto K-Wall., excluding male inflorescence and seeds, designated by De Wilde &
Duyfjes (2003)), Sylhet (Bangladesh).
Melothria trifoliolata F.Muell., Fragm. 5 (1866) 181. — Alsomitra trifoliolata (F.Muell.) K.Schum.,
Notizbl. Königl. Bot. Gart. Berlin 2, 13 (1898) 155; Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916)
15. — Neoalsomitra trifoliolata (F.Muell.) Hutch., Ann. Bot. N-S, 6 (1942) 100; I.Telford, Fl. Aus-
tralia 8 (1982) 165. — Alsomitra hookeri F.Muell., Fragm. 6 (1868) 188, nom. illeg.; (1877) 107
(not seen); Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 933. — Zanonia hookeri F.Muell.,
Syst. Census Austral. Pl. (1882) 76, nom. illeg. — Type: J. Dallachy? in herb. F. Mueller s.n. (iso
L, barcode L0128079), Australia (Rockingham Bay).
Alsomitra beccariana Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 932. — Neoalsomitra bec
cariana (Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 101. — Type: Beccari s.n. (holo FI; iso BR),
Moluccas (Kei Archipelago).
Gynostemma integrifoliola Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 916. — Alsomitra
integrifoliola (Cogn.) Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30 (1911) 121. — Neoalsomitra in
tegrifoliola (Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 99; Merr. & L.M.Perry, J. Arnold Arbot.
30 (1949) 56; M.Jacobs, Blumea 7 (1954) 622; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge,
Laos & Vietnam 15, (1975) 12. — Type: Cuming 517 (lecto K, designated by Keraudren (1975);
iso L, P), Philippines.
Gynostemma elongatum Merr., Philipp. J. Sc., Bot. 3 (1908) 267. — Hemsleya elongata (Merr.) Cogn.
in Engl., Pflanzenr. 66, 4.275.I (1916) 26. — Type: Curran FB 5474I (B†; duplicates not seen),
Philippines (Luzon).
Alsomitra muelleri Cogn., Bull. Acad. Roy. Sci. Belgique, Cl. Sci., sér. 3, 14 (1887) 363; in Engl.,
Pflanzenr. 66, 4.275.I (1916) 15. — Neoalsomitra muelleri (Cogn.) Hutch., Ann. Bot. (Oxford) 6
(1942) 100. — Type: Armit s.n. (herbarium not known, not seen), Papua New Guinea.
Alsomitra rotundifolia Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 13. — Neoalsomitra rotundifolia
(Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 100; Peekel, Fl. Bismarck Archip., translated by
E.E.Henty (1984) 541, f. 863 (2). — Type: Peekel 471 (B†), Papua New Guinea. Neotype: drawing
f. 863 (2) in Peekel (1984) (designated by De Wilde & Duyfjes (2003)).
Alsomitra pubescens Merr., Philipp. J. Sci., C. 13 (1918) 64. — Neoalsomitra pubescens (Merr.)
Hutch., Ann. Bot. (Oxford) 6 (1942) 101. — Type: Mabesa FB 26346 (PNH†), Philippines.
Alsomitra pubigera Prain var. glauca Craib, Fl. Siam. (1931) 768. — Type: Curtis 2504 (not seen),
Langkawi Island.
Alsomitra schefferiana auct. non Cogn.: Peekel, Fl. Bismarck Archip., translated by E.E.Henty (1984)
541, f. 863 (1), based on Peekel 328 (382) (B†); Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 16
(1916) 15, p.p., f. 3a–h; Harms, Bot. Jahrb. Syst. (1925) 152.
Herbaceous or (sub)woody perennial climber to 12 m long, hairy or (nearly) glabrous

all over; dioecious. Leaves: petiole 0.5 – 3(– 5) cm long; petiolules 0.4 – 2 cm long; blade
membranous, blackish on drying; 3- or (pedately) 5-foliolate (i.e. the blade is primarily
3-foliolate and the lateral ones not divided or divided into 2 leaflets, the petiolule of the
outer ones inserted on the petiolule of the inner), (5 –)10 – 20 cm across, leaflets ovate
or (narrowly) elliptic, 12(–15) by 5(– 8) cm, glands 1 or 2, flat, 1– 3 mm diam., usually
towards the base of the outer margin of the outer leaflets, base (obtuse or) attenuate,
apex obtuse or acute, 1– 2 mm mucronate; midvein distinct, lateral veins 4 –7 per side.
Male inflorescences mostly many-flowered, broadly paniculate, axillary, 10 – 30 cm
long, peduncle 1.5 – 4 cm long, branches (glandular-)pubescent or subglabrous, bracts
linear, 0.5 – 2 mm long, the basal ones larger and 3(– 5)-foliolate, to 1 cm long, flowers
up to 5 (sub)fascicled, in condensed terminal racemes. Male flowers: pedicel slender,
5 –10 mm long, (glandular-)pubescent or subglabrous; bud ovoid; perianth 5 – 8 mm
diam., receptacle shallow, 1.5 – 2 mm diam., sometimes 5-saccate below sepals; sepals
ovate-oblong, acuminate, c. 2 mm long; petals ovate, subobtuse or acute, 2.5 – 3 by
(1.5 –)2 mm, glabrous or puberulous; filaments free, c. 0.5 mm long; anthers less than
0.5 mm diameter. Female inflorescences paniculate, 2 –10 cm long, up to 10-flowered.
Female flowers: pedicel c. 10 mm, ovary subcylindrical, 7– 8(–10) mm long, finely
pubescent, styles short, stigma not lobulate. Infructescence 1- or several-fruited. Fruit
4 – 8 cm long, glabrous or pubescent, base narrowed or rounded, at apex 1.2 – 2.5(– 3)
cm wide; fruiting pedicel 1– 2 cm long. Seeds many, ± unequal-sided star-shaped, 6 –11
by 5 – 8 mm, margin (edge) of 5 –7 coarse acute or blunt teeth, faces low tuberculate,
wing 14 –17 mm long to 8 mm wide. — Fig. 8f, 53f; Plate 22.
Distribution — NE India, Bhutan, Myanmar, South China (Yunnan, Hainan), Taiwan,
Laos, Cambodia, Vietnam, Thailand, east to NE Australia (Queensland) and the Pacific
(Solomon Islands, east to Fiji); in Malesia: North Sumatra, Peninsular Malaysia (Perlis,
Kedah, and Selangor), Philippines (Luzon, Mindoro, Cebu), Moluccas (Morotai, Kei
Archipelago), New Guinea (Papua New Guinea (East Sepik, Eastern Highlands, Ma-
dang, Morobe, Central, and Northern Provinces)); not known from Java and Borneo.
Habitat & Ecology — Shaded and open places in primary evergreen montane for-
est and evergreen seasonal forest along streams, clayey soils on limestone and granite
bedrock; 200 –1300 m altitude.
Notes — 1. A polymorphic species, varying in hairiness, number and shape of leaf-
lets, and size and shape of fruit. The fruit varies in length from 4 – 8 cm and may be
(sub)glabrous or hairy including densely pubescent (N. pubigera, from Myanmar).
Fruit size varies especially in NE India and New Guinea, where large-fruited as well as
smaller-fruited specimens have been collected; for example, fruits from New Guinea
vary from 4 – 8 cm long, including fruits of c. 4 cm long with a comparatively broad
apex, to nearly 3 cm diameter, and with a broad rounded base. The largest fruits contain
the largest seeds. Alsomitra pubigera Prain var. glauca Craib is a limestone form with
somewhat succulent leaves with rather short petioles.
2. Cogniaux’s figure 3a – h (1916), apparently from Peekel 328 (B†), was errone-
ously taken for the male flowers of Alsomitra schefferiana (a species not occurring in
the Bismarck Archipelago in East New Guinea where Peekel collected), but represents
N. clavigera; the collection Peekel 328 was also depicted in Peekel (1984). The small
apical appendage of the anther obviously is exaggerated in the drawing, as normally
this is much smaller or absent.
2. Neoalsomitra hederifolia (Decne.) W.J.de Wilde & Duyfjes

Neoalsomitra hederifolia (Decne.) W.J.de Wilde & Duyfjes, Blumea 48 (2003) 108, f. 1i, 3. — Si
cyos hederifolius Decne., Nouv. Ann. Mus. Hist. Nat. 3, 3 (1834) 450; M.Roem., Fam. Nat. Syn.
Monogr. 2 (1846) 104; Span., Linnaea (1841) 206; Miq., Fl. Ned. Ind. 1, 1 (1856) 682, ‘hederae
folius’. — Gynostemma ? hederifolia (Decne.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
916, ‘hederaefolia’; M.Jacobs, Blumea 7 (1954) 617 (excluding material Hort. Bot. Bog. sub XVIII
A 45); Steenis, Webbia 11 (1956) 192, f. 1a. — Type: Anonymous 531 (3 sheets, see note 1) (lecto
P, barcode P218602, designated by De Wilde & Duyfjes (2003)), Timor.
Zanonia timorana Span., Linnaea (1841) 205, Icon. 55 (plate not found); Miq., Fl. Ned. Ind. 1, 1 (1856)
683. — Alsomitra timorana (Span.) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 117; Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 935; in Engl., Pflanzenr. 66, 4.275.I (1916) 13. — Neoalso
mitra timorana (Span.) Hutch., Ann. Bot. (Oxford) 6 (1942) 98. — Type: Spanoghe s.n., Timor;
untraced. Neotype: De Voogd 2323, male fl. (L, designated by De Wilde & Duyfjes (2003)); iso
BO, K, SING), Timor.
Gynostemma spec. C,.de Voogd, Trop. Natuur, Jubileum Uitg. (1936) 71, f. 8, photo of habit; C.van Woer-
den, Trop. Natuur 29 (1940) 6, f. 7, photo of thickened stem base (no voucher specimen collected).
Low or medium climber, perennial, densely or sparsely glandular-hairy all over (ex-
cept flowers), hairs 0.5 – 2 mm long, stem at base to 1 cm thick; dioecious; growing from
a supra-terranean tuber to 30 by 15 cm and c. 2.5 kg in weight. Leaves: petiole 1.5 – 3 cm
long; blade membranous, brown on drying, simple, unlobed or shallowly 3(– 5)-lobed,
broadly ovate or circular in outline, 5 – 6 by 4 –7.5 cm, sparsely or densely (setose)
hairy, especially so on margin and on veins beneath, glands absent, margin entire or
coarsely remotely dentate, apices acute-acuminate, 1(–2) mm mucronate; palmately
5-veined, pale. Male inflorescences paniculate, pubescent, situated occasionally close
to the base of the plant, once or twice branched, 5 –17(– 32) cm long including 1– 2.5
cm long peduncle, ultimate branches raceme-like, 1– 4 cm long, 5 –10(–15)-flowered,
with bracts and basal portion of pedicel (c. 1 mm long) persistent; bracts minute, nar-
rowly elliptic, 0.5 –1 mm long, lower bracts in the larger twice branched inflorescences
leafy, 1–1.5 cm diameter. Male flowers solitary, (sub)glabrous; pedicel 3 – 4 mm long,
articulate at c. 1 mm from the base; bud subglobose; perianth at anthesis bowl-shaped;
receptacle shallowly bowl-shaped, 0.4 – 0.5 by c. 1.5 mm; sepals long-triangular, c. 1
mm long, with out-curved apiculum; petals broadly obovate or subcircular, c. 1.5 mm
long, papillose adaxially, very minutely mucronate; filaments (0.7–)1 mm long, either
completely or for 2/3 fused (column hollow), anthers elliptic or subcircular in outline,
c. 0.5 mm long, adaxially with or without a faint brown spot. Female inflorescences
and flowers not seen. Infructescences (sub)racemose, 3 – 5-fruited or a fruit solitary
on the leafy node of lateral shoots. Fruit brittle, glabrescent, 2.5 – 3 cm long, at apex
(0.8 –)1 cm wide, base narrowly rounded, perianth scar (rim) at c. 2.5 mm distant from
the flat apex; fruiting pedicel 0.7–1 cm long, glabrous with some vestigial minute hairs.
d
3 mm 1 mm
2 cm
a 2 cm
c
e
Fig. 50. Neoalsomitra hederifolia (Decne.) W.J.de Wilde & Duyfjes. a. Habit with male inflorescences;
b. detail of male inflorescence; c. male inflorescence; d, e. male flowers (a, b, d, e: Backer 26948;
c: ‘Decaisne’ P00218601, syntype).
Seeds 15 – 20, obovate with narrowed base, c. 6 by 4.5 mm, dark brown, margin (edge)
coarsely 5-toothed, faces coarsely few-tubercled, wing 7 by 2.5 mm. — Fig. 50, 53e.
Distribution — Known from few collections: Java (Kangean Archipelago, east of
Madura), Lesser Sunda Islands (Pulau Semau (near Timor) and Timor).
Habitat & Ecology — Lowland savanna and dry forest in monsoon climate on lime-
stone; 0 – 200 m altitude; flowering: March (Kangean) and November (Timor).
Notes — 1. The three anonymous sheets with male flowers in P, syntypes of Sicyos
hederifolia, were collected around 1801–1821, either by Sautier, Riedlé, Quichenot, or
by Gaudichaud; see Van Steenis-Kruseman (Fl. Males., Ser. 1, Spermat (1950)), and
the introduction to Decaisne’s enumeration (1834).
2. The infructescences and fruits of Sicyos hederifolia are known only from an ano
nymous single collection in P, a leafless specimen annotated Zanonia indica L., Timor,
apparently collected in the same period as the type of S. hederifolius. Its assignment to
N. hederifolia is not fully certain, but the old, dead twigs with leaf scars and tendrils
agree, and the fruits and seed (one single seed only is preserved, Fig. 53e) differ from the
fruits and seeds of the other Neoalsomitras from Timor: N. sarcophylla differs in seed
and N. schefferiana subsp. podagrica differs in fruiting pedicel c. 5 mm long against c.
10 mm long in N. hederifolia.
3. Jacobs (1954) cites a note by Van Steenis on living flowering material cultivated
in Bogor, about 1939, not preserved or apparently lost, obviously not belonging to N.
hederifolia. See also the note under N. sarcophylla.
3. Neoalsomitra pilosa W.J.de Wilde & Duyfjes

Neoalsomitra pilosa W.J.de Wilde & Duyfjes, Blumea (2003) 110, f. 1j. — Type: Carr 12477, fruit
(holo CANB; iso BM, K, L, SING), Papua New Guinea (Rouna).
Herbaceous climber, possibly one or a few metres long, annual?, sparsely soft-woolly
grey glandular-hairy; monoecious. Leaves: petiole 1–1.5 cm long; blade membranous,
blackish on drying, simple, unlobed or shallowly (faintly) 3 – 5-lobed, ± ovate in out-
line, 4 –7.5 by 3.5 –7 cm, upper and lower surface sparsely villous, glands absent,
margin entire, apex rather long acute-acuminate, minutely mucronate. Inflorescences
glandular hairy, to 10 cm long (including 2 – 6 cm long peduncle), with few (uni)lateral
10 – 20-flowered male racemes, and with 1 (or 2 or 3) solitary female flowers (fruit)
at the nodes of the lower branch(es), occasionally co-axillary with the panicle an ad-
ditional male raceme; bracts lanceolate, c. 1 mm long, ± caducous. Male flowers:
pedicel 1.5 – 2 mm long, sparsely hairy; perianth ± bowl-shaped; receptacle shallowly
bowl-shaped, 0.7(–1) mm wide; sepals ± long-triangular, c. 1 mm long, sparsely hairy;
petals (ovate-)elliptic, c. 1.5 mm long, glabrous, at apex with minute papillose mucro;
filaments united into c. 0.5 mm long column; anthers broadly ellipsoid, c. 0.3 mm
long, abaxially with minute dark dot. Female flowers not known (ovary hairy). Fruit
1(– 3) per infructescence (in old otherwise male-flowered inflorescence); (2.5 –)3 – 3.5
by 1–1.3 cm, sparsely whitish villous, hairs glandular, 3 – 4 mm long, base (narrowly)
rounded, apex truncate, with 2 – 3 mm long stylar processes; fruiting pedicel 0.4 – 0.8
cm long. Seeds 15 – 20, obovate, narrowed at base, 6 –7 by (4 –)5 mm including double
6 – 8-toothed margin (edge), faces ribbed towards margin, nearly smooth in the centre,
wing c. 8 by 4 mm. — Fig. 53b.
Distribution — New Guinea (SE Papua New Guinea, in the mountains near Port
Moresby); known from 2 collections: Carr 12477 (male fl., fr.) and Carr 15873 (male
fl., immat. fr.).
Habitat & Ecology — Open places in woods on steep hillsides; 400 –1500 m altitude;
flowering and fruiting: March to June.
Note — This species is close to N. capricornica (Australia), which differs in a green
drying colour, leaf shape and more coarsely hairy capsules.
4. Neoalsomitra sarcophylla (Wall.) Hutch.

Neoalsomitra sarcophylla (Wall.) Hutch., Ann. Bot. (Oxford) 6 (1942) 100; Keraudren in Aubrév. &
J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 9, pl. 1. (p.p., excl. Neoalsomitra balansae);
W.J.de Wilde & Duyfjes, Blumea 48 (2003) 111, f. 1b; Fl. Thailand 9, 4 (2008) 489. — Zanonia
sarcophylla Wall., Numer. List (1831) 3724, nom. nud.; Pl. Asiat. Rar. (1831) 28, t. 133. — Alsomi
tra sarcophylla (Wall.) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 118; Cogn. in A.DC. & C.DC.,
Monogr. Phan. 3 (1881) 929; in Engl., Pflanzenr. 66, 4.275.I (1916) 13; Craib, Fl. Siam. (1931)
769. — Type: Wallich 3724B fruit (lecto K-Wall., designated by De Wilde & Duyfjes (2003)),
Myanmar.
Alsomitra philippinensis Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 15. — Neoalsomitra philippin
ensis (Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 100. — Type: Copeland 255 (B†), Philippines,
Luzon.
Climber 3 – 8 m long, perennial from woody rootstock, glabrous; dioecious. Tendrils

often with adhesive pads. Leaves: petiole 0.5 –1 cm long; petiolules 0.2 – 0.5(–1) cm
f
2 cm
g
2 cm
2 mm
2 cm
b
0.5 mm
1 mm
d c
Fig. 51. Neoalsomitra sarcophylla (Wall.) Hutch. a. Portion of male flowering twig; b. detail of male
inflorescence; c, d. male flower, seen from below and above; e. stamen; f. infructescence; g. node with
leaf and tendril, note tendril branched at apex. (a – e: Phonsena et al. 6342; f: Poilane 11767; g: Van
Steenis 18015).
long, after abscission leaving raised flat scars; blade carnose-leathery (± succulent),
wrinkled on drying, 3-foliolate, 5 – 20 cm diam., leaflets (broadly) ovate-elliptic to
oblong-lanceolate, to 12 by 6 cm, glands absent, base rounded or ± narrowed, apex (nar-
rowly) rounded, minutely mucronate, basal veins 3 – 5, curving towards apex, mostly
indistinct. Male inflorescences paniculate, little or much branched, sometimes branched
from the base, 10 – 20(– 30) cm long, pendent, glabrous; bracts linear, 1 mm long or
less. Female inflorescences paniculate or ± raceme-like, 5 –10 cm long. Male flowers
(1–)3 subfascicled; pedicel 2 – 5 mm long; perianth subrotate, c. 5 mm diam., glabrous,
minutely papillose; receptacle flattish, faintly 5-saccate, c. 1 mm diam.; sepals lan-
ceolate, c. 2.5 mm long, acuminate; petals (ob)ovate, c. 3 by 1.5 mm, apex acuminate,
midvein distinctly raised in lower portion, forming dissepiments between the stamens;
filaments free, spreading, c. 1 mm long, anthers less than 0.5 mm long. Female flow
ers: ovary narrow, c. 8 by 1 mm, glabrous; staminodes 0.2 – 0.5 mm long; styles short,
broad at base, stigma ± lobulate-dentate, style and stigma together c. 1 mm long. In
fructescences pendent, 5 –10 cm long, with 5 –10(– 20) fruits. Fruit much narrowed to
the base, glabrous, 3(– 4) cm long, apex truncate, 0.8(–1) cm wide; fruiting pedicel
slender, 1(–1.5) cm long. Seeds 12 – 15, subtriangular, 6 –7 by 3 – 4 mm, 2-horned at
apex, margined, edge smooth or verrucose, faces finely verrucose, wing suberect, 8 –10
by 4 mm. — Fig. 51, 53a; Plate 23.
Distribution — Myanmar, Thailand, Laos, Cambodia, Vietnam; in Malesia: rare and
scattered in East Malesia: Philippines (Luzon, Palawan), Sulawesi (Central, SW, and
Kabaena Is.), Lesser Sunda Islands (Timor). Sometimes cultivated in botanical gar-
dens.
Habitat & Ecology — Mixed, (partly) deciduous evergreen lowland and montane
forest, on basalt and granite bedrock and limestone; from sea level to 850 m altitude;
flowering: March to September.
Notes — 1. In Teijsmann 19765 (BR), Timor, the seeds lack the two horns; it is the
only known collection with seeds from East Malesia.
2. In the Bogor Botanic Garden a sterile widely creeping vegetative persisting plant,
possibly a remnant of old introduction, was still present in 1998.
5. Neoalsomitra schefferiana (Cogn.) Hutch.

Neoalsomitra schefferiana (Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 101, (excl. Peekel 328 (B†) =
Neoalsomitra clavigera.); W.J.de Wilde & Duyfjes, Blumea 48 (2003) 112, f. 1g, h. — Alsomitra
schefferiana Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 932; in Engl., Pflanzenr. 66, 4.275.
I (1916) 16, excl. descr. of male flower of Peekel 328 and f. 3a–h. — Type: Teijsmann 11854, sheet
1 of 4 (holo BO; iso BO, 3 sheets, L), Sulawesi, Pangkadjena.
Alsomitra schefferiana Cogn. var. minor Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 933; in
Engl., Pflanzenr. 66, 4.275.I (1916) 16. — Type: Teijsmann s.n. (holo BO, not seen), SW Sulawesi,
Pangkadjena.
Perennial climber to 8(– 30) m long, with one or few green stems somewhat car-
nose-woody at base, either slightly thickened, or base flask- or spindle-shaped, swol-
len, lower portion of main stem with ± hard green thorns 1– 2(– 4) cm long, apical
shoots puberulous, glabrescent, with raised leaf scars; dioecious. Tendrils on innovation
shoots, mostly caducous. Leaves: petiole 2 – 5 cm long; petiolules 0.3 – 2 cm long; blade
membranous (slightly fleshy when fresh), green on drying, (simple or) 3- or (pedately)
5-foliolate, 5 –15(– 20) cm diam., leaflets elliptic or (ob)ovate, apex roundish or acute,
minutely acuminate, the middle leaflet largest, to 11(–15) by 7(–10) cm, glands absent.
Male inflorescences paniculate, with numerous flowers, 8 – 20(– 40) cm long, once or
twice branched, ultimate branches raceme-like, pubescent, hairs c. 1 mm long; bracts
minute, c. 1 mm long, lower bracts 3-foliolate, larger, hairy. Male flowers: pedicel
c. 1.5 mm long, glabrous; perianth rotate, 2.5 – 3.5 mm diam., sparsely pubescent; recep-
tacle shallow, c. 1 mm wide; sepals ovate-acute, c. 0.5 mm long; petals broadly ovate,
acute, c. 1 mm long; filaments glabrous, fused into a partly hollow column c. 0.7 mm
long, free parts c. 0.3 mm long, anthers broadly ellipsoid, c. 0.3 mm long, dark spot
not obvious. Female inflorescences as in male, (widely) paniculate, 10–30 cm long,
flowers ultimately in racemes. Female flowers: see under subsp. podagrica. Fruit many
per infructescence, 1.5 – 2.5 by 0.7– 0.9 cm, pubescent, glabrescent, base ± attenuate or
rounded, apex truncate with minute style-remnants; fruiting pedicel 0.2 – 0.7 cm long,
glabrous or pubescent. Seeds c. 15, obovate, 4 – 5 by 3 – 3.5 mm, base attenuate, finely
or coarsely rugulose-warted, margin double 5 –7-toothed or -tuberculate, wing 6 – 8
mm long.
Distribution — SW Sulawesi, Moluccas, eastern Lesser Sunda Islands, and New
Guinea (West Papua (Vogelkop Peninsula)).
Habitat & Ecology — Limestone rocks and soil over limestone, in dry seasonal areas;
lowland; flowering: February (Timor), May (Alor), June (Seram).
Notes — 1. Neoalsomitra schefferiana is characterized by its thorny stem-base, 3- or
5-foliolate leaves, and stamens of which the filaments are c. 3/4 connate.
2. The limited fertile material available suggests the existence of only one species,
with 2 subspecies. The two subspecies chiefly differ in shape of the thorny stem-base
and fruit size; the male perianths of a specimen from Seram (Kornassi 1433) are slightly
larger than those from Timor. Because most collections are sterile and mostly the thorny
base not recorded, such specimens are tentatively assigned to subsp. schefferiana.
Subsp. podagrica is regarded as confined to Timor and the nearby islands of Semau
and Alor.
The thorns are obviously homologous with leaves. Occasionally the bases of with-
ered tendrils may resemble thorns. In the collections from Vogelkop of subsp. schef
feriana (Avé 4835) some thorns are minutely furcate at apex.
Key to the subspecies
1a. Main stem at base not or hardly thickened, 2 – 3 cm thick. Leaf blades of (fertile)
upper branches 3-foliolate. Fruit c. 2.5 cm long. — SW Sulawesi, Seram, Vogelkop
Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. subsp. schefferiana
b. Main stem at base swollen, 5 –10 cm thick. Leaf blades of (fertile) upper branches
3- or 5-foliolate. Fruit 1.5 – 2 cm long. — Timor, Alor Is. . . . . b. subsp. podagrica
a. subsp. schefferiana
Main stem base slender, 2 – 3 cm diam., thorns 1– 2 cm long. Leaf blades 3-foliolate.
Female flowers not known. Infructescences rather open, much branched, many-fruited,
short-tomentose, 20 – 30 cm long. Fruit c. 2.5 by 0.7– 0.9 cm, base shortly attenuate,
when old glabrescent except at the very base; fruiting pedicel 0.3 – 0.7 cm long, tomen-
tose. Seeds ovate, 4.5 – 5 by 3 – 3.5 mm, faces densely sharply, rather finely tuberculate,
margin (edge) finely crenulate, wing 6 –7 by c. 4 mm. — Fig. 53c.
Distribution — Known from one fruiting collection from SW Sulawesi (type); one
male flowering specimen from Moluccas (Seram); sterile specimens from SW Sulawesi
and New Guinea (Vogelkop Peninsula).
b. subsp. podagrica (Steenis) W.J.de Wilde & Duyfjes

Neoalsomitra schefferiana (Cogn.) Hutch. subsp. podagrica (Steenis) W.J.de Wilde & Duyfjes, Blumea
488 (2003) 114, f. 1h. — Neoalsomitra podagrica Steenis, Webbia 11 (1956) 192, f. 1b, 2, nom.
prov.; Blumea 8 (1955) 171, f. 1; H.Jacobsen, Sukkulenten Lex. (1970) 267, t. 112, 1; L.E.Newton
& Njoroge in Eggli, Ill. Handb. Succ. Pl.: Dicotyledons (2002) 90, f. 13e. — Type: Pleyte s.n., male
(lecto L, barcode L0128004, designated by De Wilde & Duyfjes (2003)), culta, Hort. Bot. Bogor
II-0-X-6, Timor (see note 1), 6 May 1954.
Base of main stem(s) fleshy, thickened, flask- or spindle-shaped up to 100 cm long,

by 10 cm diam., sometimes with additional superposed thickened bases of lateral stems;
thorns 1.5 – 5 cm long. Leaves: lower 3-foliolate, upper 3- or 5-foliolate. Female in
florescences paniculate, rather condensed, pubescent, the ultimate branches one-sided
racemes with up-curved flowers; bracts minute. Female flowers: pedicel c. 1 mm long,
glabrous; ovary tubular, terete, c. 3 mm long, (1–)1.5 mm wide, pubescent; sepals hardly
1 mm long; petals c. 2 by 1.5 mm, ± pubescent (some adaxially with minute appendages
at base); styles including stigma c. 1 mm long. Fruit many per infructescence, almost
glabrous, 1.5 – 2 by 0.7– 0.8 cm, base narrowed or rounded; fruiting pedicel 0.2 – 0.5 cm
long, almost glabrous. Seeds pear-shaped in outline, 4 – 5 by 3 – 3.5 mm, faces coarsely
tuberculate and with irregular radial crests, wings 6.5 – 8.5 by 3.5 – 4.5 mm. — Fig. 52,
53d.
Distribution — Eastern Lesser Sunda Islands (Alor Is., Semau Is., and Timor).
Habitat & Ecology — Lowland and hilly country on limestone.
Uses — In Timor used medicinally for its extremely bitter resin with strong haemo-
lytic action (Van Steenis, 1956). In Vogelkop the people are afraid to be hurt by the
thorns (Avé 4835).
Notes. 1. Although the subspecies seems widespread in Timor it is noteworthy that
this conspicuous plant was not seen by the old French and Dutch collectors, who col-
lected the pachypodous N. hederifolia (syn. N. timorana) several times. In the wild,
subsp. podagrica seems to flower or fruit at a not known period, under a strong seasonal
climate, but apparently not frequently.
2. This subspecies was already, though unnamed, pictured and described in a manu-
script by Father Alberti de San Tomé about 1750, in Lisbon (as was discovered there
by Koster, postmaster at Kupang (see Van Steenis (1956), Van Steenis-Kruseman, Fl.
Males., Ser. 1, Spermat (1950) 298).
2 cm
h 3 mm
3 mm f
i 3 mm
1 mm
c
1 mm
d e
a b
Fig. 52. Neoalsomitra schefferiana (Cogn.) Hutch. subsp. podagrica (Steenis) W.J.de Wilde & Duyfjes.
a. Apex of male flowering twig and apex of growing twig; b, c. male flower; d. female flower; e – g.
fruits; h. seed; i. seed, apical view (a – c: Cult. Hort. Bog. II.0.X.6; d – i: Cult. Hort. Bog. XVIII.A.45a;
drawn by Ruth van Crevel).
a b c d
2 mm
e f g
Fig. 53. Seeds of Neoalsomitra. — a. Neoalsomitra sarcophylla (Wall.) Hutch. — b. Neoalsomitra

pilosa W.J.de Wilde & Duyfjes. — c. Neoalsomitra schefferiana (Cogn.) Hutch. subsp. schefferiana.
— d. Neoalsomitra schefferiana (Cogn.) Hutch. subsp. podagrica (Steenis) W.J.de Wilde & Duyfjes.
— e. Neoalsomitra hederifolia (Decne.) W.J.de Wilde & Duyfjes. — f. Neoalsomitra clavigera (Wall.)
Hutch. — g. Neoalsomitra schultzei (Cogn.) Hutch. (a: Maxwell 00-163, Thailand; b: Carr 12477, type,
Papua New Guinea; c: Teijsmann 11854, Celebes; d: Pleyte s.n., Bogor, garden number XVIII-A-45a,
orig. Timor; e: Anonymous, barcode P218587, Timor; f: Chuang 3172, Taiwan; g: Brass 8149, Papua
New Guinea).
6. Neoalsomitra schultzei (Cogn.) Hutch.

Neoalsomitra schultzei (Cogn.) Hutch., Ann. Bot. (Oxford) 6 (1942) 98; Merr. & L.M.Perry, J. Arnold
Arbor. 30 (1949) 56; W.J.de Wilde & Duyfjes, Blumea 48 (2003) 114, f. 1k. — Alsomitra schultzei
Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 12. — Type: Schultze 170 (B†), Papua New Guinea
(Augusta River).
Small climber, annual? stem slender, subglabrous; monoecious. Leaves: petiole 1–1.5
cm long, sparsely minutely pubescent; blade membranous, green on drying, simple,
faintly to distinctly (3 –)5-lobed (up to c. 2/3 deep), ovate in outline, 3.5 –7 cm diam.,
glabrous except for a few hairs near insertion of petiole, base subtruncate or cordate,
margin smooth or coarsely sinuate, apex acute-acuminate, 1– 2 mm mucronate as are
lobe-apices. Inflorescences glabrous, all or predominantly male-flowered, sometimes
with a few female flowers co-axillary with lowermost branch(es), branches few, con-
sisting of 5 –10 male-flowered racemes; position of female flowers (from fruits) either
mixed in otherwise male flowered panicle (see above) or solitary at leafy node; bracts
oblong, 0.5 –1 mm long, subpersistent. Male flowers: pedicel c. 1.5 mm long, the lower
half persistent; perianth bowl-shaped; receptacle-tube c. 1 by 1.5 – 2 mm, inside thick-
ened into an inconspicuous disc; sepals (long-)triangular, c. 0.8(–1) by 0.7 mm, acute;
petals ovate-broadly elliptic, minutely mucronate, dotted with glands; filaments united
for 2/3 or completely united, c. 1 mm long, anthers ellipsoid, c. 0.6 mm long, without
adaxial dark blotch. Female flowers not seen (ovary subclavate, 4 – 5 mm long, glabrous,
corolla segments c. 2 mm long, see Cogniaux, 1916). Fruit 1(– 6) per infructescence,
solitary at leafy node, or peduncled, 2.5 – 3 cm long, glabrous, base rounded, 1–1.2 cm
wide at truncate apex; fruiting pedicel c. 1 cm long. Seeds 15 – 20, obovate with nar-
rowed base, 6 –7 by 4 – 5 mm, margin 5 –7 coarsely dentate or tuberculate, faces finely
warty, wing c. 9 by 4 – 5 mm. — Fig. 53g.
Distribution — New Guinea (Papua New Guinea, known from a few collections only
(Sepik, lower Fly River, Central Prov.)).
Habitat & Ecology — Lowland; margins of monsoon forest, in reed swamps. Flower-
ing and fruiting: April and October.
Notes — 1. The present description of N. schultzei, drawn from the collections Brass
8149 and Pullen 6847, is somewhat doubtful because the type, Schultze 170, got lost,
but Cogniaux’s protologue agrees very well. A designation of a neotype should wait for
a good collection from the type locality, Sepik (Augusta River).
2. Neoalsomitra schultzii is characterised within the genus by a disc-like thickened
receptacle. It forms together with N. capricornica (Australia), N. hederifolia and N.
pilosa a group with fused filaments within Neoalsomitra.
28. PAPUASICYOS
Papuasicyos Duyfjes, Blumea 48 (2003) 124. — Type species: Papuasicyos papuana (Cogn.) Duyfjes
(= Melothria papuana Cogn.).
Small climber, leafy stem c. 2 mm diam., plant subglabrous; monoecious. Probract

absent. Tendrils unbranched. Leaves: blade simple, unlobed. Flowers small; petals im-
bricate, pale yellow, free, entire. Male inflorescence a simple pedunculate raceme. Male
flowers: pedicel short, receptacle-tube bowl-shaped, expanded corolla 10(–15) mm
across; stamens 3, free, inserted c. halfway up the receptacle-tube; filaments short, anthers
all 2-thecous, free, appressed into a subglobose head, thecae sigmoid, connective broad;
disc absent or minute. Female flowers solitary, not associated with male inflorescence;
pedicel long; ovary oblong, ovules many, horizontal; perianth as in male; staminodes 3,
inserted near the throat of the receptacle-tube; disc obscure or absent; style conspicuous
with feather-like divided stigma. Fruit berry-like, (narrowly) ellipsoid, 2 – 2.6 cm long;
fruiting pedicel slender. Seeds many, foveolate, margin narrow, edge entire.
Distribution — 1 species in New Guinea.
a b
c
Plate 2. a, b. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li var. borneensis. a. Female inflorescence;
b. infructescence. — c. Alsomitra macrocarpa (Blume) M.Roem. Winged seed (a, b: Sabah; c: Penin-
sular Thailand). Photos: a, b by De Wilde; c by Kanlaya Phattarahirankanok.
a b
c d
Plate 3. Bayabusua clarkei (King) W.J.de Wilde. a. Seedling; b. detail of male flowering twig; c, d.
male flowers (all: Peninsular Malaysia, a, b. near Cameron Highlands; c. Kepong, FRIM, along canopy
walk). Photos: all by De Wilde.
a b
c d
Plate 4. a, b. Bayabusua clarkei (King) W.J.de Wilde. a. Fruit; b. fruit longitudinally opened, showing
winged seeds. — c, d. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes forma hispida (Thunb.)
W.J.de Wilde & Duyfjes. c. Male flower; d. female flower (a, b: Peninsular Malaysia, near Cameron
Highlands; c, d: Thailand). Photos: all by De Wilde.
a b
c d
Plate 5. Borneosicyos simplex W.J.de Wilde. a, c. Female inflorescences; b. male inflorescence; d. fruit
(all: Sabah, Kinabalu Park). Photos: all by De Wilde.
a b
c
Plate 6. a. Cyclanthera brachystachya (Ser.) Cogn. Fruit and male inflorescence. — b. Cucumis melo
L. forma agrestis (Naudin) W.J.de Wilde & Duyfjes. Dove egg-sized fruits and male flowers. —
c. Citrullus lanatus (Thunb.) Matsum. & Nakai. Shoot with male flowers (a: Java, Cibodas Botanical
Garden; b: Lombok; c: Java). Photos: all by De Wilde.
a b
c
Plate 7. Coccinia grandis (L.) Voigt. a. Male flowers; b. female flower; c. fruits (all Thailand). Photos:
all by De Wilde.
a b
c d
Plate 8. Diplocyclos palmatus (L.) C.Jeffrey var. palmatus. a. Male flower; b. female flower; c. fruit
and male inflorescences; d. fruits (a, b, d: Thailand; c: Java, Cibodas Botanical Garden). Photos: a – c
by De Wilde; d by D. Chusithong.
a b
c
Plate 9. Gymnopetalum chinense (Lour.) Merr. a. Male flower; b. fruit; c. creeping shoot with male
inflorescences (all: Bali). Photos: all by De Wilde.
a b
c d
Plate 10. a – c. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. scabrum. a. Seedling;
b. fruit; c. male bud. — d. Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. pectinatum
W.J.de Wilde & Duyfjes. Fruit (a, c: Thailand; b, d: Java). Photos: all by De Wilde.
a b
c d
Plate 11. a – c. Gynostemma pentaphyllum (Thunb.) Makino forma pentaphyllum. a, b. Male inflores-
cences; c. infructescence. — d. Hodgsonia macrocarpa (Blume) Cogn. Fruit (a – c: Thailand; d: Sabah).
Photos: a, b by T. Putthai; c, d by De Wilde.
a b
c d
Plate 12. a, b. Kedrostis monosperma W.J.de Wilde & Duyfjes. a. Female inflorescence; b. fruit. —
c, d. Kedrostis bennettii (Miq.) W.J.de Wilde & Duyfjes. c. Male inflorescence; d. female inflorescence
and young fruit (a, b: Peninsular Malaysia, Pahang, Ulu Krau; c: Bali; d: Sabah, Gomantong). Photos:
a, b, d by De Wilde; c by J. Bastmeijer.
a b
c d
Plate 13. a. Kedrostis bennettii (Miq.) W.J.de Wilde & Duyfjes. Infructescence. — b. Lagenaria sicer
aria (Molina) Standl. Male flower. — c, d. Luffa aegyptiaca Mill. forma aegyptiaca. c. Male flower;
d. apical part of fruit, showing operculum (a: Sabah, Gomantong; b – d: Thailand). Photos: all by De
Wilde.
a b
c d
Plate 14. a, b, d. Momordica cochinchinensis (Lour.) Spreng. a, b. Male flowers; d fruits. — c. Mo
mordica denticulata Miq. Fruit (a, b: Java, Bogor, Botanical Garden; c: Sabah; d: Sulawesi). Photos:
all by De Wilde.
a b
c d
Plate 15. a, b. Momordica charantia L. forma abbreviata (Ser.) W.J.de Wilde & Duyfjes. a. Female
flower; b. ripe fruit exposing seeds in red pulp. — c. Momordica subangulata Blume subsp. suban
gulata. Male flower. — d. Momordica clarkeana King. Fruit (a – c: Thailand; d: Peninsular Malaysia,
Perak). Photos: a – c by De Wilde; d by Imin Kamin.
a b
c d
Plate 16. a, b. Melothria pendula L. a. Male inflorescence; b. fruits. — c. Mukia javanica (Miq.)
C.Jeffrey. Fruits. — d. Mukia maderaspatana (L.) M.Roem. Fruits (a, b: Peninsular Malaysia, Kuala
Lumpur; c: Java; d: Thailand). Photos: a – c by De Wilde; d by D. Chusithong.
a b
c d
Plate 17. a, b. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. chlorocarpa. a. Male inflo-
rescence; b. fruit. — c, d. Indomelothria chlorocarpa W.J.de Wilde & Duyfjes subsp. halimunensis
W.J.de Wilde & Duyfjes. c. Male inflorescences; d. fruit and male inflorescence (a, b: Sabah; c, d: West
Java, Gn Halimun). Photos: all by De Wilde.
a b
c d
Plate 18. a – c. Neoachmandra leucocarpa (Blume) W.J.de Wilde & Duyfjes. a, b. Male and female
flowers; c. fruits. — d. Neoachmandra nesophila W.J.de Wilde & Duyfjes. Male flower (a – c: Java,
Cibodas; d: Philippines). Photos: a – c by De Wilde; d by P. Pelser.
a b
c d
Plate 19. a, b. Neoachmandra nesophila W.J.de Wilde & Duyfjes. a. Female flower; b. fruit. — c, d.
Pilogyne mucronata (Blume) W.J.de Wilde & Duyfjes. c. Male inflorescences; d. fruits (a, b: Lombok;
c, d: West Java). Photos: all by De Wilde.
a b
c d
Plate 20. a, b. Pilogyne bodinieri (H.Lév.) W.J.de Wilde & Duyfjes. a. Male inflorescence; b. fruits and
male inflorescence. — c, d. Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. pubescens
Cogn. c. Detail of male inflorescence; d. infructescence (all: Thailand). Photos: a by D. Chusithong;
b – d by De Wilde.
a b
c d
Plate 21. a, b. Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes var. marginata. a. Male in-
florescence; b. fruit and male inflorescence. — c, d. Solena heterophylla Lour. subsp. heterophylla.
c. Male inflorescence; d. fruit (all: Thailand). Photos: a, b, d by De Wilde; c by T. Putthai.
b c
Plate 22. Neoalsomitra clavigera (Wall.) Hutch. a. Male inflorescence; b. male flower; c. fruits (all:
Thailand). Photos: all by De Wilde.
a b
c d
Plate 23. Neoalsomitra sarcophylla (Wall.) Hutch. a. Male inflorescences; b. detail; c. female inflores-
cence; d. detail (all: Thailand). Photos: a by De Wilde; b – d by P. Phonsena.
a b
c
Plate 24. Thladiantha cordifolia (Blume) Cogn. a. Male inflorescence; b. male flower; c. fruit (all:
Thailand). Photos: a, b by De Wilde; c by D. Chusithong.
a b
c d
Plate 25. a – c. Trichosanthes beccariana Cogn. subsp. beccariana. a, b. Unripe and nearly ripe fruit;
c. female flower. — d. Trichosanthes elmeri Merr. Fruit (all: Sabah). Photos: all by De Wilde.
a b
c d
Plate 26. a, b. Trichosanthes elmeri Merr. a, b. Fruits. — c, d. Trichosanthes emarginata Rugayah.
c. Fruits; d. seeds (a, b: Sabah; c, d: Peninsular Malaysia, Pahang, Ulu Krau). Photos: a, b by De Wilde;
c, d by C.K. Lim.
a b
c d
Plate 27. a, b. Trichosanthes kinabaluensis Rugayah. a. Young shoot with probract; b. ripe fruit. —
c, d. Trichosanthes mucronata Rugayah. c. Male flower; d. ripe fruit (all: Sabah, near Kinabalu Park).
Photos: all by De Wilde.
b c
Plate 28. Trichosanthes pendula Rugayah. a. Male flower; b. apical portion of male inflorescence;
c. ripe fruit in female inflorescence (all: Sabah, Sepilok Forest Reserve). Photos: all by De Wilde.
a b
c
Plate 29. a, b. Trichosanthes postarii W.J.de Wilde & Duyfjes. a. Male inflorescence; b. ripe fruit. —
c. Trichosanthes pilosa Lour. var. pilosa. Male flower (a, b: Sabah, near Poring; c: Thailand). Photos:
all by De Wilde.
a b
c d
Plate 30. a. Trichosanthes montana Rugayah subsp. crassipes W.J.de Wilde & Duyfjes. Fruit. — b. Tricho
santhes sepilokensis Rugayah. Full grown fruit, not yet red-coloured. — c. Trichosanthes tricuspidata Lour.
subsp. javanica Duyfjes & Pruesapan. Infructescences. — d. Trichosanthes villosa Blume. Male inflores-
cence (a: Sabah, near Kinabalu Park; b: Sabah, near Sepilok; c, d: Thailand). Photos: all by De Wilde.
a b
c
Plate 31. a. Trichosanthes quinquangulata A.Gray. Fruit. — b. Trichosanthes wawrae Cogn. forma
wawrae. Fruit. — c. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes forma pruriens. Small-
sized ripe fruits (a: West Java; b: Thailand; c: Sabah). Photos: all by De Wilde.
a b
c
Plate 32. a, c. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes forma hispida (Thunb.) W.J.de
Wilde & Duyfjes. a. Depressed globose fruits; c. ellipsoid fruits. — b. Lagenaria siceraria (Molina)
Standl. Pear-shaped immature fruits. — c. In the background Cucurbita moschata Duchesne. Fruits
(all: Thailand, exposed on vegetable market). Photos: all by De Wilde.
b
Plate 33. a. Luffa aegyptiaca Mill. forma aegyptiaca. Fruits. — b. Luffa acutangula (L.) Roxb. (left),
and Momordica charantia L. forma charantia (right) (a: Thailand, vegetable garden; b: Thailand,
vegetable market). Photos: all by De Wilde.
1. Papuasicyos papuana (Cogn.) Duyfjes

Papuasicyos papuana (Cogn.) Duyfjes, Blumea 48 (2003) 124, f. 2, 3. — Melothria papuana Cogn.,
Bull. Acad. Roy. Sci. Belgique, Cl. Sci., sér. 3, 14 (1887) 355; in Engl., Pflanzenr. 66, 4.275.I (1916)
92; Harms, Bot. Jahrb. Syst. 60 (1925) 152. — Type: Bäuerlen 328 (holo B†; iso MEL), Papua
New Guinea (Strickland River).
Subperennial climber, c. 6 m long, with minute sparse appressed grey hairs 0.1 mm
long, glabrescent; monoecious; roots not known. Leaves: petiole 1– 2 cm long; blade
membranous, green on drying, (narrowly) ovate, 4 –15 by 1.5 – 9.5 cm, glabrescent
except for minute hairs on veins, densely set with small cystoliths above, glands ab-
sent, base rounded, truncate, or shallowly cordate, margin entire or occasionally with
2 cm
g g'
2 mm f 2 mm
1 mm
b c
Fig. 54. Papuasicyos papuana (Cogn.) Duyfjes. a. Male inflorescence; b. nearly mature male flower
bud; c. ditto, opened, showing position of stamens (disc absent); d. stamens; e. portion of branch with
a female flower; f. fruit; g, g'. seed (all: Bäuerlen 328, type).
an odd tooth at base, apex acute-acuminate, mucronate; basal veins 3 – 5(–7). Male
inflorescence a solitary pedunculate delicate 10(– 20)-flowered raceme, 5 – 8 cm long,
with minute papillose glands and appressed hairs, glabrescent; peduncle 1– 4 cm long;
flowers rather irregularly inserted; bracts absent. Male flowers: pedicel 3 – 8 mm long,
sparsely hairy, inconspicuously articulated c. 0.5 mm below apex; receptacle-tube c. 2 by
1 mm
a
1 mm
b f g
1 mm
1 mm
c d d'
Fig. 55. Papuasicyos pauana (Cogn.) Duyfjes. a. Male flower, sepals small; b. male flower, sepals
large; c. male flower (half-schematic, pilosity omitted), opened, showing position of stamens and
minute disc?; d, d'. stamens; e. female flower (somewhat schematic), opened, note staminodes; f. fe-
male receptacle-tube and sepals; g. staminode (a, c – d': Docters van Leeuwen 9873; b: Ridsdale NGF
30335; e – g: Bäuerlen 328, type).
3.5 mm, outside subglabrous, inside pilose; sepals long triangular or elliptic, 0.5 – 2
by 0.3 – 2 mm, apex acute or ± rounded, very minutely mucronate, margin minutely
fringed; petals ovate-elliptic, 5 – 6.5 by 3 – 3.5(– 5) mm, 3 – 5(–7)-veined, pilose (hairs
c. 0.2 mm long), apex subacute or rounded, margin fimbriate; filaments c. 1 mm long,
with pale shaggy hairs at base, inserted c. halfway up the receptacle, anthers closely
appressed into a nearly globose head c. 3.5 mm long, 4.5 mm wide, thecae narrow,
sigmoid, ± bilateral-symmetrical at the edges of broad rather flat connective (Fig. 54c,
d, 55c, d'); disc absent or inconspicuous. Female flowers solitary on the leafy nodes,
(sub)glabrous; pedicel 40 –100 by c. 0.5 mm; ovary oblong, c. 10 by 2.5(– 3) mm, apex
narrowed into a neck c. 1 mm long; receptacle-tube bowl-shaped, c. 1.5 by 2.5(– 3) mm,
throat hairy inside; sepals 1–1.5 by c. 0.3 mm; petals as in male flower but larger, ±
short-hairy, narrowly elliptic, c. 10 mm long, 5-veined; staminodes 3, inserted slightly
below receptacle throat, terete, c. 1 mm long, densely fine-hairy, with hairs 0.5(–1) mm
long, at apex with one or two minute transversal glabrous thickenings; receptacle below
the insertion of the staminodes inside faintly thickened (disc?); style c. 1.5 mm long,
(sub)glabrous, stigma 3-branched, with each branch 4 – 4.5 mm long, once (or twice)
forked, wholly conspicuously densely (glandular?) hairy, with hairs c. 1 mm long. Fruit
solitary, (narrowly) ellipsoid, narrowed at both ends, 2 – 2.6 by 0.9 –1.2 cm, smooth,
juicy; fruiting pedicel slender, 4 –10 cm long, glabrous. Seeds c. 40, in dry fruits vis-
ible and pressed into the transparent pericarp, ovoid-ellipsoid, c. 5 by 3.5 by 1– 2 mm,
(light) brown. — Fig. 54, 55.
Distribution — New Guinea, north and south of the Main Range: New Guinea (West
Papua and Papua New Guinea; known from 5 collections).
Habitat & Ecology — Lowland swamp forest, river banks; 0 – 500 m altitude; flower-
ing and fruiting June to December.
Notes — 1. The sepals in the few male flowers available for study are remarkably
different: those of Docters van Leeuwen 9873 (West Papua) are narrow and only c. 0.5
mm long; those of Ridsdale NGF 30335 (Papua New Guinea, Morobe Distr.) are broad
and rounded and c. 2 mm long; those of the other 3 collections are ± intermediate. In
Docters van Leeuwen 9873 possibly a small depressed central disc is present, but this
cannot be determined with certainty; however, a disc is absent in the other specimens.
2. According to molecular phylogenetic analyses Papuasicyos is congeneric with
Urceodiscus (H.Schaefer & S.S.Renner, Fam. Gen. Vasc. Pl. [Kubitzki], in press).
29. PILOGYNE
Pilogyne Schrad., Index Sem. (Gottingen) (1835) 5; Eckl. & Zeyh., Enum. Pl. Afric. Austral. 2 (1836)
408; W.J.de Wilde & Duyfjes, Reinwardtia 12 (2009) 405. — Type species: Pilogyne suavis Schrad.
(Africa).
Zehneria Endl., Prodr. Fl. Norfolk. (1833) 69, p.p, & excl. the type, Zehneria baueriana Endl.
Zehneria auct. non Endl. p.p., excl. the type, Zehneria baueriana Endl.: W.J.de Wilde & Duyfjes,
Blumea 51 (2006) 48; Fl. Thailand 9, 4 (2008) 544.
Small climbers, annual or (sub)perennial, leafy stem 1– 2 mm diam., usually black-

ish on drying; usually dioecious. Probract linear, minute, 1– 5 mm long, sometimes
caducous. Tendrils unbranched. Leaves: blade simple. Flowers small, receptacle-tube
campanulate; sepals minute, narrow; petals valvate or imbricate in bud, white or creamy,
free. Male inflorescence a short- or long-peduncled few- or many-flowered condensed
raceme, when monoecious co-axillary with female flower(s) or not; bracts absent.
Male flowers: pedicel short, 2 –10(–15) mm long, persistent; stamens 3, inserted in
the lower half of the receptacle-tube, usually near the base, filaments longer than the
anther, anthers all 2-thecous, ± included or just exserted, thecae lateral, straight or
curved, not divergent, connective narrow or broad and ± thickened adaxially, not or
little produced at apex; disc (depressed-)globose or 3-lobed. Female flowers solitary
or few at the node, or few in a peduncled cluster, when monoecious co-axillary with a
male raceme, or mixed with male flowers in a peduncled raceme; pedicel short (or long);
ovary globose with slender neck, or ellipsoid; stigma (deeply 3-lobed or) 3 on short
style-arms, papillose(-hairy); staminodes present; disc free, annular. Fruit 1 or several,
green, ripening red or purplish blackish, usually with short fruiting pedicel, globose
or ellipsoid, 0.5 – 3 cm long, juicy or pulpy; exocarp cartilaginous, minutely pitted or
tessellate. Seeds several or numerous, whitish or pale brownish, compressed, ovate or
elliptic, not sculptured, margin narrow but distinct (indistinct in P. immarginata), edge
entire, usually square; base without wing.
Distribution — A genus of c. 25 species distributed in the tropics of the Old World:
Africa and Madagascar and in SE Asia, from India, China, through Malesia, to North
Australia and far into the Pacific; c. 18 species in Asia, Malesia and the Pacific, 1 spe-
cies in Australia; in Malesia 14 species.
Note — Several species are weakly defined, sometimes only distinguishable by the
colour of the ripe fruits. For convenience, of each species the geographical distribution
is indicated as an aid. Further collecting and study is needed.
KEY TO the SPECIEs
1a. Leaf blade (narrowly) trullate, base ± rounded or broadly cuneate. [Female flowers
and fruits not known.] — Central Sulawesi . . . . . . . . . . . . . . . . . 13. P. trullifolia
b. Leaf blade ovate, cordate or (sub)circular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Ovary and fruit globose, fruit c. 1 cm diam. or less, seeds margined . . . . . . . . . . 3
b. Ovary and fruit subglobose or ellipsoid, fruit 1 cm long or longer (if smaller, then
seeds without margin) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3a. Female flower and fruit solitary, with long pedicel; fruiting pedicel 1.5 – 2.5 cm
long. — New Guinea; montane . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. P. pedicellata
b. Female flower(s) and fruit either solitary, or fascicled, or clustered in a peduncled
raceme; fruiting pedicel c. 1.5 cm long or less . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Monoecious with female flowers (and fruits) and male flowers often in one single
raceme-like inflorescence. — New Guinea; montane . . . . . . . . . . . . 8. P. pisifera
b. Mostly dioecious, male flowers in a condensed (pedunculate) cluster . . . . . . . ???
5a. Fruit mostly several on a long common peduncle (if solitary, then usually co-axil-
lary with male peduncle). — Malesia, Taiwan, montane . . . . . . . . . 9. P. repanda
b. Fruit single or few, in a sessile or short-pedunculate cluster. — Lower montane or
lowland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Fruit 1 (rarely 2 or 3) per node, c. 1 cm diam., pedicelled, but without peduncle, red
when ripe. — Widespread in South India, Sri Lanka and SE Continental Asia; rare
in Malesia (N Sumatra, Peninsular Malaysia and Sabah); lowland and montane
area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. P. bodinieri
b. Fruit 1– 5 per node, 0.6 – 0.8 cm diam., fruits on a common peduncle to 1 cm long,
greenish when ripe. — Java, Sulawesi (Salayar Is.), Lesser Sunda Islands; lower
montane area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. P. perpusilla
7a. Fruit hairy. — Philippines (Mindanao) . . . . . . . . . . . . . . . . . . 12. P. trichocarpa
b. Fruit glabrous. — Widespread (incl. Philippines) . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Fruiting pedicel in solitary fruit about as long as or (much) longer than the fruit, c.
1.5 cm long or more (or fruit few fascicled on a peduncle). Stamens inserted some-
what above the base or at about halfway in the receptacle-tube, thecae straight
with connective ± narrow or thecae curved with connective broad in the middle.
[Fruit 1– 3 cm long.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
b. Fruiting pedicel shorter than the fruit, c. 1 cm long or less. Stamens inserted at the
base of the receptacle-tube, thecae straight, with connective ± narrow. (Flowers
not known in P. rizalensis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9a. Fruit less than 1 cm long, several in a cluster on a long peduncle. Seeds 3 mm
long. Dioecious? — Philippines (Luzon) . . . . . . . . . . . . . . . . . . 10. P. rizalensis
b. Fruit 1–1.5 cm long, 1– 6 in a cluster on the node. Seeds (3 –)4 – 5 mm long.
Mostly dioecious. — Widespread: Malesia, east to New Guinea, Queensland(?),
Christmas Is. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. P. mucronata
10a. Fruit 2 – 3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
b. Fruit less than 2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11a. Fruit solitary (or with 2). Seeds c. 4 mm long. [Male inflorescences not known.]
— Lesser Sunda Islands: Lombok . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. elbertii
b. Fruit solitary or few-fascicled on a common peduncle. Seeds 5 – 6 mm long. The-
cae ± straight, vertical. Disc simple, more or less 3-lobed. Style at apex not armed,
with deeply 3-lobed stigma. — Widespread: Bismarck Archipelago, Solomon Is-
lands, New Hebrides, Fiji, Samoa . . . . . . . . . . . . . . . . . . . . . . . 11. P. samoensis
12a. Fruit (1.3 –)1.5 – 2 cm long. Seeds 4 – 5 mm long, narrowly margined. — New
Guinea; lowland up to 1000(–1750) m altitude . . . . . . . . . . 3. P. erythrobacca
b. Fruit c. 1 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13a. Plant drying brown. Seeds c. 3 mm long, unmargined, edge rounded. — Lesser
Sunda Islands: Lombok, Flores; at 1500 – 3000 m altitude . . . 4. P. immarginata
b. Plant drying green. Seeds 5 mm long, with conspicuous broad square edge, not
obviously margined. — East Papua New Guinea; at 200 m altitude . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. P. viridifolia
1. Pilogyne bodinieri (H.Lév.) W.J.de Wilde & Duyfjes

Pilogyne bodinieri (H.Lév.) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 410. — Melothria bodi
nieri H.Lév., Fl. Kouy-Tcheou (1914) 122. — Zehneria bodinieri (H.Lév.) W.J.de Wilde & Duyfjes,
Thai Forest Bull, Bot. 32 (2004) 17; Blumea 51 (2006) 51; Fl. Thailand 9, 4 (2008) 545. — Type:
Bodinier 1957 (lecto E, designated by De Wilde & Duyfjes (2004); iso P), China, Kouy-Yang.
2 mm
h
a
2 cm
g
1 mm
l
1 mm
f
i
e
1 mm
d b c k
0.5 mm
Fig. 56. Pilogyne bodinieri (H.Lév.) W.J.de Wilde & Duyfjes. a. Shoot with male and female inflo-
rescences and fruits; b, c. male flower, from outside and opened; d. stamens; e, f. female flower, from
outside and opened; g. staminodes; h. fruit; i. seed; j. node with two fruits; k. detail of upper leaf blade
surface; l. seed (a – i: De Wilde & Duyfjes 21968 (Peninsular Malaysia); j – l: Madulid et al. PPI 11592
(Philippines, Palawan)).
Melothria perpusilla (Blume) Cogn. var. subtruncata Cogn. in A.DC. & C.DC., Monogr. Phan. 3
(1881) 608. — Type: Thwaites CP 1613 (lecto K, designated by De Wilde & Duyfjes (2004); iso
L), Sri Lanka.
Zehneria hookeriana auct. non (Wight & Arn.) Arn.: C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879)
624, p.p.
Melothria perpusilla auct. non (Blume) Cogn.: Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
607, p.p.; in Engl., Pflanzenr. 66, 4.275.I (1916) 106, p.p.; Craib, Fl. Siam. (1931) 764 (incl. var.
subtruncata).
Bryonia oxyphylla, Wallich Cat. 6697, nom. nud.
Bryonia cissoides, Wallich Cat. 6698, nom. nud.
Subherbaceous 2 – 6 m long climber, leafy stem 1– 2 mm diam., plant subglabrous,

generally drying (dark) brownish; monoecious. Leaves: petiole 2 – 5 cm long; blade
rarely shallowly lobed, ovate-triangular, 4 –12 by 3 –10 cm, base subtruncate or broadly
shallowly cordate, margin denticulate. Inflorescences in male a peduncled 3 –10-flow-
ered short or sometimes ± spike-like proliferous raceme; peduncle 1– 5 cm long, usually
co-axillary with a previously developed single female flower; female flowers solitary or
rarely few, subumbellate on the node. Male flowers: pedicel 1– 4 mm long; receptacle-
tube c. 3 by 2 mm, inside with long hairs, especially at the throat; sepals 0.5 mm long;
petals ovate, c. 2 mm long, subacute, inner surface and apex hairy; stamens inserted
halfway the receptacle-tube or lower (not at the base of the tube), filaments 1– 2 mm
long, subglabrous or long-haired about the middle, anthers circular in outline, 1 mm
diam., thecae curved, the two nearly forming a ring, connective ± hairy, not produced;
disc depressed globose, 1 mm diameter. Female flowers: pedicel slender, 2 – 5 mm long;
ovary ovoid-globose, c. 3 by 2.5 mm, glabrous (except minute raised gland-dots), neck
1 mm long; perianth as in male flower but petals 2.5 – 3 mm long; style c. 3 mm long,
glabrous, stigma 3-lobed, 1.5 – 2 mm diam., papillose; disc 0.5 mm high; staminodes
slender, c. 2 mm long, the basal portion adnate with the receptacle-tube. Fruit solitary
(rarely 2 or 3), green, ripening red, globose, 0.8 –1.2 cm diam., glabrous, finely netted or
pitted when dry; fruiting pedicel 0.3 –1 cm long. Seeds several or many, pale brownish,
(narrowly) elliptic, c. 5 by 3 – 3.5 mm, narrowly margined, smooth. — Fig. 56; Plate
20a, b.
Distribution — Widespread; in Sri Lanka and South India, and from northern India to
China (including Taiwan), Indo-China, and Thailand; in Malesia: Sumatra, Peninsular
Malaysia, Sabah, and Philippines (Palawan).
Habitat & Ecology — Disturbed places, forest edges, and scrub; at 500 –1700 m
altitude; flowering and fruiting throughout the year.
2. Pilogyne elbertii (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne elbertii (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes. Reinwardtia 12, 5 (2009) 410.
— Zehneria elbertii W.J.de Wilde & Duyfjes, Blumea 51 (2006) 54. — Type: Elbert 1637 (holo L;
iso FR, not seen), Lesser Sunda Islands (Lombok).
Stoutish climber c. 5(?) m long, stem (1–)2 mm diam., glabrescent, plant greenish
brown on drying; dioecious(?). Probract linear, c. 5 mm long. Leaves: petiole 3 – 6.5 cm
long, glabrous; blade unlobed, broadly ovate, 6 –10 by 6.5 – 8.5 cm, both surfaces gla-
brous but faintly scabrous above by minute cystoliths, base broadly shallowly cordate,
margin sparsely short-dentate. Male inflorescences and male flowers not known. Female
flowers 1 (or 2) solitary at the node; pedicel c. 10 mm long; ovary ellipsoid to narrowly
ellipsoid, c. 7 by 1.5 mm, glabrous, neck 1 mm long; receptacle-tube campanulate,
c. 2.5 by 3 mm, throat densely woolly hairy, hairs 0.5 –1 mm long; sepals 0.5(–1) mm;
petals c. 3 mm long, outside glabrous, inside sparsely gland-hairy; style c. 2.5 mm long,
at apex with 3 style-arms 0.3 mm long, stigmas down-curved, (narrowly) ovoid, thick,
papillose, c. 1.5 mm long; staminodes c. 2 mm long, inserted towards the base of the
receptacle-tube, densely long woolly hairy near the apex; disc large, 0.5(–1) mm high,
c. 2 mm wide, margin somewhat irregularly sinuate. Fruit solitary (or 2), ripening
red(?), narrowly ellipsoid, (2.5 –)3 by c. 1 cm, apex apiculate; exocarp cartilaginous,
smooth (not pitted); fruiting pedicel 1–1.5 cm long. Seeds numerous, palish, nearly flat,
ovate, c. 4 mm long, smooth, unmargined.
Distribution — Lombok: north-eastern flank of Mt Rinjani, Sembalun Highlands,
known only from the type.
Habitat & Ecology — In scrub-forest; on loamy soil over volcanic breccia; at 1100 –
1300 m altitude; flowering and fruiting: May.
Notes — 1. This species, known only from one female flowering and fruiting col-
lection, is of a stout habit and obviously belongs, on account of the basal insertion of
the staminodes in the female flower, to Pilogyne. However, it also has traits of Neoach
mandra, e.g. the solitary rather long-pedicelled fruits, the smooth (not pitted) pericarp,
and the unmargined seeds.
2. Pilogyne elbertii keys out beside the widespread P. samoensis from the Pacific. The
sole specimen of P. elbertii differs in various minor, characteristics, sometimes difficult
to define and additional material is needed to prove its status as a distinct species.
3. Pilogyne erythrobacca (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne erythrobacca (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Gard. Bull. Singapore 61,
1 (2009) 210. — Zehneria erythrobacca W.J.de Wilde & Duyfjes, Blumea 51 (2006) 55, f. 13a – h.
— Type: Brass 21720 (holo L; iso A), Papua New Guinea (Menapi, Cape Vogel Peninsula).
Melothria indica auct. non Lour.: Peekel, Fl. Bismarck Archip., translated by E.E.Henty (1984) 543,
f. 867.
Diplocyclos palmatus auct. non (L.) C.Jeffrey: Peekel, Fl. Bismarck Archip., translated by E.E.Henty
(1984) 547, f. 873, p.p.
Climber or creeper, 1.5 – 5 m long, leafy stem 1(– 3) mm diam., plant sparsely minute-
ly hairy, glabrescent, brown on drying; monoecious. Probract minute. Leaves: petiole
1– 5.5 cm long, glabrous or scabrous; blade unlobed or sometimes ± hastate or lobed
to 1/5 deep, ovate, 3 – 9(–11) by 2.5 – 9(–14) cm, subglabrous or scabrous above, with
minute cystoliths, glabrous beneath but veins sometimes ± hairy, base (narrowly or)
broadly cordate, margin (sparsely) coarsely dentate, apex short or long acuminate. Male
inflorescences subsessile or peduncled racemes; peduncle up to 3 cm long; racemes
0.1– 0.5 cm long (rarely up to 2 cm long), 5 –15-flowered, flowers dense or loose, with
rather long pedicels, sometimes mixed with female flowers. Male flowers: pedicel 5 –10
mm long; expanded perianth 4 – 6(– 8) mm diam.; receptacle-tube 1.5 – 2.5 by 2 – 3 mm,
outside glabrous, inside (especially at the throat) densely white woolly hairy, hairs to
2 cm
b
2 mm
2 mm
i 1 mm 2 mm d g
h
1 mm 1 mm
j e f
Fig. 57. a – h. Pilogyne erythrobacca (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes. a. Branch
with fruits; b. ditto; c, d. nodes with male inflorescences; e, f. male flowers; g. node with female in-
florescence; h. seed. — i, j. Pilogyne viridifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes.
i, j. Female flowers (a, g, h: Widjaja EAW 6961; b: Brass 21720, type; c – f: Brass 8104; i, j: Brass
23914, type).
1 mm long; sepals 0.5 –1.5 mm long; petals 2 – 3(– 4) mm long, minutely gland-hairy
outside; stamens inserted towards the base of the receptacle-tube, filaments 1.5 – 2 mm
long, woolly hairy in upper half, anthers partly exserted, ellipsoid, 1(–1.5) mm long,
thecae ± straight, connective narrow; disc depressed globose or at apex ± concave,
0.5(–1) by 1–1.5 mm, not or faintly lobed. Female flowers few or several in a subsessile
fascicle-like short raceme, or one female flower below male flowers in a short raceme;
pedicel 5 –10 mm long; ovary ellipsoid(-fusiform), 3.5 – 6 by 1.5 – 3 mm, glabrous (ex-
cept for some glandular papillae), neck 0.5 –1 mm long; style 1.5 – 3 mm long, stigma
c. 2 mm diam., consisting of 3 parts on 0.2–0.3 mm long style-arms; disc 0.5 mm high;
staminodes inserted about halfway in the receptacle-tube, 0.5 –1.5 mm long, glabrous
or woolly hairy at apex. Fruit 1– 3(– 5) in loose fascicles, ripening orange or red; (sub-
globose-)ellipsoid, (1.3 –)1.5 – 2 by 1–1.5 cm; exocarp finely pitted or tessellated; fruit-
ing pedicel 1– 2.5 cm long. Seeds numerous, pale, ovate, 4 – 5 by 3 – 3.5 mm, not orna-
mented, margin narrow, edge square and often with a groove in the middle.
Field-notes — Milliken 1315 records that the fruit is edible. — Fig. 57a – h.
Distribution — Throughout New Guinea and Bismarck Archipelago.
Habitat & Ecology — Secondary forest, riverbanks, scrub-land; lowland to 1000
(–1750) m altitude; flowering and fruiting throughout the year.
Notes — 1. Pilogyne erythrobacca is common in New Guinea where it largely seems
to replace P. mucronata. The latter is a common lowland species in most of Malesia.
The fruits of P. mucronata, which is a rare species in New Guinea, are green when
ripe, turning red only when remaining for a longer time on the plant, a condition not
mentioned on the field labels. The fruits of P. erythrobacca are orange or red when ripe,
a condition often recorded on the labels.
2. Telford (Queensland), in litt., mentions on-going renewed studies in Zehneria-like
plants from Australia and indicates an as yet undescribed species with female flowers
on long pedicels and red fruit distributed within Australia in coastal North Queensland
and the islands in the Torres Strait, including Dauan Is., close to Papua New Guinea.
This taxon might be identical with Pilogyne erythrobacca. A better understanding of
the present species and related taxa from Australia will call into dispute the occurrence
of Pilogyne mucronata in this area as well.
4. Pilogyne immarginata (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne immarginata (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009)
410. — Zehneria immarginata W.J.de Wilde & Duyfjes, Blumea 51 (2006) 63. — Type: Loeters
1580 (holo L), Lesser Sunda Islands (Flores).
Climber 1– 3 m long, leafy stem 1– 2 mm diam., plant subglabrous, dark on drying;

monoecious or dioecious. Leaves: petiole 1– 2.5 cm long, sparsely hairy; blade nar-
rowly triangular or ovate-elliptic, 5 – 8 by 2 – 4.5 cm, glabrous above but scabrous by
numerous small cystoliths, glabrous below or (scabrid-)hairy on veins, margin finely
or coarsely dentate. Male inflorescences a peduncled raceme; peduncle 1– 3 cm long;
racemes short or long, sometimes with flowers ± tiered, to 15-flowered, up to 2 cm
long. Male flowers: pedicel 4 – 6 mm long; expanded perianth 5 – 6 mm diam.; recepta-
cle-tube c. 2 by 3 mm, outside glabrous, inside densely woolly hairy, especially in the
upper half, hairs 0.5 –1 mm long; sepals c. 0.5 mm long; petals c. 2.5 mm long, outside
papillose, inside hairy towards base; stamens inserted near the base of the receptacle-
tube, filaments c. 2 mm long, glabrous, anthers just exserted, ellipsoid, 1–1.5 mm long,
thecae somewhat curved, connective ± narrow at apex; disc c. 1 mm diam., unlobed.
Female flowers solitary or few in subsessile short raceme; pedicel (5 –)10 –15 mm long;
ovary ellipsoid-fusiform, 2.5 – 3 mm long, glabrous, neck c. 0.5 mm long; perianth as
in male flower but somewhat larger, subglabrous, style 2 – 3 mm long, style-arms c. 1
mm long, stigma-lobes longer than broad, c. 1 mm long, papillose; staminodes 2 – 3
mm long; disc c. 0.5 mm high. Fruit solitary or 2 – 5 in a subsessile fascicle, ripening
pale yellowish, subglobose or ellipsoid, 0.8 –1 by 0.6 – 0.8 cm, exocarp minutely pit-
ted; fruiting pedicel (0.5 –)1.5 – 2.5 cm long. Seeds numerous, ovate in outline, c. 3 by
2 mm, not sculptured, smooth but sometimes very short-hairy at the ends, margin not
apparent, edge rounded.
Distribution — Lesser Sunda Islands: Lombok (Mt Rinjani), Flores (Mt Kelimoetoe,
Mt Ranaha).
Habitat & Ecology — Edges of Myrica scrub, grassy places, margins of Cusuarina
forest; volcanic (lapilli) soils; at 2000 – 3500 m altitude; flowering and fruiting: March
to June.
Notes — 1. Pilogyne immarginata resembles in general habit the more widespread
P. repanda from a similar habitat; the latter differs, e.g. in its short fruiting pedicel
0.3 – 0.6 cm long. Pilogyne immarginata is characterised by the long-stalked stigma
lobes, by the long-pedicelled fruits and the unmargined small seeds, with slightly con-
vex faces. Unmargined seeds and long-pedicelled fruits are characters of the genus
Neoachmandra, but P. immarginata dries dark, and agrees furthermore in all traits,
including details of the male flowers, with the genus Pilogyne. Pilogyne immarginata
also resembles P. pedicellata from mountainous New Guinea, the latter differs in glo-
bose fruits and somewhat larger, narrowly margined seeds.
2. The fruits of Kostermans & Wirawan 733, from Flores, deviate in being small,
less than 1 cm long; the collection approaches P. perpusilla from the same area.
5. Pilogyne mucronata (Blume) W.J.de Wilde & Duyfjes

Pilogyne mucronata (Blume) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 410. — Bryonia
mucronata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 923 (incl. var.); Ser. in DC., Prodr. (1828) 304 (incl.
var. denticulata). — Zehneria mucronata (Blume) Miq., Fl. Ned. Ind. 1, 1 (1856) 656; I.Telford,
Fl. Australia 8 (1982) 182; C.M.Simmons & W.J.de Wilde, Blumea 45 (2000) 237, f. 1; W.J.de
Wilde & Duyfjes, Blumea 51 (2006) 65, f. 14, pl. 8c, d. — Melothria mucronata (Blume) Cogn.
297. — Type: Blume s.n. “Pariagengie” (lecto L, barcode L0048324, designated by Simmons & De
Wilde (2000); iso L, 2 sheets), Java.
Zehneria alba Ridl., J. Straits Branch Roy. Asiat. Soc. 45 (1906) 195. — Melothria alba (Ridl.) Cogn.
in Engl., Pflanzenr. 66, 4.275.I (1916) 109. — Type: Ridley s.n. (K, not seen), Christmas Is. (Indian
Ocean).
Melothria lobata Merr., Philipp. J. Sci., C 7 (1912) 104. — Type: Vanoverbergh 1241 (not seen),
Philippines (Luzon).
Bryonia arguta Span., Linnaea (1841) 206, nom. nud. — Voucher specimen: Zippelius s.n., barcode
L0129472 (L), Timor.
g
a 2 cm 1 mm
1 mm
2 mm
e f b c
Fig. 58. Pilogyne mucronata (Blume) W.J.de Wilde & Duyfjes. a. Twig with male inflorescences; b, c.
male flowers; d. twig with female inflorescences; e, f. female flowers; g. portion of upper leaf blade
surface, with cystoliths; h. node with sessile infructescence; i. margined seed (a – c: De Wilde & Duyfjes
21727; d – g: De Wilde & Duyfjes 21728; h, i: Backer 36679).
Climber to 5 m long, leafy stem 1– 2 mm diam., plant finely hairy, glabrescent, dark
on drying; usually dioecious, sometimes monoecious. Leaves: petiole 1– 5 cm long,
(sparsely) scabrid; blade unlobed or shallowly (deeply) lobed, triangular or ovate, or
± 5-angular, (2 –)4 – 8 by (2 –)4 –7 cm, subglabrous with cystoliths above, subglabrous
beneath, but variously hairy on the veins, margin subentire or shallowly dentate. Male
inflorescences peduncled, densely or loosely flowered racemes, occasionally prolifer-
ating; peduncle to 5 cm long. Male flowers: pedicel 2 – 5 mm long; expanded perianth
3 – 5 mm diam.; receptacle-tube 2.5 – 4 by 2 – 3.5 mm, outside glabrous or with sparse
hairs, inside variously hairy; sepals 0.5 –1 mm long; petals 2 – 3 mm long, minutely hairy
inside; stamens inserted near the base of the receptacle-tube, 2(– 3) mm long, filaments
completely hairy or at least in the lower half, anthers somewhat longer than broad, c. 1
mm diam., thecae straight or ± curved, connective narrow, ± hairy, not produced; disc
depressed globose, sometimes faintly 3-lobed, c. 1 mm diameter. Female flowers in
few- (or many-)flowered, sessile or peduncled clusters, occasionally mixed with male
flowers; pedicel 1– 2(– 5) mm long; ovary narrowly ovoid-ellipsoid, 2 – 3 mm long, at
apex narrowed into a neck c. 1 mm long, glabrous; style 2 – 2.5 mm long, stigma c. 2
mm diam., consisting of 3 papillose-hairy lobes; staminodes linear, variously hairy; disc
c. 0.5 mm high. Fruit (1–)2 – 6 per cluster, green, ultimately ripening red, subglobose,
ellipsoid or narrowly ellipsoid, 1–1.5 cm long, apex rounded with minute point; exo-
carp minutely pitted, glabrous; fruiting pedicel 0.2 – 0.5(–1) cm long. Seeds numerous,
whitish or pale brown, ovate-elliptic, (2 –)3 – 4(– 5) mm long, smooth, margin narrow
but distinct. — Fig. 58; Plate 19c, d.
Distribution — West Malesia to NE Australia (Telford, 1982, no specimens seen);
in Malesia: Sumatra, Java (also Christmas Is.), Philippines, Sulawesi, Lesser Sunda
Islands, Moluccas, New Guinea (West Papua and Papua New Guinea); no records from
Peninsular Malaysia and Borneo.
Habitat & Ecology — In scrub and forest edges, lowland to 1600(– 2000) m altitude;
flowering and fruiting throughout the year.
Notes — 1. Pilogyne mucronata resembles P. maysorensis, an insufficiently known
species from South India, similar in ellipsoid fruits. The latter differs in its usually single
fruit, co-axillary with the male inflorescence. However, this latter condition is occasion-
ally also found in P. mucronata from Papua New Guinea, e.g. in Darbyshire 197.
2. Zehneria alba was described as endemic to Christmas Is., distinct by large 4 mm
long male petals, and glabrous filaments. We have not seen the type, but in two other
collections from Christmas Is., Andrews s.n. (BO) and Mitchell M 21 (KEP), the male
flowers are smaller (petals only 2 mm long) and the filaments hairy.
6. Pilogyne pedicellata (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne pedicellata (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009)
410. — Zehneria pedicellata W.J.de Wilde & Duyfjes, Blumea 51 (2006) 69. — Type: Brass 30732
(holo L; iso K), Papua New Guinea, Eastern Highlands Province (Mt Wilhelm).
Creeper or climber, 2 – 4 m long, leafy stem 1(– 2) mm diam., plant sparsely fine-
hairy, glabrescent, blackish brown on drying; dioecious. Leaves: petiole 0.5 –1(– 2)
cm long, subglabrous; blade unlobed, narrowly triangular or (narrowly) ovate, 4 –7 by
1.5 – 3.5 cm, both surfaces glabrous or the veins variously scabrid-hairy, upper surface
with cystoliths, base shallowly cordate with broad sinus, margin shallowly (deeply) den-
tate, apex long acuminate. Male inflorescences peduncled racemes; peduncle slender,
3 –7 cm long, raceme 2 – 3 mm long, with up to 10 flowers, clustered in a condensed
subumbel. Male flowers: pedicel 5 –10 mm long; expanded perianth c. 4 mm diam.;
receptacle-tube c. 1.5 by 2 – 2.5 mm, glabrous but throat minutely sparsely hairy; sepals
0.5 mm long, suberect; petals (long-)triangular, c. 1.5 mm long, outside minutely papil-
lose-hairy; stamens inserted at the base of the receptacle-tube, filaments c. 1.5 mm long,
glabrous, anthers subglobose, c. 1.5 mm diam., thecae half-circular, almost touching at
both ends, nearly horizontal, connective broad in the middle, convex, minutely hairy
along the thecae; disc c. 1.5 mm diam., deeply 3-lobed, lobes half-globose. Female
flowers solitary; pedicel 10 – 25 mm long; ovary globose, 2 – 3 mm diam., glabrous, neck
c. 2 mm long; perianth as in male flower but longer, receptacle-tube c. 2 by 3.5(– 4)
mm, throat finely woolly hairy; style c. 2.5 mm long, glabrous, stigma c. 1.5 mm diam.,
consisting of 3 arms 0.3 mm long, each bearing a subglobose papillose stigma-lobe
0.5 – 0.7 mm diam.; staminodes distinct, subulate, c. 2 mm long, glabrous; disc c. 1 mm
high, deeply 3-lobed. Fruit solitary at the node, ripening blackish, globose, 0.6 – 0.8 cm
diam., exocarp faintly minutely pitted, glabrous; perianth not persistent but flower-neck
remaining as a minute beak at apex; fruiting pedicel 1.5 – 2.5 cm long. Seeds rather nu-
merous, ovate-elliptic, c. 4 by 3 mm, not ornamented, margin narrow but distinct.
Distribution — New Guinea (Papua New Guinea (Western and Eastern Highlands
Provinces)).
Habitat & Ecology — Forest edges and secondary growth; riverbanks; at 2000 – 2700
m altitude; flowering and fruiting throughout the year.
Notes — 1. Pilogyne pedicellata is in habit similar to P. pisifera. The latter differs
in a shorter fruiting pedicel, c. 1.5 cm long or less.
2. In Vinas UPNG 4874 (L) the leaves are more conspicuously dentate compared to
the rest of the collections.
3. Female specimens, with solitary, long-pedicelled flowers or fruits may be con-
fused with members of the genus Neoachmandra; the latter genus is monoecious, with
the male flowers few or solitary, not as in the present species in a peduncled raceme.
7. Pilogyne perpusilla (Blume) W.J.de Wilde & Duyfjes

Pilogyne perpusilla (Blume) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 410. — Cucurbita
perpusilla Blume, Cat. Gew. Buitenzorg (1823) 105. — Bryonia perpusilla (Blume) Blume, Bijdr.
Fl. Ned. Ind. 15 (1826) 926; Miq., Fl. Ned. Ind. 1, 1 (1856) 660. — Melothria perpusilla (Blume)
Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 607; Backer in Backer & Bakh.f., Fl. Java 1
(1964) 297, p.p. — Zehneria perpusilla (Blume) Bole & M.R.Almeida, J. Bombay Nat. Hist. Soc.
79, 2 (1983) 315, for the type only; C.M.Simmons & W.J.de Wilde, Blumea 45 (2000) 238; W.J.de
Wilde & Duyfjes, Blumea 51 (2006) 70. — Bryonia stipulacea Willd. var. perpusilla (Blume) Ser.
in C., Prodr. 3 (1828) 307. — Zehneria scabra auct. non (L.f.) Sond.: C.Jeffrey, Kew Bull. 15 (1962
(‘1961’)) 369, p.p., for the synonym Melothria perpusilla (Blume) Cogn. only. — Type: Blume s.n.,
(lecto L, barcode L0048312, designated by Simmons & De Wilde (2000); iso L, 4 sheets), Java.
Cucurbita scabra Blume, Cat. Gew. Buitenzorg (1823) 105, non Bryonia scabra L.f., 1781. — Bryonia
scabrata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 923; Ser. in DC., Prodr. 3 (1828) 304; Miq., Fl. Ned.
Ind. 1, 1 (1856) 659. — Type: Blume s.n. “Korreg Kottok” (lecto L, barcode L0048319, designated
by Simmons & De Wilde (2000); iso L), Java, Salak.
Climber 2 – 3 m long, leafy stem 1(– 2) mm diam., plant subglabrous, brownish black-
ish on drying; monoecious or dioecious. Leaves: petiole 1– 5 cm long, subglabrous;
blade unlobed or 3(– 5)-lobed and irregular in shape with the lobes up to halfway,
triangular or ovate, 4 –10 by 3 –7 cm, scabrous with cystoliths above, (sub)glabrous
beneath, margin coarsely or finely (serrate-)dentate. Male inflorescences subsessile or
to 1 cm (in Lesser Sunda Islands to 3 cm) long peduncled short racemes, rather few-
flowered (sometimes proliferating in Lesser Sunda Islands), mixed with few female
flowers or not, and often with a single female flower co-axillary. Male flowers: pedicel
3 – 5 mm long; expanded perianth 3 – 4 mm diam.; receptacle-tube c. 1.5 by 1.5 mm,
outside glabrous, throat and inside woolly hairy; sepals less than 0.5 mm long, erect;
petals (narrowly) triangular, 1.5 – 2 mm long, ± hairy at both surfaces; stamens inserted
near the base of the receptacle-tube, filaments c. 1.5 mm long, long-woolly hairy at
and below the middle, anthers subglobose, c. 1 mm diam., thecae curved, connective
rather broad in the middle, hairy; disc 1 mm diam., deeply 3-lobed, lobes half-globose.
Female flowers in the same way disposed as male flowers; pedicel 2 – 5 mm long; ovary
globose, c. 2 mm diam., glabrous, neck 1 mm long; style c. 2.5 mm long, slender, gla-
brous, stigma 1 mm diam., deeply 3-lobed; staminodes c. 1.5 mm long, woolly hairy
in upper half; disc 0.5 mm high, 3-parted. Fruit solitary or 2 – 5 in a sessile or to 1 cm
long peduncled cluster, ripening greenish, globose, 0.6 – 0.8 cm diam., faintly finely
pitted; fruiting pedicel c. 0.5 cm long. Seeds 5 –10, flat, elliptic, 2.5 – 3.5 mm long, with
narrow margin.
Distribution — West Java, Sulawesi (Salayar Is.), Lesser Sunda Islands: (?Bali, Lom
bok, Sumbawa, Flores).
Habitat & Ecology — Lower montane area, in forest edges and low scrub; at (300 –)
500 –1000 m altitude; flowering and fruiting throughout the year.
Note — In a previous publication on Zehneria in Java (Simmons & De Wilde, 2000)
Pilogyne perpusilla was accepted as a not well-defined species beside P. mucronata and
P. repanda. Pilogyne perpusilla seems closer to P. repanda because both have rounded
anthers with curved more or less horizontal thecae, and a more or less convex connec-
tive. In the Lesser Sunda Islands a division of Pilogyne into 3 taxa can be made, viz.
P. mucronata, P. repanda, and a more or less intermediate form. This intermediate form
differs in details from the limited material available of P. perpusilla from West Java
(Mt Salak), but still warrants its inclusion in that species. It is likely that the differences
of these 3 feebly defined species are (at least partly) ecotypic, and they may hybridize
as well. Consequently, a part of the collections cannot be determined with certainty.
Similar specimens, but with thickish (not flat) unmargined seeds have been described
as a separate species, P. immarginata (Lesser Sunda Islands).
8. Pilogyne pisifera (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne pisifera (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 410.
— Zehneria pisifera W.J.de Wilde & Duyfjes, Blumea 51 (2006) 71, f. 16, 17. — Type: Brass 29609
(holo L; iso K), Papua New Guinea, Morobe Province (Kaindi).
Melothria cissybium M.Jacobs, Blumea 7 (1954) 617, f. 2, p.p., excluding the type which is Urceo
discus belensis.
2 cm
3 mm
c
1 mm 1 mm f
d
0.5 mm
e e'
1 mm 1 mm i
h g b
Fig. 59. Pilogyne pisifera (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes. a. Twig with inflores-
cences; b. inflorescence, enlarged, showing fruits, and at apex one female flower and male flowers;
c, d. male flowers; e, e'. stamens, ab- and adaxially respectively; f, g. female flowers; h. fruit with
withered perianth; i. seed (all: Hoogland & Pullen 5926).
Climber 0.5 – 2 m long, leafy stem 1 mm diam., plant sparsely hairy, early glabres-
cent, blackish on drying; monoecious. Leaves: petiole 1– 3.5 cm long, scabrid or sub-
glabrous; blade triangular or ovate, 3 – 8 by 1.5 – 5 cm, (sub)glabrous with cystoliths
above, variously scabrid along the veins or subglabrous beneath, base subtruncate or
cordate, with broad sinus, margin entire, variously dentate, rarely (shallowly) lobed-
dentate, apex (long) acuminate. Male inflorescences 2 – 5 cm long peduncled, slender,
spike-like racemes 3 – 4 cm long (flowers in young inflorescences few-fascicled at
the end of the peduncle but later proliferating), flowers frequently 2 – 5-tiered, but
also dispersed, commonly mixed with female flowers. Male flowers: pedicel 2 – 3 mm
long, perpendicular to the rachis; expanded perianth 3 – 4 mm diam.; receptacle-tube
1.2 –1.5 by 2.2 – 3 mm, glabrous, throat minutely hairy inside; sepals 0.3 – 0.5 mm long,
erect, at base connected by a line or low rim; petals triangular, 1–1.5 by 1–1.5 mm,
outside minutely hairy; stamens inserted near the base of the receptacle-tube, filaments
1–1.5 mm long, glabrous, anthers subglobose, 0.5 mm diam., thecae curved, nearly
horizontal, connective broad in the middle, convex and with minute hairs adaxially;
disc 1 by 1.5 mm, consisting of 3 rounded lobes. Female flowers in the same way dis-
posed as the male flowers, and frequently mixed with them; pedicel 3 – 5(–7) mm long;
ovary globose, 1.5 – 2 mm diam., glabrous, neck 0.5(–1) mm long; style 1(–1.5) mm
long, glabrous, stigma consisting of 3 down-curved papillose lobes, in all 1 mm diam.;
staminodes distinct, inserted near the base of the receptacle-tube, subulate, 1 mm long,
glabrous; disc 0.3 mm high, faintly lobed. Fruit 1 (or 2) at the node or usually several
in the spike-like inflorescence, often when still proliferating and producing flowers
at the apex, green, ripening pale yellow or red, globose, 0.5 – 0.7 cm diam., glabrous,
withered perianth persisting; exocarp not or faintly pitted; fruiting pedicel 0.2 – 0.6 cm
long. Seeds c. 10, pale brown, elliptic, c. 2.5 by 1.5 mm, not ornamented, margin nar-
row, faint. — Fig. 59.
Field-notes — Corolla valvate, yellow. Fruits spherical, said to be edible; exocarp
red when ripe (Takeuchi 10518). Flowers yellow (Pullen 427).
Distribution — New Guinea (eastern part of mountainous Papua New Guinea: Cen-
tral, Eastern Highlands, Morobe, Southern Highlands, and Western Highlands Prov-
inces).
Habitat & Ecology — Grassy places, scrub and open Nothofagus forest; at 1400–
2700 m altitude; flowering and fruiting: May to December.
Note — Fruits have been recorded as pale yellow or red, also as unripe and green, but
such green fruits appear to contain mature seeds. The persistent perianth dries blackish.
The recorded yellow flower colour (Pullen 427), an aberrant colour in Pilogyne, needs
further confirmation.
9. Pilogyne repanda (Blume) W.J.de Wilde & Duyfjes

Pilogyne repanda (Blume) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 410. — Zehneria repan
da (Blume) C.M.Simmons, Blumea 45 (2000) 240, f. 2. — Bryonia repanda Blume, Bijdr. Fl. Ned.
Ind. 15 (1826) 923; Ser. in DC., Prodr. (1828) 305. — Melothria punctata (Thunb.) Cogn. in A.DC.
& C.DC., Monogr. Phan. 3 (1881) 615, p.p. — Zehneria scabra auct. non (L.f.) Sond.: C.Jeffrey,
Kew Bull. 15 (1962 (‘1961’)) 369, p.p., for the synonym Bryonia repanda Blume only. — Type:
Blume s.n. (lecto L, barcode L0048320, designated by Simmons & De Wilde (2000)), Java.
Zehneria exasperata Miq., Fl. Ned. Ind. 1, 1 (1856) 655. — Type: Horsfield s.n. (lecto U, barcode
U0122777, designated by Simmons & De Wilde (2000)), Java.
Melothria cordata auct. non (Thunb.) Cogn.: Backer in Backer & Bakh.f., Fl. Java 1 (1964) 297, p.p.
Zehneria maysorensis auct. non (Wight & Arn.) Arn.: C.Jeffrey, Kew Bull. 15 (1962 (‘1961’)) 371,
p.p., for the Sumatra record only.
Climber 2 – 4 m, leafy stem 1– 2 mm diam., plant glabrous or sparsely (or densely)

hairy, dark on drying; mostly dioecious. Leaves: petiole 1– 3 cm long, glabrous or
hairy; blade unlobed or irregularly 3(– 5)-angular, with lobes to c. 1/4 deep, triangu-
lar, ovate(-elliptic) in outline, 3 – 8 by 2 – 6 cm, scabrous with cystoliths above, lower
surface (almost) glabrous, but veins subglabrous or conspicuously bristly hairy, base
truncate or broadly cordate, margin entire or coarsely serrate-dentate (to 5 mm deep,
including mucro 0.5 mm), apex acute-acuminate. Male inflorescences 1– 2.5(– 5) cm
long peduncled few- or many-flowered clusters, or flowers dispersed or ± tiered in
racemes to 1.5 cm long, sometimes co-axillary with a shorter second inflorescence.
Male flowers: pedicel ± patent, 2 – 4(–7) mm long; expanded perianth 4 – 8 mm diam.;
receptacle-tube 1.5 – 2.5 by 2 – 3 mm, (sub)glabrous but throat usually densely woolly
hairy; sepals c. 0.5 mm long, patent; petals 1.5 – 2.5 mm long, outside minutely papil-
lose-hairy; stamens inserted at the base of the receptacle-tube, filaments 1– 2 mm long,
glabrous or (woolly) hairy at the middle, anthers ± globose, 1–1.5 mm diam., thecae
half-circular, nearly horizontal, connective broad in the middle, convex, (partly) hairy;
disc depressed globose, c. 1.5 mm diam., shallowly 3-lobed. Female flowers in the same
way disposed as male flowers but peduncle shorter, up to 1.5 cm long; pedicel 2 – 6 mm
long; ovary globose, c. 1.5 mm diam., glabrous, neck (1–)1.5 mm long; style 2 – 2.5 mm
long, glabrous, stigma deeply 3-lobed, c. 1 mm diam., papillose; staminodes subulate,
c. 1.5 mm long, glabrous; disc 0.5 mm high, 3-lobed or 2- or 3-parted. Fruit in (short)
peduncled clusters of up to 6, sometimes with a single fruit co-axillary, glaucous-green,
ultimately ripening purplish black, globose, 0.5 – 0.8(–1) cm diam., glabrous; exocarp
finely pitted; fruiting pedicel 0.3 – 0.6 cm long. Seeds numerous, elliptic, c. 3 by 2 mm,
not ornamented, margin absent or narrow and faint. — Fig. 60.
Field-notes — Perennial creeper; stem partly lianescent, at base to c. 1 cm diam.,
often widely branched and covering several square metres. Flowers white fading to pale
yellowish.
Distribution — Taiwan (1 collection, Van Steenis 20752); in Malesia: Sumatra, Java,
Philippines (Luzon, Visayas, Mindoro), Sulawesi, Lesser Sunda Islands (Bali); not
known from Borneo.
Habitat & Ecology — Scrub and forest edges; at 1400 – 2700 m altitude; flowering
and fruiting throughout the year.
Note — Several collections tend to be intermediate with Pilogyne mucronata, e.g.
in female flowers having a relatively short neck, or margined seeds. It is possible that
such specimens, from higher altitudes, are hybrids or ecotypes of P. mucronata which
are much resembling P. repanda, sharing the same harsh montane climate in the more
open places.
1 cm i
g
1 mm j
1 mm
f
1 mm
1 mm
1 mm h
d
b 1 mm e
Fig. 60. Pilogyne repanda (Blume) W.J.de Wilde & Duyfjes. a. Portion of twig with male inflores-
cences; b, c. male flowers; d. apex of stamens, anthers about as long as wide; e, f. opened male flow-
ers of different specimens, somewhat schematic; g. node with female inflorescence; h. nearly mature
female bud; i. portion of twig with infructescences; j. seed, without clear margin (a – d: Kleinhoonte
388; e: Kleinhoonte 231; f: Anon., Herb. J.J. Smith 134; g, h: De Wilde & Duyfjes 21764; i, j: Korthals
s.n., barcode L 0048302).
d
0.5 mm
a
2 cm
b
c
3 mm 1 mm
Fig. 61. Pilogyne rizalensis W.J.de Wilde & Duyfjes. a. Node with infructescence; b. detail of infruc-
tescence; c. seed, faintly hairy; d. detail of upper leaf blade surface (all: Ramos 2023, type).
10. Pilogyne rizalensis W.J.de Wilde & Duyfjes

Pilogyne rizalensis W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2008) 410. — Type: Ramos BS 2023
(holo BM; iso BRI, PNH (not seen)), Luzon.
Herbaceous climber to 3 m long, leafy stem 1– 2 mm diam., plant largely glabrous,

dark on drying; dioecious. Leaves: petiole 3 – 4.5 cm long, sparsely hairy; blade unlobed,
upper surface glabrous, veins sparsely hairy below, ovate, 5 –10 by 4 – 8 cm, base cor-
date, margin (coarsely) shallowly dentate, apex acute-acuminate. Male inflorescences,
male and female flowers not known. Infructescences with (2 –)5 –12 fruits in a cluster
on a (1–)2 –7 cm long peduncle. Fruit ellipsoid, 0.7– 0.9 cm long, 0.6 – 0.7 cm wide,
apex round with a narrow c. 0.5 mm long beak; exocarp minutely pitted, glabrous; fruit-
ing pedicel 0.2 – 0.3 cm long. Seeds numerous, pale brown, ovate-elliptic, 3 by 2 mm,
smooth or faintly appressed-hairy by partly detached cells of translucent membrane
(possibly of the mesocarp), margin narrow but distinct, forming a square edge. — Fig.
61.
Distribution — Philippines, Luzon (Rizal Province), where only known from the type.
Habitat & Ecology — Unknown; fruiting in November.
Note — Pilogyne rizalensis is close to the widespread P. mucronata. The sole collec-

tion on which the present species is based possibly represents a local ecotype. However,
it is at variance with all specimens reckoned to P. mucronata, especially in its long-
peduncled dense infructescence, small ellipsoid fruit and extremely small seeds, which
are among the smallest in the whole family Cucurbitaceae.
The type, Ramos BS 2023 has already been depicted in De Wilde & Duyfjes (2008),
and was discussed there in a note under P. trichocarpa.
11. Pilogyne samoensis (Cogn.) W.J.de Wilde & Duyfjes

Pilogyne samoensis (A. Gray) W.J.de Wilde & Duyfjes, Gard. Bull. Singapore 61 (2009) 210. — Ka
rivia samoensis A. Gray, U.S. Expl. Exped., Phan. (1854) 643. — Homotypic synonyms (see note
2): Zehneria grayana (Cogn.) Fosberg & Sachet, Smithsonian Contr. Bot. 47 (1981) 12; W.J.de
Wilde & Duyfjes, Blumea 51 (2006) 59. — Melothria grayana Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 591, p.p., excluding specimens from Tahiti. — Type: US exploring expedition s.n.
(lecto US, barcode US00147706, designated by Fosberg & Sachet, 1981), Samoa, without further
locality.
Stout 4 –10 m long vine, leafy stem 2(– 3) mm diam., plant (sub)glabrous, brown,
blackish, or obscurely green on drying; dioecious. Leaves: petiole 2 – 4 cm long,
(sub)glabrous; blade ovate or subcircular, 5 –15 by 3.5 –11 cm, unlobed or rarely shal-
lowly 3-lobed to c. 1/5 deep, upper surface scabrous or not, (sub)glabrous, cystoliths
present or absent, lower surface glabrous, base shallowly or deeply cordate, with broad
(rarely narrow, in New Ireland and New Britain) sinus, margin shallowly or deeply den-
tate, teeth up to 3 mm long, apex acute-acuminate, mucronate; veins glabrous or with
minute sparse hairs. Male inflorescences: a subsessile or short- or long-peduncled dense
short raceme, 2 – 5 mm long, with flowers clustered as in a subumbel, rarely branched;
peduncle 0.4 – 5 cm long. Male flowers: pedicel 4 – 8(–10) mm long, with a joint up to
1.5 mm below the receptacle-tube; expanded perianth 5 –7 mm diam.; receptacle-tube
(2 –)2.5 – 3.5 by 3 – 4 mm, outside glabrous, at throat and inside in the upper-half densely
hairy, hairs 0.2 – 0.5 mm long; sepals suberect (patent), 0.5 –1 mm long (sometimes
larger and petal-like, up to 2 by 1.5 mm, Solomon Islands), glabrous or sparsely hairy;
petals narrowly triangular or elliptic, 2 – 3 by 1.5 – 2 mm, subacute or blunt, densely
(papillose-)hairy outside at apex and wholly hairy inside (hairs 0.5 –1 mm long); sta-
mens inserted about halfway in the receptacle-tube, filaments 1.5 – 2 mm long, glabrous
at apex and base but densely hairy over most of their lengths, hairs 0.5 –1 mm long,
anthers subcircular or broadly or narrowly elliptic in outline, 1.5 – 2 mm long, thecae
curved and nearly touching at apex (Samoa) or ± straight, connective broad (Samoa) or
narrow, with stiff hairs lining the thecae; disc depressed half-globose, large, (0.5 –)1 by
(1–)1.5 – 2.5 mm, entire or faintly 3-lobed. Female flowers solitary or up to 5 in a loose
fascicle, sessile or up to 3 cm peduncled; pedicel 10 – 50 mm long when flowers soli-
tary, 5 – 20 mm long when flowers fascicled; ovary (only seen in Green RSNH 1281(L),
Vanuatu) narrowly ellipsoid, 8 –10 by 3 mm, glabrous, neck short and broad, c. 2 mm
long, perianth as in male flower but sepals longer, long-triangular, c. 1.5 mm long, ±
papillose-hairy; style c. 2.5 mm long, glabrous, stigma c. 2.5 mm diam., consisting of
3 papillose lobes c. 2 mm long, curved downwards, shortly connate at base; staminodes
inserted halfway the receptacle-tube, 1–1.5 mm long, densely hairy, hairs 1 mm long,
but apex glabrous; disc conspicuous, c. 1.5 mm high, irregularly shallowly 3-lobed.
Fruit solitary with long pedicel (New Ireland) or 2 – 4 fruits in a peduncled fascicle and
sometimes co-axillary with a solitary fruit, ripening red, (narrowly) ellipsoid, 2 – 4 by
1–1.5 cm, base and apex rounded or short-attenuate, glabrous; exocarp not or faintly
pitted; fruiting pedicel 0.5 – 5 cm long. Seeds numerous, whitish or pale brown, 5 – 6 by
3.5 – 4.5 mm, not ornamented, margin rather distinct, broad with rounded edge.
Distribution — Papua New Guinea east to Vanuatu, Fiji (no specimens seen), and
Samoa; in Malesia: New Guinea (Papua New Guinea (New Britain, New Ireland)), and
Solomon Islands.
Habitat & Ecology — Scrub and (shore) forest, rainforest edges, disturbed forest
on slopes and in gullies; up to 1000 m altitude; flowering: mainly August to October;
fruiting throughout the year.
Notes — 1. The sole specimen known from New Britain (Frodin NGF 26539) has
male inflorescences but no flowers. It somewhat deviates in the narrow cordate leaf
base. The sepals of Powell BSIP 19393 (Solomon Islands, San Cristόbal) are, possibly
as an exception, very large and petal-like.
2. The epithet samoensis could not be used at the time by Cogniaux (1881) in Mel
othria, because the combination was already preoccupied by M. samoensis A. Gray, a
different species, now Neoachmandra samoensis (A. Gray) W.J.de Wilde & Duyfjes.
12. Pilogyne trichocarpa (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne trichocarpa (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009)
411. — Zehneria trichocarpa W.J.de Wilde & Duyfjes, Reinwardtia 12, 4 (2008) 271, f. 2. — Type:
Ramos & Edaño BS 84954 (holo GH), Philippines, Mindanao.
Small climber, leafy stem 1–1.5 mm diam., all parts hairy, dark on drying, mono-
ecious(?). Leaves: petiole 1.5 – 2.5 cm long; blade (subcircular-) 5-angular in outline or
(3 –)5-lobed, 1/4 –1/2-way deep, 3 –7 by 4 – 8 cm, appressed-hairy above, lower surface
more densely so, cystoliths not apparent, margin coarsely dentate. Male inflorescences:
5 –10 flowers in a dense head (condensed raceme) on a 1–1.5 cm long peduncle, and
with a persistent pedicel of a solitary flower at base. Male flowers: pedicel 3 mm
long; expanded perianth 7– 8 mm diam.; receptacle-tube c. 2 by 3.5(– 4) mm, outside
(sparingly) hairy, inside subglabrous but sparse-hairy at the throat; sepals narrowly
triangular(-linear), c. 1 mm long; petals c. 3 by 2 mm, acute, glandular-hairy in upper
part; stamens inserted at base of the receptacle-tube, filaments c. 2.5 mm long, subgla-
brous, anthers somewhat longer than broad, c. 1.5 mm long, the thecae slightly curved,
nearly touching each other at apex, connective broad at base and in the middle, coarsely
hairy, not produced at apex; disc subglobose, c. 1 mm diameter. Female flowers 1 or
2 subsessile on the nodes; pedicel 1–1.5 mm long; ovary narrowly ellipsoid, c. 5 by 2
mm, densely hairy; perianth as in male flowers; style, stigmas, and disc not examined.
Fruit solitary (or 2), colour of ripe fruit not recorded, ellipsoid, c. 1.5 by 1 cm, base and
apex rounded; exocarp faintly tessellated (not pitted), sparsely hairy, hairs c. 0.5 mm
long; fruiting pedicel 0.2 – 0.4 cm long. Seeds numerous, pale, ovate-elliptic, c. 2.5 by
1.8 by 0.5 mm, smooth or possibly faintly hairy, faces flat, not ornamented, narrowly
margined. — Fig. 62.
2 mm 2 cm
c d
0.5 mm
e
3 mm
f
2 cm
3 mm
1 mm
j g
k
h 0.5 mm
2 cm
i 3 mm
l
Fig. 62. Pilogyne trichocarpa (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes. a, b. Node with male
inflorescence; c, d. male flower, from outside and opened respectively; e. detail of stamen, showing the
slightly curved thecae, mark coarsely hairy connective; f, g. node with female inflorescence; h, i. node
with one hairy fruit; j. seed, faintly hairy; k, l. detail of upper and lower leaf blade surface respectively
(all: Ramos & Edaño BS 84954, type).
Distribution — Philippines (Mindanao, Cotabato Province, Nutol) where only known

from the type, collected March –April 1932.
Notes — 1. Pilogyne trichocarpa belongs to a group of similar species, among which
P. mucronata (Blume) Miq. and P. repanda (Blume) C.M.Simmons, which are all relat-
ed to what is known in Africa as the widespread, variable Zehneria scabra (L.f.) Sond.
Pilogyne trichocarpa is unique in SE Asia by its densely hairy overall habit, including
a hairy ovary and fruit, and by its small seeds, all traits which occur regularly in Africa,
but which are absent or very rare in Asia.
2. The seeds of both Pilogyne trichocarpa and P. rizalensisi belong to the smallest
of all Cucurbitaceae of SE Asia.
13. Pilogyne trullifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne trullifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Reinwardtia 12, 5 (2009) 411.
— Zehneria trullifolia W.J.de Wilde & Duyfjes, Blumea 51 (2006) 77. — Type: De Vogel 5618
(holo L; iso BO, K), Central Sulawesi (Sopu Valley).
Climber to 6 m long, leafy stem 1–1.5 mm diam., plant subglabrous, (green-)brown

on drying; dioecious(?). Probract less than 1 mm long. Leaves: petiole 1– 3 cm long,
glabrous; blade trullate (or narrowly ovate), 5 – 8 by 3 – 4.5 cm, broadest at about 1/4
from the base, base broadly rounded or broadly cuneate, both surfaces glabrous but
somewhat scabrous with small cystoliths, margin finely sparsely dentate, occasionally
coarsely dentate. Male inflorescences 1–1.5 cm long peduncled 3 –10-flowered short
racemes, 0.1– 0.3 cm long. Male flowers: pedicel 1.5 – 2 mm long; perianth ± succu-
lent, when expanded c. 4 mm diam., early caducous; receptacle-tube c. 2.5 by 3 mm,
outside glabrous, inside in upper half and at throat finely woolly hairy, hairs less than
0.5 mm long; sepals 0.3 mm long, glabrous; petals c. 1.5 mm long, ± acute, both sur-
faces minutely papillose-hairy; stamens inserted near the base in the receptacle-tube,
filaments c. 1.5 mm long, glabrous, anthers subglobose, (1–)1.5 mm diam., thecae
curved, connective broad but narrow at apex; disc depressed, irregular in outline, large,
(0.5 –)1 by c. 2 mm. Female flowers and fruits not known.
Distribution — Central Sulawesi, near Sopu, SSE of Palu; known only from 2 collec-
tions.
Habitat & Ecology — In sloping primary mountain forest with sparse undergrowth
on clayey soil derived from granite; at 1000 –1500 m altitude; flowering in May.
14. Pilogyne viridifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes
Pilogyne viridifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes, Gard. Bull. Singapore 61
(2009) 210. — Zehneria viridifolia W.J.de Wilde & Duyfjes, Blumea 51 (2006) 77, f. 13i, j. — Type:
Brass 23914 (holo A; iso CANB, L), Papua New Guinea, Milne Bay Province.
Climber 1– 2 m long, leafy stem 1–1.5 mm diam., plant sparsely hairy, glabrescent,
green on drying; monoecious. Probract c. 1 mm long. Leaves: petiole 1.5 – 3 cm long,
rough-hairy; blade (ovate or) circular in outline, 5 – 9 cm diam., ± hastate or deeply 3(– 5)-
lobed to c. halfway, short-scabrous hairy with minute cystoliths above, subglabrous but
veins hairy beneath, base broadly shallowly cordate, margin (sparsely) dentate. Male
inflorescences: flowers in peduncled short racemes; peduncle 0.5 –1 cm long; racemes

0.1– 0.5 cm long, up to 15-flowered. Male flowers: pedicel 4(– 5) mm long; expanded
perianth c. 5 mm diam.; receptacle-tube 1.5 – 2 by 2.5 – 3 mm, outside (sub)glabrous, in
and below the throat densely woolly hairy, hairs to 0.5 mm long; sepals c. 1 mm long;
petals 2 – 2.5 mm long, both surfaces minutely finely gland-hairy; stamens inserted
at or slightly above halfway the receptacle-tube, filaments 0.5 –1 mm long, glabrous,
anthers ± exserted, broad ellipsoid, c. 1.3 by 1 mm, thecae ± curved, connective broad,
but narrow and truncate at apex; disc (depressed-)globose, c. 1.5 mm diameter. Female
flowers 1 or 2 at the nodes, sometimes co-axillary with male inflorescences; pedicel
5 –10 mm long; ovary ellipsoid-fusiform, 3.5 – 4 by c. 2 mm, glabrous, neck 1(– 2) mm
long; perianth as in male flower; style c. 2 mm long, with 3 style-arms 0.5 mm long,
each with an ovoid-ellipsoid 1 mm long stigma, papillose; staminodes inserted at c. 1/4
from the base of the receptacle-tube, 1.5(– 2) mm long, slender, ± hairy in the upper
half; disc large, 1 by 1.8 mm. Fruit 1 or 2; ellipsoid, 1 by 0.8 cm, recorded as unripe
(but see note); exocarp smooth, not finely pitted; fruiting pedicel 2 – 3 cm long. Seeds
few, 1(– 5?), ovate, c. 5 by 4 mm, not obviously margined, with a conspicuous broad
square edge nearly 2 mm wide. — Fig. 57i, j.
Distribution — New Guinea (Papua New Guinea, Milne Bay Province, Gwariu River;
Biniguni Camp (see Brass, Bull. Amer. Mus. Nat. Hist. 111 (1956) pl. 17)); known only
from the type.
Habitat & Ecology — Along second growth rainforest, near river; at c. 200 m alti-
tude; flowering and fruiting in August.
Note — This species is tentatively treated under Pilogyne, but it is not sure whether
the remarkable sole collection on which the present species is based belongs in that
genus. It deviates from most Pilogynes in its green drying colour. Within Pilogyne it
comes closest to P. erythrobacca (distributed all over New Guinea), the latter being
distinct by a brown drying colour, by the leaf blade which is hardly lobed and not or
hardly scabrous, and by the stamens which are inserted towards the base of the recep-
tacle-tube.
30. SCOPELLARIA
Scopellaria W.J.de Wilde & Duyfjes, Blumea 51 (2006) 297. — Scopella W.J.de Wilde & Duyfjes,
Blumea 51 (2006) 34, nom. illeg. — Type species: Scopellaria marginata (Blume) W.J.de Wilde
& Duyfjes.
Climbers to 6 m long, annual or (sub)perennial, leafy stem 1.5 – 2(– 3) mm diam.,

green on drying; monoecious. Probract absent. Tendrils unbranched, hairy (see note).
Leaves simple, unlobed or lobed, margin entire or dentate, palmiveined. Flowers: ex-
panded corolla 5 –10 mm diam., yellow; sepals minute, narrowly elliptic or linear,
subpatent; petals imbricate in bud, free, (ob)ovate-elliptic; receptacle-tube campanu-
late; disc present. Male inflorescence 1(–3) slender peduncled short raceme(s) with the
flowers (pedicels) crowded at apex, co-axillary with 1 (or 2) female flower(s) or not;
bracts absent. Male flowers: pedicel short, persistent; stamens 3, exserted, inserted in
the throat of the receptacle-tube, erect, filaments long, anthers all 2-thecous, thecae lat-
eral, straight, connective narrow, not produced; disc free, (depressed-)globose. Female
flowers 1 or 2 co-axillary with peduncle of male inflorescence or not; pedicel usually

long; ovary globose or ellipsoid, without slender neck; stigma consisting of 3 rather
long flat lobes, cleft or notched at apex, fringed on the margin; staminodes present;
disc annular, free from receptacle-tube. Fruit 1 (or 2), ripening red, with short or long
fruiting pedicel, globose, ellipsoid, or narrowly ellipsoid (fusiform), 1–1.5(– 3) cm long,
juicy or pulpy; exocarp filmy, membranous or cartilaginous, smooth. Seeds numerous,
moderately compressed, ovate-elliptic, faces ± convex, scrobiculate, margin distinct,
edge (sub)entire, not winged
Distribution — A genus with 2 species ranging from South China through Indo-
China to West Malesia; not known from Myanmar and Taiwan; in Malesia: east to
Sulawesi, and Philippines.
Note — In Scopellaria the tendrils are completely hairy (although sometimes sparing
ly so); in resembling genera the tendrils are glabrous or hairy only below the first spiral.
KEY TO THE SPECIES
1a. Leaf blade coriaceous, margin hard-spiny. Seeds 7(– 8) mm long. —Borneo

(Sabah) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. S. diversifolia
b. Leaf blade membranous or chartaceous, margin denticulate. Seeds 2.5 – 6 mm long.
— Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. S. marginata
1. Scopellaria diversifolia (Merr.) W.J.de Wilde & Duyfjes

Scopellaria diversifolia (Merr.) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 297. — Melothria diversi-
folia Merr., J. Malayan Branch Roy. Asiat. Soc. 1 (1923) 44. — Scopella diversifolia (Merr.) W.J.de
Wilde & Duyfjes, Blumea 51 (2006) 34, nom. illeg. — Type: Ramos 1896 (holo PNH†; lecto BO,
designated by De Wilde & Duyfjes (2006); iso UC), Sabah, near Sandakan.
Subherbaceous 2 – 6 m long climber, leafy stem 1.5 – 3 mm diam., glabrescent. Leaves:

petiole 1– 2(– 4) cm long, short scabrid-hairy, hairs 0.5 mm long; blade (sub)coriaceous,
narrowly ovate to narrowly elliptic, 7–14(– 20) by 2 – 8(–12) cm, upper surface coarsely
scabrous by conspicuous (white) cystoliths, lower surface subglabrous, base truncate,
shallowly cordate or sagittate with acute lobes to 1.5 cm long, margin entire or faintly
dentate with sparse, small but coarse 1 mm long hard teeth. Male inflorescences solitary;
peduncle 2 – 3.5 cm long, subglabrous; raceme short, rarely 2-branched, with 5(–10)
flowers; male pedicel 1– 2(– 5) mm long, female pedicel 10 – 20 mm long. Mature male
and female flowers not seen, presumably similar to those of S. marginata (see note):
anthers c. 1.5 mm long; ovary ovoid, c. 3 mm long; sepals lanceolate, c. 1.5 mm long;
petals ovate-elliptic, acuminate, c. 5 mm long; staminodes 1.2 mm long; stigma-lobes
deeply cleft, c. 2 mm long. Fruit solitary or co-axillary with (previously) developed
male inflorescence, globose, 1–1.5 cm diam.; exocarp cartilaginous, glabrous; fruiting
pedicel 2 – 4.5 cm long, subglabrous. Seeds numerous, pale yellowish brown, elliptic or
obovate in outline, 6 – 8 by 4 – 4.5 by 1–1.5 mm, base narrowed, faces almost smooth
or indistinctly scrobiculate, margin narrow but distinct.
Distribution — Borneo (Central and East Sabah).
Habitat & Ecology — In thickets and forest edges; from sea level to 250 m altitude;
flowering and fruiting throughout the year.
Note — According to Merrill (1923) who examined Ramos 1175, 1210, 1896 (mate-
rial now partly lost), the male flowers are similar to female flowers; the flower details
given are his.
2. Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes

Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 297; Fl. Thailand 9, 4
(2008) 493. — Bryonia marginata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 924; Ser. in DC., Pro-
dr. (1828) 305; M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 36; Miq., Fl. Ned. Ind. 1, 1 (1856)
660. — Melothria marginata (Blume) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 593;
Backer in Backer & Bakh.f., Fl. Java 1 (1964) 296. — Zehneria marginata (Blume) Keraudren
in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 55, pl. 9: 4; W.J.de Wilde &
Duyfjes, Thai Forest Bull, Bot. 32 (2004) 21, f. 2. — Scopella marginata (Blume) W.J.de Wilde &
Duyfjes, Blumea 51 (2006) 35, f. 9, nom. illeg. — Type: Blume 920 (holo L), Java.
Bryonia epigaea auct. non Rottl.: Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 925. — Aechmandra blumeana
M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 33, p.p; Miq., Fl. Ned. Ind. 1, 1 (1856) 657, p.p. (a
new name for Bryonia epigaea auct. non Rottl.: Blume). — Type: Blume s.n. (holo L, barcode
L0130057; iso L, 3 sheets), Java.
?Cerasiocarpum maingayi C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 629. — Type: Maingay 1268
(holo K), Peninsular Malaysia (male flower with reduced androecium; see note under var. margi
nata).
Melothria affinis King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 67 (1898) 38; Cogn. in Engl., Pflan-
zenr. 66, 4.275.I (1916) 94. — Type: Scortechini 495 (lecto K, designated by De Wilde & Duyfjes
(2006)), Peninsular Malaysia, Perak.
Melothria gracilipes Merr., Pap. Michigan Acad. Sci. 19 (1934) 199. — Type: Bartlett 7228 (holo NY;
iso MICH), Sumatra.
Annual or biennial, (sub)herbaceous, creeping or climbing, (0.5 –)1– 5 m long, vari-

ously scabrid-hairy, leafy stem 1– 2 mm thick; glabrescent. Leaves: petiole (compared
to resembling species short) 1– 5 cm long, with soft or coarse 0.5 –1(– 2) mm long hairs;
blade unlobed or shallowly or deeply (hastately) 3- or 5-lobed, variable in shape, (long)
triangular, ovate, or cordiform, 4 –16 by 3 –10 cm, upper surface rough-hairy and/or
scabrous by (coarse) cystoliths, lower surface coarsely hairy mostly only on the veins,
base truncate, rounded, or (deeply) cordate, margin entire with minute teeth or dentate,
apex acute-acuminate. Male inflorescences 1(– 3), glabrescent, usually co-axillary with
1 (or 2) previously developed female flower(s); peduncle 1.5 – 6 cm long, 0.5 mm thick;
raceme 0.3 –1(– 2) cm long, (3 –)5–10(– 25)-flowered; pedicels half-patent, persistent.
Male flowers: pedicel 2 –7 mm long; receptacle-tube 1.5 – 2 mm diam., sparsely coarsely
hairy outside, throat hairy; sepals patent or ± out-curved, 1 mm long, with sparse hairs;
petals (2 –)3 – 4 by 2.5 – 3 mm, rounded or (sub)obtuse-acuminate, (papillose) hairy;
filaments 1–1.5 mm long, glabrous, anthers ellipsoid, 1 mm long, thecae 1 mm long;
disc c. 1 mm diameter. Female flowers: pedicel (5 –)10 – 30 mm long; ovary ellipsoid,
3 – 5 mm long, with sparse hairs, glabrescent; receptacle-tube shallow; perianth as in
male flower but petals larger, 4 – 5 mm long; style c. 2 mm long, glabrous; stigma con-
sisting of 3 feather-like arms, shallowly forked at apex, each c. 2 mm long; staminodes
linear, 1 mm long, inserted at the base of the receptacle-tube; disc less than 1 mm high.
Fruit (remarkably variable in shape and texture, see note under var. marginata), either
globose or ellipsoid, 1–1.5 cm long with base and apex rounded, or fruits narrowly
1 cm
l
c
d
2 cm
1 mm
a e
f
1 mm
k
1 mm
2 mm
j
i h g
Fig. 63. Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes var. marginata. a. Habit; b, c, d.
leaves; e, f. male flowers; g, h, i:. female flowers, from outside, opened, and with expanded stigma
respectively; j. fruit (form with globose fruit); k. seed; l. fruit (form with fusiform fruit) (a: Awong
Kaya s.n., barcode L0130026 (Brunei); b: De Wilde & Duyfjes 12614 (Sumatra); c: De Wilde & Duyfjes
21794 (Sumatra); d: Iwatsuki c.s. 1732 (Sumatra); e – j: De Wilde & Duyfjes 21756 (Sulawesi); k: De
Wilde & Duyfjes 21699 (Java); l: De Wilde & Duyfjes 22182 (Thailand)).
ellipsoid, tapering at base and apex, 1.5 – 2(– 3) by c. 0.5 cm, glabrescent; exocarp
membranous or ± leathery, leaving the seeds visible or not on drying; ripe fruits juicy
or with yellowish pulp; fruiting pedicel short or long, (0.5 –)1– 5 cm long (short when
fruit narrowly ellipsoid). Seeds 5 – 35 per fruit, whitish, (somewhat) compressed or faces
± convex, narrowly ovate in outline, 3 – 5(– 6) by1.5 – 3.5 by c. 1.5 mm, faces (deeply)
finely or coarsely scrobiculate, margined, edge distinctly square.
Distribution — East Myanmar (Wallich Cat. 6713, only photo seen), China (Yunnan,
no material seen), Thailand, Laos, Cambodia, Vietnam; in Malesia: Sumatra, Peninsular
Malaysia, Borneo (mainly Sabah), West Java, Philippines, and Central Sulawesi.
Habitat & Ecology — Open and shaded places, damp sites, mostly in forest edges,
roadsides; sea level to 1500 m altitude; flowering and fruiting mostly June to Decem-
ber.
Note — Both Cogniaux (1881) and King (1898) described the fruits as velvety, but
this aspect obviously is caused by drying; in fact the fruits are early glabrescent.
1a. Leaf blade unlobed, narrowly ovate, base cordate or short-sagittate. Seeds 4 – 5(– 6)
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. penangense
b. Leaf blade ± unlobed, or shallowly or deeply 3(– 5)-lobed, ovate or triangular, base
rounded, cordate or hastate. Seeds 2.5 – 4 mm long . . . . . . . . . a. var. marginata
a. var. marginata
Climber 0.5 – 3 m long. Leaves: blade usually angular or lobed. Fruit globose, el-
lipsoid or long-fusiform. Seeds few or numerous, small, 2.5 – 4 mm long, the faces flat
or somewhat convex, coarsely scrobiculate. — Fig. 63; Plate 21a, b.
Note — Apart from a variable habit of the plant, mainly due to variation in leaf-shape
and indumentum, the variety marginata exhibits a remarkable variation in fruit-shape
and length of fruiting pedicel. Most specimens have globose fruits (as the type of Mel
othria affinis) with long pedicels, some have (narrowly) ellipsoid-fusiform fruits (as
the type of Bryonia marginata) with short pedicels (see Fig. 63a, j, l). Although most
herbarium specimens have a constant fruit form, there are several collections indicating
that both forms may occur in one plant. The type-specimen of the present species is the
less frequent form with the slender fusiform fruits. The form with elongate, fusiform
fruits with comparatively small male flowers, often with reduced petals and stamens,
may be related to small plants from poor soils that flower precociously.
b. var. penangense (C.B.Clarke) W.J.de Wilde & Duyfjes

Scopellaria marginata (Blume) W.J.de Wilde & Duyfjes var. penangense (C.B.Clarke) W.J.de Wilde &
Duyfjes, Blumea 51 (2006) 296. — ?Cerasiocarpum penangense C.B.Clarke in Hook.f., Fl. Brit. Ind.
2 (1879) 629. — Scopella marginata (Blume) W.J.de Wilde & Duyfjes var. penangense (C.B.Clarke)
W.J.de Wilde & Duyfjes, Blumea 51 (2006) 38, nom. illeg. — Bryonia heterophylla auct. non Blume:
Wallich Cat. 6704. — Type: Wallich Cat. 6704 (holo K-W), Peninsular Malaysia, Penang.
Climber 1– 5 m long. Leaves: blade entire, ovate or narrowly ovate, base sometimes
sagittate. Fruit short-ellipsoid or globose, 1–1.5 cm long. Seeds numerous, 4 – 5(– 6)
mm long, the faces flat, finely scrobiculate.
Distribution — In the western half of the species area where it occurs beside the
type-variety: South China, Myanmar (see note), Thailand, Laos, ViêtNam, Cambodia;
in Malesia: Sumatra and Peninsular Malaysia.
Note — The specimen Wallich Cat. 6713 (photo seen only) is mentioned under
Kedrostis (Rhynchocarpa) by various previous authors; it is the only collection from
Myanmar known to us which may represent the present species.
31. SECHIUM
Sechium P.Browne, Civ. Nat. Hist. Jamaica (1756) 355., nom. cons.; C.Jeffrey in Hanelt, Mansfeld’s
encycl. agric. hort. crops 3 (2001) 1555. — Type species: Sechium edule (Jacq.) Sw. (Sicyos edulis
Jacq.), typ. cons.
Medium or stout climbers, perennial, plants (almost) glabrous; monoecious. Probract

absent. Tendrils 2 – 5-branched. Leaves: blade simple, angular or shallowly lobed, base
cordate. Flowers rather small, short-pedicelled; male flowers in racemes, female flow-
ers solitary, or 2 or 3, often co-axillary with male inflorescence; petals pale greenish
yellow, free. Male flowers: receptacle-tube shallow; sepals linear; petals elliptic-oblong,
(sub)acute, margin entire; stamens 3, filaments united into a column, inserted at base
of receptacle-tube, ± free at apex, anthers two 2-thecous, one 1-thecous, thecae plicate;
disc (pistillode) absent. Female flowers short- or long-pedicelled; perianth as in male
flowers; ovary ovoid(-oblong), smooth or with few (coarse) hairs, unilocular, ovule one,
style slender, short, stigma capitate, 2-lobed, lobes entire or lobulate, recurved; stami-
nodes absent, but base of receptacle-tube sometimes with 10 shallow pouches. Fruit
indehiscent, fleshy (or fibrous, not in Asia), ± obovoid, smooth or sulcate, glabrous,
setose or with soft spines. Seed one, large, ovoid, sometimes ± compressed, testa woody,
glabrous, margin absent, edge entire, sharp.
Distribution — About 5 species in 2 sections in tropical America. One species,
Sechium edule, widely cultivated.
Note — Schaefer & Renner (Fam. Gen. Vasc. Pl. [Kubitzki], in press) found with
molecular analysis that the genus is nested in Sicyos.
1. Sechium edule (Jacq.) Sw.

Sechium edule (Jacq.) Sw., Fl. Ind. Occid. (1800) 1150; Cogn. in A.DC. & C.DC., Monogr. Phan. 3
(1881) 901; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 306; C.Jeffrey, Cucurbitaceae E. Asia
(1980) 59; J.M.M.Engels & C.Jeffrey, PROSEA 8 (1993) 246; W.J.de Wilde & Duyfjes, Sandakania
17 (2008 ‘2007’) 77, f. 10; Fl. Thailand 9, 4 (2008) 496. — Sicyos edulis Jacq., Enum. Syst. Pl.
(1760) 32. — Chayota edulis (Jacq.) Jacq., Select. Stirp. Amer. Hist., ed. 2 (1780) 245. — Type:
Jacquin (BM?, not seen, designated by Jeffrey (1980)), Cuba.
Stout perennial herb, 5 –10(–15) m long, leafy stem 2 – 5 mm diam., plant almost
glabrous with tuber-like rootstock; cultivated. Leaves: petiole 5 –15 cm long; blade
usually 5-angular or shallowly lobed, subcircular in outline, 10 – 20 cm diam., finely
scabrous especially on upper surface, margin finely sparsely dentate, lobes acute; dis-
tinctly reticulately veined. Male racemes 8 – 30 cm long, 10 – 30-flowered, with flowers
2(– 6) subfasciculate. Male flowers: pedicel 1– 6 mm long; receptacle-tube c. 5 mm
wide; sepals narrowly elliptic to linear, 5 –7 mm long; petals subglabrous, 10 –17 mm
long; staminal column 1– 2 mm long, spreading free parts c. 1 mm long, anthers c. 2 mm
diameter. Female flowers: ovary c. 5 mm long, glabrous or ± hairy, style 3 – 4 mm long,
stigma c. 3 mm diam., papillose. Fruit solitary, ripening green or whitish, 7–15(– 20)
cm long, often with irregular surface, low-warted or irregularly sulcate, smooth or (soft)
spiny; later on the embryo germinating within the seed and emerging at apex of fruit;
fruiting pedicel 2 – 3 cm long. Seed 50 –100 mm long.
Distribution — Originally wild and cultivated in Central America, now widely cul-
tivated (usually over trellis) in the tropics and subtropics all over the world, and com-
monly running wild.
Vernacular name — Chayote.
Uses — The shoots and fruits are used as a vegetable. The nectariferous male flowers
attract bees.
Note — Sechium edule was already known cultivated in pre-Columbian times and
is now known from numerous cultivars or land-races.
32. SIRAITIA
Siraitia Merr., Pap. Michigan Acad. Sci. 19 (1934) 200; W.J.de Wilde & Duyfjes, Blumea 51 (2006)
499. — Type species: Siraitia silomaradjae Merr.
Perennial herbaceous climbers with sub or supra-terraneous tuber (always?), leaves

and stems with blackish glandular hairs; dioecious. Probract absent or not obvious. Ten
drils distally 2-branched, spiralling both below and above point of branching, although
basal portion straight. Leaves: blade ovate, simple, unlobed, green on drying. Flowers:
petals yellow, free. Male flowers in racemes or panicles; bracts absent or present; re-
ceptacle-tube rather shallow, conspicuously coarsely anastomosing-veined externally;
without obvious disc, although receptacle-tube with thickened bottom; pedicels articu-
late towards apex; sepals ± triangular; petals (narrowly) elliptic or (narrowly) ovate, two
or all with an incurved basal adaxial scale; stamens 5, inserted near the throat of shallow
receptacle-tube, in two pairs and one solitary, filaments long, anthers 1-thecous, thecae
sigmoid, the connective frequently with conspicuous papillae. Female flowers solitary
(or elsewhere few in a short raceme); style-arms largely free, stigmas ± 2-lobed or
reniform; staminodes 5, glandular. Fruit ± fleshy, indehiscent, subglobose to cylindric,
rather large (5 –11 cm long), smooth; pulp sometimes sweet. Seeds numerous, broadly
ovate or subcircular in outline, compressed, faces smooth, with broad finely radiatingly
ribbed, corky, double or 3-ridged margin, edge (sub)entire.
Distribution — A genus of three or four species distributed in NE India, South &
Central China, Thailand, Indochina and West Malesia; in Malesia 1 species.
Note — Siraitia grosvenorii (Swingle) A.M.Lu & Zhi Y.Zhang does not occur in our
region, but its fruits are sold in Chinese shops and local markets in Malesia for its sweet
medicinally used fruits (Fig. 65c–e). The female flowers depicted are taken from Chinese
material, because female flowers of the genus from the Malesian area are not known.
1. Siraitia siamensis (Craib) S.Q.Zhong & D.Fang

Siraitia siamensis (Craib) S.Q.Zhong & D.Fang, Guihaia 4 (1984) 23; C.Jeffrey, Cucurbitaceae E.
Asia (1980) 26, comb. nov. provis.; W.J.de Wilde & Duyfjes, Blumea 51 (2006) 501; Fl. Thailand
9, 4 (2008) 501. — Thladiantha siamensis Craib, Bull. Misc. Inform. Kew (1914) 7; Fl. Siam.
(1931) 759; Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 51; Keraudren in Aubrév. & J.-F.Leroy,
Fl. Cambodge, Laos & Vietnam 15 (1975) 34, f. 10: 5–7. — Type: Kerr 1171 (holo K; iso E),
Thailand, Doi Suthep.
Climber with one (or few) shoots to 5(–10) m long from a supra-terranean club-
shaped or ovoid tuber to 25 cm diam.; leafy stem subterete or ± angular, c. 3 mm diam.,
with long or short hairs. Tendrils finely pubescent. Leaves: petiole 4 –10 cm long,
long-hairy; blade ovate, 9 – 30 by 8 – 22 cm, upper surface sparsely appressed-hairy,
lower surface variously hairy and with many blackish glandular hairs, cystoliths not
obvious, base deeply cordate, margin sparsely minutely dentate, apex acute-acuminate.
Male inflorescences: a simple or once branched raceme, (10 –)20 cm long, the flowers
± crowded at the apex of the inflorescence branches; bracts minute, 1(– 2) mm long or
mostly absent; peduncle 5 –13 cm long; all parts finely hairy; female flowers solitary (or
2 in a reduced raceme to 1 cm long). Male flowers finely hairy; pedicel 15 – 30 mm long;
receptacle-tube bowl-shaped, c. 10 by 5 mm; disc not obvious, but base of receptacle-
tube ± thickened; sepals triangular, c. 5 mm long, 5 mm wide at base, apex obtuse or
subacute, receptacle and sepals outside with conspicuous anastomosing-netted veins;
petals broadly elliptic, (10 –)15 mm long, 4 – 8 mm wide at base, apex rounded, inner
surface finely papillose, at base with 2 large and 2 or 3 smaller scales, these together
with the thickened bases of the filaments concealing a hollow in the basal portion of
the receptacle-tube; stamens erect or ± curved, filaments 5 –7 mm long, anthers c. 4 mm
long, thecae curved with one arm shorter. Female flowers: pedicel 10(– 20) mm long;
ovary ovoid-ellipsoid, c. 10 mm long, hairy; perianth as in male flowers; style (China)
4 – 5 mm long, 3-armed. Fruit solitary (or 2), ripening greenish yellow, subglobose,
5 – 8 cm diam., hairy, partly glabrescent; pericarp thin, c. 1 mm thick, woody; fruit pulp
whitish, enclosing numerous densely packed seeds. Seeds c. 15 mm diam., pale brown,
corky, with broad 3-layered margin, the middle layer largest, the proper seed (excluding
margin) elliptic in outline, c. 8 mm long.
Field-notes — Young leaves and twig apices purplish reddish.
Distribution — Widespread, from South China through Indochina into West Malesia;
in Malesia: Sumatra, Peninsular Malaysia, Borneo, and Java; rare and seldom flowering.
1a. Male inflorescences 2 (or 3) times branched, c. 20 cm long. Sepals of male flowers
acute-acuminate, rather open in bud. Petals narrowly triangular, c. 4 mm wide at
base. — North Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. silomaradjae
b. Male inflorescences simple or once branched, 10 – 20 cm long. Sepals of male
flowers obtuse or subacute, connivent and closing the bud before anthesis. Petals
elliptic, 4 – 8 mm wide at base with rounded or blunt apex. — Widespread . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. var. siamensis
1 mm c
5 mm 2 cm
f a
g
1 mm
e 5 mm
2 cm
b d
Fig. 64. Siraitia siamensis (Craib) S.Q.Zhong & D.Fang var. siamensis. a. Male flowering branch;
b. apex of male inflorescence; c. male flower bud; d, e. open male flower, seen from beneath and above
respectively; f. inner petals with petal-scale at base; g. stamens (all: Phonsena 4225 (Thailand)).
2 mm
2 cm
1 mm
a
d
3 mm
e b
Fig. 65. a, b. Siraitia siamensis (Craib) S.Q.Zhong & D.Fang var. siamensis. a. Portion of branch with
fruit; b. seed, face and side view. — c – e. Siraitia grosvenorii (Swingle) A.M.Lu & Zhi Y.Zhang. c. Im
mature female flower; d. ditto, opened; e. seed, face and side view (a: Larsen et al. 32487 (Thailand);
b: Van Beusekom & Phenghklai 918 (Thailand); c, d: G. Zhang 445 (China); e: De Wilde et al. 22306
(from fruit imported from China).
a. var. siamensis
Male inflorescences simple, or once-branched, 10 – 20 cm long. Sepals of male flow-
ers obtuse (rounded) or subacute at apex; petals broadly elliptic, apex rounded. — Fig.
64, 65a, b.
Field-notes — Mature fruits green. Lörzing 5595 gives the following observations
(translated): “Climbing herb 8 m, profusely branched; rare; male flowers: calyx bright
red-brown, with bright green veins, when in flower the red colour vanishes for the
greater part; corolla deeply 5-lobed, yellow, with pale yellow-green veins; anthers two
with each 2 free thecae, and one with 1 theca and with two scale-like staminodia; always
1 stamen or staminodia basally adnate to a petal; stamens bright yellow or green-yellow,
at the base yolk-yellow; 5 petals, all the same size, broadly obovoid, dorsally somewhat
rounded; upper surface of leaves green, somewhat glossy, lower surface dull pale green;
lower surface of young leaves chocolate with grey-green veins.”
Distribution — Possibly as the species, but see the note under var. silomaradjae.
Habitat & Ecology — Open or disturbed places in lowland forest, forest edges, scrub;
also on limestone; from sea level to 800 m altitude. It is extremely rarely collected
fertile. From Java one fruiting collection from 1912 is only known (Backer 5937, BO).
From Peninsular Malaysia and Borneo sterile collections are only known.
b. var. silomaradjae (Merr.) W.J.de Wilde & Duyfjes

Siraitia siamensis (Craib) S.Q.Zhong & D.Fang var. silomaradjae (Merr.) W.J.de Wilde & Duyfjes,
Blumea 51 (2006) 503. — Siraitia silomaradjae Merr., Pap. Michigan Acad. Sci. 19 (1934) 200;
A.M.Lu & J.Q.Li in Li, Acta Phytotax. Sin. 31 (1993) 54, f. 2. — Type: Bartlett 8702a (holo US;
iso L), male flowers in June 1927, Sumatra.
Male inflorescences long-peduncled, 15 – 25 cm long, (1 or) 2 (or 3) times branched

panicles. Sepals of male flowers long-acute-acuminate, the apices in bud free and
slightly out-curved; petals narrow, long-triangular, at base c. 4 mm wide, apex long-
acute (see Lu & Li, 1993: f. 2).
Distribution — Northern Sumatra, Asahan area. Only known from the type.
Notes — 1. The only other flowering collection of the species known from the Male-
sian area is Lörzing 5595 (BO, L) from Sibolangit, not far from Asahan. Although the
flowers of Lörzing 5595 are in a bad state of conservation, they are more similar to
those of var. siamensis than to those of var. silomaradjae.
2. We cannot decide on the distribution of var. silomaradjae beyond the type-locality
as all other collections from Malesia (Sumatra, Peninsular Malaysia, Java, Borneo) are
sterile, except the old collection from Java in fruit.
33. SOLENA
Solena Lour., Fl. Cochinch. 2 (1790) 514; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos
& Vietnam 15 (1975) 62. — Melothria sect. Solena (Lour.) Cogn. in A.DC. & C.DC., Monogr.
Phan. 3 (1881) 604, p.p.; in Engl., Pflanzenr. 66, 4.275.I (1916) 104, p.p. — Type species: Solena
heterophylla Lour.
Karivia Arn., Madras J. Lit. Sci. 12 (1840) 50; J. Bot. (Hooker) 3 (1841) 275; Miq., Fl. Ned. Ind. 1, 1
(1856) 660. — Type species: Karivia amplexicaulis (Lam.) Arn. (Bryonia amplexicaulis Lam.).
Herbaceous or at base subwoody climbers, with perennial rootstock or tubers, in

Malesia largely glabrous; dioecious (elsewhere monoecious). Probract minute, linear,
often caducous or absent. Tendrils unbranched. Leaves: petiole usually short, render-
ing the leaves frequently semi-amplexicaul; blade simple, variable in shape. Flowers:
petals valvate-induplicate, white or cream, free. Male flowers in sessile or peduncled
condensed racemes; pedicel suberect, persistent, often with bract at base or inserted
about halfway; receptacle-tube cup-shaped; sepals minute; petals small, hairy; stamens
3, two 2-thecous, one 1-thecous, filaments free, long, inserted at base of the recepta-
cle-tube, anthers with swollen connective (elsewhere with narrow connective), not
produced, thecae oblique and (slightly) curved, or somewhate sigmoid (elsewhere erect
and straight); disc 3- (or 4-)lobed, conspicuous, carnose. Female flowers single, solitary
or co-axillary with male raceme; perianth as in male. Ovary narrowly ellipsoid, glabrous
or hairy; ovules few or several; staminodes 3 (or 4), inserted on the receptacle; disc a
deeply lobed cup or consisting of 3(– 5) lobes, at base of style. Fruit carnose, glabrous,
hairy or scabrous. Seeds several, little compressed or nearly globose, smooth (elsewhere
sculptured).
Distribution — An Asian genus of 3 species; in Malesia: 1 species.
1. Solena heterophylla Lour.

Solena heterophylla Lour., Fl. Cochinch. 2 (1790) 514; ed. 2, 2 (1793) 629; Keraudren in Aubrév. &
J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 63, pl. 10, f. 1– 5; W.J.de Wilde & Duyfjes,
Blumea 49 (2004) 73; Fl. Thailand 9, 4 (2008) 504. — Bryonia heterophylla (Lour.) Raeusch.,
Nomencl. bot. (1797) 282; Steud., Nomencl. bot., ed. 1 (1821) 123; ed. 2 (1841) 232. — Melothria
heterophylla (Lour.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 618; Backer in Backer &
Bakh. (1964) 297. — Type: Loureiro s.n. (BM), Cochinchina and China.
Bryonia hastata Lour., Fl. Cochinch. 2 (1790) 594; Merr., Trans. Amer. Philos. Soc. 24 (1935) 377.
— Type: Loureiro s.n., China.
Bryonia rheedii Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 925; Ser. in DC., Prodr. (1828) 306. — Karivia
rheedii (Blume) M.Roem., Fam. Nat. Syn. Monogr. 2 (1846) 45; Miq., Fl. Ned. Ind. 1, 1 (1856)
661. — Type: Blume s.n. (holo L, barcode L0127474), Java.
Bryonia sagittata Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 925; Ser. in DC., Prodr. (1828) 305. — Type:
Blume s.n. (holo L, barcode L0127475), Java.
Zehneria connivens Miq., Fl. Ned. Ind. 1, 1 (1856) 656. — Type: Horsfield s.n. (holo U, barcode
U0226802; iso K), Java.
Zehneria hastata Miq., Fl. Ned. Ind. 1, 1 (1856) 656, nom. illeg. — Voucher specimens: Zollinger 669
(L), Horsfield s.n. (K), both Java.
Melothria ovata Cogn. in Engl., Pflanzenr. 66, 4.275.I (1916) 114. — Type: Anon. (? Thomson) (G,
not seen), Sikkim.
Zehneria umbellata auct. non (Willd.) Thwaites: C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 625,
p.p.; Ridl., Fl. Malay Penin. (1922) 851.
Note — De Wilde & Duyfjes (Blumea, 2004) distinguished in Solena heterophylla

two subspecies, subsp. napaulensis (Ser.) W.J.de Wilde & Duyfjes (northern India, Ne-
pal, China (Yunnan) and Myanmar) and subsp. heterophylla; only subsp. heterophylla
occurs in Malesia.
2 cm
2 cm
e
1 mm
2 mm
i
1 mm
3 mm
d g
f
Fig. 66. Solena heterophylla Lour. subsp. heterophylla. a, b. Habit of male flowering specimens;
c. ditto, a specimen with atypical more or less compound inflorescences resulting from axillary short-
shoots; d. ditto, note gland-bearing bracts and bracteoles; e, f. male flower; g. stamens; h. fruit; i. seed
(a: Koorders 41273; b: Zollinger 669; c – g: Garrett 1192; h – i: Whitmore FRI 20266).
subsp. heterophylla
Climber 2 – 6 m long, perennial, leafy stem 2 – 3 mm diam., plant sparingly pubescent
in younger parts, glabrescent; dioecious. Probract 1– 5(–10) mm long, caducous. Leaves:
petiole 0.5 –1(–1.5) cm long; blade subcoriaceous, either unlobed, or 3- or 5-angular,
or (deeply) 3- or 5-lobed, the lobes broad or (very) narrow, the lower ones frequently
hastate, ovate or hastate, 3 –10(– 22) cm long, to 15 cm wide, glands several or many,
small, scattered, but most glands near the insertion of the petiole, margin sparsely 1– 2
mm mucronate-dentate; palmately 3- or 5-veined, reticulation distinct on both surfaces.
Male inflorescences 1 (or 2) sessile or to 4 cm peduncled, persistent subumbellate dense
raceme(s), sparsely hairy, pedicels persistent; bracts persistent or caducous, narrowly
elliptic or linear, 2 – 5 mm long, inserted on the pedicel (sometimes towards the apex),
± carnose, with or without glands, often a single (abortive) flower co-axillary. Male
flowers: pedicel 3 – 6(–10) mm long, articulate at apex; receptacle-tube 3 – 4(– 5) by 2 – 3
mm; sepals triangular, 0.2 – 0.5 mm long, wide apart; petals triangular, 1– 2 mm long,
finely papillose-hairy (without gland-hairs); filaments 2 – 3 mm long, glabrous, inserted
at or slightly above the base of the receptacle-tube; anthers subglobose, 1– 2 mm long,
thecae in an oblique or (sub)horizontal position, forming a nearly closed ring below the
broad convex, papillose-hairy connective; disc lobes papillose-hairy, 1– 2 mm long. Fe
male flowers solitary (rarely a few in a short-shoot); pedicel 3 – 5(–10) mm long; ovary
c. 10 mm long, (densely) short hairy, sometimes with few irregular blotches; perianth
resembling male but slightly larger, (to 2.5 mm long) ± persistent in young fruit; style
c. 2 mm long; stigma consisting of 3 subglobose long-papillose lobes; staminodes 3 (or
4), elongate, 0.5 –1 mm long, glabrous, inserted towards the base of receptacle-tube;
disc deeply lobed or consisting of 3 (or 4) free lobes, 1– 2 mm long. Fruit green, paler
striped, ripening yellow or red, (narrowly-)ellipsoid, 2 – 5 by 1.5 – 2.5 cm, velutinous
or scabrous; pulp yellowish; fruiting pedicel 0.5 –1.5 cm long. Seeds (5 –)10 – 20, grey,
nearly globose or somewhat elongate, little compressed, (5 –)6 –7 mm long, c. 5 mm
thick, smooth, margin faint, edge entire. — Fig. 66; Plate 21c, d.
Distribution — Widely distributed, from NE Afghanistan, the Himalayas, through
northern India to South & East China, and SE to Myanmar, Thailand, and Indochina;
in Malesia: Peninsular Malaysia (Perak, Terengganu, and Pahang), and Java; not in
Sumatra, Borneo, and East Malesia.
Habitat & Ecology — Prefers a seasonal climate, in open forest, scrub, and along
streams; on sandstone and granite derived soils; lower and medium altitudes.
Notes — 1. The forms according to leaf shape, as proposed by Cogniaux (1916), and
repeated by Keraudren (1975) have no taxonomic value.
2. In living specimens the lower surface of the leaves is conspicuously pale, glau-
cous. Unripe fruits are green with irregular whitish stripes, ripe fruits red.
34. THLADIANTHA
Thladiantha Bunge, Enum. Pl. Chin. Bor. (1833) 29; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
421; in Engl., Pflanzenr. 66, 4.275.I (1916) 40; Craib, Fl. Siam. (1931) 759; Backer in Backer &
Bakh.f., Fl. Java 1 (1964) 296; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam
15 (1975) 29. — Type species: Thladiantha dubia Bunge.
Perennial herbaceous climbers, usually with tuberous rootstock, stem and leaf blades
without black glandular hairs; dioecious. Probract absent in sterile shoots, in inflores-
cences present or absent. Tendrils unbranched or 2-branched and spiralling only above
the point of branching. Leaves: blade simple, unlobed (elsewhere palmately lobed or
3 – 9-foliolate), ovate, green or brown on drying. Flowers: petals yellow, free. Male
inflorescences: bracteate racemes or panicles, rarely flowers solitary. Male flowers:
receptacle-tube shallow, with eccentric half-globose basal disc; sepals linear to (nar-
rowly) ovate; petals 1– 3 with an adaxial basal scale; stamens (3 –)5, inserted near base
or throat of receptacle-tube, often in two pairs and one solitary, filaments long, anthers
1-thecous, thecae short, not curved, erect. Female flowers solitary (or few in short
racemes or panicles), perianth as in male; style single with 3 style arms, stigmas sub-
globose; staminodes 5. Fruits solitary, (narrowly) ellipsoid, carnose, indehiscent, ribbed
or verrucose, or smooth, hairy. Seeds numerous, ovoid or ellipsoid, finely sculptured,
with or without narrow margin, edge entire.
Distribution — A genus of about 25 species in SE Asia; in Malesia: 2 species.
KEY TO THE SPECIES
1a. Male raceme bracteate, bracts 10 –15 mm long, incised. Fruit fenestrately pitted.
— Java, Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. T. cordifolia
b. Male raceme without bracts, bracts absent or minute, caducous. Fruit smooth.
— Mindanao . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. T. nudiflora
1. Thladiantha cordifolia (Blume) Cogn.

Thladiantha cordifolia (Blume) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 424; Backer in
Backer & Bakh.f., Fl. Java 1 (1964) 296; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos
& Vietnam 15 (1975) 30, p.p.; W.J.de Wilde & Duyfjes, Blumea 51 (2006) 508, f. 6a – d, 7, pl. 2a;
Fl. Thailand 9, 4 (2008) 508. — Luffa cordifolia Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 929; Ser. in
DC., Prodr. (1828) 302; Miq., Fl. Ned. Ind. 1, 1 (1856) 666. — Type: Blume 1464, fruit (lecto L,
barcode L0001624, designated by De Wilde & Duyfjes (2006); iso L), Java.
Thladiantha cordifolia (Blume) Cogn. forma glabrescens Hochr., Candollea (1934) 287. — Type:
Hochreutiner 1255 (iso L), Java.
Thladiantha calcarata Cogn. in Hook.f., Fl. Brit. India 2, 3 (1880) Errata, without page; in A.DC. &
C.DC., Monogr. Phan. 3 (1881) 423; Craib, Fl. Siam. (1931) 759; C.Jeffrey, Kew Bull. 34 (1980)
790. — Type: Wallich Cat. 6740A collection Gomez (‘Momordica calcarata’) (lecto K-W, desig-
nated by Jeffrey, Kew Bull. 34 (1980)), Sylhet.
Trichosanthes javanica Miq., Fl. Ned. Ind. 1, 1 (1856) 678. — Type: Junghuhn 875 (holo U; iso L),
Java.
Gymnopetalum piperifolium Miq., Fl. Ned. Ind. 1, 1 (1856) 680. — Type: Horsfield s.n. (K, not seen),
Java.
Gymnopetalum horsfieldii Miq., Fl. Ned. Ind. 1, 1 (1856) 680. — Type: Horsfield s.n. (K, not seen),
Java.
Perennial climber to 6 m long, leafy stem 2(– 3) mm diam., subglabrous or hairy.

Probract usually present. Tendrils spiralling throughout their length or only in the up-
per part. Leaves: petiole 3 – 6 cm long, finely hairy or subglabrous; blade simple, ovate,
6 –10 by 4 – 9 cm, upper surface appressed-hairy, scabrous by cystoliths, lower surface
1.5 cm
a
3 mm
b c
Fig. 67. Thladiantha cordifolia (Blume) Cogn. a. Node with male inflorescence; b. male flower;
c. idem, opened, partly schematic, note eccentrically situated disc-gland; d. node with 1-flowered
female inflorescence (a – c: Wieringa & Janzen 3417; d: Docters van Leeuwen s.n., 28-09-1910).
hairy or subglabrous, base deeply cordate, margin coarsely or finely dentate. Male in
florescence a 5 –10 cm long peduncled bracteate raceme (sometimes with a previously
developed solitary flower at base), sparsely minutely pubescent; peduncle 2 – 5 cm long,
at or close to the base with a rhombiform or subelliptic (5 –)10 mm long irregularly
incised probract; flower bracts conspicuous, close together, in older inflorescences the
rachis thickened by the scars of fallen flowers and bracts; bracts obovate or obtriangular,
2 mm
3 mm
1 cm
d
a
1 mm
c
Fig. 68. Thladiantha cordifolia (Blume) Cogn. a. Node with 1-flowered female inflorescence; b. female
flower, perianth partly removed, disc absent; c. fruit, note fenestrately pitted outer surface; d. seeds
(a: Docters van Leeuwen s.n., 28-09-1910; b: Wongprasert s.n. (SN 120885); c, d: Maxwell 99-96).
10 –15 mm long, in the apical half (irregularly) dentate or incised to c. 1/4 deep. Male
flowers: pedicel c. 10 mm long; receptacle-tube obliquely cup-shaped, tapering, 3 – 4
mm diam., throat short-hairy; sepals long-triangular or narrowly elliptic, 8 –12 by 3 – 5
mm, acute, 3(– 5-)veined, sometimes reflexed; petals obovate-elliptic, 15 – 20 by c. 15
mm, apex rounded and faintly dentate; filaments 3 – 5 mm long, ± curved, ± dilated
towards the base, anthers 3 – 5 mm long, slightly curved, the median(?) petal with con-
spicuous curved scale, c. 4 mm long, concealing the disc at base of the receptacle-tube;
disc inserted slightly laterally, large, ellipsoid, c. 4 mm long. Female flowers: solitary;
pedicel 10 – 40 mm long, with probract at or near base; perianth as in male flowers;
ovary ovoid-ellipsoid, densely woolly grey-hairy, c. 10 mm long; style-column c. 2 mm
long, arms 4 – 5 mm long, stigmas broadly reniform, each c. 3 mm diam.; staminodes
c. 1.5 mm long, erect, disc not obvious. Fruit solitary; narrowly ellipsoid, (2 –)3 – 6
by 2 – 2.5 cm, broadly rounded at both ends, sparsely or densely bristly hairy, outer
surface fenestrately pitted in c. 15 rows; fruiting pedicel 1.5 – 4 cm long. Seeds ovoid,
5 – 6 mm long, somewhat flattened, rugose, edge shallowly 2-grooved. — Fig. 67, 68;
Plate 24.
Distribution — [Eastern?] East India, Myanmar, Thailand to South China; in Malesia:
Sumatra and Java; not known from Peninsular Malaysia.
Habitat & Ecology — Primary forest and forest edges, open places in disturbed
forest, scrub, along rivers and near waterfalls; to 1500 m altitude.
2. Thladiantha nudiflora Hemsl.

Thladiantha nudiflora Hemsl. in F.B.Forbes & Hemsl., J. Linn. Soc. Bot. 23 (1887) 316, pl. 8; C.Jeffrey,
Cucurbitaceae E. Asia (1980) 28; W.J.de Wilde & Duyfjes, Reinwardtia 12 (2009) 412. — Type
(syntype, Jeffrey 1980): Henry 3253, 2937, 2208, 2937A, 2005 (all K, lectotype not designated),
China (Hupeh and Nanto).
Thladiantha formosana Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25, Art. 19 (1908) 100, pl. 11. — Type:
Nakahara s.n., October 1906 (holo TI?, not seen), Taiwan (Tozan).
Perennial climber to 5(?) m long, leafy stem 2 – 3 mm diam., plant pubescent-hispid.

Probract absent. Tendrils 2-branched or unbranched, spiralling in the upper 2/3. Leaves:
petiole 3 – 5(–12) cm long, hispid or soft hairy; blade simple, broadly ovate, 6 –13(–15)
by 5.5 –10(–13) cm, upper surface scabrous hairy and with minute cystoliths, lower
surface densely yellowish or grey hispid, especially on the veinlets, margin serrate,
sometimes faintly lobed. Male inflorescences a simple raceme or few-branched, with
small leaves at base of ramifications, 4 –10 cm long, pubescent; peduncle 2 – 3 cm long,
raceme 1.5 – 5 cm long, (5 –)10-flowered, pedicels persistent, bracts minute, caducous.
Flowers densely yellowish grey long-pubescent. Male flowers: pedicel 3 – 5 mm long
(to 10 mm in solitary flower at base of raceme); receptacle tube shallow, 4(– 5) mm
wide; sepals oblong, acute, 3 – 5(– 6) mm long, 3-nerved; petals ovate-oblong, acute,
8 –12(–15) by 6 –7 mm, basal petal scale distinct; stamens free, inserted at base of re-
ceptacle, filaments c. 4 mm long, anthers c. 2.5 mm long; basal disc globose, not seen.
Female flowers solitary; pedicel 10 – 20 mm long, villose; probract not seen; perianth
as in male flowers; ovary oblong, 12 –15 by 4 – 5 mm, villose-hispid; style 3-armed,
stigmas large, reniform (2-lobed); staminodes small. Fruit (brown-)red, oblong, 3 – 5
by 3 – 3.5(– 4) cm, apex obtuse, smooth, pubescent, glabrescent. Seeds c. 5 by 3.5 – 4
mm, reticulate.
Distribution — South China, Taiwan; in Malesia: Philippines (Mindanao), where
known from one collection, Schwabe s.n. (B), of which the precise location and habitat
are unknown.
Habitat & Ecology — (In Taiwan) montane.
Note — In the original description (Hemsley in Forbes & Hemsley, 1887) the inflo-
rescences are described as racemose, but figured (plate. 8) as paniculate and glabrous.
Hayata (1908, plate 11), for the synonym T. formosana, depicts a good habit and male
flower details.
35. TRICHOSANTHES
Trichosanthes L., Sp. Pl. 1 (1753) 1008; Lour., Fl. Cochinch. (1790) 588; Blume, Bijdr. 15 (1826)
932; Miq., Fl. Ned. Ind. 1, 1 (1856) 674; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 606; Cogn.
302; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos & Vietnam 15 (1975) 75; Rugayah,
Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999); W.J.de Wilde & Duyfjes, Fl. Thailand 9,
4 (2008) 514. — Lectotype (Green, Prop. Brit. Bot. (1929) 190): Trichosanthes anguina L.
Cucumeroides Gaertn. Fruct. Sem. 2 (1791) 485, t. 4.
Involucraria Ser., Mém. Soc. Phys. Genèv. 3, 1 (1825) 25, t. 5.
Climber, wild or cultivated, perennial (annual), 3 – 30 m long, leafy stem 2 – 5 mm

diam.; dioecious, less frequently monoecious Probract present or absent. Tendrils un-
branched or 2 – 5(– 9)-branched below halfway. Leaves: membranous or coriaceous,
glabrous, glabrescent, scabrous, or hairy; blade simple or foliolate, unlobed or lobed,
margin entire or dentate. Petals white, sometimes reddish veined, free, long-fringed,
the fringes sometimes reddish or yellow. Male inflorescences: bracteate racemes. Male
flowers: receptacle-tube elongate; sepals much shorter than petals; stamens 3, inserted
in the receptacle-tube mostly towards the throat, included; filaments short, free, mostly
shorter than anthers; anthers two 2-thecous, one 1-thecous, mostly united into an elon-
gate truncate synandrium, the thecae narrow, S-shaped; disc present as 3 mostly linear
bodies at base of receptacle-tube. Female inflorescences: flowers solitary (sometimes
co-axillary with male raceme). Female flowers: style one, slender, stigmas 1 (deeply)
lobed, or 3, entire or 2-lobed. Fruits solitary (or few), ripening (orange-)red or green,
often lighter striped, berry-like, exocarp leathery or woody, indehiscent, ellipsoid, glo-
bose or fusiform, 5 – 20 cm long (elongate, 100 cm long, in cultivated T. cucumerina var.
anguina), smooth, hairy or glabrous; pulp (mesocarp) green-black, or white or (orange-
)red, bitter or sweet of taste, sometimes fibrous. Seeds numerous, light or dark brown,
grey (or black), compressed or tumid (with inflated sides), variable of shape, faces not or
little sculptured, margin usually present, edge usually smooth, or crenate or undulate.
Distribution — About 100 species, throughout subtropical and tropical eastern Asia:
from India, South China and Japan, east to tropical Australia and Fiji; in Malesia 43
species in 6 not well-defined sections.
Notes — 1. Flower characters: detailed characteristics of the flowers are sometimes
left out because frequently only incomplete material is available. Moreover, the flow-
ers are fragile, and not easy to analyse from boiled dry material. In Trichosanthes the
general shape of the petals is either narrowly elliptic with long fringes all along the
margins, in bud folded spherical, e.g. in T. pilosa; in other species general shape of the
petals is obtriangular with the fringes shorter to the top, and the petals in bud folded in
a more elongate body, e.g. in T. quinquangulata. We feel that this character state could
be used for the distinction of sections.
2. Sepals: these are free, narrowly triangular, ovate, narrowly ovate, or narrowly
elliptic, often with a long-acute apex; the margin is entire, or dentate, or (coarsely)
laciniate, or lobed. These characters are of important taxonomic value, but one should
be aware that in some species the sepals of female flowers are entire, whereas in male
flowers they are dentate or laciniate.
3. Receptacle-tube and pseudo-ovary: the receptacle is of a tubular shape, widened

towards the apex, usually with white (long) hairs inside. In some species, e.g. T. villosa,
and T. intermedia, the receptacle-tube of the male flowers forms a pseudo-ovary, the
swollen base of the tube, which contains disk-like structures.
4. Peduncle: the male racemes are pedunculate, and the peduncle may be slender or
(very) stout. In older male inflorescences the rachis may be thickened, bearing persist-
ent old bracts and pedicels or pedicel scars. In some species the rachis is zigzag, e.g. in
T. refracta.
5. Straw-like appendage: in the description of several Trichosanthes species from
New Guinea a ‘straw-like appendage’ at the node is mentioned. This appendage origi-
nates either from the peduncle of the male raceme or from the pedicel of a solitary
co-axillary male flower.
NOTE TO THE KEYS
Eleven keys are presented. The general keys 1 and 2 are for male flowering and female
flowering/fruiting specimens respectively. The remaining keys, 3 –11, are regional keys
applicable for all collections.
1 – GENERAL KEY TO THE SPECIES

(for male flowering specimens, also based on vegetative characters and distribution)
1a. Leaf blade foliolate (in T. celebica and T. wawrae leaves sometimes partly or all
simple). The 5 species normally with foliolate leaves are insufficiently known in
the male flowering state, and hence cannot be keyed out.
– Plants stout. Probract concave. — a. Peninsular Malaysia, Singapore, Sumatra,
Borneo, Palawan: 13. T. elmeri; b. Moluccas, New Guinea: 27. T. papuana
– Plants more delicate. Probract ± flat. — a. Peninsular Malaysia, Sumatra, Borneo,
Java: 43. T. wawrae (2 forms); b. Sulawesi, Moluccas: 5. T. celebica; c. Lesser
Sunda Islands: 15. T. floresana
b. Leaf blade simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Plant monoecious. Bracts 1(– 2) mm long, caducous. Flowering diurnal. Corolla
2 0(– 30) mm diameter. — Widespread . . . . . . . . . . 7. T. cucumerina (2 varieties)
b. Plant usually dioecious. Bracts 1 mm long or (much) longer, (late) caducous or
persistent. Flowering nocturnal. Corolla 30 mm diam. or more . . . . . . . . . . . . . . 3
3a. Younger twig and leaves densely soft-hairy, hairs 2 mm long or more . . . . . . . . . 4
b. Younger twig and leaves scabrous, (sparsely) short-hairy or glabrous (glabres-
cent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4a. Bracts large, 20 – 50(– 60) mm long. — Borneo, Java, Philippines, Lesser Sunda
Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42. T. villosa (2 subspecies)
b. Bracts small, c. 1 mm long. — Sabah . . . . . . . . . . . . . . . . . . . . 25. T. mucronata
5a. Blade base at transition to the petiole with 2 bulgings or auricles with glands be-
neath (sect. Asterosperma) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
b. Blade base at transition to the petiole without gland-bearing bulgings or auricles . 8
6a. Flowers solitary, axillary to leaves, conspicuously brown-hairy. — Sabah . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. T. fusca
b. Flowers in peduncled racemes, minutely grey- or brown-hairy . . . . . . . . . . . . . 7

7a. Inflorescences inserted on older stems, far below the leaves. Bracts narrowly el-
liptic, entire. — Sabah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32. T. postarii
b. Inflorescences axillary to leaves. Bracts elliptic, dentate. — Borneo (Sarawak,
Sabah, West, Central, and East Kalimantan) . . . . . . . . . . . . . . . . 2. T. auriculata
8a. Leafy stem sharply ridged or winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
b. Leafy stem striate or grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9a. Inflorescences axillary to leaves. Bracts large, broad. — Sabah . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20. T. kinabaluensis
b. Inflorescences pendent, mostly below the leaves. Bracts small, narrow. — Sabah
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. T. pendula
10a. Plant not from New Guinea. [Specimens with simple unlobed leaves may belong
to a species usually with foliolate leaves, 5. T. celebica, or 43. T. wawrae; see
lead 1] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
b. Plant from New Guinea (including widespread 30. T. pilosa, 35. T. quinquangu-
lata; see the regional key for New Guinea) . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
11a. Bracts large, (10 –)15 mm long or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
b. Bracts small, 1–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Leaf blade unlobed, in outline circular, glabrous. — Sumatra . 37. T. rotundifolia
b. Leaf blade unlobed or lobed, in outline broadly ovate, glabrous or hairy . . . . . 13
13a. Leaf blade finely hairy beneath, at least along the (finer) veins. [Bracts (2 –)5–15
mm long.] — Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Leaf blade scabrous-hairy or (sub)glabrous beneath . . . . . . . . . . . . . . . . . . . . 14
14a. Leaf blade unlobed, large, (8 –)12 – 23 cm long, when older coriaceous, with re-
curved margin. [Bracts 5 –7 mm long.] — Sumatra, West Java . . . 6. T. coriacea
b. Leaf blade unlobed and often smaller, or (deeply) lobed . . . . . . . . . . . . . . . . . 15
15a. Veins raised beneath. Bracts minute, 1– 3 mm long, caducous . . . . . . . . . . . . 16
b. Veins not or hardly raised beneath. Bracts larger, persistent . . . . . . . . . . . . . . . 17
16a. Leaf blades almost adherent to supporting tree trunk, scabrous. — Sabah . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. T. adhaerens
b. Leaf blades free (plant climbing freely), faintly scabrous or glabrous . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. beccariana (2 subspecies)
17a. Leaf blade unlobed or deeply 3(– 5)-lobed. Pedicel remnants on rachis short,
shorter than the bracts, bracts 6 –15 mm long. — Sabah . . . . . 19. T. intermedia
b. Leaf blade deeply 5-lobed. Pedicel remnants on rachis about as long as bracts,
bracts 7–12 mm long. — Northern Kalimantan, Sarawak, and Sabah . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. T. longispicata
18a. Raceme including bracts finely yellowish soft-hairy; bracts entire, 35 –70 mm
long. Blade base cuneate-decurrent. Tendrils simple . . . . . . . . . . 17. T. globosa
b. Raceme including bracts brown-rusty hairy, glabrous or glabrescent. Tendrils
branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19a. Bracts (ob)ovate, margin (sub)entire. — Widespread . . . 35. T. quinquangulata
b. Bracts incised . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20a. Leaf blade unlobed, margin coarsely dentate, blade glands few, large, 2–3 mm
diameter. — Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31. T. planiglans
b. Leaf blade unlobed or lobed, margin finely remotely denticulate, blade glands
smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Rachis zigzag. — Borneo, Philippines; peat swamp forest . . . . . . 36. T. refracta
b. Rachis straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22a. Bracts broad, deeply fan-shaped incised. Plant dull blackish on drying. — Bor-
neo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. T. obscura
b. Bracts variously incised, but not deeply fan-shaped incised. Plant green, (reddish )
brown or blackish on drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23a. Younger parts reddish on drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
b. Younger parts green, brown or blackish on drying . . . . . . . . . . . . . . . . . . . . . . 25
24a. Probract narrowly elliptic or linear, 10 – 20 by c. 2 mm. Leaf blade finely hairy
beneath. — Western Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. T. pubera
b. Probract obovate or narrowly elliptic, 10 –15 by 5 – 8 mm. Leaf blade ± scabrous
beneath. — North Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. T. leuserensis
25a. Bracts (deeply) finely incised. Leaf blade 3-lobed to about halfway deep, greenish
on drying. — Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . 29. T. philippinensis
b. Bracts coarsely incised, i.e. at apex dentate. Leaf blade unlobed or variously lobed,
greenish (brown) or brown on drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26a. Rachis stout, 3 –7 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
b. Rachis more slender, 1.5 – 3 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
27a. Leaf blade unlobed or shallowly (2- or)3-lobed, to 1/3 deep 39. T. sepilokensis
b. Leaf blade deeply 3 –7-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28a. Blade glands scattered or at base only, 1– 2 mm diameter. Peduncle 9 –18 cm long.
Bracts 40–5 0 mm long. — East Malesia . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Blade glands absent or few near blade base, 1– 3 mm diameter. Peduncle 2 – 6 cm
long. Bracts 40 – 60 mm long. — West Malesia . 24. T. montana (2 subspecies)
29a. Leaf blade unlobed. — Sarawak, Philippines . . . . . . . . . . . . . . 12. T. ellipsoidea
b. Leaf blade lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Leaf blade 3-lobed, to 1/3–2/3 deep (leaves unlobed in forma siberutensis). — Wide-
spread in West Malesia [Sumatra, Peninsular Malaysia, Singapore, Borneo, Java,
Sulawesi, Lesser Sunda Islands, and Moluccas] . . 40. T. tricuspidata (4 forms)
b. Leaf blade more deeply 5-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31a. Probract c. 4 by 2 mm, with few glands. — North Sumatra, Peninsular Malaysia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. T. emarginata
b. Probract 5 –18 by 1– 2 mm, glands absent. — Sumatra, Peninsular Malaysia, Singa
pore, Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
32a. Leaf blade finely hairy beneath, especially along the smaller veins. Petals (exclud-
ing fringes) (narrowly) elliptic or (narrowly) ovate. — Widespread .30. T. pilosa
b. Leaf blade variously hairy or glabrous beneath. Petals obtriangular (cuneate) . 33
33a. Bracts entire, 10 – 30 mm long. — Widespread . . . . . . . . 35. T. quinquangulata
b. Bracts dentate (incised), sometimes at very apex only, or bracts much smaller.
Eight species endemic to New Guinea (male flowers not known in T. dentifera;
see regional key for New Guinea, lead 11 onwards: T. dentifera, T. dieniensis,
T. densiflora, T. edulis (3 varieties), T. hastata, T. laeoica (4 forms), T. pulleana,
T. schlechteri).
2 – GENERAL KEY TO THE SPECIES

(applicable for fruiting specimens with emphasis on seed characters, also based on vegetative char-
acters and distribution)
Note — Excluded from this key are 5 species of which fruit and seeds are not known:
T. adhaerens (Sabah), T. dieniensis (Papua New Guinea), T. fusca (Sabah), T. longispicata
(Sarawak), and T. pulleana (Moluccas, West Guinea).
1a. Leaf blade (2 –)3 – 5-foliolate (in T. celebica and T. wawrae blades occasionally
simple) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Leaf blade simple, lobed or unlobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2a. Probract chartaceous, ± flat. Texture of leaf blade various. Fruit 10 cm long or
less, when fruit larger, then exocarp woody . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
b. Probract thick, ± concave. Leaf blade chartaceous or coriaceous. Fruit (narrowly)
elliptic, (9 –)10 cm long or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Probract with glands. Exocarp leathery. Seeds 16 – 20 mm long. — West
Malesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. T. elmeri
b. Probract without glands. Exocarp woody. Seeds 20 – 27 mm long. — Moluccas,
New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. T. papuana
4a. Seeds ± ovate, 15 – 21 by 8 –14 mm. — Sumatra, Peninsular Malaysia, Borneo,
Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Seeds narrower, (narrowly) elliptic, 9 –17 by 3 – 6 mm . . . . . . . . . . . . . . . . . . . . 5
5a. Fruiting pedicel 5 –7 cm long; seeds elliptic, 9 –10 by 4 – 4.5 mm. — Sulawesi,
Moluccas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
b. Fruiting pedicel 1–1.5 cm long; seeds (narrowly) obovate, 11–16 by 6 –10
mm. — Flores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. T. floresana
6a. Plant (sub)annual; monoecious; diurnal. [Male bracts 1(– 2) mm long, caducous.]
Corolla 20(–30) mm diam. or less. Seeds thickish, compressed, with coarsely
sinuate (undulate) edge. [Fruit small, or very much elongated in cultivated var.
anguina.] — Widespread . . . . . . . . . . . . . . . . . . . 7. T. cucumerina (2 varieties)
b. Plant (sub)perennial; monoecious or dioecious; largely nocturnal. Corolla 30
mm diam. or more. Seeds either tumid, with median belt, or seeds compressed
with edge smooth or ± crenulate (sometimes ± truncate or notched at one or both
ends) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Seeds tumid (i.e. as if 3-parted, with the embryo in the middle) . . . . . . . . . . . . . 8
b. Seeds compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
8a. Seeds fusiform (barrel-shaped), lateral sides ± conical (Fig. 91b). — Wide-
spread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b Seeds butterfly-like, with inflated sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9a. Stem angular. [Leaf blade scabrous, apex cuspidate. Seeds 10 –12 by 17– 20 by c.
5 mm.] — Sabah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. T. pendula
b. Stem terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Apex of leaf blade (obtuse or) acute-acuminate, often with a short mucro. Seeds
6–10 by 7–12 by 2–3 mm. — Sumatra, Borneo . . 3. T. beccariana (2 subspecies)
b. Apex of leaf blade acute-acuminate, with a 13 – 20(– 25) mm long acumen. Seeds
c. 10 by 18 by 4 mm. — Sabah . . . . . . . . . . . . . . . . . . . . . . . . . . 25. T. mucronata
11a. Seeds subcircular, with broad margin, radiately striate or not. Leaf blade un-
lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
b. Seeds (ob)ovate or (narrowly) elliptic, margin narrower or absent. Leaf blade lobed
or unlobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12a. Leaf blade subcircular in outline, auricles absent. — Sumatra . . 37. T. rotundifolia
b. Leaf blade (narrowly) ovate in outline, with at transition to petiole 2 gland-bearing
bulges or auricles. — Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13a. Fruiting among or close to the leaves . . . . . . . . . . . . . . . . . . . . . . . . 2. T. auriculata
b. Fruiting far below the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32. T. postarii
14a. Seeds notched or undulate-truncate at one or two ends . . . . . . . . . . . . . . . . . . . . . 15
b. Seeds neither notched nor undulate-truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
15a. Fruiting pedicel 2-coloured (with a smooth portion to the fruit). Seeds embedded in
green-black pulp. — North Sumatra, Peninsular Malaysia . . . . . 14. T. emarginata
b. Fruiting pedicel evenly coloured; seeds embedded in white or pink pulp (fruit of
T. pulleana not known). — New Guinea [sect. Edulis] . . . . . . . . . . . . . . . . . . . . . 16
16a. Leaf blade unlobed, subcoriaceous, ± hastate, mostly with some spines towards the
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. T. schlechteri
b. Leaf blade unlobed or lobed, ± membranous, chartaceous or (sub)coriaceous, margin
not spiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a. Seeds with cuneate (acute) base, margin faint. Leaf blade unlobed, margin entire.
Fruit with exocarp longitudinally grooved, ± woody, smooth. — Papua New Guinea
(Woodlark Is., Bougainville Is.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. T. dentifera
b. Seeds truncate, without margin. Leaf blade lobed or unlobed. Fruit with exocarp
leathery, drying ± winkled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Seeds subtriangular, at apex much broader than at base . . . . . . . . . . . . . . . . . . . . . 19
b. Seeds narrowly elliptic, ± parallel-sided, ± equally broad at both ends (resembling
seeds of the genus Lagenaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19a. Fruit smooth on drying; fruiting pedicel 1.5 – 2.5 cm long, 2 – 3 mm thick. Seeds
8 – 10 mm long. [Male rachis not thickened; bracts dentate.] . . . . . . 18. T. hastata
b. Fruit rough on drying; fruiting pedicel c. 3 cm long, 3 mm thick, finely brown-hairy.
Seeds 10–13 mm long. [Male rachis thickened; bracts (sub)entire.] . 8. T. densiflora
20a. Leaf blade chartaceous or coriaceous, glabrous. Fruiting pedicel 3–10 cm long . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. T. edulis (3 varieties)
b. Leaf blade membranous or chartaceous, rough hairy on upper surface. Fruiting pedi-
cel 1.5–3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. T. laeoica (4 forms)
21a. Seeds at base chisel-shaped pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
b. Seeds not pointed (base rounded or truncate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
22a. Leaf blade broadly ovate in outline, unlobed, but margin coarsely dentate. Blade
glands large, 2 – 5 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . 31. T. planiglans
b. Leaf blade subcircular in outline, 5(–7)-angular or shallowly lobed. Blade glands
small, c. 1 mm diam. or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23a. Stem sharply angular; young shoots reddish. Leaf blade brown or greenish on
drying. Fruit ripening orange, paler speckled. — Sabah . . . 20. T. kinabaluensis
b. Stem grooved; young shoots green. Leaf blade blackish on drying. Fruit ripening
evenly red. — Widespread . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
24a. Seeds, at least partly, (8 –)10 mm wide or more . . . . . . . . . . . . . . . . . . . . . . . . 25

b. Seeds less than (8 –)10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
25a. Plant villose all over. Fruit pulp white. Tendril 3–9-branched . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42. T. villosa (2 subspecies)
b. Plant subglabrous. Fruit pulp green-black. Tendril 2- or 3-branched . . . . . . . . . . . 26
26a. Leaf blade sometimes simple, shallowly or deeply lobed (usually 3 – 5-foliol-
ate). Fruit ovoid, evenly orange-red or yellow-striped; pericarp c. 5 mm thick . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae (2 forms)
b. Leaf blade 3–5-lobed. Fruit globose, whitish green-striped or not; pericarp 10–18
mm thick. — North Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. T. leuserensis
27a. Seeds (13–)15–21(–25) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
b. Seeds less than 15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
28a. Fruiting pedicel 2-coloured, with a smooth part at the side of the fruit, the remaining
part grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
b. Fruiting pedicel evenly coloured and smooth overall . . . . . . . . . . . . . . . . . . . . . . . 29
29a. Fruiting pedicel 3–5 mm thick. Leaf blade drying dull, ± scabrous . . . . . . . . . . . 30
b. Fruiting pedicel 10 mm thick or more. Leaf blade not scabrous . . . . . . . . . . . . . . 31
30a. Fruit globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. T. obscura
b. Fruit ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. T. intermedia
31a. Tendril simple. Blade base at transition to petiole ± cuneate . . . . . . . 17. T. globosa
b. Tendril 2–4-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32a. Seeds ± rounded at both ends, 17–18 by 5–6 mm . . . . . . . . . . . . . . . . 41. T. valida
b. Seeds subtruncate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33a. Leaf blade deeply (3–)5-lobed (in juvenile stages entire). — Sumatra, Borneo, Java
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana (2 subspecies)
b. Leaf blade entire or 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34a. Fruit ellipsoid, c. 5 cm long, fruiting pedicel c. 2 cm long. — Sarawak, Philippines
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. T. ellipsoidea
b. Fruit subglobose, 10–13 cm long, fruiting pedicel c. 4 cm long. — Sabah
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39. T. sepilokensis
35a. Leaf blade unlobed, ± coriaceous, margin recurved. — Central & South Sumatra and
West Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. T. coriacea
b. Leaf blade lobed (or unlobed), chartaceous or (sub)coriaceous, margin flat . . . . . 36
36a. Leaf blade reddish brown on drying. Probract narrowly elliptic or linear, 10–20 by
c. 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
b. Leaf blade green-brown on drying. Probract (ob)ovate-elliptic, 3–15 by 4–8 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
37a. Fruit ovoid-ellipsoid; seeds 1–2 mm thick . . . . . . . . . . . . . . . . . . . . . 33. T. pubera
b. Fruit subglobose; seeds 3–4 mm thick . . . . . . . . . . . . . . . . . . . . . 22. T. leuserensis
38a. Fruit more than 10 cm long. — Sulawesi, Timor, Philippines, Moluccas . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Fruit (much) less than 10 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39a. Fruiting pedicel 5 –7 cm long. [Leaf blade simple, but usually 5-foliolate.] —
Sulawesi, Moluccas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
b. Fruiting pedicel 1.5–2.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40a. Fruit ripening (possibly) green. — Borneo, Philippines; peat swamp forest . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36. T. refracta
b. Fruit ripening red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41a. Fruit broadly ovoid, 6 –7 cm long; dry pericarp 10 –15 mm thick; fruiting pedicel
c. 2 mm thick. Leaf blade green on drying, margin usually conspicuously dentate.
— Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. T. philippinensis
b. Fruit (narrowly) ovoid, 7– 9 cm long; dry pericarp 5 –10 mm thick; fruiting pedicel
3 – 9 mm thick. Leaf blade (green-)brown on drying, margin usually entire or shal-
lowly dentate (dentate in Sulawesi). — Widespread, not in the Philippines . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata (4 forms)
3 – KEY TO THE SPECIES OF SUMATRA (15 species)
3. T. beccariana Cogn. 24. T. montana Rugayah

4. T. borneensis Cogn. 30. T. pilosa Lour.
6. T. coriacea Blume 33. T. pubera Blume
7. T. cucumerina L. 35. T. quinquangulata A.Gray
13. T. elmeri Merr. 37. T. rotundifolia Rugayah
14. T. emarginata Rugayah 40. T. tricuspidata Lour.
17. T. globosa Blume 43. T. wawrae Cogn.
22. T. leuserensis Rugayah
1a. Leaf blade foliolate (sometimes partly or all simple) . . . . . . . . . . . . . . . . . . . . . 2
b. Leaf blade simple (unlobed or lobed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Plant slender, 2 – 5 m high. Leaf blade 3 foliolate, chartaceous; probract membra-
nous, flat (sometimes caducous). Tendrils simple or 2-branched. Fruit 6 – 9 cm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Plant stouter. Leaf blade 3 – 5 foliolate, coriaceous; probract chartaceous, in-
curved-concave. Tendrils 2- or 3-branched. Fruit more than 10 cm long . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13.T. elmeri
3a. Leaf blade unlobed. [Leaf blade subcircular or ovate, margin entire, minute sparse
dents excepted.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
b. Leaf blade lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4a. Blade margin ± recurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. T. coriacea
b. Blade margin flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Leaf blade (usually) minutely hairy beneath, at least along the veinlets . 30. T. pilosa
b. Leaf blade glabrous beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Veinlets raised beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. beccariana
b. Veinlets not or but little raised beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Leaf blade chartaceous, ovate; venation ± raised, distinct. Seeds 8 –10 mm long.
— Siberut Is. . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata forma siberutensis
b. Leaf blade ± membranous; venation flat, faint beneath. Seeds 15 – 20 mm long 8
8a. Leaf blade subcircular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37. T. rotundifolia
b. Leaf blade ovate-hastate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
9a. Plant with fruit (and female flowers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
b. Plant with male flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
10a. Fruit ovoid. Seeds tumid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

b. Fruit globose, ellipsoid, (ob)ovoid, or long-cylindrical. Seeds compressed . . . 12
11a. Fruit pulp not fibrous. Seeds ± fusiform (barrel-shaped) (Fig. 91b) . 30. T. pilosa
b. Fruit pulp fibrous. Seeds butterfly-shaped with inflated lateral parts. (Lobed leaf
blades not seen in Sumatra) . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. beccariana
12a. Seed margin undulate or coarsely notched. Fruit ± ellipsoid, 2.5 – 5 cm long, or
long-cylindrical in cultivated var. anguina . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Seed margin entire. Fruit globose, ellipsoid, or (ob)ovoid . . . . . . . . . . . . . . . . 13
13a. Fruiting pedicel with 2 parts with differing surface colour and texture . . . . . . 14
b. Fruiting pedicel evenly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14a. Fruit ellipsoid. Seeds notched at one end . . . . . . . . . . . . . . . . 14. T. emarginata
b. Fruit globose. Seeds not notched . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
15a. Fruit (sub)globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
b. Fruit longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
16a. Fruiting pedicel thick, 10 –20 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
b. Fruiting pedicel c. 5 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17a. Tendrils simple. Fruit globose, 5.5–12 cm diameter . . . . . . . . . . 17. T. globosa
b. Tendrils 2- or 3-branched. Fruit ellipsoid, more than 10 cm diameter . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana subsp. montana
18a. Leaf blade deeply 3 – 5-lobed. Seeds 3 – 4 mm thick, base and apex rounded . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. T. leuserensis
b. Leaf blade shallowly 5-lobed or ± 5-angular. Seeds 1–2 mm thick, chisel-shaped
pointed at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
19a. Fruit more than 10 cm long; fruiting pedicel c. 10 mm thick . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana subsp. montana
b. Fruit less than 10 cm long; fruiting pedicel 5 mm thick or less . . . . . . . . . . . . 20
20a. Pericarp thin, less than 5 mm thick. Seeds (narrowly or) broadly ovate, 15 – 20
mm long, 2 – 3 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Pericarp thick, c. 5 mm or more. Seeds obovate, 10 –13 mm long, less than 2 mm
thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Leaf blade hairy beneath. Stem and leaf blade in young state reddish, reddish
brown on drying. Probract narrowly elliptic or linear . . . . . . . . . . 33. T. pubera
b. Leaf blade glabrous. Stem and leaf blade green, (dark) brown on drying. Probract
obovate or elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
22a. Bracts 1(– 2) mm long, often caducous (including cultivated var. anguina) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Bracts more than 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23a. Rachis 1– 2 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
b. Rachis 2 –10 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24a. Persistent basal part of pedicel (1–)2–15 mm long. Leaf blade beneath finely hairy,
at least along the veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Persistent basal part of pedicel usually (nearly) absent. Leaf blade scabrous . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
25a. Rachis of raceme 5 –10 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
b. Rachis of raceme 2 – 5 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
26a. Tendrils simple. Bracts membranous, with short soft yellowish hairs, margin en-
tire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. T. globosa
b. Tendrils 3-branched. Bracts chartaceous, with minute brown hairs, apical part
coarsely dentate . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana subsp. montana
27a. Younger parts of shoots reddish tinged on drying . . . . . . . . . . . . . . . . . . . . . . . 28
b. Whole plant green, brown, or blackish on drying . . . . . . . . . . . . . . . . . . . . . . . 29
28a. Leaf blade glabrous. Petals white . . . . . . . . . . . . . . . . . . . . . . 22. T. leuserensis
b. Leaf blade hairy beneath. Petals with pinkish veining and/or fringes . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. T. pubera
29a. Leaf blade ± 5-angular, with several minute glands close to the insertion of the
petiole. Bracts entire. [Sepals with slender sidelobes.] . . 35. T. quinquangulata
b. Leaf blade (deeply) 3 – 5 lobed, c. 1/3 or deeper; glands fewer, not close to the
insertion of the petiole. Bracts dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30a. Leaf blade 3-lobed to c. 1/2-way deep, lobes ± ovate-triangular. Bracts without
glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
b. Leaf blade deeply 5-lobed, lobes narrowly ellipsoid. Bracts with glands . . . . . . 31
31a. Probract ± elliptic, c. 4 mm long. [Seeds notched at apex] . . . 14. T. emarginata
b. Probract narrowly elliptic, 5 –18 mm long. [Seeds ± rounded at both ends] . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
4 – KEY TO THE SPECIES OF PENINSULAR MALAYSIA & SINGAPORE (7 species)
4. T. borneensis Cogn. 35. T. quinquangulata A.Gray

7. T. cucumerina L. 40. T. tricuspidata Lour.
13. T. elmeri Merr. 43. T. wawrae Cogn.
30. T. pilosa Lour.
1a. Leaf blade foliolate (in T. wawrae sometimes partly or all simple) . . . . . . . . . . 2
b. Leaf blade simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Leaf blade 3-foliolate, membranous or chartaceous. Probract membranous, flat,
sometimes caducous. Tendrils 1- or 2-branched. Rachis of male raceme 1–1.5 mm
thick. Fruit ovoid, 6 – 9 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Leaf blade 3 – 5-foliolate, chartaceous or coriaceous. Probract chartaceous, con-
cave. Tendrils 2- or 3-branched. Male rachis 2 – 3 mm thick. Fruit narrowly ovoid,
more than 10 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13.T. elmeri
3a. Plant (sub)annual; monoecious. Leaf blade membranous, green-yellow on drying.
Corolla 20(–30) mm diameter. Male bract 0.5– 2 mm long, persistent or caducous.
[Fruit small, ovoid, or much elongated, cylindrical in cultivated var. anguina.]
Seeds with undulate margin . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious. Leaf blade membranous or charta-
ceous, green, brown, or blackish on drying. Corolla more than 30 mm diameter.
Male bract 3 mm long or more. Seeds without undulate margin . . . . . . . . . . . . . 4
4a. Plant with fruit (and female flowers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
b. Plant with male racemes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5a. Fruit globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
b. Fruit ovoid or ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6a. Fruiting pedicel of 2 differently structured and coloured parts. Leaf blade deeply
5(–7)-lobed. Seeds ± rounded at both ends . . . . . . . . . . . . . . . . . 4. T. borneensis
b. Fruiting pedicel of even colour and structure. Leaf blade ± 5-angular. Seeds chisel-
shaped pointed at base . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
7a. Seeds tumid (barrel-shaped or (sub)fusiform) (Fig. 91b) . . . . . . . . . 30. T. pilosa
b. Seeds compressed, rounded at both ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Leaf blade 3-lobed, (1/4 –)1/2 deep. Fruit ripening evenly red. Seeds 10–13 mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
b. Leaf blade variously lobed. Fruit ripening red, mostly yellow striped. Seeds 15 – 20
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
9a. Rachis 1– 2 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
b. Rachis 2 – 4 mm thick or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10a. Leaf blade usually finely hairy beneath. [Seeds tumid (barrel-shaped or (sub)-
fusiform) (Fig. 91b).] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Leaf blade glabrous beneath. [Seeds compressed.] . . . . . . . . . . . . 43. T. wawrae
11a. Leaf blade ± 5-angular, with small glands near the insertion of the petiole. Bracts
entire. Sepals (mostly) with narrow lateral lobes . . . . . . 35. T. quinquangulata
b. Leaf blade unlobed or variously lobed. Bracts incised. Sepals entire, rarely few-
lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Leaf blade deeply 5 –7-lobed, usually with few glands 1– 3 mm diameter . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
b. Leaf blade 3-lobed, 1/4 –1/2 deep; glands scattered, less than 1 mm diameter . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
5 – KEY TO THE SPECIES OF BORNEO (25 species)
1. T. adhaerens W.J.de Wilde & Duyfjes 25. T. mucronata Rugayah

2. T. auriculata Rugayah 26. T. obscura Rugayah
3. T. beccariana Cogn. 28. T. pendula Rugayah
4. T. borneensis Cogn. 30. T. pilosa Lour.
7. T. cucumerina L. 32. T. postari W.J.de Wilde & Duyfjes
12. T. ellipsoidea Merr. 33. T. pubera Blume
13. T. elmeri Merr. 35. T. quinquangulata A.Gray
14. T. globosa Blume 36. T. refracta C.H.Yueh
16. T. fusca W.J.de Wilde & Duyfjes 39. T. sepilokensis Rugayah
19. T. intermedia W.J.de Wilde & Duyfjes 40. T. tricuspidata Lour.
20. T. kinabaluensis Rugayah 42. T. villosa Blume
23. T. longispicata Rugayah 43. T. wawrae Cogn.
24. T. montana Rugayah
1a. Leaf blade foliolate (in T. wawrae sometimes, in T. elmeri rarely, partly or all
simple) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
b. Leaf blade simple (unlobed or lobed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2a. Leaf blade 3-foliolate, membranous or chartaceous; probract membranous, flat
(sometimes caducous). Tendrils simple or 2-branched. Male rachis 1–1.5 mm
thick. Fruit 6 –9 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Leaf blade 3 – 5-foliolate, chartaceous or coriaceous; probract chartaceous, con-

cave. Tendrils 2- or 3-branched. Male rachis 2 – 3 mm thick. Fruit more than 10
cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. T. elmeri
3a. Plant (sub)annual; monoecious. [Leaf blade membranous, green-yellow on dry-
ing.] Corolla 20(– 30) mm diameter. Male bracts 1(– 2) mm long, often caducous.
[Fruit ovoid, or long-cylindrical in cultivated var. anguina.] Seeds with undulate
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious. Corolla usually more than 30
mm diam. (smaller in T. postari). Male bract 3 mm long or more. Seeds tumid or
compressed, not with undulate margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Whole plant conspicuously (densely) soft hairy; hairs 1 mm long or more . . . . 5
b. Whole plant glabrous or (sparsely) hairy; hairs less than 1 mm long . . . . . . . . . 6
5a. Indumentum grey-rusty villose. Tendrils 5 –15-branched, point of branching 1– 2
cm from the base. Leaf blade without conspicuous mucro. Fruit broad ellipsoid
(-globose), 9 –15 cm long. Seeds compressed . . . . . . . . . . . . . . . . . 42. T. villosa
b. Indumentum grey, hirsute. Tendrils 3 – 5-branched, point of branching 2 – 4 cm
from the base. Leaf blade with conspicuous mucro, 13 – 25 mm long. Fruit el-
lipsoid, 9 –10 cm long. Seeds tumid . . . . . . . . . . . . . . . . . . . . 25. T. mucronata
6a. Leaf blade at transition to petiole with 2 gland-bearing auricles. [Male bracts
linear. Fruit in a raceme, globose.] Seeds subcircular with broad radially striate
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Leaf blade at base without auricles. Seeds tumid or compressed, without radially
striate margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7a. Flowers conspicuously brown hairy (fruit and seed not known) . . . . 16. T. fusca
b. Flowers greyish hairy or glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Inflorescences borne on leafy twigs . . . . . . . . . . . . . . . . . . . . . . 2. T. auriculata
b. Inflorescences borne on older stems . . . . . . . . . . . . . . . . . . . . . . . . 32. T postari
9a. Stem sharply angular or ridged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
b. Stem striate or grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10a. Stem and leaf blade brown on drying, reddish tinged. Leaf apex without acu-
men. Probract narrowly elliptic. Male raceme solitary. Seeds compressed, with
± cuneate base, and broadly rounded apex . . . . . . . . . . . . . 20. T. kinabaluensis
b. Stem and leaf blade green on drying, not reddish tinged. Leaf apex often with
a 10 – 25 mm long acumen. Probract absent. Male (and female) racemes mostly
fascicled, pendent. Seeds tumid . . . . . . . . . . . . . . . . . . . . . . . . . . 28. T. pendula
11a. Plant with fruit (and/or female flowers). Fruit of T. longispicata not known . . 12
12a. Seeds tumid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
b. Seeds compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
13a. Leaf venation ± flat beneath, usually minutely hairy. Fruit pulp not fibrous. Seeds
barrel-shaped or (sub)fusiform (Fig. 91b) . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Leaf venation much raised beneath, glabrous or hairy. Fruit pulp coarsely fibrous.
Seeds butterfly-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a. Plant free-climbing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. beccariana
b. Plant growing adherent to tree trunks. (Fruit not known) . . . . . . 1. T. adhaerens
15a. Fruit globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

b. Fruit ovoid or ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
16a. Fruiting pedicel with two parts of different surface and colour . 4. T. borneensis
b. Fruiting pedicel evenly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a. Fruiting pedicel less than 10 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
b. Fruiting pedicel 10 –15 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18a. Leaf blade ± 5-angular, blackish on drying; glands several, 0.5 –1 mm diam., close
to the insertion of the petiole. Seeds 9 –14 by 4 – 5 mm, chisel-shaped pointed at
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Leaf blade unlobed or (deeply) lobed, brown-blackish on drying; glands absent
or few, 1– 2 mm diam., scattered but mainly towards the base of the blade. Seeds
18 – 22 by 7– 8 mm, rounded at both ends . . . . . . . . . . . . . . . . . . . 26. T. obscura
19a. Leaf blade pale green on drying. Tendrils simple . . . . . . . . . . . . . 17. T. globosa
b. Leaf blade green or brown on drying. Tendrils (usually) 2 – 4(– 5)-branched . 20
20a. Leaf blade unlobed or shallowly 3-lobed, green on drying. Fruiting pedicel c. 4
cm long. Seeds 13 –18 by 7– 9 mm . . . . . . . . . . . . . . . . . . . . 39. T. sepilokensis
b. Leaf blade deeply 5 –7-lobed, greenish brown or blackish on drying. Fruiting
pedicel c. 3 cm long. Seeds 17– 20 mm long . . . 24. T. montana subsp. crassipes
21a. Plant slender, leafy stem 2 – 3 mm diameter. Pericarp ± leathery, thin . . . . . . . 22
b. Plant stouter, leafy stem 3 – 4 mm diameter or more. Pericarp carnose, c. 5 mm
thick when dry. [Seeds c. 10 mm long. Leaf blade 3- or 5-lobed.] . . . . . . . . . . 23
22a. Leaf blade ovate-hastate, usually ± lobed; glands c. 0.5 mm diameter. Fruit 6 –10
cm long, yellow striped. Seeds 15 –18 mm long . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae and 19. T. intermedia
b. Leaf blade (narrowly) ovate, unlobed; glands 1– 2 mm diameter. Fruit c. 5 cm long
(ripening colour not known). Seeds c. 13 mm long . . . . . . . . . 12. T. ellipsoidea
23a. Fruit (possibly) ripening green. — Peat swamp forest . . . . . . . . . 36. T. refracta
b. Fruit ripening red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24a. Leaf blade (±) hairy, 3 – 5-lobed, margin (coarsely) dentate. Young parts of shoots
reddish brown on drying. Probract narrowly elliptic or linear, 10 – 20 mm long.
Petal fringes tinged pinkish. Fruiting pedicel (1.5 –)2 – 5 cm long . 33. T. pubera
b. Leaf blade glabrous or early glabrescent, 3-lobed to 1/4 –1/2 deep, margin entire
or dentate. Young parts of shoots brown on drying. Probract obovate or elliptic,
5(–10) mm long. Petals fringes white. Fruiting pedicel 0.5 – 2.5 cm long . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
25a. Rachis of male raceme 1–1.5(– 2) mm thick. Leaf blade frequently unlobed . . 26
b. Rachis of male raceme 2 –10 mm thick. Leaf blade mostly lobed . . . . . . . . . . . 29
26a. Blade glands c. 2 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . . 12. T. ellipsoidea
b. Blade glands less than 1 mm diameter, or absent . . . . . . . . . . . . . . . . . . . . . . . 27
27a. Leaf blade glabrous beneath (hairy in forma hirsuta). Persistent part of pedicel
almost absent (sometimes persistent) . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae
b. Leaf blade ± hairy beneath. Persistent part of pedicel 2 mm long or more . . . . 28
28a. Venation on lower leaf surface raised. Peduncle short, 0.5 – 3 cm long . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. T. beccariana
b. Venation on lower leaf surface ± flat. Peduncle longer, (2 –)5 – 8(–17) cm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
29a. Leaf blade ± 5-angular. Bracts entire. Sepals mostly with narrow sidelobes . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Leaf blade various. Bracts incised. Sepals entire or lobed . . . . . . . . . . . . . . . . 30
30a. Persistent part of pedicel 5 –10 mm long. Bracts 7–12 mm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. T. longispicata
b. Persistent part of pedicel shorter or absent. Bracts usually larger . . . . . . . . . . . 31
31a. Rachis 2 – 4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
b. Rachis 5 –10(– 20) mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
32a. Rachis ± zigzag . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36. T. refracta
b. Rachis straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33a. Bracts less than 15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . 19. T. intermedia
b. Bracts more than 15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34a. Bracts broad, deeply fan-shaped incised. Plant dull blackish on drying . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. T. obscura
b. Bracts incised, but not fan-shaped. Plant green or (reddish) brown on drying . 35
35a. Leaf blade ± hairy. Shoots reddish tinged. Petal fringes tinged pinkish . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. T. pubera
b. Leaf blade glabrous or early glabrescent. Shoots green or brown on drying. Petal
fringes white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36a. Leaf blade deeply 5 –7-lobed. Blade glands few (or absent), near the insertion of
the petiole, 1– 3 mm diameter. Probract narrowly elliptic, without glands . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. borneensis
b. Leaf blade 3-lobed to 1/4 –1/3 of the blade. Blade glands scattered, less than 1 mm
diameter. Probract ± (ob)ovate or elliptic, with conspicuous glands . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
37a. Bracts entire, pale yellow soft hairy. Tendrils simple . . . . . . . . . . 17. T. globosa
b. Bracts incised. Tendrils 2 – 4-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38a. Leaf blade unlobed or shallowly (2- or) 3-lobed. — Lowland . 39. T. sepilokensis
b. Leaf blade deeply 5(–7)-lobed. — Lowland and montane . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana subsp. crassipes
6 – KEY TO THE SPECIES OF JAVA (10 species)
6. T. coriacea Blume 33. T. pubera Blume

17. T. globosa Blume 40. T. tricuspidata Lour.
24. T. montana Rugayah 42. T. villosa Blume
30. T. pilosa Lour. 43. T. wawrae Cogn.
1a. Leaf blade 3-foliolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43. T. wawrae

b. Leaf blade simple, lobed or unlobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Leaf blade (membranous) coriaceous, unlobed, margin entire, somewhat recurved.
[Male bracts small, 5–7 mm long, shifted upwards on the pedicel.] . 6. T. coriacea
b. Leaf blade (except in old stages) generally not coriaceous, unlobed or lobed,
margin various, flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Leaf blade and young stem conspicuously villose . . . . . . . . . . . . . 42. T. villosa
b. Leaf blade and stem hairy, or scabrous, not villose . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Plant mostly annual. Leaf blade membranous. Flowers monoecious, nocturnal
and diurnal; corolla small, 20(– 30) mm diameter. Bracts of male inflorescences
minute, 2 mm long or less, mostly caducous. Probract absent. Seeds compressed
with undulate edge. (Fruit much elongated in cultivated var. anguina) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial. Leaf blade generally thicker. Flowers mostly dioecious
(sometimes monoecious in T. quinquangulata), nocturnal; corolla 30 mm diameter
or more. Bracts of male inflorescences 3 mm long or more, persistent or caducous.
Probract present. Seeds tumid or compressed, edge not undulate . . . . . . . . . . . . 5
5a. Bracts of male inflorescences less than 15 mm long. Seeds tumid (barrel-shaped
or (sub)fusiform) (Fig. 91b); fruit ± ellipsoid-fusiform . . . . . . . . . . 30. T. pilosa
b. Bracts of male inflorescences generally much larger. Seeds (rather) compressed,
obovate or narrowly elliptic with truncate or cuneate base; fruit ovoid, ellipsoid
or globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Tendrils unbranched. Blade base mostly cuneate at transition to the petiole. Rachis of
male inflorescences conspicuously thickened, bracts large, finely hairy. Fruit glo-
bose; fruiting pedicel stout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. T. globosa
b. Tendrils 3 – 5-branched. Blade base cordate. Rachis of male inflorescences thick-
ened or not. Fruit (ob)ovoid or globose; fruiting pedicel generally more slender 7
7a. Rachis of male (or hermaphroditic) inflorescences conspicuously thickened and
set with persistent dark-drying bracts. Leaf blade glabrous, deeply lobed. — Mon-
tane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. T. montana subsp. montana
b. Rachis not thickened. Leaf blade finely hairy beneath and deeply lobed, or gla-
brous and shallowly lobed. — Lowland or montane . . . . . . . . . . . . . . . . . . . . . . 8
8a. Growing twig tinged reddish at apex. Leaf blade ± scabrid-hairy beneath, lobed
to ± halfway or more. Probract narrowly elliptic or linear, entire or ± incised at
apex. [Male bracts finely incised. Sepals mostly few-lobed. Fruit ovoid.] . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. T. pubera
b. Growing twig green. Leaf blade glabrous (glabrescent) or scabrous, lobed to
c. 1/3 deep. Probract (narrowly) ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9a. Male bracts incised. Sepals in male flowers (sub)entire. Fruit ovoid. Leaf blade
3-lobed, glands scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata
b. Male bracts (almost) entire. Sepals in male flowers usually with few slender lobes.
Fruit (depressed) globose. Leaf blade 5-angular or shallowly 5-lobed, glands usu-
ally several, close to the insertion of the petiole . . . . . . . 35. T. quinquangulata
7 – KEY TO THE SPECIES OF SULAWESI (6 species)

5. T. celebica Cogn. 35. T. quinquangulata A.Gray
30. T. pilosa Lour. 41. T. valida Rugayah
1a. Leaf blade 3-foliolate (or partly simple) . . . . . . . . . . . . . . . . . . . . . 5. T. celebica

2a. Plant (sub)annual; monoecious. Probract absent. Male bracts 1(– 2) mm long,
often caducous. Corolla 20(– 30) mm diameter. [Fruit small, ovoid, or long-
cylindrical in cultivated var. anguina.] Seeds compressed with undulate edge . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious. Probract present (absent in
T. pilosa). Corolla 30 mm diameter or more. Seeds tumid or compressed, not with
undulate edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Leaf blade ± 5-angular, [blackish on drying, with minute glands close to the in-
sertion of the petiole]. Male bracts entire. [Sepals usually with distinct, narrow,
lateral lobes.] Fruit (depressed) globose. Seeds compressed, (narrowly) elliptic,
chisel-shaped pointed at base . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Leaf blade various. Male bracts entire or mostly incised. Fruit (short-)ovoid, ob
ovoid, or ellipsoid. Seeds tumid or compressed, not pointed at base . . . . . . . . . 4
b. Plant with male flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
5a. Fruit ± ellipsoid, 10 –18 cm long; fruiting pedicel 10 – 20 mm thick. [Seeds com-
pressed, obovate or elliptic, (8 –)9 –11(– 25) mm long.] . . . . . . . . . 41. T. valida
b. Fruit ovoid or ellipsoid; fruiting pedicel 2 – 4 mm thick . . . . . . . . . . . . . . . . . . 6
6a. Seeds tumid (barrel-shaped or (sub)fusiform) (Fig. 91b). Fruit mostly striped . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Seeds compressed. Fruit evenly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Leaf blade subhastate (or 3-foliolate). Fruiting pedicel 2– 7 cm long. Seeds 9 –13
by 4 – 7 by 2 – 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
b. Leaf blade suborbicular, 3-lobed to halfway deep. Fruiting pedicel 0.5 – 2.5 cm
long. Seeds 10 –13 by 5 –10 by 1– 3 mm . . . . . . . . . . . . . . . 40. T. tricuspidata
8a. Leaf blade deeply 5-lobed. Peduncle 9 –18.5 cm long; rachis 5 –10 mm thick . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Leaf blade (sub)unlobed or 3-lobed. Rachis (1–)2 – 3 mm thick, peduncle usually
shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9a. Plant (dark) brown on drying. Bracts obovate or rhomboid, 15 – 25 mm long. Leaf
blade ± glabrous, 3-lobed to c. halfway the blade . . . . . . . . . 40. T. tricuspidata
b. Plant greenish on drying. Bracts (narrowly) elliptic, 25 mm long or less. [Leaf
blade various.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Leaf blade usually ± hairy beneath. Persistent part of pedicel 2 mm long or
more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Leaf blade glabrous. Persistent part of pedicel less than 2 mm long . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
8 – KEY TO THE SPECIES OF PHILIPPINES (11 species)
7. T. cucumerina L. 31. T. planiglans Rugayah

11. T. cf. edulis Rugayah (not in the key) 35. T. quinquangulata A.Gray
12. T. ellipsoidea Merr. 36. T. refracta C.H.Yueh
13. T. elmeri Merr. 41. T. valida Rugayah
29. T. philippinensis Rugayah 42. T. villosa Blume
30. T. pilosa Lour.
1a. Leaf blade 3- or 5-foliolate. — Palawan . . . . . . . . . . . . . . . . . . . . . 13. T. elmeri
2a. Plant (sub)annual; monoecious; diurnal. Corolla 20(– 30) mm diameter. Male bracts
1(– 2) mm long, mostly caducous. Seeds compressed, with undulate edge. [Pro-
bract absent. Fruit small, ovoid, or long cylindrical in cultivated var. anguina.] .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious. Corolla 30 mm diam. or more.
Male bracts larger. Seeds tumid or compressed, edge not undulate . . . . . . . . . . 3
3a. Plant densely (grey-)brown villose. [Male bracts large, membranous, ± entire.] .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42. T. villosa
b. Plant thinly hairy or glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Leaf blade unlobed (broadly ovate); glands (1–)2 –5 mm diameter or more . . . . 5
b. Leaf blade unlobed or lobed; glands 1(– 2) mm diameter or less, or absent . . . . 6
5a. Blade margin coarsely dentate; glands 2 – 5 mm diameter, situated near the inser-
tion of the petiole. Seeds 13 –16 mm long, cuneate at one end . 31. T. planiglans
b. Blade margin entire; glands 1– 2 mm diam., scattered at about the centre or to-
wards the base. Seeds c. 13 mm long, not cuneate at one end . 12. T. ellipsoidea
7a. Fruit (depressed) globose. [Leaf blade ± 5-angular, blackish on drying, with sev-
eral small glands close to the insertion of the petiole. Seeds narrowly elliptic,
chisel-shaped pointed at base.] . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Fruit ± (ob)ovoid or ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Fruit c. 10 cm long. Fruiting pedicel 10 – 20 mm thick . . . . . . . . . . 41. T. valida
b. Fruit smaller. Fruiting pedicel less than 5 mm thick . . . . . . . . . . . . . . . . . . . . . . 9
9a. Seeds tumid (barrel-shaped or (sub)fusiform) (Fig. 91b) . . . . . . . . . 30. T. pilosa
b. Seeds compressed, (narrowly) ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Fruit ripening (possibly) green. — Peat swamp forest . . . . . . . . . 36. T. refracta
b. Fruit ripening red. — Dryland areas . . . . . . . . . . . . . . . . . . 29. T. philippinensis
11a. Bracts 10 – 30 mm long, entire. Leaf blade 5-angular, blackish on drying . . . . . .
b. Bracts incised (or much smaller). Leaf blade various, not blackish on drying . 12
12a. Male raceme with stout rachis, 5 –10 mm thick. Leaf blade deeply 5-lobed . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Male raceme with slender rachis, 1– 2 mm thick. Leaf blade unlobed or 3(– 5)-
lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13a. Leaf blade usually ± hairy beneath. Corolla c. 30 mm diameter . . . . 30. T. pilosa
b. Leaf blade glabrous (early glabrescent). Corolla larger . . . . . . . . . . . . . . . . . . 14
14a. Raceme zigzag; bracts 10 – 40 mm long, shallowly incised to 3 mm deep, without
glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36. T. refracta
b. Raceme straight; bracts 25 – 45 mm long, deeply finely incised, 5–15 mm deep,
with few glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. T. philippinensis
9 – KEY TO THE SPECIES OF LESSE SUNDA ISLANDS (7 species)

15. T. floresana Rugayah 41. T. valida Rugayah
30. T. pilosa Lour. 42. T. villosa Blume
35. T. quinquangulata A.Gray (not yet
recorded for Lesser Sunda Islands)
1a. Leaf blade 3(– 5)-foliolate (or partly unlobed ?) . . . . . . . . . . . . . . 15. T. floresana
2a. Plant (sub)annual; monoecious; diurnal. Male bracts 1(– 2) mm long, mostly ca-
ducous. Corolla 20(–30) mm diameter. [Fruit small, ovoid, or long-cylindrical in
cultivated var. anguina.] Seeds compressed, pale, with coarsely undulate edge . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial. Corolla c. 30 mm diam. or more. Seeds tumid or compressed,
edge not undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Whole plant densely (grey-)brown villose . . . . . . . . . . . . . . . . . . . . 42. T. villosa
b. Plant thinly hairy or glabrous (glabrescent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Plant with fruit (or female flowers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
b. Plant with male flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
5a. Fruit globose. Seeds narrowly elliptic, chisel-shaped pointed at base. [Leaf blade
± 5-angular, blackish on drying; with several small glands close to the insertion of
the petiole.] (Not yet recorded for Lesser Sunda Islands.) 35. T. quinquangulata
b. Fruit ovoid(-ellipsoid). Seeds tumid or compressed, not pointed. [Leaf blade un-
lobed or (deeply) lobed.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Seeds tumid (barrel-shaped or (sub)fusiform) (Fig. 91b). Leaf blade (sub)glabrous
or hairy, at least on smaller veins beneath . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Seeds compressed, (narrowly) ovate, rounded at both ends. Leaf blade glabrous . 7
7a. Fruit large, 10 cm long or more. Leaves not scabrid . . . . . . . . . . . . . 41. T. valida
b. Fruit smaller. Leaf blade scabrid . . . . . . . . . 40. T. tricuspidata forma asperifolia
8a. Leaf blade glabrous, ± 5-angular [blackish on drying, with several small glands
near the insertion of the petiole.] Bracts c. 20 mm long, entire. Sepals with narrow
sidelobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Leaf blade glabrous, hairy, or scabrous, unlobed or (deeply) lobed. Bracts (coarsely)
indented, or much smaller, (sub)entire. Sepals (mostly) entire . . . . . . . . . . . . . . . 9
9a. Leaf blade subglabrous or hairy, at least on smaller veins beneath, not scabrous.
Bracts 15 mm long or less, coarsely dentate or entire, without glands . 30. T. pilosa
b. Leaf blade scabrous. Bracts c. 20 mm long, deeply indented, with glands . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata forma asperifolia
10 – KEY TO THE SPECIES OF MOLUCCAS (8 species)
5. T. celebica Cogn. 34. T. pulleana Harms

27. T. papuana F.M.Bailey 40. T. tricuspidata Lour.
30. T. pilosa Lour. 41. T. valida Rugayah
1a. Leaf blade 3-foliolate (or partly or all simple) . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Fruit c. 6.5 cm long. Fruiting pedicel 5 –7 cm long; exocarp thick leathery. Seeds
9 –10 by 4 – 4.5 mm. — Buru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
b. Fruit 9 –16(– 20) cm long. Fruiting pedicel 1– 3 cm long; exocarp woody. Seeds
20 – 27 by 11–15 mm. — Aru Islands . . . . . . . . . . . . . . . . . . . . . 27. T. papuana
3a. Plant (sub)annual; monoecious; diurnal. [Probract absent.] Male bracts 1(– 2) mm
long, mostly fugacious. Corolla 20(–30) mm diameter. Seeds compressed, pale,
with coarsely undulate edge. [Fruit small, ovoid, or long-cylindrical in cultivated
var. anguina.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious. [Probract present or absent.] Male
bracts larger. Corolla c. 30 mm diam. or more. Seeds tumid or compressed, not
with undulate edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Plant with fruit (or female flowers). [Fruit not known in T. pulleana, Aru Islands.]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
b. Plant with male flowers. [Male flowers not known in T. tricuspidata forma sera
mensis.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5a. Seeds tumid (barrel-shaped or (sub)fusiform) (Fig. 91b). [Probract absent.] . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Seeds compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Fruit globose. Leaf blade ± 5-angular; with several small glands close to the inser-
tion of the petiole. Seeds narrowly elliptic, chisel-shaped pointed at base . . . . . .
b. Fruit obovoid or ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Fruit large, ± obovoid, 10 –18 cm long; fruiting pedicel 10 – 20 mm thick. — Halma
heira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Fruit smaller; fruiting pedicel more slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Leaf blade ± hastate (or 3-foliolate). Fruiting pedicel 5 –7 cm long. — Buru . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. T. celebica
b. Leaf blade 3-lobed to halfway deep. Fruiting pedicel c. 2.5 cm long. — Seram
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. T. tricuspidata (forma seramensis)
9a. Bracts 10 – 30 mm long. [Sepals mostly with narrow lateral lobes. Leaf blade 5-
angular, blackish on drying; with small glands close to the insertion of the petiole.]
b. Bracts either large (c. 20 mm long or more), incised (sometimes only shallowly
so at apex), or bracts much smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Leaf blade deeply 5(–7)-lobed. Rachis of raceme 5 –10 mm thick . 41. T. valida
b. Leaf blade unlobed or lobed. Rachis 1– 3 mm thick . . . . . . . . . . . . . . . . . . . . . 11
11a. Leaf margin finely dentate. Bracts and flowers (inflorescence) densely brown
short-hairy. — Aru Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. T. pulleana
b. Leaf margin entire. Bracts and flowers (sub)glabrous . . . . . . . . . . . . . . . . . . . 12
12a. Persistent part of pedicel 2 mm long or more . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Persistent part of pedicel less than 2 mm long or absent (flowers from Moluccas
not seen) . . . . . . . . . . . . . . . 5. T. celebica (Buru), 27. T. papuana (Aru Islands)
11 – KEY TO THE SPECIES OF NEW GUINEA (13 species)

(see note at the end of this key)
7. T. cucumerina L. 27. T. papuana F.M.Bailey

8. T. densiflora Rugayah 30. T. pilosa Lour.
9. T. dentifera Rugayah 34. T. pulleana Harms
10. T. dieniensis Merr. & L.M.Perry 35. T. quinquangulata A.Gray
11. T. edulis Rugayah 38. T. schlechteri Harms
18. T. hastata Harms 41. T. valida Rugayah
21. T. laeoica C.Y.Cheng & Lu Q.Huang
1a. Leaf blade 3 – 5-foliolate (sometimes simple in juvenile stages) . . 27. T. papuana
2a. Plant (sub)annual; monoecious; diurnal. Probract absent. Male bracts 1(– 2) mm
long, mostly fugacious. Corolla 20(–30) mm diameter. Seeds compressed, pale,
with coarsely undulate edge. [Fruit small, ovoid, or long-cylindrical in cultivated
var. anguina.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. cucumerina
b. Plant (sub)perennial; monoecious or dioecious; largely nocturnal. Male bracts 5
mm long or more. Corolla 30 mm diam. or more. Seeds compressed (tumid in
T. pilosa), not with undulate edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Entire portion of petal narrowly elliptic. Seeds tumid (barrel-shaped or (sub)fusi
form) (Fig. 91b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. T. pilosa
b. Entire portion of petal broader, ± obovate (apex ± truncate). Seeds compressed . 4
4a. Male bracts c. 20 mm long, entire. Fruit depressed globose; [seeds embedded in
green-black pulp, (narrowly) elliptic, chisel-shaped pointed at base. Leaf blade ±
5-angular.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. T. quinquangulata
b. Male bracts longer or shorter, usually dentate. Fruit subglobose or ellipsoid . . . . 5
5a. Male bracts 40 – 50 mm long, entire or shallowly lobed. Fruit pulp green-black.
Seeds elliptic, rounded at both ends . . . . . . . . . . . . . . . . . . . . . . . . . 41. T. valida
b. Male bracts smaller, dentate (sometimes only at very apex), or bracts much smaller.
Fruit subglobose (T. laeoica, p.p.) or (narrowly) ellipsoid; pulp not green-black;
seeds various, usually densely packed. Leaf blade of various shape, unlobed or
(deeply) lobed. [Old peduncles and/or pedicels often persistent, straw-like. Sect.
Edulis (see note at the end of this key).] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6a. Plant with fruit (or female flowers) (fruit not known in T. dieniensis, T. pulleana)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
b. Plant with male flowers (male flowers not known in T. dentifera) . . . . . . . . . . . 14
7a. Leaf blade entire or lobed, broadly ovate or circular in outline . . . . . . . . . . . . 12

b. Leaf blade entire, ovate-hastate or triangular in outline . . . . . . . . . . . . . . . . . . . 8
8a. Plant delicate. Leaf blade membranous, margin entire . . . . . . 10. T. dieniensis
b. Plant stouter. Leaf blade various, margin (very) finely or coarsely dentate . . . . 9
9a. Petiole hairy, at least in apical part near the base of the blade . . . 18. T. hastata
b. Petiole glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Plant from montane area, 1700 – 2500 m altitude. Blade base cordate . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. T. densiflora
b. Plant from lowland or lower montane area, up to 1500 m altitude. Blade base
truncate or subhastate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11a. Margin of leaf blade dentate, usually with some prickly teeth towards the base
only. Blade base truncate or with shallow sinus . . . . . . . . . . . 38. T. schlechteri
b. Margin of leaf blade dentate with minute teeth only. Blade base with deep sinus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. T. pulleana
12a. Leaf blade entire. Young twig apices minutely sparsely hairy, early glabrescent.
Seeds not truncate or but faintly notched at apex, cuneate at base. — Papua New
Guinea (Woodlark Is., Bougainville Is.), and western Pacific . . . . 9. T. dentifera
b. Leaf blade entire or (deeply) lobed. Young twig apices and male inflorescences
hairy. Seeds truncate or notched at apex. — Whole of New Guinea . . . . . . . . . 13
13a. Leaf blade often scabrous above, short hairy or glabrescent beneath . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. T. laeoica
b. Leaf blade glabrous, not scabrous, above, glabrous beneath (densely brown hairy
in T. edulis var. septemloba) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. T. edulis
14a. Inflorescences (and flowers) densely hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
b. Inflorescences (sub)glabrous (glabrescent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15a. Hairs of inflorescences and flowers woolly, ± curly, dark brown, more than 2 mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. T. edulis
b. Hairs of inflorescences and flowers ± straight, light brown, 1– 2 mm long . . . 16
16a. Leaf blade (narrowly) ovate or subhastate, unlobed, (sub)glabrous, not scabrous
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. T. pulleana
b. Leaf blade ovate, unlobed or 3-lobed, hairy or glabrescent, scabrous above . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. T. laeoica
17a. Plant delicate. Leaf blade membranous, ovate, margin entire. Bracts about 10 mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. T. dieniensis
b. Plant stouter. Leaf blade chartaceous or coriaceous, ± hastate, margin usually ±
dentate. Bracts 10 mm long or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Rachis 3 – 5 mm thick. Bracts 30 – 45 mm long. — Montane at 1500 – 2000 m . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. T. densiflora
b. Rachis c. 2 mm thick. Bracts smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19a. Leaf blade (sub)coriaceous, margin usually with some prickles towards the base.
Petiole glabrous. Leafy stem and rachis ± zigzag. Bracts 8 –13 mm long . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. T. schlechteri
b. Leaf blade chartaceous, margin remotely (minutely) dentate. Petiole hairy or late
glabrescent. Leafy stem and rachis straight. Bracts 8 – 20 mm long . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. T. hastata
2 cm
2 mm
2 cm g'
3 mm
2 mm
d e f
Fig. 69. Trichosanthes adhaerens W.J.de Wilde & Duyfjes. a. Male flowering leafy twig. b, c. leaves;
d. detail of lower leaf blade surface; e. male flower; f. ditto, opened, showing disc elements and sta-
mens; g. stamen, 2-thecous; g'. stamen, 1-thecous (all: Lim, Postar & Markus SAN 143273, type).
Note on section Edulis Rugayah

This section consists of 8 species mainly in New Guinea, of which 6 species seem to
be mutually more closely related. The species are inadequately defined: T. densiflora,
T. dieniensis, T. hastata, T. laeoica, T. pulleana (also in Moluccas), and T. schlechteri,
largely due to lack of complete material. Especially the species circumscriptions of
T. hastata and T. schlechteri are unclear. Both species were described by Harms (in Bot.
Jahrb. Syst. 60 (1925) 159, 160), largely based on material collected by Schlechter, lost
in Berlin. The present description of both species is composed from extant material and
differs in some characters from the descriptions by Harms, notably the length of the
bracts (in T. schlechteri according to Harms 20 – 30 mm long, in the present treatment
8 –13 mm long) and the margin of the male sepals (in T. hastata according to Harms
entire, in the present treatment entire or incised, or usually slenderly lobed or dentate).
Of recent years only few new collections are known, but more material is needed to
explain the discrepancies. Moreover, the available fruits of all species in the group are
too scanty to frame a key based on fruit and seed characters; instead, for fruiting speci-
mens, a provisional key mainly based on vegetative characters is presented.
1. Trichosanthes adhaerens W.J.de Wilde & Duyfjes

Trichosanthes adhaerens W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 11, f. 1. — Type: Lim, Postar
& Markus SAN 143273 (holo SAN; iso L), Borneo (Sabah, Sipitang, Sg Melabid).
Climber, 4 – 6 m long, clinging to tree trunk with adhesive pads on the tips of short
tendrils, plant green-white harshly hairy, hairs 1– 2 mm long, leafy stem 2 – 4 mm diam.;
dioecious. Probract absent. Tendrils hairy, 5- or 6-branched. Leaves: petiole c. 0.5 cm
long; blade green on drying, coriaceous, simple, shallowly or deeply 3(– 5)-lobed, in
outline broadly ovate or circular, 4 –12 by 4 – 8 cm, scabrous, coarsely rough hairy, hairs
c. 2 mm long, with glands scattered above and beneath, c. 0.5 mm diam., cystoliths ab-
sent, margin coarsely lobulate-dentate to c. 1 cm deep, apex acuminate-mucronate, 2 – 4
mm long mucronate; veins distinctly raised beneath. Male raceme erect; peduncle 1.5 – 2
cm long, 1–1.5 mm thick; rachis not thickened, 4 – 8 cm long, 8 –10(–15)-flowered;
bracts to 2 mm shifted upwards on the pedicel, caducous, (narrowly) elliptic, 1– 2 mm
long, entire, glands absent. Male flowers: pedicel 10 –12 mm long, persistent, thickened
and somewhat longer, c. 15 mm, when perianth is shed; receptacle-tube c. 30 mm long,
faintly curved, slightly constricted in the middle, at throat c. 8 mm wide; sepals linear,
± recurved, c. 5 mm long, margin entire; petals ± elliptic, c. 15 mm long, threads 10(–15)
mm long; synandrium c. 4 by 3 mm, anthers connivent (not fused), filaments free, slen-
der, glabrous, c. 4 mm long; disc consisting of 3 firm-carnose elements, c. 15 by 2.5 mm,
partly connate with the tube. Female flowers and fruit not known. — Fig. 69.
Distribution — Malesia: Borneo (Sabah; known only from the type collection).
Habitat & Ecology — Open disturbed area in lower montane fagaceous forest; flow-
ering April.
Notes — 1. This species is close to T. beccariana, especially to subsp. pusilla, but
the latter differs in leaves with shorter hairs, smaller male flowers with a c. 15 mm long
receptacle-tube.
a'
2 cm
c g
a h
3 mm
2 mm
2 cm
f
3 mm b'
2 mm
b
d e
Fig. 70. Trichosanthes auriculata Rugayah. a, a'. Male flowering leafy twig; b, b'. base of leaf blade
showing auricles, note glands on lower surface; c. male bract; d. male flower bud; e. ditto, opened;
f. infructescence; g. fruit; h. seed (a – e: Church et al. 702; f – h: Chai S 36193, type).
2. Deeply divided leaves more or les appressed to a substrate occur commonly in ju-
venile stages of several Trichosanthes species, e.g. T. tricuspidata. The growth habit of
T. adhaerens is possibly typical and unique for the species, and could be a neotenic trait.
2. Trichosanthes auriculata Rugayah

Trichosanthes auriculata Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 3 (1998) 216; Rugayah
& W.J.de Wilde, Reinwardtia 11, 4 (1999) 251; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia,
thesis (1999) 159, pl. 21; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 14, f. 2. — Type: Chai S
36193 (holo L; iso K, KL, S), Borneo (Sarawak, Sg Apa).
Climber, sometimes creeping and rooting at the nodes, 15 m long, at first with minute
hairs, glabrescent, leafy stem 3 – 5 mm diam.; dioecious or monoecious. Probract ab-
sent. Tendrils 2-branched. Leaves: petiole (3 –)4.5 – 8 cm long; blade green on drying,
membranous, or chartaceous, simple, unlobed, in outline (narrowly) ovate, (5.5 –)10 – 26
by 9 –19.5 cm, scabrous above, glands several, scattered, 0.5 –1 mm diam., base with
2 concave auricles c. 5 mm diam. laterally at apex of petiole, each with 2 –7 glands,
margin entire or shallowly sinuate towards the base, with few small teeth c. 0.5 mm
long, apex sometimes rounded. Male raceme shortly brown hairy, partly glabrescent;
peduncle 1– 5 cm long, 2 – 3 mm thick; rachis not thickened, 10 –14 cm long, 15 – 35-
flowered; bracts subpersistent, to halfway shifted upwards on the pedicel, lanceolate,
10 – 20 mm long, entire, glands few, small. Male flowers: pedicel 10 – 20 mm long,
persistent or caducous; receptacle-tube 5-ribbed, lower part tapered to the base, 18 – 27
mm long, at throat c. 5 mm wide; sepals linear, 4 –7 mm long, c. 1 mm wide at base,
entire; petals in bud forming an 8 –12 mm long elongate body, petals at anthesis obovate,
15 – 22 mm long, threads 7–10 mm long; synandrium 6 –7 mm long, anthers fused in the
upper part only, filaments glabrous, free, slender, 1–1.5 mm long, inserted at c. 1/3 of
the tube below the throat, each continuing into a rib ending in a low 3-lobed ring in the
basal part of the tube; disc absent. Female flowers: in a raceme (always?); pedicel c. 20
mm long; ovary ellipsoid, c. 8 mm long; perianth incompletely known. Fruit 1– 4 per
infructescence, ripening green with paler or yellowish streaks, broad-ellipsoid, c. 10 by
7 cm, apex c. 6 mm beaked; dry pericarp leathery, c. 6 mm thick, smooth; pulp white,
finely fibrous; fruiting pedicel c. 2.5 cm long, 7 mm thick. Seeds brown, compressed,
subcircular, 8 –10 by c. 1 mm, margin broad, radiately ribbed, entire. — Fig. 70.
Distribution — Malesia: Borneo (Sarawak; Sabah; West, Central, East Kalimantan).
Habitat & Ecology — Forest edges and recently disturbed places; to 600 m altitude.
Flowering & fruiting throughout the year.
Notes — 1. The limited material available suggests that this species may be (partly)
monoecious. There is one collection of a twig with male racemes and a detached fruit,
and one other collection with stout racemes with two fruits and numerous persistent
pedicels possibly of male flowers. The elongate shape of the folded petals in male
buds and the habit of creeping and rooting at the nodes are reminiscent of the genus
Gymnopetalum.
2. The present species occupies an isolated position within the genus Trichosanthes,
particularly by its seeds with radiately ribbed margin, and the gland-bearing auricles at
the blade base; the latter characters are also found in T. postarii.
3. Trichosanthes beccariana Cogn.

Trichosanthes beccariana Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 380; Rugayah & W.J.de
Wilde, Reinwardtia 11, 4 (1999) 251; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis
(1999) 77; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 15. — Type: Beccari 802 bis (lecto FI,
designated by Rugayah (1999); iso K, MEL), West Sumatra, Padang Province, Ajer Mantjoer.
Climber to 12 m long, grey-brown short-hairy, mostly glabrescent, leafy stem

1.5 – 3 mm diam.; dioecious or monoecious. Probract absent. Tendrils unbranched
or 2- or 3-branched. Leaves: petiole 1– 3 cm long; blade green on drying, membra-
nous, thinly chartaceous or (sub)coriaceous, simple, unlobed or 3 – 5-lobed to 1/2
deep, in outline ovate-oblong, 4 – 21 by 2.5 – 9 cm, not or faintly scabrous above
and beneath, glands few to many, scattered, 0.5 mm diam., cystoliths sometimes on
veins beneath, margin entire or irregularly coarsely sinuate, rarely ± dentate, apex
rarely obtuse, often with a short mucro; veins 3 – 5, curved, veinlets raised beneath.
Male raceme grey-rusty pubescent, sometimes mixed with a few female flow-
ers towards the base; peduncle 0.5 – 3 cm long, c. 1 mm thick; rachis not thickened,
3.5 –12.5 cm long, 3 – 25-flowered; bracts subpersistent or caducous, on the rachis
or to 5 mm shifted upwards on the pedicel, elliptic, 2 – 3 by 0.5 – 2 mm, entire, with
minute glands. Male flowers: pedicel 3 –10 mm long, persistent; receptacle-tube 15 –
20 mm long, at throat 3 – 4 mm wide; sepals narrowly triangular or linear, c. 2 mm
long, 1– 2 mm wide at base, entire; petals narrowly elliptic, c. 10 mm long, threads
c. 10 mm long; synandrium c. 3 mm long, filaments glabrous, 1.5 mm long. Female
flowers: pedicel 10 – 20 mm long (when solitary at the node), 5 –10 mm long (when
mixed in male raceme); ovary hairy, oblong-fusiform, c. 10 mm long, stigma 4- or 5-
lobed (always?). Fruit ripening orange-red, paler striped, ovoid-ellipsoid or (narrowly)
ellipoid, 2 –10 by 2 – 6.5 cm, apex subacute and shortly beaked; exocarp leathery or
papery, 0.5 –1 mm thick; pericarp ± carnose, when dry 5 mm thick; pulp orange-red,
conspicuously fibrous; fruiting pedicel 2 – 3 cm long (at the nodes), 0.5 – 2 cm long (in
the raceme), c. 2 mm thick. Seeds dark brown, tumid, with two lateral inflated parts,
6 –10 by 7–12 by 2 – 3 mm, edge entire.
Distribution — Malesia: West Central Sumatra and all over Borneo; 2 subspecies.
1a. Leaves glabrous above (except the 1 mm long hairy midrib); fruit 6 –10 by 3.5 – 6.5
cm; seeds 8 –10 by 10 –12 mm . . . . . . . . . . . . . . . . . . . . . . . a. subsp. beccariana
b. Leaves usually short soft-hairy above; fruit 2 – 4.5(– 6) by 2 – 3 cm; seeds 6 –7 by
7–10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. pusilla
a. subsp. beccariana
Climber to 12 m long. Leaves: blade chartaceous, unlobed, in outline ovate-elliptic,
7–19 by 4 –12 cm, glabrous above (except hairy midrib), glabrescent beneath. Male
raceme 3 – 9 cm long; peduncle 1.5 – 2.5 cm long. Fruit ovoid or narrowly ellipsoid,
6 –10 by 3.5 – 6.5 cm; exocarp leathery. Seeds 8 –10 by 10 –12 mm. — Plate 25a – c.
3 mm
2 cm
3 mm
a j i c
2 cm
3 mm
2 mm
g e h' f h
Fig. 71. Trichosanthes beccariana Cogn. subsp pusilla Rugayah. a. Juvenile leafy twig, note adhesive
discs at apex of tendrils; b. male flowering leafy twig; c. leaf; d. part of male inflorescence; e. male
flower bud; f. male flower opened showing disc elements and androecium; g. petal of male flower;
h. stamen, 2-thecous; h'. stamen, 1-thecous; i. fruit; j. seed (a: De Wilde, Postar & Ubaldus SAN 144010;
b – h': De Wilde, Postar & Ubaldus SAN 144006; i, j: Fedilis Krispinus SAN 95877).
Distribution — West Central Sumatra (where only known from the type); North
Borneo (Kalimantan and Sabah).
Habitat & Ecology — Primary forest edges, lower montane forest, roadsides; 300 –
1500 m altitude.
b. subsp. pusilla Rugayah

Trichosanthes beccariana Cogn. subsp. pusilla Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11,
3 (1998) 217; 11, 4 (1999) 251; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999)
79. — Type: Mogea & de Wilde 4372 (holo BO; iso K, L), Central Kalimantan.
Climber to 15 m long. Leaves: blade membranous, unlobed or 3(– 5)-lobed, or margin

deeply sinuate, in outline (narrowly) ovate, 4 –11 by 1.5 – 6 cm, usually densely soft-
hairy above. Male raceme: wholly grey soft-hairy, 1.5 – 5 cm long; peduncle 0.5 – 3 cm
long. Fruit ovoid or ellipsoid, 2 – 4.5(– 6) cm long, 2 – 3 cm wide; exocarp papery. Seeds
6 –7 by 7–10 mm. — Fig. 71.
Distribution — Malesia: Borneo (Sarawak, Sabah, and Central & East Kalimantan).
Habitat & Ecology — Primary forest, hill forest, forest edges, logged over areas,
open areas in newly disturbed forest, roadside scrub, along riversides, on yellow clay
soil; from sea level to 350 m altitude. Flowering and fruiting throughout the year.
4. Trichosanthes borneensis Cogn.

Trichosanthes borneensis Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 369; Rugayah & W.J.de
(1999) 135; W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu (2001) 210; Sandakania
14 (2004) 17. — Type: Korthals s.n. (‘54’) (holo L, barcode L0130152), Borneo (Kalimantan, Mt
Sakoembang).
Trichosanthes sumatrana Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 373, excluding var.
obtusiloba. — Type: Beccari s.n. (lecto FI, acq. no. FI 4427, designated by De Wilde & Rugayah
(Reinwardtia 11 (1999)), West Sumatra, Padang Province, Ajer Mantjoer.
Trichosanthes wallichiana auct. non (Ser.) Wight: Ridl., Fl. Malay Penin. I (1922) 845.
Climber, 6(– 20) m long, finely brown hairy, early glabrescent, leafy stem 2 – 5 mm
diam.; dioecious. Probract narrowly elliptic, 5 –18 by 1– 2 mm, glands absent. Tendrils
2- or 3-branched. Leaves: petiole 2.5 – 8 cm long; blade brown or green on drying,
membranous, simple, deeply 3 – 5(–7)-lobed, (1/2 –)9/10 deep, in outline broadly ovate
or circular, 7– 23 by 7.5 – 23 cm, scabrous above, usually smooth beneath, glands few
(sometimes absent), situated at the base near the insertion of the petiole, 1– 3 mm diam.,
cystoliths obvious, margin entire or towards the base of the outer lobes coarsely dentate
or dentate-sinuate; veins 3 – 5(–7), straight. Male raceme occasionally with a single
male flower co-axillary at the node, finely pubescent; peduncle 2 –11(–13) cm long, 3 – 5
mm thick; rachis somewhat thickened, 6 – 30 cm long, 3 – 5(–10) mm thick, including
thickened bract-scars, 10 – 20(– 50)-flowered; bracts late-caducous, narrowly or broadly
ovate, or rhomboid, (10 –)20 – 50 by 6 – 30 mm, margin irregularly sharply incised or
dentate to 5 mm deep, glands absent. Male flowers: pedicel 3 – 5 mm long (c. 25 mm
long in solitary flowers), caducous; receptacle-tube 40 –70 mm long, at throat 5 – 9 mm
wide; sepals narrowly triangular or lanceolate, 6 –16 mm long, 1.5 – 3 mm wide at base,
Fig. 72. Trichosanthes borneensis Cogn. a. Node with male inflo-

rescence; b, node with fruit; c. seed (a: Sidiyassa 2652; b: Church
et al. 782; c: Haegens et al. 382).
2 cm
2 mm
b c
entire or minutely narrowly lobed; petals obovate or rhomboid, c. 15 mm long, threads

7–11 mm long; synandrium 7–11 mm long, anthers fused, filaments glabrous, 3 – 4 mm
long. Female flowers not known. Fruit ripening orange-red, paler speckled or faintly
striped, (depressed-)globose, 3.5 –10 by 5 – 9 cm, apex obtuse (not beaked); exocarp
woody, c. 1 mm thick, smooth; pericarp juicy, c. 10 mm thick (dry pericarp ± vanished);

pulp green-black, bitter; fruiting pedicel (2.5 –)3.5 – 8 cm long, 5 –15 mm thick, 2-col-
oured, the distal part orange-red and smooth, (0.5 –)1(– 2) cm long, the remaining part
green and grooved. Seeds black, compressed, (narrowly) elliptic, 13 – 20 by 5 –7 by 3
mm, base truncate or ± obtuse, apex obtuse, margin faint, edge entire. — Fig. 72.
Field-notes — Flowers white with highly divided petals. Fruits yellowish green to
yellow or bright orange to red when ripe, fruit stalk thick, orange for the distal 1 cm,
green below, pulp green, with cucumber smell, very bitter and foetid.
Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo (Kali-
mantan, Sarawak, Sabah).
Habitat & Ecology — Open places and edges of primary and degraded forest, on
sandy clay; lowland to 900 m altitude.
Uses — Used as a medicine (Gimlette s.n., Peninsular Malaysia); the tuberous root
is incorporated in dart poison (Gadoh anak Umbai KL 1771).
Note — The distal part of the fruiting pedicel apparently belongs morphologically
to the fruit.
5. Trichosanthes celebica Cogn.

Trichosanthes celebica Cogn. in A.DC. & C.DC. Mongraph. Phan. 3 (1881) 385; Rugayah & W.J.de
(1999) 99. — Type: Beccari 51 (holo FI), SE Sulawesi, near Kendari, Lepo-Lepo.
Climber to 10 m long, early glabrescent, at first with sparse minute hairs, leafy stem
1.5 – 3 mm diam.; dioecious. Probract ovate, ± flat, 3 – 5 by 3 mm, glands present.
Tendrils (unbranched or) 2-branched. Leaves: petiole 2 – 5 cm long; petiolules 0.3 – 0.7
cm long; blade green on drying, subchartaceous or membranous, simple and unlobed
or 3-foliolate; unlobed blade in outline ovate-oblong or hastate, 10 – 20 by 6 –10 cm,
foliolate blade in outline circular, middle leaflet 10 –12 by 3 – 3.5 cm, base cuneate,
faintly scabrous above, glands several, scattered, 0.5(–1) mm diam., cystoliths obvious,
margin entire or sparsely minutely dentate; unlobed blades with 3(– 5) curved veins,
leaflets pinniveined. Male raceme minutely rust-hairy; peduncle 2.5 – 4 cm long, c. 3
mm thick; rachis somewhat thickened, with bract-scars, 7–11 cm long, 8 –15-flowered;
bracts subpersistent, narrowly elliptic, 15 – 25 mm long, margin deeply incised to c. 1/3
deep, with glands. Male flowers (from buds): pedicel 2 – 3 mm long; sepals narrowly
triangular, 5 –7 mm long, entire. Female flowers not known. Fruit ripening red (possibly
not paler striped), ovoid, 6.5 by 4.5 cm, apex c. 2 mm beaked; exocarp thick-leathery,
smooth; dry pericarp c. 5 mm thick; pulp greenish black; fruiting pedicel 2–7 cm long,
3– 4 mm thick. Seeds blackish, compressed, elliptic, 12–13 by 4 – 7 by 2 – 3 mm, margin
absent, edge entire.
Distribution — Malesia: Sulawesi (Kendari and Lake Matano area), Moluccas (NW
Buru).
Habitat & Ecology — Open areas in secondary forest or disturbed primary forest,
river sides, on limestone and clayey soil; 50 – 400 m altitude.
Notes — 1. The fruits of T. celebica resemble those of T. wawrae, but in the latter
species the exocarp is thinner and the seed larger, 15 – 20 by 8 –15 mm.
2. The few specimens on which the present description of T. celebica is based are
variable, e.g. in the leaf blade glands, fruiting pedicel, and seeds, so possibly a second
species may be involved; a redescription is in preparation..
6. Trichosanthes coriacea Blume

Trichosanthes coriaceae Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 935; Miq., Fl. Ned. Ind. 1, 1 (1856)
674; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 355; Backer in Backer & Bakh.f., Fl. Java
1 (1964) 303; Rugayah & W.J.de Wilde, Blumea 42, 2 (1997) 477; Reinwardtia 11, 4 (1999) 252;
Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 162, pl. 22. — Type: Blume s.n.
(holo L, barcode L0130310; iso L, barcode L0130311), West Java, Gunung Salak.
Climber to 10 m long, short brown hairy, partly glabrescent, leafy stem 2 – 4 mm

diam.; dioecious (or monoecious?). Probract obovate, 3 by 2 mm, glands absent. Ten
drils 2-branched (the second branch reduced). Leaves: petiole 1.5 – 3 cm long; blade
brown on drying, (membranous-)coriaceous, simple, unlobed, in outline (narrowly) or
broadly ovate, (8 –)12 – 23 by (6.5 –)10 –18 cm, finely bullate and somewhat shiny above,
glands few, 0.5 –1 mm diam., cystoliths not obvious, base shallowly cordate-truncate,
margin entire, somewhat recurved; veins (3 –)5 from the base, curved, and few smaller
veins from the midrib, ± sunken above. Male raceme densely short-hairy; peduncle
(1.5 –)3 – 4 cm long; rachis not thickened, 6 – 21 cm long, 3 – 4 mm thick, (10 –)20 – 35-
flowered, with conspicuously thickened persistent pedicels; bracts caducous or subper-
sistent, somewhat shifted upwards the pedicel, (ob)ovate, 5 –7 by c. 3 mm, obtuse, entire,
with glands. Male flowers (from buds): pedicel c. 2 mm long, persistent (after the flowers
are shed becoming thicker and longer, giving the raceme a branched aspect); sepals nar-
rowly triangular, c. 3 mm long, entire. Female flowers not known. Fruit ripening evenly
orange-red, ovoid-ellipsoid, c. 10 by 7 cm, apex shortly beaked; dry pericarp possibly
c. 10 mm thick; exocarp thick-leathery, smooth; pulp not known; fruiting pedicel c. 1.5
cm long, 5 mm thick. Seeds brown, compressed, obovate, 13 –15 by 8 – 9 by 1–2 mm,
base cuneate, apex broadly rounded, margin c. 1.5 mm broad, edge entire.
Distribution — Malesia: Sumatra, Java (where known only from the type collection).
Habitat & Ecology — Primary hill forest; 400 m altitude.
Note — Trichosanthes coriacea is a rare species. Only 8 collections are known, of
which 7 were made in the 19th century; Laumonier TFB 2170 is the only recent one,
collected in 1983, in Bukit Sebelah Nature Reserve, Sumatra.
7. Trichosanthes cucumerina L.
Trichosanthes cucumerina L., Sp. Pl. 2 (1753) 1008; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge,
Laos & Vietnam 15 (1975) 91; Rugayah & W.J.de Wilde, Blumea 42 (1997) 478; Reinwardtia 11,
4 (1999) 252; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 66; Duyfjes &
Pruesapan, Thai Forest Bull, Bot. 32 (2004) 82; W.J.de Wilde & Duyfjes, Fl. Thailand, 9, 4 (2008)
516. — Type: “Padavalam” in Rheede, Hort. Malab. 8 (1688) 39, t. 15 (lecto, designated by Ker-
audren (1975)), habitat in India.
Trichosanthes reniformis Miq., Fl. Ned. Ind. 1, 1 (1856) 675. — Type: Horsfield s.n. Java (holo BM;
iso U, barcode U0008346), Java.
Trichosanthes pedatifolia Miq., Fl. Ind. Bat. 1, 1 (1856) 677. — Type: Horsfield s.n. (holo BM; iso
U, barcode U0008345), Java.
Climber, 5 – 8 m long, with sparse (dense) minute hairs, partly glabrescent, leafy
stem 1.5 – 2(– 5) mm diam.; monoecious. Probract absent. Tendrils 2- (or 3-)branched.
Leaves: petiole 2 –12 cm long; blade green on drying, membranous, simple, unlobed, or
shallowly or deeply 3 –7-angular or -lobed, 2/3 deep, in outline subcircular, or broad-
ly reniform, 5 –12(– 20) by 9 –12(– 25) cm, finely hairy, sometimes faintly scabrous,
without or with few scattered minute glands, cystoliths not obvious, margin entire or
sparsely shallowly dentate-undulate; veins 3(– 5). Male raceme sometimes with co-ax-
illary a solitary male flower or a solitary female flower (the female flower developing
before the male raceme), hairy or glabrescent; peduncle 5 –15 cm long, 1(– 2) mm thick;
rachis not thickened, 3 –10 cm long, 5 –10 (or more)-flowered; bracts subpersistent or
caducous, elliptic, 0.5 – 2 mm long, margin (sub)entire, glands absent. Male flowers:
pedicel 5 – 20 mm long, persistent; receptacle-tube 15 – 20 mm long, at throat 3 – 4(– 5)
mm wide; sepals linear, 2 – 3 mm long, c. 1 mm wide at base, entire; petals (narrowly)
ovate, 6 –10 mm long, threads c. 10 mm long; synandrium 2 – 3 mm long, filaments
short. Female flowers: pedicel 5 –12 mm long or longer; ovary hairy, (narrowly) el-
lipsoid, 3 –10(– 30) mm long. Fruit ripening bright orange, paler speckled or striped,
ellipsoid or long-cylindrical, 2.5 – 5(–150) by 1.5 – 4 cm, apex beaked; exocarp (thinly)
leathery, smooth; pericarp ± juicy, dry pericarp 3 – 5 mm thick; pulp orange, not fibrous,
slightly bitter; fruiting pedicel 1– 2 cm long, 2 mm thick. Seeds pale or dark brown,
compressed, (narrowly) elliptic, 6 –18 by 4 – 9 by 2.5 – 3.5 mm, margin broad, distinct
or faint, edge undulate.
Distribution — Widely distributed from India, Sri Lanka, South China, and Thailand,
through Malesia into West, North and NE Australia; in Malesia: Java, Philippines,
Sulawesi, Lesser Sunda Islands; 2 varieties.
Note — The two varieties flower during daytime.
1a. Plant delicate, growing wild. Leaves 5 –14 cm diameter. Fruit (2.5 –)4 – 6 cm long,
containing to 10 seeds; seeds 6 – 8(–10) mm long . . . . . . . . . . a. var. cucumerina
b. Plant more robust in all parts, cultivated. Leaves to 25 cm diameter. Fruit much
elongated, snake-like, 35 –100(–150) cm long, containing many seeds; seeds 14 –18
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. anguina
a. var. cucumerina
Climber, annual, growing wild; leafy stem delicate, 1– 2 mm diam., with or without
scattered pale coarse hairs, 1 mm long. Leaves: petiole 2 – 6 cm long; blade 5 –14 cm
diameter. Fruit (2.5 –)4 – 6 cm long, containing few (to 10) seeds; pulp bitter (always?).
Seeds 6 – 8(–10) mm long. — Fig. 91a.
Distribution — The variety cucumerina is the wild variety and is widely distributed
from India through Malesia into West, North and NE Australia; in Malesia: most collec-
tions from Java and Madura, one from Philippines (Luzon), one from Sulawesi (Maros),
few from Lesser Sunda Islands (Flores, Sumbawa, and Sumba).
Habitat & Ecology — Forest edges, scrub, disturbed areas; apparently solely in areas
with a seasonal climate; sea level to 1200 m altitude. Flowering and fruiting in and after
the wet season.
b. var. anguina (L.) Haines

Trichosanthes cucumerina L. var. anguina (L.) Haines, Bot. Bihar Orissa (1922) 388; Rugayah &
W.J.de Wilde, Blumea 42, 2 (1997) 478; W.J.de Wilde & Duyfjes, Sandakania 17 (2008 ‘2007’)
79. — Trichosanthes anguina L., Sp. Pl. 2 (1753) 1008; Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 933;
A.DC. & C.DC., Mon. Phan. Prodr. 3 (1881) 359; Backer in Backer & Bakh.f., Fl. Java 1 (1963)
304. — Type: “Anguina Sinensis, flore albo, elegantissimo, capillamentis tenuissimis ornato, fructu
longo intorto, sub initium ex albo, & viridi variegato, per maturitatem prorsus rubro” in Micheli,
Nov. Pl. Gen., 12, t. 9, 1729 (lecto, designated by Jeffrey in Jarvis & al. (ed.), Regnum Veg. 127
(1993) 95), habitat in China.
Climber, subperennial, cultivated, leafy stem 2 – 5 mm diameter. Leaves: petiole 2 –12

cm long; blade to 25 cm diameter. Fruit snake-like, 30 –100(–150) cm long, containing
to 50 seeds; pulp rather sweet. Seeds 14 –18 mm long.
Distribution — Widespread in cultivation; immature fruit used as vegetable.
8. Trichosanthes densiflora Rugayah

Trichosanthes densiflora Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 252; Ruga
yah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 180. pl. 25. — Type: Von Roemer 706
(holo BO), south-western West Papua.
Trichosanthes bracteata auct. non Voigt: Pulle, Nova Guinea 8 (1912) 689; Harms, Bot. Jahrb. Syst.
60 (1925) 160, p.p.
Climber to 6 m long, at first with short hairs, glabrescent, leafy stem 2 – 4 mm diam.;
dioecious. Probract ± ovate, 5 mm long, coarsely dentate, glands absent. Tendrils 2-
branched. Leaves: petiole 2.5 – 6 cm long; blade green on drying, subcoriaceous, simple,
unlobed, in outline (narrowly) ovate, 9 –12 by 6.5 –10.5 cm, glands several, at the base
towards the insertion of the petiole, 0.5 mm diam., cystoliths present, base cordate, mar-
gin shallowly spiny dentate; veins 5, curved. Male raceme glabrescent; peduncle 3 – 6
cm long, 2(– 3) mm thick; rachis stout, thickened, with thickened bract-scars, 11–13
cm long, 3 – 5 mm thick, to 30-flowered; bracts distinctly veined, (narrowly) obovate,
30 – 45 by 15 – 20 mm, margin entire, sometimes shallowly lobed, glands scattered in
upper half, 0.5 –1 mm diameter. Male flowers: pedicel 2 mm long, caducous; recepta-
cle-tube c. 27 mm long, at throat 6 – 8 mm wide; sepals, petals and stamens not known.
Female flowers not known. Fruit ripening green, (narrowly) ellipsoid, 6 – 8 by c. 4.5 cm,
apex c. 5 mm beaked; exocarp papery; pericarp possibly carnose, drying thin and firm,
irregularly ornamented (see note 2), pulp possibly creamy, not red; fruiting pedicel c.
3 cm long, 3 mm thick, finely brown hairy. Seeds brown, densely packed, compressed,
(narrowly) elliptic, 10 –13 by (4 –)5 – 6 by c. 1 mm, apex shallowly notched, margin
faint, edge entire.
Distribution — Malesia: New Guinea (south-western West Papua (Lorenz River);
Papua New Guinea (Western Highlands and West Sepik Provinces)).
Habitat & Ecology — Secondary growth and regrowth; 1500(?)– 2400 m altitude.
Notes — 1. Trichosanthes densiflora is known only from three collections, identical

on vegetative characters, although found wide apart. A fourth collection, Eyma 5123,
from Wissel lake region, West Papua, has immature female flowers with a coarse woolly
rufous indumentum. It seems most related to T. densiflora but may represent a separate
species.
2. In Henty NGF 41616, the dried fruit has a narrowed base, apparently caused by
the drying of the thick juicy-carnose pericarp, becoming thin when dry; the irregularly
ornamented surface indicates the presence of the seeds underneath.
9. Trichosanthes dentifera Rugayah

Trichosanthes dentifera Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 253; Rugayah,
Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 182. — Type: McGregor s.n., anno 1890
(holo MEL), Papua New Guinea.
Tall climber, with sparse minute grey hairs, early glabrescent, leafy stem 3 – 4 mm
diam.; dioecious. Probract ovate or elliptic, 3 – 4 by c. 2 mm, without glands. Tendrils
2- or 3-branched. Leaves: petiole 2 – 4 cm long; blade greenish brown on drying, charta-
ceous, simple, unlobed, in outline broadly ovate, 8 –15 by 8 –12 cm, glands several to
many, at the blade base or scattered, 0.5 mm diam., cystoliths occasionally on old stem,
petiole and veins, margin entire or sparsely minutely dentate; veins 5(–7), arching. Male
raceme and male flowers not known. Female flowers (from bud): sparsely brown hairy;
pedicel c. 15 mm long; ovary narrowly ellipsoid, with c. 10 ribs; sepals linear, c. 12 mm
long, 0.5 mm wide at base, margin entire; mature petals not seen (see note). Fruit ripen-
ing red, ellipsoid or (sub)globose, 10 –12 by 5.5 –7 cm, apex c. 2 mm beaked; exocarp
woody, c. 1 mm thick, smooth with c. 10 shallow grooves; fruiting pedicel 2 – 2.5 cm
long, c. 5 mm thick. Seeds brown, densely packed, compressed, ± narrowly elliptic, ±
parallel-sided, 12 –13 by 5 by 2 mm, base cuneate, apex ± notched, margin faint, edge
(sub)entire.
Field-notes — Mature female flowers 6 –7 cm long, corolla 6 –7 cm diam.; petals
fimbriate, dirty white. Fruits round, red when ripe.
Distribution — Vanuatu; in Malesia: New Guinea (Papua New Guinea (Woodlark
Is., Bougainville Is.)).
Habitat & Ecology — Rain forest, riverside forest, climbing over the topmost branch-
es of tall forest trees; to 240 m altitude.
Uses — The oily seeds are edible, roasted or cooked.
Notes — 1. The pericarp is possibly juicy and shows on drying 15 –18 vascular
strands.
2. Trichosanthes dentifera seems related to the Australian T. subvelutina F.Muell. ex
Cogn. and T. species A, both treated by Telford (Fl. Austr. 8, 1982), having also grooved
fruits.
10. Trichosanthes dieniensis Merr. & L.M.Perry

Trichosanthes dieniensis Merr. & L.M.Perry, J. Arnold Arbor. 30 (1949) 59; Rugayah & W.J.de Wilde,
Reinwardtia 11, 4 (1999) 254; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999)172,
pl. 23. — Type: Brass 3898 (holo A), Papua New Guinea, Dieni, Ononga Road.
Climber, early glabrescent, leafy stem 1–1.5 mm diam.; dioecious. Probract ab-
sent(?). Tendrils 2-branched. Leaves: petiole 2 – 4.5 cm long; blade green on drying,
membranous, simple, unlobed, in outline ovate, 4.5 –12 by 2.5 – 8 cm, scabrous above,
but at apex minutely hairy, glands few, along the margin of the basal sinus, c. 0.5
mm diam. or less, cystoliths obvious, margin sparsely minutely dentate; veins 3(– 5),
curved. Male raceme glabrous; peduncle 3– 5.5 cm long, c. 1 mm thick; rachis slender,
not thickened, 3.5–6 cm long, 5 –12-flowered; bracts narrowly or broadly (ob)ovate,
5 –13 by 2 –10 mm, margin shallowly lobed or dentate, glands few, c. 0.5 mm diameter.
Male flowers: in the raceme and one solitary at base of the raceme; pedicel in raceme
2 –7 mm long, persistent or caducous, when solitary 40 – 50 mm long, withering into a
straw-like persistent appendage; receptacle-tube c. 40 mm long, at throat c. 6 mm wide;
sepals narrowly triangular or linear, c. 10 mm long, entire or few-dentate at base; petals
obovate-rhomboid, c. 10 mm long, threads 10 – 20 mm long; synandrium c. 6 mm long,
filaments short. Female flowers, fruit and seeds not known.
Distribution — Malesia: New Guinea (Papua New Guinea (Central Province, Dieni,
known only from the type collection).
Habitat & Ecology — Massed on dead tree trunk; 500 m altitude.
Note — Trichosanthes dieniensis is similar to T. hastata, and possibly only a tiny
form of this. The latter differs in having flowers with larger petals (c. 2.5 cm long, ex-
cluding threads), larger male bracts without glands, and rough-hairy petioles (glabrous
in T. dieniensis).
11. Trichosanthes edulis Rugayah

Trichosanthes edulis Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 254, f. 2; Ru-
gayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 184, pl. 26. — Type: Streimann
NGF 30731 (holo L; iso BRI, CANB, LAE), Papua New Guinea, Morobe Province.
Trichosanthes trifida Lu Q.Huang, B.Yang & C.H.Yueh, Act. Phytotax. Sin. 35, 2 (1997) 130, pl. 3:
45– 47, nom. nud. — Voucher specimen: Kalkman 4401, West Papua.
Climber or creeper, sometimes rooting at the nodes, to 10 m long, brown hairy,

late glabrescent, leafy stem 3 – 5 mm diam.; dioecious. Probract sometimes caducous,
(narrowly) elliptic, 5 –15 by 1– 8 mm, apex subacute, with many glands. Tendrils 2 –
4-branched. Leaves: petiole 3 – 9 cm long; blade brown-green or blackish brown on
drying, (thinly) chartaceous-coriaceous, simple, shallowly or deeply 3 – 5-lobed or -
angular, lobes to 1/2 – 5/6 deep, rarely unlobed, in outline broadly ovate or circular,
9 – 25 by 9 – 23 cm, above and beneath glabrous or (densely) brown hairy, glands few
or many, scattered, 0.5 –1 mm diam., or absent, cystoliths sometimes obvious on stem
and leaves, margin entire, or sparsely minutely spiny-dentate or (coarsely) dentate;
veins 3 –7, straight. Male raceme occasionally with a solitary male flower at base, dark
brown woolly-hairy, hairs dark brown, curly, more than 2 mm long, partly glabrescent;
peduncle 4.5 –16(– 20) cm long; rachis not thickened or thickened in var. septemloba,
at base with thickened bract-scars, 5 – 20(– 35) cm long, 2 – 5(–7) mm thick, (3 –)5 – 30-
flowered or more; bracts broadly ovate-rhomboid, 10 – 40 by (5 –)12 – 25 mm, margin
entire or undulate-dentate, with small glands. Male flowers: pedicel 5 –10 mm long, in
solitary flowers much longer, caducous; receptacle-tube 30(– 50) mm long, at throat
7–15 mm wide; sepals narrowly triangular or ovate, acute, 5 –15 mm long, 2 – 5 mm

wide at base, entire or incised; petals obovate- rhomboid, c. 20 mm long, threads c. 20
mm long; synandrium c. 10 mm long, filaments glabrous, 1– 2 mm long. Female flow
ers: pedicel 30 – 50 mm long; ovary (narrowly) ellipsoid, 20 – 35 by 6 – 8 mm, densely
brown hairy; receptacle-tube 30 – 50(– 60) mm long; sepals 5 –10 mm long, 2 – 5 mm
wide at base, incised (or entire); petals as in male; style c. 45 mm long. Fruit ripening
evenly orange or red, narrowly ellipsoid or (long) pyriform, 10 – 25 by 5 –7 cm, apex ±
acute; pericarp carnose, smooth, dry pericarp woody, rough; pulp red; fruiting pedicel
3 –10 cm long, (4 –)5 –10 mm thick. Seeds densely packed, compressed, narrowly el-
liptic and ± parallel-sided or obtriangular (broad at apex), 10 –15 by 4 – 8 by 1– 2 mm,
± truncate or notched at one or both ends, with or without a longitudinal depression in
the middle, margin broad, edge entire or sometimes finely crenulate.
Distribution — Malesia: New Guinea; 3 varieties; also Philippines (see note).
Note — One out-reaching collection from the Philippines, Luzon, PP1 3044, with
immature male flowers belongs to T. edulis as at present defined. It seems to link up
with deviating specimens from Vogelkop as discussed under var. edulis, and its status
needs further study.
1a. Leaves deeply (5 –)7-lobed, densely brown hairy on both surfaces . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c. var. septemloba
b. Leaves 3 – 5-lobed, glabrous or glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Leaves shallowly 3-lobed or 3 – 5-angular, to halfway deep, rarely unlobed, drying
dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. sativa
b. Leaves deeply 3 – 5-lobed, 1/2 – 3/4 deep, drying brown-green . . . . a. var. edulis
a. var. edulis
Plant glabrescent. Probract 5 –10 mm long. Leaves: blade chartaceous-coriaceous,
drying brown-green, 3 – 5 lobed, lobes 1/2 – 3/4 deep, blade subcircular in outline, to
25 cm diam., glabrous or glabrescent above and beneath, margin entire or minutely
spiny-dentate. Male raceme dark brown woolly hairy, partly glabrescent, 20 – 25(– 55)
cm long; peduncle 5 – 20 cm long; bracts broadly obovate-rhomboid, c. 20 by 15 – 25
mm, entire or undulate. Female flowers not known. Fruit long-ellipsoid or pyriform,
15 – 22 by 5 –7 cm; fruiting pedicel 6 – 9 cm long. Seeds brown-black, narrowly oblong,
± parallel-sided, 12 –15 by 4 – 5 by 1– 2 mm, notched on one or both ends, narrowly
lengthwise grooved in the middle, edge entire or sometimes faintly crenulate.
Field-notes — Leaves coriaceous, slightly glossy dark green above, light green be-
neath. Fruit red when ripe; pulp bright red; seeds black. Cooked fruit edible.
Distribution — Malesia: New Guinea (West Papua and Papua New Guinea).
Habitat & Ecology — Secondary growth, degraded forest in logging areas, swampy
places, river side forest; to 1500 m altitude.
Note — Kalkman 4401 (West Papua) differs in smaller male flowers and in thin
chartaceous densely arranged male bracts. Vink 17576, Avé 4076 and Polak 850, all
from low altitude in West Papua, Vogelkop, differ in ± obtriangular seeds, truncate at
apex, resembling those of var. sativa.
b. var. sativa Rugayah

Trichosanthes edulis Rugayah var. sativa Rugayah in Rugayah & W.J.de Wilde, Reinwardtia, 11, 4
(1999) 256; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 188. — Type: Vink
16350 (holo L; iso CAN, LAE), Papua New Guinea, Western Highlands Province.
Plant glabrescent. Probract 5 –10 mm long. Leaves: blade chartaceous, drying dark
brown, shallowly 3-lobed or 3 – 5-angular, to halfway deep, rarely unlobed, broadly
ovate or subcircular in outline, to 20 cm long, glabrous or glabrescent above and be-
neath, margin entire, or finely or coarsely dentate; veins 5. Male raceme to 20 cm long,
dark brown short-woolly hairy; peduncle not seen; bracts narrowly elliptic or rhomboid,
10–20 mm long, irregularly dentate. Female flowers: as the species. Fruit pyriform-
ellipsoid, to c. 25 cm long when fresh; fruiting pedicel c. 4.5 cm long, c. 5 mm thick.
Seeds brown, compressed, narrowly (elliptic) or obtriangular, 10 –12 by 5 – 8 by c. 1
mm, shallowly or deeply notched on one or both ends, edge entire or finely crenulate,
not grooved in the middle.
Field-notes — Fruit apex dark green, base orange, bright red when ripe. Fruit edible,
cooked in ashes of fire.
Distribution — Malesia: New Guinea (montane areas of Central West Papua and
Papua New Guinea (Western Highlands, Eastern Highlands, and Simbu Provinces, and
Bismarck Archipelago)).
Habitat & Ecology — Cultivated in gardens or running wild in secondary forest
edges; 1000 – 2500 m altitude.
Vernacular name — Gin bogl (Kuma language), Kisang.
c. var. septemloba Rugayah

Trichosanthes edulis Rugayah var. septemloba Rugayah in Rugayah & W.J.de Wilde, Reinwardtia
11, 4 (1999) 256; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 189. — Type:
Stevens LAE 54823 (holo L; iso CAN, LAE), Papua New Guinea (Morobe Province).
Plant densely brown hairy. Probract 10 –15 mm long. Leaves: blade thinly charta-
ceous, drying brown-green, 5 –7-lobed, to 3/4 – 5/6 deep, subcircular in outline, to 22 cm
diam., densely brown hairy above and beneath, margin coarsely dentate. Male raceme
(immature) 10 –17 cm long; peduncle to 15 cm long, rachis 3 –7 mm thick, sometimes
with a solitary woolly hairy male flower at base, hairs c. 1 mm long; bracts (narrowly)
elliptic, 20 – 30(– 40) by 15 – 20 mm, entire. Female flowers and fruit not known.
Field-notes — Leaves dark green above, paler beneath, venation very clear. Flowers
large.
Distribution — Malesia: Papua New Guinea (Morobe, Southern Highlands, and Cen-
tral Provinces).
Habitat & Ecology — Open places in rain forest, regrowth brush, in old farm land;
800 –1600 m altitude.
12. Trichosanthes ellipsoidea Merr.

Trichosanthes ellipsoidea Merr., Philipp. J. Sc., C 13 (1918) 332; Rugayah & W.J.de Wilde, Rein-
wardtia 11, 4 (1999) 258; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 113,
pl. 8. — Type: Ramos BS 30364 (holo PNH, lost; iso K, US), Philippines, Catanduanes, slopes of
Mt Mariguidon.
Climber to 5 m long, early glabrescent, at first with minute powdery hairs, leafy stem
1.5 – 3 mm diam.; dioecious. Probract (narrowly) elliptic, c. 5 mm long, entire. Tendrils
unbranched or 2-branched. Leaves: petiole 3 – 4.5 cm long; blade greenish on drying,
membranous or chartaceous, simple, unlobed, in outline (narrowly) ovate, 10 –15 by
6 – 9.5 cm, ± scabrous above, glands few or several, scattered at about the centre or to-
wards the base, 1– 2 mm diam., margin entire but at base faintly repand, apex slenderly
acute-acuminate; veins (3 –)5, curved. Male raceme finely hairy; peduncle 3 – 6 cm long,
c. 2 mm thick; rachis not thickened, c. 10 cm long, c. 2 mm thick, 20 – 25-flowered;
bracts rhomboid or broadly (ob)ovate, 15 – 20 by c. 15 mm, margin deeply incised or
dentate, to 1/3 –1/2 deep, sometimes with 3 main lobes, glands present. Male flowers:
pedicel 1– 3 mm long, persistent or caducous; receptacle-tube (from large bud) some-
what swollen at base, 30 mm long, at throat 7– 8 mm wide; sepals narrowly ovate, 7
mm long, 3 mm wide at base, entire; synandrium c. 7 mm long, filaments sparsely hairy,
c. 3 mm long. Female flowers: pedicel 15 – 20(– 30) mm long; receptacle-tube 60 –70
mm long; sepals lanceolate, 6 –7 mm long; petals obovate, including threads 30 mm
long. Fruit: ripening colour not known, ellipsoid, c. 5 cm long; exocarp thick-leathery,
smooth, dark brown when dry; pericarp and pulp not known; fruiting pedicel 1.5 – 2
cm long, 2–3 mm thick. Seeds compressed, c. 13 by 6–7 by 1.5–2 mm, base narrowly
truncate, apex pointed, not margined, edge entire.
Field-notes — The two collections known from Sarawak (Ashton S 17757 and Chai S
34116) are annotated as having dark green leaves, variegated with pale silvery patches.
Distribution — Malesia: Borneo (Sarawak: Bt Raya (Kapit District) and Ulu Sg
Engkari (Lubok Antu District)), Philippines, Luzon (Catanduanes, where known from
3 collections).
Habitat & Ecology — Mixed dipterocarp forest on yellow clay rich soil, secondary
forest along river banks; 200–800 m altitude.
13. Trichosanthes elmeri Merr.

Trichosanthes elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 299; Rugayah & W.J.de Wilde, Reinward-
tia 11, 4 (1999) 258; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 105; W.J.de
Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu (2001) 210; Sandakania 14 (2004) 17. — Type:
Elmer 20298 (holo PNH, not seen; iso BR, L, NSW, U), Borneo, Sabah, Sandakan.
Climber, 5 –10 m long, early glabrescent, at first minutely brown hairy, stem 2 – 4 mm
diam.; dioecious. Probract concave, 5 –7 by 3 – 6 mm, glands present. Tendrils 2- (or
3-)branched. Leaves: petiole 3 – 6 cm long; petiolules 0.3 –1 cm long; blade greenish
on drying, chartaceous or subcoriaceous, rarely simple, 3- or 5-foliolate, middle leaflet
obovate-oblong, 8.5 –18 by 3.5 – 8.5 cm, scabrous above, glands few or several, scat-
tered, c. 0.5 mm diam., cystoliths obvious, base cuneate, margin ± recurved, entire or
obscurely or sometimes coarsely dentate, lateral leaflets ± smaller and unequal-sided;
2 cm
2 cm
3 mm
b d
c
Fig. 73. Trichosanthes elmeri Merr. a. Leafy node with male inflorescence, note probract at base of
peduncle; b. male bract; c. fruit; d. seed (a, b: Amin & Donggop SAN 113625; c, d: De Wilde, Postar
& Ubaldus SAN 144005).
veins few, pinnate, or in simple leaves 3 – 5 from the base. Male raceme 10 – 30 cm long,
minutely brown hairy; peduncle 3 –10 cm long, 2 – 3 mm thick; rachis not thickened (ex-
cept enlarged nodes), 7– 20 cm long, 5 –15-flowered; bracts late-caducous, (ob)ovate,
15 – 30 by (5 –)8 –12 mm, margin coarsely incised to c. 1/4 deep, glands present. Male
flowers: pedicel 3 – 5 mm long, caducous; receptacle-tube c. 50 mm long, at throat
7– 8 mm wide; sepals narrowly elliptic, 10 –14 mm long, 3 – 4 mm wide at base, entire;
expanded petals not seen; synandrium 13 –14 mm long, filaments glabrous, c. 3 mm

long. Female flowers: pedicel 2 –10 mm long; ovary glabrous, (narrowly) ellipsoid, ±
fusiform, c. 25 by 6 mm; receptacle-tube c. 30 mm long, at throat 6 mm wide; sepals
narrowly elliptic, c. 10 mm long, 2 – 3 mm wide at base, entire; expanded petals not
seen. Fruit ripening orange-red, often paler striped, (narrowly) ovoid or ovoid-ellip-
soid, 12 – 20 by 5 –10 cm, apex c. 4 mm beaked; exocarp leathery, smooth; pericarp
firm-carnose, dry pericarp c. 10 mm thick; pulp blackish green, bitter; fruiting pedicel
0.5 –1.5(– 3) cm long, 4 – 9 mm thick. Seeds blackish brown, compressed, (narrowly) or
broadly ovate, or subcircular, 16 – 20 by 11– 20 by 3 – 4.5 mm, base subtruncate, apex
± acute or obtuse, margin broad but obscure, edge entire. — Fig. 73; Plate 25d, 26a, b.
Distribution — Malesia: Sumatra, whole of Peninsular Malaysia, Singapore, whole
of Borneo, and Philippines (Palawan).
Habitat & Ecology — Edges of primary and secondary forests, margins of swamp
forests, river sides; on limestone and clay soil; from sea level to 900 m altitude.
Note — Trichosanthes elmeri is a variable species, with less robust specimens some-
times linking up with T. wawrae. The collections Chai S 34181, Mamit S 32647 and
Othman et al. S 37534, all from West Sarawak, have thinner leaves and long fruiting
pedicels, 1.5 – 3 cm long.
14. Trichosanthes emarginata Rugayah

Trichosanthes emarginata Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 258, f. 4;
Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 138, pl. 15; C.K.Lim, Folia
Malaysiana 10, 2 (2009) 153, pl. 1–5. — Type: De Wilde & Duyfjes 14862 (holo L; iso BO, K),
Sumatra, Aceh (Ketambe).
Climber, 5 – 20 m long, finely brown hairy, glabrescent, leafy stem 5 mm diam.;

dioecious(?). Probract narrowly elliptic, c. 4 by 2 mm, with few glands. Tendrils
2-branched. Leaves: petiole 5 –7 cm long; blade brown on drying, membranous or
(sub)chartaceous, simple, deeply 5-lobed to 3/4 – 4/5 deep (in juvenile stage unlobed),
in outline circular, 18 – 22 by 20 – 23 cm, puberulent beneath, glands few, at leaf base,
1 mm diam. or absent, cystoliths not obvious, margin entire; veins 5(–7), straight
(curved in juvenile leaves). Male raceme brown puberulous; peduncle 13 –18 cm long,
c. 5 mm thick; rachis not thickened, 2 – 5 cm long, 3 mm thick, at least 10-flowered;
bracts persistent, membranous, broadly obovate-rhomboid, 40 – 55 mm long, margin
shallowly irregularly dentate, glands absent. Male flowers: pedicel c. 10 mm long, ca-
ducous; receptacle-tube 55 – 60 mm long, at throat 10(–12) mm wide; sepals narrowly
elliptic, 13 –15 mm long, entire; petals broadly ovate (size not known); synandrium c.
12 mm long, filaments possibly glabrous, c. 4 mm long. Female flowers not known.
Fruit green, whitish striped, ripening evenly red, broad-ellipsoid, 10 –14 by 6 – 8 cm;
exocarp woody-leathery, smooth; pericarp carnose, dry pericarp (5 –)10 –15 mm thick;
pulp green-black; fruiting pedicel 2-coloured, 3.5 – 5 cm long, 10 mm thick, with a
slightly narrower smooth red part, c. 1 cm long, towards the fruit. Seeds brown-black,
± compressed, narrowly elliptic, often ± parallel-sided, 15 –18 by 7–10 by 3 – 5 mm,
base irregularly truncate, apex conspicuously notched, margin faint, edge entire. — Fig.
74; Plate 26c, d.
2 cm a
3 mm
b
Fig. 74. Trichosanthes emarginata Rugayah. a. Died off plant with fruit; b. seed (both: De Wilde &
Duyfjes 14862, type).
Distribution — Malesia: Sumatra (North Sumatra (Sibolangit) and Aceh (Ketambe))

and Peninsular Malaysia (Pahang, Krau).
Habitat & Ecology — Secondary forest, and forest edges; 200 – 500 m altitude.
Notes — 1. Known from three collections: the type-collection, de Wilde & Duyfjes

14862, is monocarpous, with mature fruits on a died off plant; the second collection,
Lörzing 8347, is a leafy plant with male flowers; the third collection (Peninsular Ma-
laysia) consists of fruit on leafy twigs; the seeds were erroneously drawn as emarginate
at both sides.
2. Trichosanthes emarginata superficially resembles T. montana, the latter differing
in a shorter peduncle, 2.5 – 3 cm long, subcoriaceous bracts, and a thickened rachis.
3. The 2-coloured fruiting pedicel reminds that of T. borneensis, a species with glo-
bose fruit, with seeds not notched at apex.
15. Trichosanthes floresana Rugayah

Trichosanthes floresana Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 260, f. 5;
Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 101, pl. 6. — Type: Schmutz
4614 (holo L), Flores.
Climber, early glabrescent, leafy stem 2 – 4 mm diam.; dioecious. Probract ovate-el-

liptic, ± flat, 5 –10 mm long, glands absent. Tendrils 2- or 3-branched. Leaves: petiole
4 –7 cm long; petiolules 0.2 – 0.5 cm long; blade greenish brown on drying, chartaceous-
coriaceous, 3- or 5-foliolate, ± scabrous with minute pale dots, glands not obvious,
cystoliths abundant, especially on stem, petioles and veins beneath, middle leaflet
(obovate-)oblong, 9 –12 by 4 – 6 cm, lateral leaflets smaller, sometimes 2-lobed and
unequal-sided, base cuneate, margin entire; veins ± pinnate. Male raceme sparsely short
brown hairy, glabrescent; peduncle 8 cm long; rachis not thickened, 2 – 8 cm long, 3
mm thick, 10 –15-flowered; bracts obovate-rhomboid, 20 – 35 by 10 –15 mm, margin
entire or with few dents or slender lobes to 5 mm long, glands present. Male flowers
(from immature bud): pedicel c. 1 mm long, caducous; receptacle-tube c. 7 mm long;
sepals narrowly triangular, 15 mm long, 2 – 3 mm wide at base, margin entire (once
with a narrow sidelobe); petals and synandrium not seen. Female flowers not known.
Fruit ripening evenly orange-red, subglobose-ovoid, 4 – 8 by 3.5 – 6 cm, apex c. 3 mm
beaked; exocarp ± leathery, or thinly woody; pericarp firm-carnose, dry pericarp 8 –10
mm thick; pulp green-black; fruiting pedicel (incompletely known) 1–1.5 cm long,
3 – 4 mm thick. Seeds blackish brown, rather compressed, (narrowly) obovate, 11–16
by 6 –10 by 3 – 5 mm, base and apex ± obtuse, margin absent, edge entire.
Distribution — Malesia: Lesser Sunda Islands (Flores).
Habitat & Ecology — Secondary scrub; 400 – 650 m altitude.
Notes — 1. The dry fruit in the herbarium shows up as ± ovoid, but on the fieldlabels
it is described as globose.
2. The three fruiting collections known vary markedly in size of the mature fruits
and seeds. The fruit of Schmutz 4385 is c. 8 cm long, with seeds 15 –16 by 8 –10 mm,
whereas the fruit of Schmutz 4614 is only 4 cm long, with seeds 10 –11 by 6 –7 mm; the
fruit and seeds of the third specimen, Verheijen 4285, are intermediate.
3. According to Telford (in litt.) T. floresana may be identical to T. pentaphylla Benth.
from Australia (Telford, Fl. Australiana 8 (1982) 198). Further collecting is necessary
to make a decision.
16. Trichosanthes fusca W.J.de Wilde & Duyfjes

Trichosanthes fusca W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 17, f. 4. — Type: Kato, Okamoto
& Walujo 11279 (holo L), Kalimantan, Mt Buduk Rakit, north of Long Bawan, Krayan, 115°47' E,
4°3' N (close to the Sabah border).
Scrambler, wholly sparsely rusty hairy, hairs 0.5 mm long, leafy stem 4 – 5 mm diam.;
dioecious(?). Probract absent (or inconspicuous?). Tendrils hairy, 2-branched. Leaves:
petiole (3 –)4 – 8 cm long; blade green on drying, membranous, simple, unlobed, in
outline (narrowly) ovate or (faintly) hastate, c. 15 by 8 –12 cm, scabrous, glands absent
or few and minute beneath, cystoliths not obvious, base with at the transition to the
petiole at each side a small shallowly bulging auricle, each with 1– 3 crowded glands,
margin irregularly shallowly lobulate-dentate or sinuate, apex acuminate-mucronate.
Male raceme with a single flower, terminal of a short branch 1–1.5 cm long with a few
scattered bracts, co-axillary with the initial of a lateral sterile shoot or possibly a male
raceme; bracts 2 – 3 mm long, conspicuously densely harshly hairy, hairs c. 1 mm long,
dark rusty-brown. Male flowers (from mature bud): pedicel c. 6 mm long; receptacle-
tube long-cupshaped, narrowed in the lower half, c. 10 mm long, 8 mm wide at throat;
sepals long-triangular, c. 8 mm long, 1 mm wide at base, margin entire; petals in bud
folded into a conical body with free apices, at anthesis presumably c. 8 by 6 mm, threads
c. 10 mm long; synandrium subtruncate at both ends, c. 4 by 3.5(– 4) mm, consisting of
3 free anthers, tightly appressed, but not fused, filaments c. 1 mm long, slender, terete,
inserted halfway the tube where passing into strongly brown hairy thickenings of the
tube c. 2 mm long; disc absent. Female flowers and fruit not known. — Fig. 75.
Fig. 75. Trichosanthes fusca W.J.de Wilde & Duyfjes. a. Male flowering
leafy twig; b. male flower bud; c. ditto, opened showing stamens (all:
Kato, Okamoto & Walujo 11279, type).
3 mm 2 cm
b c a
Distribution — Malesia: Borneo (Kalimantan).

Habitat & Ecology — On gravel river bank; 1200 m altitude. Flowering in August.
Note — The flower of Trichosanthes fusca, with free anthers, is quite unique. It
seems most related to T. postarii and more distantly to T. auriculata. All three species
are similar in their leaves having two more or less well-developed auricles, with small
glands inside, at both sides of the blade base at the transition to the petiole. Tricho
santhes fusca lacks the narrowing in the receptacle-tube below the insertion of the
filaments as present in both T. auriculata and T. postarii. The three species are placed
in section Asterosperma W.J.de Wilde & Duyfjes (2004). The collection Clemens 21110
(sterile, BO) from Sarawak, Kapit, upper Rejan River, obviously belonging in sect.
Asterosperma, differs in having much coarser, brittle stiff hairs and cystoliths all over,
and may represent a fourth species in this section. SAN (Postar et al.) 144100, sterile,
from Sabah, Luasong, possibly belongs here as well.
17. Trichosanthes globosa Blume

Trichosanthes globosa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 936; Backer in Backer & Bakh.f., Fl.
Java 1 (1964) 936; Rugayah & W.J.de Wilde, Blumea 42 (1997) 478; Reinwardtia 11, 4 (1999)
260; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 140, pl. 16; W.J.de Wilde
& Duyfjes, Sandakania 14 (2004) 19. — Involucraria globosa (Blume) M.Roem., Fam. Nat. Syn.
Monogr. 2 (1846) 99. — Type: Blume 717 (holo L; iso BR, L, P), Java.
Trichosanthes grandiflora Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 934; Miq., Fl. Ned. Ind. 1, 1 (1856)
674; Cogn. in A.DC. & C.DC., Monogr. Prodr. 3 (1881) 364. — Type: Blume s.n. (holo L, barcode
L0130288; iso BR, P), Java.
Trichosanthes sumatrana Cogn. var. obtusiloba Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 373.
— Type: Beccari FI 4428, FI 4428A, FI 4428B, FI s.n. (syntype, all FI, lectotype not indicated),
all Sumatra.
Climber to 10 m long, at first with sparse grey-brown hairs, early glabrescent (ex-
cept male bracts), leafy stem 2 – 5 mm diam.; dioecious. Probract (narrowly) ovate,
4 –7 by 1.5 – 2.5 mm, glands absent. Tendrils unbranched. Leaves: petiole 3 – 8 cm
long; blade pale greenish brown on drying, (sub)chartaceous, simple, deeply (2 –)3 – 5-
lobed to 9/10 deep, or unlobed (variously lobed in young stages), in outline broadly
(ob)ovate or subcircular, 10 – 20(– 30) by 5 –15(– 40) cm, glabrous above and beneath,
sometimes scabrous, glands few to several, scattered, 0.5 mm diam., cystoliths not
obvious, base cuneate-decurrent near the insertion of the petiole, margin entire, rarely
undulate-dentate, apex (sub)obtuse or acute-acuminate; 3 – 5(–7) subpalmately veined,
veins (nearly) straight; lobes triangular or (lanceolate) oblong. Male raceme: peduncle
3 –13 cm long, 3 – 5 mm thick; rachis thickened, 7– 9 cm long, 10 – 20 mm thick, includ-
ing conspicuous bract scars, many-flowered; bracts caducous, pale yellow on drying,
membranous, obovate, (35 –)50 –70 by 25 – 30 mm, densely soft hairy, margin entire or
shallowly few-lobed at apex, glands minute. Male flowers: pedicel (10 –)30 – 45 mm
long, caducous; receptacle-tube 60 –70 mm long, at throat 7– 8 mm wide; sepals nar-
rowly ovate-triangular, 10 –15 mm long, 2 – 3(– 5) mm wide at base, margin entire, apex
long-acuminate; petals broadly obovate-rhomboid, 13 – 20 mm long, threads (yellow
in Sabah, once seen), 11–15 mm long, synandrium 8 –11 mm long, filaments glabrous,
2 – 3.5 mm long. Female flowers: pedicel 20 – 22 mm long; ovary glabrous, ellipsoid,
2 cm
1 cm
c
d
b
Fig. 76. Trichosanthes globosa Blume. a. Stem node with tendril, leaf, lateral shoot, and male inflo-
rescence; b. apical part of male flower; c. petal; d. leaf of immature plant (a – c: SAN 144003; d: SAN
159462).
Fig. 77. Trichosanthes globosa Blume. a. Fruit; b. seed (both:

Kern 8523).
a 2 cm 3 mm b
c. 15 by 7.5 mm; perianth not known. Fruit ripening evenly bright red, globose, 5.5 –12
by 6 –12 cm, apex not beaked; exocarp thinly woody, smooth; dry pericarp 10 mm thick;
pulp green-black; fruiting pedicel 2 – 6.5 cm long, 10 – 20 mm thick. Seeds brown or
black, compressed, ovate or narrowly elliptic, 15 – 20 by 7– 9 by c. 3 mm, base and apex
rounded, margin present but inconspicuous, edge entire. — Fig. 76, 77.
Distribution — Malesia: Sumatra (Aceh, Padang, Bengkulu), Borneo (Sabah), West
and Central Java.
Habitat & Ecology — Primary and degraded forest edges, riverine forest; 100 – 900
m altitude.
18. Trichosanthes hastata Harms

Trichosanthes hastata Harms, Bot. Jahrb. Syst. 60 (1925) 160; Rugayah & W.J.de Wilde, Reinwardtia
11, 4 (1999) 262; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999)169. — Type:
Schlechter 19286 (holo B, lost; iso BR, drawing and fragment of stem), Papua New Guinea, Kaiser
Wilhelmsland, along Waria River, near Jaduna.
Climber, 5 –10 m long, late glabrescent or with (sparse) rigid pale hairs, leafy stem
1– 4 mm diam.; dioecious. Probract (sometimes caducous), ovate or narrowly elliptic,
3 –7 by 1– 5 mm, glands absent. Tendrils unbranched or 2-branched. Leaves: petiole
2 – 6 cm long, hairy; blade greenish on drying, chartaceous, simple, unlobed, in outline
(narrowly) ovate or sagittate, 6 –18 by 4 –14 cm, scabrous and sometimes finely bul-
late above, glabrous beneath, glands few, c. 0.5 mm diam. or absent, cystoliths usually
obvious, margin remotely (minutely) dentate; veins 3 – 5 from the base, curved, also
a few pinnate ones. Male raceme thinly hairy, flowers solitary at the node, or in the
raceme and often with a solitary flower at base, its pedicel often later on showing as
a persistent straw-like appendage; peduncle 1– 5 cm long, 1– 2 mm thick; rachis not
thickened, 4 –10(–15) cm long, 5 –15-flowered; bracts (narrowly) obovate or (narrowly)
e
1 mm
d
2 mm
2 cm
3 mm
c b
Fig. 78. Trichosanthes hastata Harms. a. Branch with male inflorescences; b. node with male inflo-
rescence in an early stage; c. node with leaf and fruit; d. seed; e. detail of upper leaf blade surface
(a, b: Van Royen NGF 16077; c – e: Womersley NGF 19152).
rhomboid, 8 – 20 by 6 –15 mm, margin sharply dentate to 1/3 deep (or entire), glands
absent or few, 0.5 mm diameter. Male flowers: pedicel 2 – 5 mm long, caducous, in
solitary flowers 10 – 30 mm long, subpersistent; receptacle-tube 25 – 45 mm long, at
throat 5 –10 mm wide; sepals long-triangular or narrowly elliptic, 6 –10 mm long, 1– 2
mm wide at base, entire or incised, or usually slenderly lobed or dentate, lobes 1– 2
mm long; petals obovate-rhomboid, 15 – 25 by 10 – 25 mm, threads 10 – 20 mm long;
synandrium 5 –7 mm long, filaments glabrous, 2 – 3 mm long. Female flowers: pedicel
15 – 25 mm long; ovary (sub)glabrous, (narrowly) ellipsoid, 15 – 25 by 3 – 6 mm. Fruit
ripening evenly orange-red, but greenish at apex, (narrowly) ovoid, c. 7.5 by 4.5 cm,
apex subacute; exocarp thin, woody, smooth; dry pericarp not seen; pulp red; fruiting
pedicel 1.5 – 2.5 cm long, 2 – 3 mm thick. Seeds brown, compressed, ± parallel-sided,
8 –10 by 5 –7 by 1– 2 mm, base subacute or narrowly notched, apex broadly notched,
margin broad but faint, edge ± grooved, coarsely crenulate. — Fig. 78.
Distribution — Malesia: New Guinea (West Papua (Freeport); Papua New Guinea
(Morobe, Northern, and Central Provinces)).
Habitat & Ecology — Riversides, swampy and regrowth forests, forest edges, shrub-
bery and old gardens, on yellow sandy clay soil and river gravel; lowland to 600 m
altitude. Flowering throughout the year.
Notes — 1. Because most herbarium specimens show expanded flowers, it is sup-
posed that T. hastata is mainly diurnal.
2. Ridsdale NGF 31736 and Carr 16334 deviate in having leaves not or hardly sca
brous above, petioles short-hairy only at apex, and hardly incised male bracts.
3. See also the note under the regional key.
19. Trichosanthes intermedia W.J.de Wilde & Duyfjes

Trichosanthes intermedia W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 19, f. 5, 6. — Type: Wood
SAN 16077 (holo SAN; iso A, BO, BRI, K, KEP, L, SING), Borneo (Sabah).
Trichosanthes sp. 2, W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu (2001) 212.
Subherbaceous climber, 2 – 8 m long, minutely hairy (except inflorescences), glabres-

cent, leafy stem 1.5 – 3.5 mm diam.; dioecious or monoecious. Probract absent or very
small. Tendrils unbranched (rarely 2-branched). Leaves: petiole (1.5 –)4 cm long; blade
brown-green on drying, simple, unlobed or 3(– 5)-lobed occasionally to the base, never
foliolate, in outline (narrowly) ovate, subcircular when lobed, 8 –15 by 4 –7(–19) cm,
faintly scabrous, glands several to many, scattered, cystoliths abundant, but small, base
cordate, margin entire, apex of lobes acute-acuminate. Inflorescences: either racemes
male, or racemes of male flowers mixed with some female ones, or few female flowers
in a short raceme (1– 3.5 cm long) or female flowers single at the nodes, wholly densely
hairy, especially bracts and flowers, hairs 0.1 mm long, brown. Male raceme: peduncle
5 – 20 cm long; rachis 4 –16.5 cm long, 2 – 3 mm thick, 5 – 40-flowered; bracts persistent,
(narrowly) ovate, 6 –15 mm long, margin coarsely dentate, apex acute, glands several to
many. Male flowers: pedicel 3 – 6 mm long, persistent; receptacle-tube 30 – 40 mm long,
at throat 6 –7 mm wide, at base broadened into a semi-globose pseudo-ovary 3 – 5 mm
diam.; sepals narrowly elliptic, 6–8 by 2–2.5 mm, entire; petals ± obdeltoid, 10 mm long,
threads 10 mm long; synandrium slightly exserted, truncate at apex, ± narrower at base
2 cm
a
3 mm
e
2 cm
d
Fig. 79. Trichosanthes intermedia W.J.de Wilde & Duyfjes. a, b. Male flowering leafy twigs, note
flowers with pseudo ovaries; c. male flower; d. female flowering leafy twig (flowers exceptionally in
short raceme); e. fruit; f. seed (a – c: De Wilde SAN 116493; d: De Wilde & Postar SAN 141936; e, f:
Lassan & Ampon SAN 71525).
c
1 mm
d 2 mm
2 mm
f
a b
Fig. 80. Trichosanthes intermedia W.J.de Wilde & Duyfjes. a. Male flower bud; b. ditto, opened, note
synandrium and semiglobose disc at base; c. synandrium, cross section; d. female flower; e. ditto,
opened; f. ditto, ovary, longitudinal section (a – c: Asik Mantor SAN 115882; d – f: De Wilde & Postar
SAN 141936).
and entering into the narrowed portion of the receptacle-tube, c. 9 by 3.5 mm, anthers
fused, filaments glabrous, c. 1 mm long, inserted c. 5 mm below the receptacle-throat
(attached to the synandrium well above its base); disc c. 3 by 3 – 4 mm, largely fused
with the receptacle-tube, in dry specimens causing the pseudo-ovary (see note 2). Female
flowers: pedicel 6 –10 mm long, ovary (sub)glabrous, c. 9 by 5 mm; receptacle-tube
c. 30 mm long; sepals narrow, 5 mm long. Fruit mostly solitary, green, ripening evenly
red, ovoid or globose, 6 – 9 by 6 –7 cm, pericarp when fresh 5(–10) mm thick, much thin-
ner when dry; fruiting pedicel 1.5(– 4) cm long, c. 3 mm thick. Seeds brown, compressed,
(narrowly) elliptic, 15(– 20) mm long (see note 3), 7– 9 mm wide, base truncate-rounded,
margin narrow and a low ridge in the middle, edge entire. — Fig. 79, 80.
Distribution — Malesia: Borneo (Sarawak (Sg Sipayan) and Sabah).
Habitat & Ecology — Open forest and forest edges, often on damp sites; sea level
to 1300 m altitude. This species starts flowering already in the late afternoon.
Notes — 1. Trichosanthes intermedia can be confused with T. wawrae, T. longispi
cata, and T. refracta. The latter, a species from peaty forest in Brunei and Philippines,
differs in sinuate male racemes and lack of glands on the bracts. Trichosanthes long
ispicata from Sarawak is similar, but stouter, and differs in larger 5 –7-lobed leaves and
male pedicels about as long as the bracts. In T. wawrae (Peninsular Malaysia, Sumatra,
West Java, not known with certainty from Sabah) the leaves may be entire, lobed or
compound, the leaflets distinctly petioluled, its male racemes are generally more slender
and the male flowers lack the pseudo-ovary, its seeds are generally broader. The differ-
ences between T. intermedia and T. wawrae need more study, however.
2. The phenomenon of a pseudo-ovary in male flowers is caused by the presence of
a thick cup- or cushion-shaped disc at the bottom of the receptacle-tube, and becomes
apparent on drying, resembling an ovary, which in the female flowers, of course, is
inferior below the receptacle-tube. A pseudo-ovary is absent in the resembling T. longi-
spicata, but occurs in T. refracta, and in the non-resembling T. villosa.
3. The collection Chew, Corner & Stainton RSBN 1204 (Kinabalu) differs in having
a long fruiting pedicel, c. 4 cm long, and larger seeds, c. 20 mm long.
20. Trichosanthes kinabaluensis Rugayah

Trichosanthes kinabaluensis Rugayah, Reinwardtia 11, 5 (2000) 419; Rugayah in Rugayah & W.J.de
Wilde, Reinwardtia 11, 4 (1999) 262, f. 6, nom. inval.; Rugayah, Trichosanthes (Cucurbitaceae) in
Malesia, thesis (1999) 90, pl. 2; W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu (2001)
211, pl. 16: d, e; Sandakania 14 (2004) 23. — Type: Chew, Corner & Stainton RSBN 2830 (holo
BO; iso K, L, SAN), Borneo, Sabah, Tenompok.
Climber or creeper, 5 –10 m long, glabrous, leafy stem 3 – 5 mm diam., sharply angu-
lar or winged, young shoots reddish; dioecious. Probract somewhat succulent, entire,
narrowly elliptic, 10 –15 mm long, glands few or absent. Tendrils 2- (or 3-)branched.
Leaves: petiole 5 –12 cm long; blade brown or greenish on drying, membranous or
chartaceous, simple, 3 – 5-lobed or 5-angular, in outline circular, 6 – 22 by 6 – 21 cm,
scabrous, glands few, towards the base of the midrib, (0.5 –)1– 2 mm diam., cystoliths
present, margin entire or somewhat undulate-dentate; veins 5, straight, prominent on
both surfaces. Male raceme sparsely minutely hairy, glabrescent, peduncle 7–16 cm
long, 2 – 4 mm thick; rachis not thickened, 4 –12 cm long or more, 10 – 20-flowered;

bracts obovate-rhomboid, 30 – 45 by 20 – 45 mm, margin irregularly deeply incised to c.
1/3 deep, glands few or absent. Male flowers: pedicel 10 – 35 mm long, caducous; recep-
tacle-tube 50 –70 mm long, at throat 10 –12 mm wide; sepals long-triangular, 10 –17 mm
long, 3 – 5 mm wide at base, margin usually coarsely dentate or entire; petals obovate-
rhomboid, c. 25 mm long, threads c. 20 mm long; synandrium 10 –12 mm long, with
spine-like hairs between the thecae, filaments glabrous, 3 – 4 mm long. Female flowers:
pedicel c. 20 mm long, ovary glabrous, ellipsoid, 10 –14 mm long; receptacle-tube nar-
row, 60 – 65 mm long, at throat 7– 9 mm diam., style hairy, c. 60 mm long, stigma c. 5
mm long. Fruit ripening orange, paler speckled, globose, 5.5 –7.5 cm diam.; exocarp
leathery, or thinly woody; dry pericarp 5 –10 mm thick; pulp green-black, bitter; fruit-
ing pedicel 1.5 – 2.5 cm long, 4 –7 mm thick. Seeds dark brown, compressed, elliptic, c.
10 by 3 – 4(– 5) by 1– 2 mm, base ± cuneate, apex broadly rounded, margin faint, edge
entire. — Plate 27a, b.
Distribution — Malesia: Borneo (Sabah (Kinabalu and northern Crocker Range area)).
Habitat & Ecology — Moist places in secondary roadside vegetation, forest and
scrub edges; 1000 –1600 m altitude.
Note — Young shoots in living plants are reddish purplish.
21. Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang

Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang, Bull. Bot. Res., Harbin 16, 4 (1996) 503, f. 2; Ru-
gayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 262; Rugayah, Trichosanthes (Cucurbitaceae) in
Malesia, thesis (1999) 173. — Type: Coode et al. NGF 32585 (holo CANB; iso BO, K, L, LAE),
Papua New Guinea, Eastern Highlands Province, Kassam Pass.
Climber, 3 – 6 m long, glabrescent or pale brown hairy, leafy stem 3 – 5 mm diam.; di-
oecious. Probract elliptic or linear, 2 – 5 mm long, glands present. Tendrils unbranched
or 2-branched. Leaves: petiole (3 –)5 –12 cm long; blade (brown) or green on drying,
membranous or chartaceous, simple, unlobed, or shallowly or deeply 3 –7-lobed to
1/3(– 2/3) deep, or 3 – 5-angular, in outline subcircular or broadly ovate, 10 – 22 by
8 – 20 cm, often scabrous, especially above, short hairy or glabrescent beneath, glands
few to several, scattered but usually close to the insertion of the petiole, 0.5 –1(– 2) mm
diam., margin entire, or sparsely minutely dentate or sinuate; veins 5(–7), straight or
arching. Male raceme usually with co-axillary a solitary male flower or a straw-like
appendage, brown hairy, hairs 1–2 mm long, peduncle 5 –16 cm long, c. 2 mm thick;
rachis not thickened, 5 –10 cm long, 5 –10-flowered; bracts ovate or (narrowly) obovate,
4 –10 by 3 – 5 mm, acute, margin entire or finely dentate, glands present. Male flowers:
pedicel in the raceme c. 5 mm long and caducous, in solitary flower c. 75 mm long and
persisting as a straw-like appendage; receptacle-tube 35 – 80 mm long, at throat 8 –12
mm wide; sepals narrowly triangular, 4 – 5 mm long, 1– 2 mm wide at base, margin en-
tire; petals obovate or rhomboid, 10 – 25 mm long, threads c. 10 mm long; androecium
not seen. Female flowers: mature flowers not known; ovary finely hairy, (narrowly)
ellipsoid; sepals as in male. Fruit ripening evenly red, (sub)globose, (narrowly) ellip-
soid or pyriform, 10 – 20 by 5 – 9.5 cm, glabrous or sparsely hairy; exocarp woody and
smooth or leathery and wrinkled, pericarp carnose, dry pericarp (2 –)5 –10 mm thick;
pulp orange-red or yellow; fruiting pedicel 1.5 – 3.5 cm long, 4 – 8 mm thick. Seeds pale
or dark brown, compressed, narrowly elliptic or ± parallel-sided, 12 –17 by 4 – 6(–7) by
2 mm, notched at one or both ends, margin faint, edge entire or faintly undulate.
Distribution — Malesia: New Guinea; 4 forms.
KEY TO THE FORMS
1a. Leaves unlobed, glabrous beneath. Fruit (narrowly) pyriform. — eastern Papua

New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. forma sicyocarpa
b. Leaves unlobed or shallowly 3-lobed or 3-angular, densely hairy or glabrescent
beneath. Fruit globose or (narrowly) ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a. Fruit (sub)globose or short-ellipsoid; exocarp woody, smooth. — West Papua, So-
rong area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c. forma sorongensis
b. Fruit (narrowly) ellipsoid; exocarp wrinkled (not completely known in forma lae
oica) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Male receptacle-tube 40 – 80 mm long. — Papua New Guinea, Eastern Highlands
Province . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. forma laeoica
b. Male receptacle-tube c. 35 mm long. — Western West Papua d. forma yapenensis
a. forma laeoica
Leaves: blade unlobed, 3–5-angular or 3-lobed, greenish brown on drying, hairy be-
neath. Male raceme with a solitary flower at the node, the pedicel often as a straw-like
remnant at base. Male flowers: pedicel in raceme c. 5 mm long, in solitary flowers c. 75
mm long; receptacle-tube (40–)50–80 mm long. Fruit (incompletely known): c. 5 cm
wide; exocarp finely wrinkled, glabrous.
Field-notes — Petals white with a green line abaxially. The flowers probably are open
at night only, wilting quickly at sunrise. Fruit said to be red when ripe, edible.
Distribution — Malesia: New Guinea (Papua New Guinea (Eastern Highlands Prov-
ince)).
Habitat & Ecology — Secondary forest; c. 1400 m altitude.
b. forma sicyocarpa Rugayah

Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang forma sicyocarpa Rugayah in Rugayah & W.J.de Wil-
de, Reinwardtia 11, 4 (1999) 264; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999)
176. — Type: Brass 5347 (holo BO; iso BRI), Papua New Guinea (Central Province, Mafulu).
Leaves: blade unlobed, green or dark brown on drying, broadly ovate, glabrous
beneath, margin entire or shallowly undulate. Male raceme, male and female flowers
not known. Fruit (narrowly) pyriform, c. 20 by 3 cm; exocarp thinly woody, smooth;
fruiting pedicel c. 3 cm long, c. 8 mm thick.
Field-notes — Fruit green, ripening orange to red, cucumber-like, edible; sometimes
cultivated.
Distribution — Malesia: New Guinea (Papua New Guinea (Western Highlands, Mo-
robe, and Central Provinces)).
Habitat & Ecology — Secondary regrowth forest, along logging roads; 900 –1400 m
altitude.
Note — The precise size and shape of the fruit is not known.
c. forma sorongensis Rugayah

Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang forma sorongensis Rugayah in Rugayah & W.J.de
(1999) 177. — Type: Avé 4782 (holo L), West Papua, Sorong (Ayawasi).
Leaves: blade unlobed or 3-lobed to 1/4 deep, greenish on drying, densely hairy be-
neath, glands many, scattered. Male raceme, male and female flowers not known. Fruit
(sub)globose or short-ellipsoid, 7.5–12.5 by 8–10 cm (spirit); exocarp thin, woody, smooth,
sparsely hairy; fruiting pedicel 1.5–3 cm long, 4–6 mm thick.
Field-notes — Climber 6 m long Fruit c. 12.5 cm long; inside orange red, edible.
Distribution — Malesia: New Guinea (West Papua: Vogelkop (Sorong: Ayawasi)).
Habitat & Ecology — Secondary forest, margin of gardens; on deep brown-grey clay;
c. 450 m altitude.
d. forma yapenensis Rugayah

Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang forma yapenensis Rugayah in Rugayah & W.J.de
(1999) 178. — Type: Widjaja 6882 (holo BO; iso L), West Papua, Yapen Is.
Leaves: blade unlobed or 3-lobed, 1/3–2/3 deep, green or brown on drying, scabrous
hairy or glabrous beneath, scabrous above. Male raceme at base with a straw-like append-
age (withered peduncle or pedicel of solitary flower). Male flowers: pedicel c. 5 mm long;
receptacle-tube c. 35 mm long. Fruit (narrowly) ellipsoid, c. 10 by 5 cm; exocarp leathery,
finely wrinkled; fruiting pedicel c. 3.5 cm long, 4 mm wide.
Field-notes — Creeping on the ground, leaves rough; fruit ripening red, edible.
Distribution — Malesia: New Guinea (West Papua (Yapen Is.)).
Habitat & Ecology — Primary and secondary forest, creeping along roadsides; sea level
to 1000 m altitude.
Note — Known only from a few incomplete collections; the fruit in spirit is 12 cm long
and 6 cm wide.
22. Trichosanthes leuserensis Rugayah

Trichosanthes leuserensis Rugayah, in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 265; Ru-
gayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 127. — Type: de Wilde & Duyfjes
18605 (holo BO; iso K, L), Aceh, Gunung Leuser Park, Ketambe.
Climber, 10 – 20 m long, glabrous (minute powdery hairs on growing shoots ex-

cepted), leafy stem 3 – 6 mm diam.; monoecious. Probract obovate or narrowly elliptic,
10 –15 by 5 – 8 mm. Tendrils 2- or 3-branched. Leaves: petiole 3 –10 cm long; blade
reddish brown on drying, membranous or chartaceous, simple, deeply 3 – 5-lobed, in
outline subcircular, 8 –18 by 7–18 cm, ± scabrous, glands few or several, towards the
insertion of the petiole and sometimes scattered, 0.5 –1 mm diam., cystoliths present,
margin (minutely) serrate-dentate, more coarsely so towards the base; veins 5(–7),
straight. Male raceme thinly minutely brown hairy, glabrescent; peduncle 6 –11 cm long,
2 – 4 mm thick; rachis not thickened, 4 –10 cm long, up to 20-flowered; bracts persist-
ent, broadly ovate-rhomboid, 30 – 45 by 15 – 20 mm, margin finely dentate or incised
2 – 5(–10) mm deep, glands absent. Male flowers: pedicel 5(–10) mm long, caducous;
receptacle-tube c. 50 mm long, at throat 8 mm wide; sepals narrowly elliptic, 12 –15 mm
long, 3 – 4 mm wide at base, margin dentate, rarely entire; petals obovate-rhomboid, c.
10 mm long, papillose hairy, threads short; synandrium c. 10 mm long, filaments c. 2
mm long, glabrous. Female flowers (from bud): solitary or 1 or 2 at base in male raceme,
with smaller bract; pedicel 5 –10 mm long; ovary glabrous, (narrowly) ellipsoid, c. 10
by 3 mm. Fruit ripening orange, with or without whitish greenish stripes, subglobose,
6.5 – 8 by 5 –7.5 cm, apex c. 2 mm beaked; exocarp leathery, smooth; pericarp firm-
carnose, dry pericarp 10 –18 mm thick; pulp greenish black, bitter; fruiting pedicel in
the raceme 0.5 –1.5 cm long, when solitary on the node c. 2 cm long, c. 4 mm thick.
Seeds dark brown, ± compressed, (narrowly) elliptic, base and apex broadly rounded,
15 –18 by 8 –10 by 3 – 4 mm, margin absent, edge entire.
Field-notes — Herbaceous climber 8 –15 m tall. Fruit (sub)globose, 9 –10 cm diam.,
bright orange, with or without whitish greenish stripes, pericarp 2.5 cm thick, pale yel-
low to yellowish green inside, seeds brown, imbedded in blackish pulp. Immature fruit
when cut smelling of cucumber.
Distribution — Malesia: Sumatra: Aceh (Kluet, Ketambe); North Sumatra (Sibolan-
git).
Habitat & Ecology — Rain forest, along rivers; on yellow-red loamy soil; 60 – 600 m
altitude.
Notes — 1. Trichosanthes leuserensis is similar to T. pubera; the latter is dioecious,
and differs in reddish tinged shoots, a much more elongated probract, leaves hairy be-
neath, and ellipsoid (not subglobose) fruits always at the nodes, generally with a less
thick pericarp, and smaller and compressed seeds.
2. Trichosanthes leuserensis is probably entirely or mostly monoecious, because the
collections contain male and fruiting (or female flowering) elements, although the fruit
is sometimes mounted detached on the sheets.
23. Trichosanthes longispicata Rugayah

Trichosanthes longispicata Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 266, f. 7,
p.p., for the type only; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 96, pl.
4, p.p., for the type only. — Type: Yii & Othman S 46270 (holo L; iso K, SAR), Borneo, Sarawak,
1st Div., Mt Hujan.
Climber to 5 m long, minutely brown hairy, glabrescent, leafy stem 4– 6 mm diam.;

dioecious. Probract not seen. Tendrils 2- or 3-branched. Leaves: petiole 4 – 5 cm long;
blade greenish brown on drying, membranous or chartaceous, simple, deeply 5(–7)-
lobed 7/8 deep, in outline circular, c. 20 by 23 cm, faintly scabrous above, glands few
or absent, situated towards the insertion of the petiole, c. 1 mm diam., cystoliths in-
conspicuous or absent, margin entire; veins 5(–7), straight. Male raceme finely brown
2 cm
Fig. 81. Trichosanthes longispicata Rugayah. Twig with male racemes (Yii & Othman S 46270, type).
(-yellow) hairy, glabrescent, peduncle 4– 5 cm long, c. 3 mm thick; rachis not thickened,

2 0– 27 cm long, 25 – 50-flowered or more; bracts persistent, (narrowly) (ob)ovate, 9–12
by 3 –7 mm, margin conspicuously dentate, glands numerous, obscure. Male flowers
(from buds): pedicel 9–12 mm long, persistent; receptacle-tube c. 10 mm long, at throat
5 mm wide, at base without a pseudo-ovary (see note under T. intermedia); sepals nar-
rowly triangular, 4 – 5 mm long, c. 1.5 mm wide at base, entire; petals and androecium
not seen. Female flowers and fruit not known. — Fig. 81.
Distribution — Malesia: Borneo (Sarawak, where known only from the type).
Habitat & Ecology — Logged over dipterocarp forest; at low altitude.
24. Trichosanthes montana Rugayah

Trichosanthes montana Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 3 (1998) 218; Rugayah
& W.J.de Wilde, Reinwardtia 11, 4 (1999) 268; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia,
thesis (1999) 143, pl. 17. — Type: De Wilde & Duyfjes 21841 (holo BO; iso L), West Java, Telaga
Warna near Cibodas.
Trichosanthes sumatrana auct. non Cogn.: Rugayah & W.J.de Wilde, Blumea 42, 2 (1997) 479, f. 4, p.p.
Climber to 12 m long, minutely brown hairy, hairs 1 mm long, glabrescent, leafy

stem 3 – 6 mm diam.; monoecious or dioecious. Probract membranous or somewhat
fleshy, (narrowly) elliptic, 5 –10(–15) mm long, glands absent or present. Tendrils
2 – 4-branched. Leaves: petiole 3.5 –14 cm long; blade greenish brown on drying, mem-
branous, chartaceous or subcoriaceous, simple, 5 –7-lobed to c. 3/4 deep (unlobed in
juvenile stages), in outline circular, 10 – 28 by 11– 26 cm, not or little scabrous, often
finely bullate above, glands few or absent, generally towards the base, 1– 3 mm diam.,
cystoliths not obvious, margin sometimes recurved, entire or irregularly shallowly
dentate; veins 5 –7, straight, ± sunken above. Male raceme thinly brown hairy; pedun-
cle 2 – 6 cm long, 3 –7 mm thick; rachis thickened, 13 – 35 cm long, 5 – 20 mm thick,
20-flowered (or more); bracts late-caducous or persistent, subcoriaceous, rhomboid or
broadly obovate, 40– 60 by 20 – 30 mm, margin coarsely dentate, sometimes with 3 main
lobes, glands absent. Male flowers: pedicel 3 –10 mm long, caducous; receptacle-tube
30 – 45 mm long, at throat 7– 8(–10) mm wide; sepals narrowly triangular, 10(–17) mm
long, 3 – 4 mm wide at base, entire or with few narrow lobes; petals obovate-rhomboid,
17– 20 mm long, threads 5 –10 mm long; synandrium 12 –15 mm long, inserted at c. 15
mm below throat, filaments 3 – 4 mm long, hairy; disc close to the base of the recepta-
cle-tube, consisting of 3 transverse pads. Female flowers: sometimes monoecious with
flower co-axillary with the male raceme, or some flowers mixed in the male raceme;
pedicel 15 – 30 mm long (when in male raceme shorter), ovary short brown hairy or
subglabrous, subglobose or ovoid, 10 –15 by 8 –10 mm; receptacle-tube and perianth
as in male. Fruit solitary at the nodes or in old male raceme, ripening evenly orange or
bright red, (depressed) globose or ellipsoid, 10 –14 by 8 –14 cm, apex rounded; exocarp
woody, c. 1 mm thick, smooth; dry pericarp firm-carnose, 15 mm thick; pulp green-
black, bitter; fruiting pedicel 3 – 5 cm long, 10 – 20(– 25) mm thick. Seeds brown-black,
compressed, ovate-oblong, 17– 20 by 6 – 8 by 2 – 3 mm, base and apex narrowly rounded,
margin present, distinct or faint, edge entire.
Distribution — Malesia: Sumatra, Borneo, and Java; 2 subspecies.
2 cm
2 cm c
3 mm
2 cm
2 cm e
b
Fig. 82. Trichosanthes montana Rugayah subsp. crassipes W.J.de Wilde & Duyfjes. a. Apex of growing
shoot; b. node with mature leaf; c. fruit; d. seed; e. young plant, cotyledons still present (a, b: De Wilde
& Duyfjes 21997; c, d: Sugau JBS 106; e: De Wilde¸ Postar & Pereira SAN 141925).
c
6
4 2
3
1
a
3 cm
Fig. 83. Trichosanthes montana Rugayah subsp. montana. a. Node with male inflorescence in its natural
up-curved position; b. detail of male inflorescence with also a half-grown fruit; c. node, showing 1:
leaf, 2: female flower, perianth removed, 3: developing male inflorescence, 4: tendril; 5: sterile lateral
shoot, 6: probract (a, b: De Wilde & Duyfjes 21841; c. De Wilde & Duyfjes 21865).
1a. Fruits solitary at the node or in old male raceme, ripening orange, ellipsoid. Fruiting
pedicel 4 – 5 cm long, c. 10 mm thick, smooth or shallowly fissured. — Sumatra,
Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. subsp. montana
b. Fruits solitary at the node, ripening bright red, (depressed) globose. Fruiting pedicel
c. 3 cm long, 15 – 20(– 25) mm thick, deeply fissured and brown corky. — Borneo
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. crassipes
a. subsp. montana
Fruit solitary or developed in the male raceme, ripening orange, ellipsoid, 10 –14 cm
long, c. 8 cm wide; fruiting pedicel 4 – 5 cm long, c. 10 cm thick, smooth or shallowly
fissured. — Fig. 83.
Distribution — Malesia: West Sumatra (Gn Sago); South Sumatra (Lampung (Gn
Tanggamus)); West Java.
Habitat & Ecology — Primary montane forest; 900 –1350 m altitude.
b. subsp. crassipes W.J.de Wilde & Duyfjes

Trichosanthes montana subsp. crassipes W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 23. — Type:
De Wilde, Tajuddin & Good SAN 143916 (holo SAN; iso L), Sabah (Tongod: Imbak River).
Fruit solitary at the node, ripening bright red, (depressed)globose, 10 –14 cm diam.;
fruiting pedicel c. 3 cm long, 15 – 20(– 25) mm thick, deeply fissured and brown corky.
— Fig. 82; Plate 30a.
Distribution — Malesia: Borneo (Sarawak: 1st Division (Gn Pueh); Sabah: Tenom-
pok (near Ranau), Tawau Hill Forest Reserve, and Lungmanis Forest Reserve; East
Kalimantan (near Balikpapan and Berau)).
Habitat & Ecology — Along roadsides and forest edges; 100 –1600 m altitude.
25. Trichosanthes mucronata Rugayah

Trichosanthes mucronata Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 268; Ru-
gayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 80, pl. 1; W.J.de Wilde & Duyfjes
in Beaman et al., Pl. Mt. Kinabalu (2001) 211, pl. 17: a, b; W.J.de Wilde & Duyfjes, Sandakania
14 (2004) 25. — Type: De Wilde & Duyfjes SAN 139464 (holo SAN; iso BO, L), Borneo, Sabah,
Mt Kinabalu area.
Climber to 5 m long, whitish or brown soft hairy, hairs 1– 2 mm long, leafy stem 2 – 4
mm diam.; dioecious. Probract absent. Tendrils 3 – 5-branched, point of branching 2 – 4
cm from the base. Leaves: petiole 3 –7 cm long; blade green on drying, membranous,
simple, unlobed, in outline broadly ovate, 6 –19(– 21) by 5 –13 cm, finely scabrous
above and beneath, glands several, scattered, 0.5 mm diam. or less, cystoliths present
but indistinct, margin minutely dentate or dentate-serrate, apex acute-acuminate, with
a 13 – 20(– 25) mm long acumen; veins 5 (or 7), ± arching. Male raceme (sparsely or)
densely brown soft hairy; peduncle 4 – 8(– 9) cm long, 2 – 3 mm thick; rachis not thick-
ened, 4 –7 cm long, 5 –12-flowered, occasionally male flowers single; bracts caducous,
on the rachis, c. 1 mm long. Male flowers densely dark brown hairy, including small
gland-hairs; pedicel 20 – 40 (in single flowers to 45) mm long, (sub)persistent, peduncle
or solitary pedicel often straw-like withering; receptacle-tube narrow, tubular, 35 – 65
mm long, at throat 5 – 6(–7) mm wide; sepals narrowly triangular or linear, 11–16 mm
long, (1.5 –)2 – 2.5 mm wide at base, entire; petals narrowly elliptic, c. 15 mm long,
threads much longer; synandrium 7– 8 mm long, at apex with 3 processes 1–1.5 mm
long, filaments 4 – 5 mm long, glabrous; disc consisting of 3 fleshy, line-shaped ele-
ments, c. 25 mm long. Female flowers solitary, resembling male flowers; pedicel 15 – 25
mm long; ovary densely hairy, narrowly ellipsoid, 10 –15 by c. 5 mm. Fruit ripening
(yellow-)green with whitish stripes, ellipsoid, 9 –10 by c. 6 cm, apex shortly beaked;
exocarp leathery, thin, smooth; pericarp carnose, dry pericarp 5 mm thick; pulp whitish,
slightly bitter; fruiting pedicel c. 2 cm long, 4 mm thick. Seeds tumid, broad, with two
inflated lateral parts, c. 10 by 18 by 4 mm, margin absent, edge entire, smooth. — Plate
27c, d.
Field-notes — The male flowers open at night and stay open until noon. The space
between the anthers is densely set with stiff white hairs, c. 0.5 mm long.
Distribution — Malesia: Borneo (endemic to Sabah, in a restricted area below the
entrance of Kinabalu Park).
Habitat & Ecology — Wet places in forest edges and shrubby roadsides; 1000 –1400
m altitude. Flowering in December; fruiting December to March.
26. Trichosanthes obscura Rugayah

Trichosanthes obscura Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 269; Ru-
gayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 92, pl. 13; W.J.de Wilde & Duyfjes,
Sandakania 14 (2004) 25. — Type: Hansen 968 (holo L; iso C, S), Borneo, Sarawak, Ulu Belaga,
Sepakau logging camp.
Climber, 6 – 8 m long, finely greyish brown hairy, glabrescent, leafy stem 3 – 4 mm

diam.; dioecious. Probract (ob)ovate, 4 – 8 mm long, subentire, minutely papillose
hairy, glands absent. Tendrils 2- or 3-branched. Leaves: petiole 3 – 6 cm long; blade dull
blackish on drying, membranous, simple, unlobed or 3-lobed, 1/3 – 2/3 deep, in outline
ovate or circular, 7– 20 by 8 –18 cm, finely scabrous above or papillose hairy, glands few
or absent, mainly towards the base, 1– 2 mm diam., cystoliths occasionally on petiole
and veins, margin entire or undulate-dentate, apex slenderly acuminate; veins 5(–7),
straight or curved. Male raceme finely brown hairy, to 30 cm long, including the 8 –14
cm long and 2 – 3 mm thick peduncle; rachis not thickened, c. 4 cm long, 5 –10-flowered;
bracts broadly rhomboid or obovate, 15 – 20 by c. 15 mm, margin finely fan-shaped
incised to nearly halfway, glands absent. Male flowers (from bud): pedicel c. 5 mm
long, caducous; receptacle-tube c. 15 mm long, at throat 5 –7 mm wide; sepals narrowly
ovate, 5 –7 mm long, c. 3 mm wide at base, margin narrowly lobed; petals and stamens
not seen. Female flowers: pedicel c. 15 mm long; ovary glabrescent, subglobose, 10 –15
mm long; receptacle-tube c. 20 mm long; sepals narrowly elliptic, 5 mm long. Fruit
pale green with whitish dots, ripening evenly bright orange, (depressed-)globose, c.
7 by 9 cm; exocarp woody, smooth, c. 1 mm thick; pericarp juicy, dry pericarp 1 mm
thick; pulp green-black; fruiting pedicel 2 – 3 cm long, c. 5 mm thick. Seeds compressed,
(narrowly) elliptic, with a faint lengthwise ridge in the middle, 18 – 22 by 7– 8 by 2 mm,
base and apex obtuse-truncate, margin broad but faint, edge entire.
Field-notes — Mature fruit globose, green, spotted with light green. Bygrave 49
(Brunei: Belait) noted: “Climber growing through the lower canopy. Leaves covered
in glandular hairs giving them a sticky texture. Fruit bright orange on short, green
stalk.”
Distribution — Malesia: Borneo (Brunei (Labi Hills Forest Reserve and Belait);
Sarawak (4th Div.: Bintulu, 7th Div.: Ulu Belaga); Sabah (Pensiangan Kayu Forest
Reserve, Mendalom Forest Reserve) and North Kalimantan (Sg Magne)).
Habitat & Ecology — Mixed dipterocarp forest, riversides and hillsides, also keran-
gas forest; at low altitudes.
27. Trichosanthes papuana F.M.Bailey

Trichosanthes papuana F.M.Bailey, Queensland Agric. J. 7 (1900) 349; Harms, Bot. Jahrb. Syst. 60
(1925) 161; Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 271; Rugayah, Trichosanthes (Cu-
curbitaceae) in Malesia, thesis (1999) 103, pl. 7. — Type: Le Hunte s.n. (holo BRI), south-eastern
Papua New Guinea.
Large climber, glabrous, leafy stem 2 – 5 mm diam., sometimes lenticellate; dioe-

cious. Probract rather thick, broadly ovate, concave, 3 – 8 by 2 – 5 mm, glands absent.
Tendrils 2- or 3-branched. Leaves: petiole 3 – 6.5 cm long; petiolules 0.5 –1.5 cm long;
blade green on drying, membranous or chartaceous, 3 – 5-foliolate (simple and hastate
in juvenile plants), in outline circular, c. 30 cm diam., not or little scabrous, with minute
pale or dark dots, glands few or absent, scattered, 0.5 mm diam., cystoliths obvious;
middle leaflet 9 –18 by 3.5 –10 cm, base rounded or cuneate, margin (sub)entire, apex
acute-acuminate. Male raceme minutely brown hairy, glabrescent; peduncle 5 –10 cm
long, c. 2 mm thick; rachis not thickened, c. 15 cm long, c. 10-flowered; bracts per-
sistent, rhomboid or broadly ovate, 20 – 30 by 15 – 25 mm, margin irregularly sharply
incised to 4 mm deep, glands not obvious. Male flowers (from subsessile bud): sepals
long-triangular, c. 5 mm long, 2 mm wide at base, subentire. Female flowers not known.
Fruit ripening bright red, possibly somewhat striped at apex, broadly ovoid-ellipsoid,
9 –16(– 20) by 7–13(–17) cm; exocarp woody, smooth, 2 – 3 mm thick; pulp green-black;
fruiting pedicel 1– 3 cm long, 7–15 mm thick. Seeds blackish, compressed, ovate or nar-
rowly elliptic, 20 – 27 by 11–15 by 4 – 6 mm, base subtruncate, apex narrowly rounded,
margin broad but faint, edge entire.
Field-notes — Fruit large, orange or bright scarlet. The plant is propagated by cut-
tings. Takeuchi & Ama 17069 (Morobe Province, Lae) noted: “Fruits 16 cm long, 12 cm
wide; pulp yellow, seeds flat, black. Fruits are roasted in open fire and eaten by local
settlers.”
Distribution — Malesia: Moluccas (Aru Islands, Pulau Wokam); New Guinea (West
Papua (Jayapura Province: Waisiniwai), and Papua New Guinea (Morobe Province: Lae
and Western Province)).
Habitat & Ecology — Primary and secondary forest, scrambling in deep roadside
vegetation and in margins of old gardens; at low altitudes.
Notes — 1. The yellow colour of the pulp as mentioned in Takeuchi & Ama 17069
is at variance with the fruit in the herbarium collection and also with the pulp colour
in related species, e.g., T. elmeri. Possibly the yellow pulpy pericarp is meant, whereas
the blackish seeds are embedded in green blackish watery pulp.
2. Trichosanthes papuana is similar to the Australian T. pentaphylla as well as to the
Malesian T. celebica, T. elmeri and T. floresana. The fruit in T. papuana is large and has
a thin, woody pericarp. In the resembling species the pericarp is leathery, collapsing on
drying.
28. Trichosanthes pendula Rugayah

Trichosanthes pendula Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 3 (1998) 219, f. 1– 3;
Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 271; Rugayah, Trichosanthes (Cucurbitaceae)
in Malesia, thesis (1999) 83; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 25. — Type: Joseph
B. et al. SAN 120771 (holo SAN; iso KEP, L), Borneo, Sabah, Lahad Datu.
Climber, 10 – 20 m long, harshly grey or rusty hairy, hairs 0.5 – 2 mm long, partly late-
glabrescent, leafy stem 2 – 5(–15) mm diam., conspicuously 4- or 5-angular or ribbed,
the ribs green and hairy; dioecious or possibly monoecious. Probract absent. Tendrils
2 – 5-branched. Leaves: petiole 3.5 – 6 cm long; blade green on drying, membranous,
simple, shallowly 3 – 5-lobed or -angular or unlobed, in outline subcircular or broadly
ovate, 8 – 20 by 7– 20 cm, scabrous above and beneath, glands several, scattered, 0.5
mm diam. or less, margin sparsely minutely dentate-serrate, apex acute-acuminate,
often with a 10 – 25 mm long acumen; veins 5(–7), straight. Inflorescences unisexual
or possibly sometimes male and female flowers mixed. Male raceme pendent, among
the leaves or on the older stem close to the forest floor, single, unbranched or branched
at the base forming several racemes in a bundle, dark brown hairy; peduncle 0.3(– 2)
cm long, 1.5 – 3 mm thick; rachis not thickened, to 7 cm long, to 30-flowered; bracts
caducous, inserted on the pedicel towards the base, c. 1 mm long. Male flowers: con-
spicuously rusty hairy (pale when living); pedicel 20 – 40 mm long, persistent, above
the insertion of the bract withering into straw-like appendages; receptacle-tube tubular,
50 – 60 mm long, at throat 5(–7) mm wide; sepals linear, 10 –15 mm long, c. 1 mm wide
at base, entire; petals narrowly elliptic, c. 20 mm long, threads c. 25 mm long; synan-
drium 5 – 6 mm long, with c. 0.5 mm long coarse white acute hairs between the thecae,
filaments 3 – 4 mm long, glabrous; disc consisting of 3 line-shaped thickenings at base
of the tube. Female flowers: solitary or several in a simple or compound raceme to 12
cm long; pedicel c. 25 mm long; ovary hairy, (narrowly) fusiform, 10 –12 by 3 – 4 mm,
style c. 35 mm long, glabrous, stigma (4- or)5-lobed, c. 3 mm long. Fruit solitary, ripen-
ing yellow-green, whitish striped, (narrowly) ovoid, 10 –11.5 by 6 –7 cm, apex c. 2 mm
beaked; exocarp leathery; pericarp carnose (c. 15 mm thick when fresh), dry pericarp 5
mm thick; pulp white, finely fibrous, not bitter; fruiting pedicel 3 – 5 cm long, c. 3 mm
thick. Seeds whitish, pink, cream or pale brown, tumid, with two inflated lateral parts,
10 –12 by 17– 20 by c. 5 mm, margin absent, edge entire. — Plate 28.
Field-notes — Flowers white, pendent, in female flowers the receptacle-tube is hori-
zontally curved at anthesis.
Distribution — Malesia: Borneo, endemic to Sabah: Sandakan (Sepilok Forest Re-
serve), near Tawau (Lahat Datu), Luasong).
Habitat & Ecology — In primary and disturbed lowland forest, in depressions, damp
slopes and forest edges; on clayey soil; 50 – 600 m altitude. The flowers open at night and
stay open until noon; flowering and fruiting January to February, and July to August.
Notes — 1. Trichosanthes pendula is isolated within Trichosanthes because of the
prominently ribbed stem, the absent probract, the flowers which are in both male and fe-
male borne in compound, pendulous racemes, forming tassels, the synandrium with many
whitish hairs between the thecae (also in T. mucronata), the (4- or) 5-lobed stigma, the
presence of a disk consisting of 3 line-shaped thickenings adnate to the base of the recep-
tacle tube, and the broad tumid seeds with inflated lateral parts (also in T. mucronata).
2. The older tassel-like pendulous inflorescences frequently appear densely bunched

caused by numerous persistent pedicels, which are withered into straw-like appendages.
Similar appendages, derived from peduncle or pedicel, are found in T. mucronata and
in some species from New Guinea.
3. The whole inflorescence, including the flowers, dries dark rusty brown, but in
living plants all parts, including the hairs, are whitish and the petals pure white.
29. Trichosanthes philippinensis Rugayah

Trichosanthes philippinensis Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 271, f. 8;
Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 115, pl. 9. — Type: Elmer 11067
(holo L; iso BM, K, U), Philippines, Mindanao, Mt Apo.
Climber to 5 m long, glabrescent, leafy stem 2 – 5 mm diam.; dioecious. Probract

ovate or elliptic, 5 – 8 mm long, subentire, glands present. Tendrils 2- or 3-branched.
Leaves: petiole 3 –7 cm long; blade greenish on drying, membranous or chartaceous,
simple, 3(– 5)-lobed, 1/3 – 3/4 deep, in outline ovate or subcircular, 10 –18 by 10 – 20
cm, sometimes faintly scabrous above and beneath, glands few, scattered or towards the
insertion to the petiole, 0.5 –1.5 mm diam., cystoliths obvious, base deeply cordate(-
hastate), margin inconspicuously or coarsely serrate-dentate; veins 5(–7), ± straight,
venation conspicuous beneath. Male raceme at first densely brown hairy; peduncle
5 –7.5 cm long, 2 – 3 mm thick; rachis not thickened but sometimes stout, 5 –12 cm long,
10 – 20-flowered; bracts persistent, broadly obovate-rhomboid, 25 – 45 by 20 – 25 mm,
margin irregularly and finely incised, 5 –15 mm deep, glands numerous, of different
size. Male flowers: pedicel c. 5 mm long, caducous; receptacle-tube c. 30 mm long,
at throat 7–10 mm wide; sepals (narrowly) ovate, 10 –14 mm long, 2 – 5 mm wide at
base, finely and deeply irregularly incised, with few glands; petals (including threads)
c. 30 mm long; synandrium not seen. Female flowers not known. Fruit ripening evenly
red or red and yellow striped, broadly ovoid, 6 –7.5 by 5 – 6 cm, apex c. 1 mm beaked;
exocarp leathery, thin, smooth; pericarp carnose, dry pericarp 10 –15 mm thick; pulp
green-black; fruiting pedicel 2 – 2.5 cm long, 2 – 3 mm thick. Seeds (from PPI 5024)
compressed, obovate or narrowly elliptic, 10 –12 by 4.5 –7 by 2(– 3) mm, base narrowly
truncate with small midlobe, apex pointed, margin faint or absent, edge entire.
Field-notes — Fruit apple-like, red and yellow.
Distribution — Malesia: Philippines: Luzon (Sorsogon; Albay (Mayon Volcano);
Benguet); Panay; Mindanao (Davao (Mt Apo)).
Habitat & Ecology — (Degraded) forest, secondary growth; on clay, clay loam, and
limestone; 500 –1280 m altitude.
Note — Trichosanthes philippinensis vegetatively resembles T. tricuspidata (not in
the Philippines); in T. tricuspidata the male bracts are less deeply incised and the sepals
entire.
30. Trichosanthes pilosa Lour.

De Wilde & Duyfjes (Reinwardtia, 2008) distinguished in T. pilosa two varieties, var.
roseipulpa W.J.de Wilde & Duyfjes (Thailand) and var. pilosa; only var. pilosa occurs
in Malesia.
var. pilosa
Trichosanthes pilosa Lour., Fl. Cochinch. 1 (1790) 588; W.J.de Wilde & Duyfjes, Reinwardtia 12
(2008) 270; Fl. Thailand 9, 4 (2008) 526. — Type lost, Vietnam. Neotype: Bon 4019 (holo P, des-
ignated by De Wilde & Duyfjes, Reinwardtia (2008)), Vietnam.
Trichosanthes ovigera Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 934; Miq., Fl. Ned. Ind. 1, 1 (1856) 674;
(1964) 303; Rugayah & W.J.de Wilde, Blumea 42, 2 (1997) 478; Rugayah, Trichosanthes (Cucur-
bitaceae) in Malesia, thesis (1999) 72; W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu
(2001) 211; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 25; Duyfjes & Pruesapan, Thai Forest
Bull., Bot 32 (2004) 89. — Type: Blume s.n. (holo L, barcode L0130442; iso L (barcode L0130439),
P), Java, Gunung Salak.
Trichosanthes horsfieldii Miq., Fl. Ned. Ind. 1, 1 (1856) 677. — Type: Horsfield s.n. (holo BM; iso
K, U (barcode U0001472)), Java.
Trichosanthes vanoverberghii Merr., Philipp. J. Sci., C 9 (1915) 458. — Type: Vanoverbergh 3662 bis
(iso K), Luzon, subprovince of Bontoc, Bauco.
Trichosanthes mafuluensis Merr. & L.M.Perry, J. Arnold Arbor. 30 (1949) 58. — Type: Brass 5257
(iso BO), Papua New Guinea, Central Province, Mafulu.
Climber to 10 m long, variably hairy to various degree (see note 1 and 2), rarely gla-
brous, leafy stem 1– 3(– 5) mm diam., grooved; dioecious (occasionally monoecious).
Probract absent. Tendrils 2- (or 3-)branched. Leaves: petiole 3 – 8 cm long; blade green
on drying, membranous or chartaceous, simple, unlobed or 3 – 5(–7)-lobed, to 3/4
deep, in outline subcircular, or (broadly) ovate or ± triangular, 5.5 – 21 by 4 – 20 cm,
short-hairy beneath, at least along the (finer) veins (rarely glabrous), glands several or
absent, scattered, 0.5 mm diam., cystoliths sometimes obvious on petiole and veins
beneath, base rarely hastate, margin entire, finely sparsely dentate, or coarsely dentate
or undulate, apex obtuse or acute-acuminate; veins 5, straight or curved. Male raceme
sometimes with a solitary male flower co-axillary at the node, densely or sparsely short
hairy; peduncle (2 –)5 – 8(–17) cm long, 1– 2 mm thick; rachis not thickened, 3 – 4(– 8)
cm long, 5 –10-flowered; bracts (sub)persistent, linear, or narrowly or broadly (ob)ovate
or narrowly elliptic, (2 –)5 –15 by (1–)5 –10 mm (or larger), margin entire, few-lobed
or dentate, glands absent. Male flowers: pedicel (1–)2 –15 mm long, (in solitary flow-
ers much longer), usually persistent; receptacle-tube 20 – 25(– 30) mm long, at throat
3 – 5(– 6) mm wide; sepals narrowly triangular, 3 –7(– 9) mm long, 1– 2 mm wide at base,
entire; petals (narrowly) ovate or (narrowly) elliptic, c. 10 by 4 mm, threads 7–11(–17)
mm long; synandrium 3 – 4 mm long, filaments glabrous (always?), 1– 2 mm long. Fe
male flowers: resembling male flowers, solitary, rarely in a short female raceme; pedicel
5 –17 mm long; ovary hairy, (narrowly) ellipsoid, 10(–15) by c. 3 mm. Fruit ripening
red, usually paler striped, (subglobose to) usually (narrowly) ovoid or narrowly ellip-
soid, 3 –10(–14) by 2.5 – 3.5(– 6) cm, apex acute and 3 – 5 mm beaked; exocarp leathery,
rarely woody, smooth or hispid, c. 0.5 mm thick; pericarp firm-carnose, dry pericarp
5 – 8 mm thick; pulp white (or bright orange-red, not in Malesia), not bitter; fruiting
pedicel 1.5 – 4.5 cm long, 1– 3 mm thick. Seeds tumid by lateral swellings, variously
barrel-shaped, 7–11 by 6 – 8 by 3 – 4 mm, margin absent. — Fig. 91b; Plate 29c.
Distribution — Widespread in SE Asia from NE India and China south-east to Aus-
tralia and the Solomon Islands; in Malesia: Sumatra, Peninsular Malaysia, Singapore,
Borneo (Sabah, eastern Kalimantan), Java, Philippines (Luzon, Mindoro), Sulawesi,
Lesser Sunda Islands (Bali, Flores, Timor), Moluccas (few collections), and all over
New Guinea.
Habitat & Ecology — Primary and secondary forest edges, on hillsides, in grassland,
along riverbanks, in neglected gardens, growing in soil over a variety of bedrock, in-
cluding limestone; lowland to 2000 m altitude.
Uses — Young fruits are edible.
Notes — 1. Trichosanthes pilosa is widespread and particularly variable in indu-
ment, leaf shape, size of bract, male raceme and fruit. The variation is largest in eastern
Malesia and culminates in New Guinea. In general this species is easily recognised by
the tumid seeds and the hairy lower leaf surface (rarely glabrous). The specimens Vano
verbergh 3662-bis, and Merrill 4864 (both from Luzon and in fruit) have the fruits in
short racemes. These fruits are almost globose, 3 – 3.5 by 2.5 – 3 cm, with only c. 2 mm
long fruiting pedicels. Kooy 1359, from Timor, has scabrous leaves, and small fruits
with a rather woody and partly hispid-hairy exocarp.
2. The variability of T. pilosa in New Guinea is unusually large, but extremes appear
not readily to be segregated. Some are represented by the following choice of speci-
mens:
Karenga & Baker LAE 56930 from high altitude (1960 m) is illustrative of specimens
with very scabrous, deeply lobed leaves, and large elongate fruit, 10 –14 cm long.
Höft 2104, Streimann 8301 and Streimann & Kairo NGF 27797 (1250, 850, and 1500
m respectively) are illustrative of similar plants, with smaller fruits which are ± hispid
hairy.
Koster BW 4326 from lowland Jayapura area, West Papua, deviates by large, densely
packed, coarsely dentate, male bracts, measuring 25 by 25 mm.
Henty & Lelean NGF 49912, from lowland (400 m), has remarkably crenate-dentate
leaf margins.
Brass 5257 from eastern Papua New Guinea, 1250 m altitude, is the type of T.
mafuluensis, and represents a form with unlobed leaves and very narrow male bracts.
Similar linear male bracts have e.g. Stevens & Veldkamp LAE 55741 or Womersley &
Millar NGF 8472, both from high altitudes (c. 1700 m), but with deeply lobed leaves.
3. The root is sometimes tuberous.
31. Trichosanthes planiglans Rugayah

Trichosanthes planiglans Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 273, f. 9;
Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 133, pl. 14. — Type: Ramos &
Edaño BS 29110 (holo US; iso K), Philippines, Luzon, Mt Tulaog.
Climber, glabrous, leafy stem 2 – 3 mm diam.; dioecious. Probract ovate, somewhat

concave, c. 5 mm long, entire, with few glands. Tendrils 2-branched. Leaves: petiole
2.5 – 5 cm long; blade green on drying, membranous, simple, unlobed, in outline broadly
ovate, 9 – 20 by 8 –16 cm, glands 4 – 8, large, 2 – 5 mm diam., situated towards the inser-
tion of the petiole, cystoliths on stem, petioles and veins beneath, margin conspicuously
coarsely serrate-dentate, teeth 5 –12 mm long, apex with slender acumen to 5 mm long;
veins 5, arching. Male raceme and female flowers not known. Fruit ovoid? (precise
shape and size not known); exocarp ± leathery, smooth; pericarp rather thin, pulp
possibly green-black; fruiting pedicel c. 2 cm long, c. 4 mm thick. Seeds light brown,

compressed, narrowly elliptic, 13 –16 by 6 –7 by 1.5 mm, base cuneate, apex rounded,
margin faint, edge entire or indistinctly crenulate.
Distribution — Malesia: Philippines (Luzon, where known only from the type col-
lection).
Habitat & Ecology — Unknown. The specimen was collected in May.
Notes — 1. Trichosanthes planiglans seems related to T. quinquangulata, with simi-
lar seeds with a cuneate base, but the latter differs in the leaves with ± an entire margin,
smaller and more numerous glands, and a blackish drying colour.
2. Ramos & Edaño BS 49161 from Mindanao, which apparently belongs to T. quin
quangulata resembles T. planiglans in its coarsely dentate leaf margin; the specimen
also deviates from T. quinquangulata in ± lobed bracts.
32. Trichosanthes postarii W.J.de Wilde & Duyfjes

Trichosanthes postarii W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 26, f. 7– 9. — Type: Postar et
al. SAN 144066 (holo SAN; iso L), Borneo, Sabah, Maliau Valley.
Trichosanthes sp. 1, W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu (2001) 212, pl. 17: C.
Much branched spacious liana, 10 – 20 m long, scabrous pale brown hairy, hairs 1 mm
long; leafy stem 5 mm diam.; dioecious. Probract absent. Tendrils 2- (or 3-)branched,
finely harshly hairy. Leaves: petiole 4 – 4.5 cm long; blade green on drying, membra-
nous, simple, unlobed or in juvenile or shooting leaves 5-angular or -lobed, to c. 1/4
deep, in outline ovate-oblong, 9 –22 by 6 –13 cm, finely scabrous, glands many, small,
scattered, cystoliths often present, base at the transition to the petiole at each side with a
small bulging auricle, 3 – 4 mm diam., each inside with 3 – 5 crowded glands, blade mar-
gin finely dentate, teeth 0.5 –1 mm long; basal veins 5(–7). Inflorescences on leafless
shoots hanging down close to the forest floor. Male inflorescence usually formed below
the leaves, consisting of 1 or 2 (or 3) racemes, fascicled, horizontal or drooping, pedun-
cle short, rachis slender, 4.5 –12 cm long, 1(–1.5) mm thick, 10 – 20-flowered; bracts
subpersistent, shifted upwards on the pedicel for 1– 3 mm, narrowly elliptic, 5 – 8 mm
long, margin coarsely shallowly dentate, glands several, near the margin. Male flowers:
pedicel 6 –10 mm long, persistent; receptacle-tube ± urceolate, 15(– 20) mm long, at
throat 6(– 8) mm wide, consisting of a ± campanulate upper half and a swollen lower
half, inside demarcated by a diaphragm-like raised rim; sepals (long-)triangular, 2 – 3
mm long; petals in bud folded into a ± conical body with free apices, 10 –12 mm long,
expanded petals broadly cuneate, c. 10 mm long, threads 6 – 8 mm long; synandrium
truncate, c. 3.5 by 2.5 mm, anthers tightly appressed, but not fused, filaments glabrous,
with few hairs on thickened base, inserted halfway on the receptacle-tube where it is ±
contracted; disc absent. Female flowers 1 or 2 in short raceme; pedicel c. 10 mm long;
ovary subglabrous, ellipsoid, c. 15 by 6 mm; receptacle-tube c. 12 mm long; sepals tri-
angular, c. 2 mm long. Fruit on leafless shoots, one or two per infructescence, ripening
green, white-striped, narrowly ellipsoid, 13 –16 by 4.5 – 5.5 cm, base subobtuse, apex
acute, pericarp thin, pulp white, bitter; fruiting pedicel 2.5 – 3 cm long, 4 – 5 mm thick.
Seeds compressed, circular, 10 mm diam., margin broad, radiately ribbed. — Fig. 84,
85; Plate 29a, b.
2 cm 2 cm
e
2 mm
2 mm
d
2 mm
g c 2 mm
j j'
1 cm
3 mm
f h i
Fig. 84. Trichosanthes postarii W.J.de Wilde & Duyfjes. a, b. Leafy twigs; c, d. base of leaf blades
showing auricles, note glands on lower surface; e. young developing leaf; f. male inflorescences on
older twig; g. male bract; h. male flower; i. ditto, opened showing free stamens; j. stamen, 2-thecous;
j'. stamen, 1-thecous (all: De Wilde & Duyfjes SAN 139470).
b a 2 mm
1 cm
2 mm 1 cm
c d
Fig. 85. Trichosanthes postarii W.J.de Wilde & Duyfjes. a. Female inflorescences; b. infructescence;
c. seed; d. seedling (a, c – d: Postar, Ubaldus & De Wilde SAN 144110; b: Postar, Ubaldus & De Wilde
SAN 144066, type).
Distribution — Malesia: Borneo (endemic to Sabah where known from Poring (Lan-

ganan Waterfall) and Maliau Valley).
Habitat & Ecology — A species of primary forest and stony slopes with half open
forest on wet soil; 50 –1300 m altitude. Flowering specimens were collected in July and
December, fruiting specimens in July.
Notes — 1. Pendulous shoots, mostly without leaves but with only flowers, are found
in the shady zone near the forest floor. Sterile shoots often creep for some distance over
the ground, a habit also known from other species, e.g. T. villosa.
2. The single or 2- (or 3-)fascicled male racemes at a node indicate that these racemes
actually may be borne on a very short lateral shoot, similar to that in T. pendula. In most
Trichosanthes species there is only one single male raceme at a node, with or without
a single co-axillary male flower.
33. Trichosanthes pubera Blume

Duyfjes & Pruesapan (2004) distinguished in T. pubera two subspecies, subsp. rubiflos
(Cayla) Duyfjes & Pruesapan (Indochina) and subsp. pubera; only subsp. pubera oc-
curs in Malesia.
subsp. pubera
Trichosanthes pubera Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 936; Rugayah & W.J.de Wilde, Blumea 42,
2 (1997) 479, f. 1c, 2c, 3c; Reinwardtia 11, 4 (1999) 273; Rugayah, Trichosanthes (Cucurbitaceae)
in Malesia, thesis (1999) 117, map 14; Duyfjes & Pruesapan, Thai Forest Bull, Bot. 32 (2004) 92;
W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 30; Fl. Thailand, 9, 4 (2008) 531. — Trichosanthes
bracteata (Lam.) Voigt var. pubera (Blume) Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 377.
— Type: Blume s.n. (lecto L, barcode L0130546, here designated; iso P), Java.
Climber to 20 m long, grey-rusty hairy, partly glabrescent, leafy stem 2 – 5 mm diam.,

5 –7-ribbed, growing stem apex reddish tinged; dioecious. Probract narrowly elliptic or
linear, 10 – 20 by c. 2 mm, at apex entire or finely lobed and ± hooded, glands absent.
Tendrils 2- or 3-branched. Leaves: petiole 3 – 9.5 cm long; blade reddish brown on
b'
1 cm a'
1 cm
1 cm
b
1 cm 1 cm
c a
c'
1 cm
Fig. 86. Leafy stem nodes and corresponding flower bracts of male inflorescences of three species of
Trichosanthes in Java. — a, a'. T. quinquangulata A.Gray; b, b'. T. tricuspidata Lour. subsp. javanica
Duyfjes & Pruesapan; c, c'. T. pubera Blume subsp. pubera (a, a': De Wilde & Duyfjes 21772; b, b':
De Wilde & Duyfjes 21777; c, c': Blume s.n., barcode L0130546).
drying, membranous, simple, shallowly or deeply 3 – 5(–7)-lobed to 1/3 – 4/5 deep, in

outline circular or broadly ovate, 9 – 23 by 8 – 22 cm, scabrous above, finely grey-rusty
hairy beneath, glands few or several, towards the insertion of the petiole, 0.5 –1 mm
diam., cystoliths abundant on petiole and blade, margin sparsely minutely dentate, oc-
casionally at base coarsely dentate; veins 5(–7), palmate, straight. Male raceme rusty
hairy; peduncle 7–15 cm long, 2 – 3 mm thick; rachis not thickened, 7– 8 cm long,
5 –15-flowered; bracts subpersistent, obovate-rhomboid, (20 –)30 – 50 by 20 – 35 mm,
margin irregularly incised or slenderly lobed to (2 –)5 –10 mm deep, glands absent.
Male flowers: pedicel 5 –10 mm long, subpersistent; receptacle-tube 40 – 50 mm long,
at throat 8 –10 mm wide; sepals long-triangular or narrowly elliptic, 15 – 22 mm long,
5 –7 mm wide at base, with few sidelobes to 5 mm long (entire in female flowers); petals
white, ± obovate, 15 – 20 mm long, threads pinkish at apex, 5 –10 mm long; synandrium
10 –13 mm long, filaments glabrous, c. 4 mm long. Female flowers: pedicel (10 –)25 – 50
mm long; ovary densely hairy, (narrowly) ellipsoid, c. 15 by 4 – 5 mm; receptacle-tube
30 – 40 mm long, at throat 6 –7 mm wide; sepals narrowly triangular, (10 –)15 mm long,
entire; petals 12 –14 mm long, threads c. 5 mm long; style c. 25 mm long, glabrous,
stigma c. 15 mm long. Fruit ripening evenly orange-red, early glabrescent, broadly
ovoid-ellipsoid, 5 –10 by 4 – 6(– 8) cm, apex 2 – 5 mm beaked; exocarp leathery; pericarp
firm-carnose, dry pericarp 5 –10 mm thick; pulp green-black, bitter; fruiting pedicel
(1.5 –)2 – 5.5 cm long, 3 – 4 mm thick. Seeds blackish, compressed, (narrowly) elliptic,
or obliquely obovate, 8 –13 by 4 – 5.5(– 8) by 1– 2 mm (but see note), base rounded or
slightly cuneate, apex rounded, margin absent, edge entire. — Fig. 86c, c'.
Field-notes — Young shoots purple-red, stem angular, leaves bright green. Flowers
open at night, strongly fragrant; petals white with pale-pink threads. Fruit pendent, el-
lipsoid, orange-red, seeds surrounded by greenish black pulp.
Distribution — Malesia: Sumatra, Borneo (Sabah (Maliau Valley) and East Kali-
mantan), Java.
Habitat & Ecology — Primary or secondary forest edges, ravines, riversides, scrub
in wet sites; 50 –1200 m altitude. Flowering throughout the year.
Notes — 1. The collection Koorders 23006 (East Java), is strongly deviating. It con-
sist of one large fruit (c. 10 by 8 cm with a 5.5 cm long fruiting pedicel), with exceed-
ingly large seeds (18 –19 by 8 –10 by c. 4 mm). It may represent a separate taxon.
2. The collections Korthals 23 (slender plants from Central Sumatra), and Korthals
97 (stout plants from East Kalimantan), somewhat deviate in having glabrous leaves.
34. Trichosanthes pulleana Harms

Trichosanthes pulleana Harms, Bot. Jahrb. Syst. 60 (1925) 160; Rugayah & W.J.de Wilde, Rein-
pl. 24. — Trichosanthes papuana Pulle, Nova Guinea 8 (1910) 406, nom. inval. (non T. papuana
F.M.Bailey); Ridl., Trans. Linn. Soc. London Bot. 9 (1916) 58 (record based on Boden Kloss s.n.,
not seen). — Type: Versteeg 1116 (holo L; iso BO, K, U), West Papua, Noord Rivier.
Climber, possibly few metres long only, at first with short brown hairs, glabrescent,
leafy stem 2 – 4 mm diam.; dioecious. Probract elliptic, 2 – 3 mm long, with or without
glands. Tendrils 2-branched. Leaves: petiole 2.5 – 5 cm long; blade green on drying,
chartaceous or subcoriaceous, simple, unlobed, in outline (narrowly) ovate or sub-

hastate, 12 – 22 by 8 –14 cm, finely scabrous above, glands several, scattered, small, at
the leaf base and near the midrib or apex, cystoliths obvious but small, base with deep
sinus, margin finely dentate; veins 5, basally, and few curved ones from the midrib.
Male raceme densely short brown hairy, hairs 1–2 mm long; peduncle 2 – 5 cm long,
2 – 3 mm thick; rachis not thickened but stout, with bract-scars, 8 –13 cm long, 3 mm
thick, more than 20-flowered; bracts late-caducous ovate or narrowly elliptic, 6 –10
by 3 – 5 mm, entire, glands present. Male flowers in the raceme and usually with one
solitary flower at base; pedicel 2 – 5(–10) mm long (in solitary flower 25 – 40 mm long
and withering into a straw-like appendage), subpersistent or caducous; receptacle-tube
40 – 55 mm long, at throat 7– 8 mm wide; sepals (narrowly) triangular, 4 – 6 mm long,
2 – 3 mm wide at base, acute, entire; petals obovate- rhomboid, strongly nerved, c. 13
by 6 –10 mm, threads 7–10 mm long; synandrium 7 mm long, filaments glabrous, 1 mm
long. Female flowers and fruit not known.
Distribution — Malesia: Moluccas (Aru Islands), New Guinea (West Papua).
Habitat & Ecology — Primary forest, on coral stone; at low altitudes.
35. Trichosanthes quinquangulata A.Gray

Trichosanthes quinquangulata A.Gray, U.S. Expl. Exped., Phan. 1 (1854) 645; Rugayah & W.J.de
Wilde, Blumea 42, 2 (1997) 479, f. 1a, 2a, 3a; Reinwardtia 11, 4 (1999) 273; Rugayah, Trichosan-
thes (Cucurbitaceae) in Malesia, thesis (1999) 130, pl 3; W.J.de Wilde & Duyfjes in Beaman et
al., Pl. Mt. Kinabalu (2001) 212; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 30; Duyfjes &
Pruesapan, Thai Forest Bull., Bot. 32 (2004) 96; W.J.de Wilde & Duyfjes, Fl. Thailand, 9, 4 (2008)
532. — Type: Wilkes s.n. (holo GH-A), Philippines, Mangsi Is., Sulu Sea.
Trichosanthes longiflora Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 374. — Type: Beccari 223
(holo FI), West Papua (Sorong).
Trichosanthes reineckeana Cogn., Bot. Jahrb. 25 (1898) 691. — Type: Reinecke 84 (lecto W, here
designated), Samoa, Upolu.
Climber to 20 m long, glabrous, leafy stem 2 – 5 mm diam., young shoots green; dio-
ecious or occasionally monoecious. Probract (narrowly) obovate, 5 –10(– 20) mm long,
entire, glands present. Tendrils (2 –)3 – 5-branched. Leaves: petiole 2.5 –11 cm long;
blade blackish on drying, membranous, simple, unlobed or shallowly 5(–7)-angular or
(shallowly) lobed, lobes (shallowly) triangular, blade in outline subcircular, 8.5 – 24 by
8 – 21 cm, ± scabrous above, glands several to many, close to the vein axils, particularly
several in the axils of the main veins at the very base of the blade, 0.5 –1 mm diam.,
cystoliths sometimes obvious above, margin entire or coarsely remotely dentate; veins
5 –7, straight. Male raceme sometimes at base with 1 or 2 female or hermaphroditic
flowers (see note in Rugayah 1999), glabrescent; peduncle 2 – 8(–12) cm long, 2 – 3
mm thick; rachis not thickened, 3 –10 cm long, 2 – 3 mm thick, 5 –10-flowered; bracts
subpersistent, (ob)ovate-elliptic or ± rhomboid, 10 – 30 mm long, margin (sub)entire
(rarely ± dentate in East Malesia), with glands near the midrib. Male flowers: pedicel
2 – 5(– 25) mm long, (sub)caducous; receptacle-tube 40 – 50 mm long, at throat 10 mm
wide; sepals long-triangular, 12 – 20(– 30) mm long, 2 – 3 mm wide at base, usually with
few slender sidelobes; petals obovate or broadly cuneate, 20(– 30) mm long, threads
10(–15) mm long; synandrium 8 mm long, filaments glabrous, 3 mm long. Female
flowers: pedicel (5 –)10 – 20 mm long; ovary glabrous, (ovoid-)ellipsoid, 8 –10 by 5 – 6

mm; receptacle-tube 50 – 60 mm long; sepals smaller and narrower than in male, entire.
Fruit ripening evenly red, (depressed) globose, rarely slightly ellipsoid, (4.5 –)6 – 8.5
cm diam., apex not beaked; exocarp leathery; pericarp firm-carnose, dry pericarp 8 –15
mm thick; pulp green-black, bitter; fruiting pedicel 1.5 – 2.5 cm long, 3 – 5 mm thick,
usually curved. Seeds light brown, compressed, (narrowly) elliptic, 9 –14 by 4 – 5 by
1– 2 mm, base cuneate (chisel-shaped), apex rounded or acute, margin absent, edge
entire. — Fig. 86a, a', 91c; Plate 31a.
Field-notes — Flowers with sweet smell, calyx green, bracts pale green, fruit glo-
bose, juicy, red-orange when ripe, frequently found hanging on the already died-off
trailing shoots.
Distribution — South China, Myanmar, Thailand, Vietnam, Cambodia, Laos; in
Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Philippines, Moluc-
cas, New Guinea (West Papua and Papua New Guinea where known from 1 collection,
Morren 68).
Habitat & Ecology — Secondary or disturbed forest edges, scrub, open area in mixed
dipterocarp forest, logged over forest, along lake shores, in swampy areas, on loamy and
sandy soil and beaches, also on limestone; from sea level to 1250 m altitude. Flowering
and fruiting throughout the year.
Notes — 1. Ramos & Edaño BS 49161, from Mindanao, somewhat resembles T. pla
niglans; see the note under that species.
2. In most Trichosanthes species the corolla in bud is hemi-globose, but in T. quin
quangulata, and in some other species, it is ± elongate, like in the related genus Gym
nopetalum.
3. Specimens from limestone area West of Sandakan (Sabah) deviate by having larger
probracts (15 – 20 mm long), and slightly elongate fruit.
36. Trichosanthes refracta C.H.Yueh

Trichosanthes refracta C.H.Yueh, Bull. Bot. Res., Harbin 10, 4 (1996) 500, f. 1; Rugayah & W.J.de
(1999) 94. — Type: Ramos BS 43279 (holo UC; iso K, L), Philippines, Bohol, Valencia.
Climber to 4 m long, glabrous, leafy stem 2 – 4 mm diam.; dioecious. Probract linear,

5 mm long, glands absent. Tendrils 2-branched. Leaves: petiole 3 – 4 cm long; blade
green on drying, membranous, simple, deeply 5 –7-lobed, in outline subcircular, 7– 21
by 8 – 22 cm, not or little scabrous above, glands few (1– 3), ± close to the insertion of
the petiole, 0.5 – 3 mm diam., cystoliths absent, margin entire or remotely shallowly or
coarsely dentate; veins 5(–7), straight. Male raceme minutely brown hairy; peduncle
5 –10 cm long, 2 – 3 mm thick; rachis not thickened, zigzag between the bracts, 4 –13
cm long, 1.5 – 3 mm thick, 10 –15-flowered or more; bracts persistent or caducous,
obovate-rhomboid, 10 – 40 by 8 –15 mm, margin sharply incised to 3 mm deep, glands
absent. Male flowers: pedicel 1– 3 mm long, persistent; receptacle-tube 40 –70 mm long,
at throat 10 –13 mm wide, at base with an ovary-like swelling c. 5 mm long (pseudo-
ovary, see note under T. intermedia); sepals narrowly elliptic, 4.5 –7 mm long, 1.5 – 2
mm wide at base, entire or with few narrow sidelobes; petals short, fully expanded not
seen; synandrium c. 9 mm long, filaments glabrous, c. 3 mm long. Female flowers:

pedicel c. 20 mm long; ovary glabrous, ellipsoid, c. 8 by 6 mm; perianth not known.
Fruit ripening (possibly) green, paler or yellowish striped, ellipsoid, c. 6 by 4.5 – 5 cm,
apex not beaked; exocarp leathery, colour of fruit pulp not known; fruiting pedicel c. 1.4
cm long, 2 – 3 mm thick. Seeds compressed, elliptic, c. 12 by 6 by 2 mm, base truncate,
apex rounded, margin faint, edge entire.
Distribution — Malesia: Borneo (Brunei (Belait, Tutong); Sarawak (Daro Forest Re-
serve)), Philippines (Bohol).
Habitat & Ecology — Peat swamp forest, edges of swampy forest; lowland.
37. Trichosanthes rotundifolia Rugayah

Trichosanthes rotundifolia Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 3 (1998) 223; Ru-
gayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 275, f. 10; Rugayah, Trichosanthes (Cucurbita-
ceae) in Malesia, thesis (1999) 156, pl. 20. — Type: Lörzing 5284 (holo BO; iso L) North Sumatra,
Sibolangit.
Climber, 5 –15 m long, glabrescent, at first with few hairs, leafy stem 2 – 3 mm diam.;
dioecious. Probract narrowly elliptic, 5 –7 mm long, acute, subentire, glands present.
Tendrils 2-branched. Leaves: petiole 3.5 – 8 cm long; blade (green-)brown on drying,
membranous, simple, unlobed, in outline subcircular or broadly ovate, 6 – 21 by 6 –18
cm, faintly scabrous, glands few or several, scattered at the blade base and towards the
midrib, c. 0.5 mm diam., cystoliths obvious on blade and veins, margin entire or sparsely
minutely dentate. Male raceme at first minutely grey-brown hairy, glabrescent; peduncle
3 – 6.5 cm long, 1.5 mm thick; rachis not thickened, 1– 4 cm long, 5 –10-flowered; bracts
subpersistent, (narrowly) obovate, (3 –)7–10 mm long, margin entire or shallowly few-
lobed, glands present. Male flowers in the raceme and sometimes one solitary flower
co-axillary; pedicel 5 –15 mm long, (in solitary flower 50 – 80 mm long), persistent;
receptacle-tube 15 – 25 mm long, at throat c. 5 mm wide; sepals narrowly triangular,
c. 4 mm long, entire; petals obovate-rhomboid, c. 10 mm long, threads c. 5 mm long;
synandrium 3 – 4 mm long, filaments glabrous, c. 1.5 mm long. Female flowers: resem-
bling male, pedicel 10 – 30 mm long; ovary minutely hairy, c. 10 by 3 mm, style c. 13
mm long, stigma c. 2 mm long. Fruit green, pale yellow striped and speckled, ripen-
ing evenly(?) red, (narrowly) ovoid, 8 –10 by 5 – 6 cm, apex c. 3 mm beaked; exocarp
thick-leathery, smooth; pericarp firm-carnose, dry pericarp c. 10 mm thick; pulp not
known; fruiting pedicel 1– 3.2 cm long, 2 – 3 mm thick. Seeds pale brown, compressed,
subcircular or broadly elliptic, c. 20 by 15 by 4 mm, base and apex broadly rounded,
margin faint, inside hollow, chambered, edge entire. — Fig. 87.
Field-notes — Fruit ovoid, with green or pale lengthwise stripes and many bright
green speckles; eaten by orang utan.
Distribution — Malesia: Sumatra.
Habitat & Ecology — Primary forest, river sides, scrub, in foot hill and lower mon-
tane forest; 250 –1200 m altitude. Flowering March–September; fruiting July & De-
cember.
2 cm
Fig. 87. Trichosanthes rotundifolia Rugayah. Portion of twig with male inflorescences, mark the pro-
bracts at the nodes (De Wilde & Duyfjes 12412).
38. Trichosanthes schlechteri Harms

Trichosanthes schlechteri Harms, Bot. Jahrb. Syst. 60 (1925) 159; Rugayah & W.J.de Wilde, Rein-
map 25. — Type: Gjellerup 718 (lecto BO, designated by Rugayah & De Wilde (1999); iso L),
West Papua, River Tor.
d 1 cm
1 cm
1 mm
b
1 cm
a
Fig. 88. Trichosanthes schlechteri Harms. a. Portion of leafy stem with fruit; b. leaf; c. node with male
inflorescence, with co-axillary a male flower; d. ditto, with co-axillary a male flower pedicel developed
into a straw-like appendage; e. seed (a, e: Takeuchi & Ama 16269; b: Takeuchi & Ama 15663; c: Shea
1464; d: Van Royen & Sleumer 6183).
Climber, 4 –7 m long, glabrous, leafy stem 2 – 4 mm diam.; dioecious. Probract (nar-

rowly) obovate, 4 –7 by 2 – 4 mm, dentate or entire, glands present. Tendrils (unbranched)
or 2-branched. Leaves: petiole (1–)2 – 3(– 5) cm long; blade green on drying, chartaceous
or coriaceous, simple, unlobed, in outline (narrowly) ovate or hastate, (6 –)9 – 20 by 9(–
11) cm, scabrous above, glands absent or few at the leaf base, 0.5 mm diam., cystoliths
obvious or not, base broadly rounded, or truncate or hastate, margin spiny dentate, espe-
cially towards the base; veins 3(– 5), basal, and few from the midrib, curved, veins raised
beneath. Male raceme with a solitary flower co-axillary, glabrous (corolla bud minutely
hairy); peduncle 1.5 – 4 cm long, 1– 2 mm thick, together with the long pedicel of the sol-
itary flower withering into straw-like appendages, 2 – 4 mm thick; rachis somewhat zig-
zag, not thickened, 3 – 4 cm long, 1 mm thick, 5 – 8-flowered; bracts obovate-rhomboid
or narrowly elliptic, 8 –13 by 5 – 8 mm, margin sharply toothed or irregularly few-lobed,
glands present. Male flowers: pedicel 3 –10 mm long, caducous; receptacle-tube c. 50
mm long, at throat 7(–10) mm wide; sepals narrowly triangular or narrowly elliptic, 3 –10
mm long, 1– 3 mm wide at base, entire; petals obovate, 15 – 20 mm long, threads short,
c. 5 mm long; synandrium c. 8 mm long, filaments not seen. Female flowers not known.
Fruit green, paler striped, ripening orange-red, ovoid or ellipsoid, 6.5 – 20 by 4.5 –7.5
cm; exocarp leathery or woody, smooth; pericarp firm-carnose, dry pericarp 10 –13 mm
thick; pulp red; fruiting pedicel c. 2 cm long, 3 – 5 mm thick. Seeds blackish, compressed,
(narrowly) elliptic, 16 –18 by 8 –10 by 3 – 4 mm, base and apex subtruncate, sometimes
± notched or undulate, margin broad, c. 3 mm wide, edge entire. — Fig. 88.
Field-notes — Leaves yellowish green above, light green beneath. Flowers large,
open at night only. Fruit orange or red when ripe, smooth, ellipsoid, pericarp inside
yellow. Seeds blackish, immersed in scarlet pulp.
Distribution — Malesia: New Guinea (West Papua (Bird’s Head, Cycloop Mts, PT.
Freeport); Papua New Guinea (Morobe, East Sepik and Central Provinces)).
Habitat & Ecology — Rain and riverine forests, lowland regrowth forest, open
stream sides, hill forest, secondary forest, roadside vegetation, muddy and sandy soils;
lowland up to 1300 m altitude.
Uses — Fruits edible.
Notes — 1. The shape of the corolla in bud is elongated, like in T. quinquangulata
and in the related genus Gymnopetalum; in other Trichosanthes species the corolla in
bud is hemi-globose.
2. See also the footnote under the regional key.
39. Trichosanthes sepilokensis Rugayah

Trichosanthes sepilokensis Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 275; Ru-
gayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 146; W.J.de Wilde & Duyfjes in
Beaman et al., Pl. Mt. Kinabalu (2001) 212; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 30.
— Type: De Wilde & Duyfjes SAN 139042 (holo SAN; iso L), Borneo, Sabah, Sepilok area.
Climber, 15 – 30 m long, thinly grey-brown hairy, early glabrescent, leafy stem 2 – 4

mm diam.; monoecious (always?). Probract chartaceous, (narrowly) elliptic, 3 – 4 by
c. 2 mm, with few glands. Tendrils 2 – 6-branched. Leaves: petiole 5 – 8 cm long; blade
(dark) brown on drying, membranous, finely bullate above, simple, shallowly 3-lobed
to 1/3 deep or unlobed, in outline ovate, 11–15 by 6 –11 cm, glands few, scattered, 0.5
mm diam., base cordate, sometimes cordate-hastate, margin entire; veins 5(–7), arching
or straight. Male raceme thinly brown hairy; peduncle 3 – 6 cm long, c. 3 mm thick;
rachis thickened, 12 –15 cm long, 5 – 6 mm thick, 5 – 20-flowered or more; bracts sub-
persistent, withering, ± obovate, 25 – 30 by c. 20 mm, apex coarsely irregularly incised,
glands absent. Male flowers: pedicel 2 mm long, caducous; receptacle-tube (30 –)40
mm long, at throat 10 mm wide; sepals narrowly triangular, 10(–12) mm long, 4 – 5
mm wide at base, acute, entire; petals ± obovate, c. 15 mm long, with slender threads;
synandrium and filaments not seen. Female flowers: pedicel c. 20 mm long; ovary
glabrous, ellipsoid, c. 8 mm long; receptacle-tube slender; sepals and petals as in male.
Fruit yellow-green, finely paler speckled, ripening evenly red, subglobose, 10 –13 cm
long; exocarp woody, nearly 1 mm thick, smooth; pericarp firm-carnose, creamy, dry
pericarp c. 15 mm thick; pulp green-black, bitter; fruiting pedicel 3.5 – 4.5 cm long,
15 – 20 mm thick. Seeds brown or blackish, compressed, (narrowly) elliptic, 13 –18 by
7– 9 by 3 – 4 mm, margin absent, edge entire. — Plate 30b.
Field-notes — Tall climber, main stem to 2.5 cm diameter. Foliage dense, drooping
in festoons high-up from the supporting tree. Leaves with dull greyish haze, finely
bullate. Fruits bright red when ripe. Seeds near-black, embedded in black pulpy very
bitter mass. In Maliau Valley a tree, bordering an old logging road and supporting this
climber with ripe fruits, was blown down in a thunderstorm and all fruits were eaten
by banteng according to the footprints.
Distribution — Malesia: Borneo (Sabah (Sepilok Forest Reserve and Maliau Valley)).
Habitat & Ecology — Climber in tall primary forest trees, where flowering and
fruiting high-up in the canopy; at low altitudes. Fruiting in January, February and July,
flowering in July.
40. Trichosanthes tricuspidata Lour.

Trichosanthes tricuspidata Lour., Fl. Cochinch. 2 (1790) 589; Keraudren in Aubrév. & J.-F.Leroy, Fl.
Cambodge, Laos & Vietnam 15 (1975) 81, p.p. (excl. T. pubera, T. bracteata, T. palmata, T. quin
quangulata), pl. 14, f. 1, 4 –7. — Type: Loureiro s.n., lost, Vietnam. Neotype: J. & M.S. Clemens
3267 (holo P, designated by Keraudren (1975); iso BM), Vietnam, Quang Nam, Da Nang.
Duyfjes & Pruesapan (Thai Forest Bull, Bot. (2004) distinguished in T. tricuspidata two
subspecies, subsp. tricuspidata (Indochina) and subsp. javanica Duyfjes & Pruesapan;
only subsp. javanica occurs in Malesia.
subsp. javanica Duyfjes & Pruesapan

Trichosanthes tricuspidata Lour. subsp. javanica Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32
(2004) 99; W.J.de Wilde & Duyfjes, Fl. Thailand 9, 4 (2008) 535. — Type: De Wilde & Duyfjes
21777 (holo L; iso BO, K, L), Java, Gunung Bunder.
Trichosanthes tricuspidata Lour., p.p. (excluding subsp. tricuspidata): Blume, Bijdr. Fl. Ned. Ind. 15
(1826) 935; Miq., Fl. Ned. Ind.1, 1 (1856) 676; Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881)
374; Ridl., Fl. Malay Penins. 1 (1922) 844; Rugayah & W.J.de Wilde, Blumea 42, 2 (1997) 481, f.
1c & c', 2b, 3b; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 120, pl. 10;
Trichosanthes tricuspis Miq., Fl. Ned. Ind. 1, 1 (1856) 679. — Type: Horsfield s.n. (holo K), Java.
Trichosanthes bracteata auct. non (Lam.) Voigt: Backer in Backer & Bakh.f., Fl. Java 1 (1964) 304, p.p.
2 cm g
c 5 mm
f
b
2 mm
2 cm
Fig. 89. Trichosanthes tricuspidata Lour. subsp. javanica Duyfjes & Pruesapan forma javanica. a. Portion
of branch with male inflorescences; b, c. male flower from outside and opened; d. node with female flower
bud; e. node with fruit; f. seed, face and side view; g. leaves of juvenile branch; h. portion of older stem
(a: De Wilde & Duyfjes 21944 (Lombok); b, c: De Wilde & Duyfjes 21811 (Java); d: De Wilde & Duyfjes
21863 (Java); e, f, h: De Wilde & Duyfjes 21660 (Java); g: Korthals s.n., barcode L0130970 (Java)).
Climber to 20 m long, glabrous, leafy stem 2 – 4 mm diam.; dioecious or occasionally

monoecious. Probract obovate or elliptic, 5(–10) mm long, glands present. Tendrils 2- or
3-branched. Leaves: petiole 2 –7.5 cm long; blade (dark) brown on drying, membranous
or chartaceous, simple, 3(– 5)-lobed, 1/3 – 3/4 deep (unlobed in forma siberutensis), in
outline circular or broadly ovate, 4 –17 by 4 –18.5 cm, glabrous, usually scabrous above,
glands several, scattered, 0.5 –1 mm diam., cystoliths obvious or not, margin entire,
sparsely with very small dents or ± coarsely dentate; veins 3(– 5), straight; lobes gen-
erally triangular, lateral lobes often somewhat out-curved. Male raceme thinly brown
hairy, partly glabrescent; peduncle 5 –10 cm long; rachis not thickened, 10 –15 cm long,
1.5 – 3 mm thick, 3 – 20-flowered; bracts late-caducous, obovate-elliptic or rhomboid,
(10 – )20–30 mm long, margin dentate or coarsely shallowly (deeply) laciniate, 2 –7 mm
deep, glands present. Male flowers: pedicel c. 5 mm long, caducous; receptacle-tube
40 – 60 mm long, at throat 6 –10 mm wide; sepals narrowly triangular, 8 –12 mm long,
3 – 4 mm wide at base, (sub)entire, rarely (Peninsular Malaysia) with narrow sidelobes
or broadly triangular (see note); petals broadly obovate-rhomboid, 20 mm long, threads
20 – 25 mm long; synandrium (Java) c. 15 mm long, filaments subglabrous, c. 5 mm
long. Female flowers occasionally some in the male raceme; pedicel 5 –12 mm long;
ovary glabrescent, (narrowly) ellipsoid, 10 –12 by 4 – 5 mm; sepals and petals as in male,
but shorter. Fruit ripening evenly (orange-)red, (narrowly) ovoid, (5 –)7– 9 by (4 –)5 – 6
cm, apex c. 5 mm beaked; exocarp leathery or woody, smooth; pericarp firm-carnose,
dry pericarp 5 –10 mm thick; pulp green-black, bitter; fruiting pedicel 0.5 – 2.5 cm long,
3 – 9 mm thick. Seeds dark brown, compressed, obovate or narrowly elliptic, smooth,
10 –13 by (4–)5 –10 by 1– 3 mm, margin absent, edge entire.
Field-notes — Fruit bright orange-red, pericarp inside pale creamy yellow, c. 1 cm
thick. Seeds black in green-black bitter pulp.
Distribution — Thailand; in Malesia: Sumatra, Peninsular Malaysia, Singapore,
Borneo, Java, Sulawesi, Lesser Sunda Islands, and Moluccas; to allow for regional
variation 4 forms are accepted.
Habitat & Ecology — Secondary forest, forest edges, ravines, also in shrubby marsh-
land; sea level to 1400 m altitude. Flowering and fruiting throughout the year.
Note — De Wilde & Duyfjes 21971 (Peninsular Malaysia) has exceptionally broad,
broadly triangular, male sepals, c. 10 by 6 mm. Henderson 19550, also from Peninsular
Malaysia, has exceptionally large male sepals with narrow sidelobes, like in T. quin
quangulata. The collection De Wilde & Duyfjes 21848 (SW Sulawesi) deviates from
the Java material by coarsely dentate leaf margins, and smaller fruit (c. 5.5 cm long)
with small seeds.
KEY TO THE FORMS
1a. Leaves 3(– 5)-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

b. Leaves unlobed, ovate-cordate. — Siberut Is. . . . . . . . . . . . d. forma siberutensis
2a. Leaves deeply 3(– 5)-lobed, 2/3 – 3/4 deep, conspicuously scabrous. — Lesser
Sunda Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. forma asperifolia
b. Leaves 3-lobed to 2/3 deep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Seeds c. 3 mm thick. Exocarp woody. — Seram . . . . . . . . . . c. forma seramensis

b. Seeds 1– 2 mm thick. Exocarp leathery. — Western and central Malesia . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. forma javanica
a. forma javanica
Leaves: petiole 3 –7 cm long; blade chartaceous, 3-lobed to 2/3 deep, in outline
broadly ovate or circular, to 17 cm long, glands many, scattered, 0.5 –1 mm diam.,
margin entire or coarsely dentate; veins 3 – 5, straight. Male raceme to 20 cm long,
peduncle to 10 cm long. Fruit (narrowly) ovoid, 7– 9 by 5 – 6 cm; exocarp leathery;
fruiting pedicel 1.5 – 2.5 cm long, 3 – 4 mm thick. Seeds (narrowly) obovate, 10 –12 by
5 – 6 by 1– 2 mm. — Fig. 86 b, b', 89; Plate 30c.
Distribution — Malesia: as the subspecies, but not in Siberut Is., Timor and Seram.
b. forma asperifolia Rugayah

Trichosanthes tricuspidata Lour. forma asperifolia Rugayah in Rugayah & W.J.de Wilde, Reinwardtia
11, 4 (1999) 276; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 125, pl. 11.
— Type: Zippel 14 (holo L), Lesser Sunda Islands (Timor).
Leaves: petiole 3 – 3.5 cm long; blade chartaceous, 3(– 5)-lobed, 2/3(– 3/4) deep, in
outline broadly ovate or circular, 5 –7 by 4 – 8 cm, conspicuously scabrous with whit-
ish cystoliths above, glabrous beneath, glands few, scattered, c. 0.5 mm diam., margin
shallowly dentate; veins 3 – 5, straight. Male raceme 6 – 8.3 cm long; peduncle 5 – 6 cm
long. Fruit not known.
Distribution — Malesia: Lesser Sunda Island (Flores, Timor).
c. forma seramensis Rugayah

Trichosanthes tricuspidata Lour. forma seramensis Rugayah in Rugayah & W.J.de Wilde, Reinwardtia
— Type: Mogea & Ramlanto 803 (holo BO; iso L), Moluccas (Seram).
Leaves: petiole 5 – 6.5 cm long; blade subcoriaceous, 3 lobed to 2/3 deep, in outline
broadly ovate or circular, 14 –17 by c. 15 –18.5 cm, smooth or hardly scabrous above,
glands c. 0.5 mm diam., few, scattered, margin minutely dentate; veins 3 – 5, straight;
lobes (narrowly) ovate. Flowers not known. Fruit short-ellipsoid, 6.5 –7 by 5.5 – 6 cm;
exocarp c. 1 mm thick, woody; dry pericarp c. 8 mm thick; fruiting pedicel c. 2.5 cm
long, 8 – 9 mm thick. Seeds broadly obovate, 10 –13 by 8 –10 by c. 3 mm.
Distribution — Malesia: Moluccas (Seram, known only from the type collection).
d. forma siberutensis Rugayah

Trichosanthes tricuspidata Lour. forma siberutensis Rugayah in Rugayah & W.J.de Wilde, Reinwardtia
— Type: Wiriadinata 6874 (holo BO; iso L), Sumatra (Siberut Is.).
Leaves: petiole 2 – 3 cm long; blade unlobed, chartaceous, in outline ovate-cordate,

6.5 – 9 by 4 –7 cm, glands few or several, scattered, 0.5 –1 mm diam., margin obscurely
minutely dentate; veins 3(– 5), curved. Flowers not known. Fruit (broadly) ovoid, 5 – 6
by 4 – 5 cm; exocarp leathery; pericarp c. 7 mm thick; fruiting pedicel c. 0.5 cm long,
c. 4 mm thick. Seeds obovate 8 –10 by 5 – 6 by c. 2 mm.
Distribution — Malesia: Sumatra (Siberut Is., known only from the type collection).
41. Trichosanthes valida Rugayah

Trichosanthes valida Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4 (1999) 277; Rugayah,
Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999)148, pl. 18. — Type: Koorders 16611 (holo
BO; iso L), Sulawesi, Minahasa.
Climber to 10 m long, minutely grey-brown hairy, late glabrescent, leafy stem 4 – 9

mm diam.; dioecious. Probract (narrowly) elliptic, 3 – 4 mm long, subentire, with few
small glands. Tendrils 2- or 3-branched. Leaves: petiole 4 – 9 cm long; blade greenish
on drying, membranous or chartaceous, simple, deeply 3 – 5(–7)-lobed to 3/4 – 4/5 deep,
in outline circular, 14 – 25 by 15 – 22 cm, glands several, scattered or towards the base
only, 1– 2 mm diam., cystoliths absent, margin (sub)entire; veins 5(–7), straight. Male
raceme occasionally with a solitary male flower co-axillary, minutely brown hairy;
peduncle 9 –13(–18) cm long, 3 – 5 mm thick; rachis thickened, 5 –12 cm long, 5 –10
mm thick, with coarse bract-scars, 10-flowered or more; bracts persistent, broadly obo-
vate-rhomboid, 43 – 50 by 15 – 22 mm, margin entire or somewhat shallowly lobed at
apex, with few obscure glands. Male flowers: pedicel c. 10 mm long or less, caducous,
in solitary flowers to 100 mm long; receptacle-tube c. 45 mm long, at throat 7–10 mm
wide; sepals long-triangular, 8 –12 mm long, 2 – 3 mm wide at base, entire; petals c. 12
mm long (excluding threads); synandrium c. 10 mm long, filaments not seen. Female
flowers: pedicel 15 – 20 mm long; ovary short brown hairy, on drying at apex with a
cap-shaped thickening originating from the base of the receptacle-tube, (narrowly)
ellipsoid, c. 15 by 7 mm; perianth as in male. Fruit green with pale yellow markings,
ripening evenly(?) red, ellipsoid, 10 –18 by 8 – 9 cm; exocarp woody, c. 0.5 mm thick,
smooth; pericarp firm-carnose, dry pericarp 15 – 20 mm thick; pulp green-black; fruit-
ing pedicel 4 –7 cm long, 10 – 20 mm thick. Seeds dark brown, compressed, obovate or
elliptic, (8 –)9 –11(– 25) by (5 –)6 – 9 by 2 mm, base and apex rounded, margin absent,
edge entire.
Field-notes — Male corolla lobes with prominent green veins below. Fruit dark red,
shiny, ovoid, 10 by 8 cm, seeds black or brown, c. 15 by 5 mm, pulp dark green, pericarp
hard, yellow-white.
Distribution — Malesia: Philippines (Luzon, Cebu), Sulawesi (Gn Masarang; Du-
moga Bone National Park; Kendari), Lesser Sunda Islands (Timor Leste), Moluccas
(Halmahera: Dodinga), New Guinea (West Papua (Vogelkop: Andai Forest Reserve)).
Habitat & Ecology — Primary forest remnants, secondary scrub, on muddy soil of
secondary roadside forest, deep clayey volcanic soil and fertile volcanic sand; sea level
to 1200 m altitude.
Notes — 1. Bicknell 666 (with fruit, reported as 6 cm long (only photo seen))
and 1344 (male flowers), both from Cebu, may represent a different yet undescribed
taxon.
2. Besides the leaf blade glands, the lower leaf surface has in all specimens very
minute regularly spaced pale brown to blackish points, possibly of a glandular
nature.
3. Trichosanthes valida is close to T. montana, a species confined to West Malesia
in mountainous area, differing in e.g. globose (or ellipsoid, Java) fruit, and shorter
male peduncles. The taxonomy of both species needs further study.
42. Trichosanthes villosa Blume

Trichosanthes villosa Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 934; Miq., Fl. Ned. Ind. 1, 1 (1856) 675;
(1964) 304; Rugayah & W.J.de Wilde (1997) 481, (1999) 278; Rugayah, Trichosanthes (Cucurbita-
ceae) in Malesia, thesis (1999) 152; W.J.de Wilde & Duyfjes in Beaman et al., Pl. Mt. Kinabalu
(2001) 212; Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32 (2004) 101, f. 4, 5D; W.J.de Wilde
& Duyfjes, Sandakania 14 (2004) 31; Fl. Thailand 9, 4 (2008) 539. — Type: Blume s.n. (holo L,
barcode L0130803; iso P), West Java.
Climber to 24 m long, (grey-)brown or yellowish brown villose, hairs on stem and

flowers to 3 mm long, leafy stem 3 – 5 mm diam.; dioecious. Probract absent. Tendrils
5 –15-branched, point of branching 1– 2 cm from the base. Leaves: petiole 3 –11 cm
long; blade (dark) brown on drying, membranous, simple, shallowly 3 – 5-lobed or angu-
lar, or unlobed, in outline subcircular or broadly ovate, 8 – 20(– 26) by 8 –19 cm, densely
villose, particularly beneath, glands few to several, scattered, 0.5 –1 mm diam., margin
entire or sparsely minutely dentate, apex acute-acuminate, with a slender apiculum to 12
mm long; veins 5(–7), straight. Male raceme sometimes co-axillary with a solitary male
flower, hairy; peduncle 5 –12 cm long, 2 – 3 mm thick; rachis not thickened, 5 –18 cm
long, c. 10-flowered; bracts subpersistent, inserted on the rachis or to 20 mm shifted up-
wards on the pedicel, broadly rhomboid or obovate, 20 – 50(– 60) by 30 – 40 mm, margin
entire or shallowly few-dentate, mostly with small glands. Male flowers: pedicel 30 – 90
mm long, to 120 mm in solitary flowers, persistent; receptacle-tube 20 – 30 mm long, at
throat 8 –10 mm wide, at base often swollen, forming a pseudo-ovary (see note under
T. intermedia); sepals narrowly triangular, 10 –16 mm long, 2 – 3(– 5) mm wide at base,
entire; petals obovate-rhomboid, 15 – 20 mm long, threads 7–15 mm long; synandrium
c. 10 mm long, filaments glabrous, c. 2 mm long. Female flowers: pedicel 30 – 60 mm
long; ovary brown hairy, ellipsoid, c. 10 mm long; receptacle-tube and perianth as in
male. Fruit ripening green, whitish or pale yellow striped, ellipsoid-globose, 9 –15 by
8–11 cm; exocarp woody-leathery, less than 1 mm thick, smooth; pericarp firm-carnose,
dry pericarp 10 mm thick; pulp white, fibrous, sweet; fruiting pedicel 3.5 – 5 cm long,
3 – 5(–10) mm thick, possibly sometimes thickened at the distal end. Seeds brown, com-
pressed, (narrowly) ovate, 14 – 25 by 8 –15 by 2 – 5 mm, base truncate, apex narrowly
rounded, margin 2 – 3 mm broad, edge entire.
Distribution — South China (Yunnan, no material seen), Indochina (no material
seen), Thailand; in Malesia: Borneo, Java, Philippines, Lesser Sunda Islands; not known
from Peninsular Malesia and Sumatra; 2 subspecies.
2 cm
e
5 mm
d b d' c d''
2 cm 3 mm
Fig. 90. Trichosanthes villosa Blume. a. Part of twig with male inflorescence; b. male flower; c. opened
male flower; d. stamen, 1- thecous; d', d''. stamens, 2-thecous, showing ab- and adaxial side respec-
tively; e. fruit (a – d'': Phonsena et al. 4000; e: Phonsena et al. 3518).
1a. Fruiting pedicel slender, c. 3 mm thick. Seeds 14 –18 by 8 – 9 by 2 – 3 mm . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. subsp. mindorensis
b. Fruiting pedicel stout, 5(–10) mm thick. Seeds 20 – 25 by 10 –15 by 5 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. subsp. villosa
a. subsp. villosa
Bracts c. 60 mm long. Fruit broadly ellipsoid, 9 –13 by 8 –10 cm; fruiting pedicel
c. 5 cm long, 5(–10) mm thick. Seeds 20 – 25 by 10 –15 by (3 –)5 mm. — Fig. 90, 91e;
Plate 30d.
b
c
3 mm
Fig. 91. Seeds of Trichosanthes. — a. T. cucumerina L. var. cu

cumerina. — b. T. pilosa Lour. — c. T. quinquangulata A.Gray.
— d. T. wawrae Cogn. forma wawrae. — e. T. villosa Blume subsp.
villosa (a: Meebold 3171; b: Phonsena et al. 3521; c: Koonkhun
thod et al. 326; d: Avé 94; e: Phonsena et al 3518). e
Field-notes — Climber to 24 m long.
Distribution — South China (Yunnan, no material seen), Indochina (no material
seen), Thailand; in Malesia: Borneo (Sabah (Kinabalu area); Kalimantan (one doubtful
collection)), all over Java, Lesser Sunda Islands (Bali, Lombok, Sumba, and Flores).
Habitat & Ecology — Forest edges, secondary growths, often in damp places; sea
level to 1500 m altitude.
b. subsp. mindorensis Rugayah

Trichosanthes villosa Blume subsp. mindorensis Rugayah, in Rugayah & W.J.de Wilde, Reinwardtia
— Type: Conklin PNH 19132 (holo L), Philippines (Mindoro).
Bracts c. 20 mm long. Fruit subglobose, c. 8 cm diam.; fruiting pedicel c. 3.5 cm

long, 3(– 5) mm thick. Seeds 14 –18 by 8 – 9 by 2 – 3 mm.
Field-notes — Herbaceous climber, 3 – 4 m high. Fruit globose, melon-like, dark
green with longitudinal white stripes, white inside.
Distribution — Malesia: Philippines (Luzon, Mindoro, Palawan, Cebu, Leyte, and
Bohol).
Habitat & Ecology — Secondary forest; 300 –700 m altitude.
43. Trichosanthes wawrae Cogn.

Trichosanthes wawrae Cogn. in A.DC. & C.DC., Monogr. Phan. 3 (1881) 384 (‘wawraei’); Ridl., Fl.
Malay Penin. I (1922) 845; Rugayah & W.J.de Wilde, Blumea 42 (1997) 481; Rugayah, Trichosan-
thes (Cucurbitaceae) in Malesia, thesis (1999) 108; Duyfjes & Pruesapan, Thai Forest Bull., Bot.
32 (2004) 103; W.J.de Wilde & Duyfjes, Sandakania 14 (2004) 31; Fl. Thailand 9, 4 (2008) 539.
— Type: Wawra 241 (holo W), Singapore.
Trichosanthes trifolia auct. non (L.) Blume: Blume, Bijdr. Fl. Ned. Ind. 15 (1826) 936 (‘trifoliata’);
Ser. in A.DC., Prodr. 3 (1828) 316; Miq. Fl. Ned. Ind. 1, 1 (1856) 679; Cogn. in A.DC. & C.DC.,
Monogr. Phan. 3 (1881) 383; Merr., Interpr. Herb. Amboin. (1917) 494; Backer in Backer & Bakh.
f., Fl. Java 1 (1964) 303. — Involucraria trifolia auct. non (L.) M.Roem.: M.Roem., Fam. Nat.
Syn. Monogr. 2 (1846) 99 (‘trifoliata’), based on Blume s.n., “Crescit in provincia, Krawang juxta
Tjiradjas” (L, barcode L0130880), West Java.
Trichosanthes azurea C.K.Lim & Theseira, Folia Malaysiana 8, 1 (2007) 35. — Type: Lim L8602 (holo
UKMB; iso SING), Peninsular Malaysia, Pahang (Tekam Forest Reserve).
Trichosanthes azurea C.K.Lim & Theseira var. merahputih C.K.Lim & Theseira, Folia Malaysiana
10 (2009) 42, pl. 1, 2, 4 –7. — Type: Lim L10061/FRI56671 (holo KEP; iso UKMB), Peninsular
Malaysia, Pahang (Tekam Forest Reserve).
Climber, 5 –10 m long, glabrescent or densely brown short hairy, leafy stem 1.5 – 3
mm diam.; dioecious, occasionally monoecious. Probract (narrowly) ovate, flat, 3 – 5 by
1.5 – 3 mm, entire or somewhat dentate, glands present. Tendrils (simple or) 2-branched.
Leaves: petiole (1.5 –)2.5 –7 cm long; petiolules 0.5 –1 cm long; blade green on drying,
membranous or chartaceous, simple or 3-foliolate, ± finely scabrous above, glands few
to several, scattered, 0.5 mm diam., cystoliths obvious or not; simple blade shallowly
or deeply lobed, grading into leaflets, in outline (narrowly) ovate or ± hastate, 7–17 by
4.5 – 9 cm, base cordate; veins 3 – 5, ± arched; middle leaflet (narrowly) obovate, 4.5 –10
by 2 – 5 cm, base cuneate, margin entire, or sparsely minutely dentate or distinctly
serrate-dentate (Java); pinnately veined. Male raceme thinly hairy; peduncle 1– 5 cm

long, (1.5 –)2 – 3 mm thick; rachis not or hardly thickened, 6 –12 cm long, 1.5 – 2 mm
thick, 5 – 30-flowered or more; bracts subpersistent, (ob)ovate-rhomboid, variable in
size (see note 1 under forma wawrae), 7– 25 by 4 –12 mm, margin shallowly or deeply
dentate-laciniate, sometimes with 3(– 5) deep main lobes, glands present. Male flow
ers: pedicel 2 – 5(–10) mm long, sometimes persistent; receptacle-tube (20 –)40 –70
mm long, at throat 5 –7 mm wide; sepals narrowly triangular, 4 – 6 mm long, 1– 2 mm
wide at base, entire or few-dentate (lobed); expanded petals not seen; synandrium 6 – 8
mm long, filaments glabrous, 3.5 mm long. Female flowers: occasionally in the male
raceme; pedicel c. 15 mm long; ovary glabrous, ovoid, c. 10 by 5 mm; receptacle-tube,
sepals and petals as in male. Fruit ripening (orange-)red, mostly yellow-striped, ovoid,
6– 8(– 9) by 5 –7 cm, apex 2 – 5 mm beaked; exocarp mostly leathery, smooth; pericarp
juicy-fleshy, dry pericarp c. 5 mm thick; pulp green-black, bitter; fruiting pedicel 1– 2.5
cm long, 2 – 3 mm thick. Seeds brown or black, compressed, (narrowly or) broadly
ovate, 15 – 20 by 8 –15 by 2 – 3 mm, base ± truncate, apex obtuse or subacute, margin
broad but faint, edge entire.
Distribution — Thailand; in Malesia: Sumatra, Peninsular Malaysia, Singapore, Bor-
neo, and Java; 2 forms.
KEY TO THE FORMS
1a. Plant densely brown short hairy. Fruiting pedicel c. 1 cm long. — Sarawak . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. forma hirsuta
b. Plant glabrous (early glabrescent). Fruiting pedicel 1.5 – 2.5 cm long. — Sumatra,
Peninsular Malaysia, Singapore, Borneo, Java . . . . . . . . . . . . . . a. forma wawrae
a. forma wawrae
Plant glabrous (glabrescent). Probract (narrowly) ovate, 3 – 5 by 1.5 – 3 mm, entire.
Leaves: petiole (1.5 –)2.5 –7 cm long; blade membranous or chartaceous, simple or
3-foliolate. Fruiting pedicel 1.5 – 2.5 cm long, 2 – 3 mm thick. — Fig. 91d; Plate 31b.
Field-notes — Fruit ovoid, (orange-)red with yellow blotches or stripes; fruit eaten
by birds. Tuber elongate.
Distribution — Malesia: Thailand and Laos; in Malesia: Sumatra, Peninsular Ma-
laysia, Singapore, Borneo (Central and East Kalimantan; Sabah (Kinabalu area, only
sterile specimens seen)), and Java.
Habitat & Ecology — Primary and degraded forest, swamp forest, montane forest;
lowland to 1300(–1700) m altitude.
Notes — 1. In Peninsular Malaysia two groups of specimens can be recognized, viz.
lowland specimens, from Singapore and lowland Malaya, with large male bracts (c. 20
mm long), deeply and finely laciniate, and montane specimens with small male bracts
(c. 10 mm long or less), which are more shallowly laciniate. The taxonomic significance
of these differences is still unclear.
2. All specimens from Java are from (lower) montane area in the western part and
differ from the remainder of the material by a slender habit (stem 1–2 mm diam.), thin
leaves, mostly with rather distinct serrate-dentate margin. The male bracts are 10–15
mm long, deeply and slenderly incised, and approach the larger (stouter) deeply incised
bracts as found in lowland specimens from Singapore and Peninsular Malaysia, which
match the type-collection. In Java the sepals seem to be frequently ± lobed. All Javan
specimens have truly 3-foliolate leaves, apparently simple leaves never have been found
there.
Some collections from lowland Sabah, discussed under T. longispicata, with short
male bracts, come close to specimens at present treated under T. wawrae from montane
area in Peninsular Malaysia.
Lack of material from all areas prevents a satisfactory further subdivision of
T. wawrae.
3. The type specimens of T. azurea (incl. varieties) agree with T. wawrae; the bluish
or red and white colours of the fruit are possibly due to some infection.
b. forma hirsuta Rugayah

Trichosanthes wawrae Cogn. forma hirsuta Rugayah in Rugayah & W.J.de Wilde, Reinwardtia 11, 4
(1999) 279; Rugayah, Trichosanthes (Cucurbitaceae) in Malesia, thesis (1999) 113. — Type: Bayeng
S 44194 (holo L), Borneo (Sarawak, 1st Division, Gn Buri).
Plant densely brown short hairy. Probract elliptic, c. 4 mm long, somewhat dentate.
Leaves: petiole c. 2 cm long; blade chartaceous, 3-foliolate. Fruiting pedicel c. 1 cm long,
3 mm thick.
Field-notes — Creepers along logging road. Fruits green with green stripes, turning red
when ripe.
Distribution — Malesia: Borneo (Sarawak, known only from the type collection).
Habitat & Ecology — Mixed dipterocarp forest.
36. URCEODISCUS
Urceodiscus W.J.de Wilde & Duyfjes, Blumea 51 (2006) 38. — Type species: Urceodiscus belensis
(Merr. & L.M.Perry) W.J.de Wilde & Duyfjes (Melothria belensis Merr. & L.M.Perry).
Small annual or subperennial climbers, leafy stem 1– 2 mm diam., green on drying;

monoecious. Probract absent. Tendrils unbranched. Leaves simple, unlobed or lobed.
Flowers small or medium, 5 – 20 mm diam.; petals (light green or) yellow, free or to
1/3 connate; the segments imbricate in bud; receptacle-tube urceolate or campanulate;
disc in both sexes urceolate, carnose, wholly or largely fused with the base of the re-
ceptacle-tube. Male inflorescence a peduncled raceme, with flowers arranged zigzag
or not, without or with co-axillary a solitary male or female flower. Bracts absent or
minute. Male flowers: pedicel short or long, 2 – 20(– 30) mm long, persistent; stamens 3,
inserted at or to halfway below the throat of the receptacle-tube, filaments much longer
than the anther, anthers all 2-thecous, ± exserted, thecae lateral, straight (or ± curved),
parallel or ± divergent, connective narrow or broad and then at apex broad, produced or
not. Female flower solitary, single at the node or co-axillary with male raceme; pedicel
long; ovary subglobose or short-ellipsoid or fusiform, with a narrow neck, glabrous;
stigma deeply 3-lobed or 3, papillose-hairy; staminodes present; disc more shallow than
in male. Fruit solitary, with long fruiting pedicel, globose or ellipsoid, 0.8–3 cm long,
glabrous, red, juicy; exocarp ± membranous, ± translucent, minutely pustulate. Seeds
c. 10 or numerous, little compressed or almost spherical, ovoid, scrobiculate or pitted,
margin faint or narrow, edge entire.
Distribution — A genus of 7 species confined to New Guinea.
Note — The species much resemble each other vegetatively, but they are distinct by
characters of the male inflorescences and male flowers.
KEY TO THE SPECIES
1a. Male pedicels 1– 3 mm long; rachis usually straight . . . . . . . . . . . . . . . . . . . . . . . 2

b. Male pedicels 5 mm long or more; rachis frequently zigzag. [Male inflorescences
and flowers not known in U. arfakensis.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2a. Male perianth 5 – 6 mm long; filaments hairy, in-curved at apex. Leaves lobed to
1/3 to nearly to the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. U. scabridula
b. Male perianth 5 –15 mm long; filaments glabrous, straight. Leaves finely or coarsely
dentate or up to 1/3 lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Male receptacle-tube shorter than broad or about as long as broad, 2 – 3 mm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
b. Male receptacle-tube longer than broad, c. 4 mm long. [Fruit not known.] — West
Papua, Vogelkop; lowland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. U. viridis
4a. Male petals 10 mm long or more . . . . . . . . . . . . . . . . 2. U. belensis var. conferta
b. Male petals 6 – 9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Male perianth c. 6 mm diameter. — Eastern Papua New Guinea; at c. 2000 m alti-
tude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. U. carrii
b. Male perianth c. 4 mm diameter. — Northern West Papua, Cycloop Mountains; at
c. 1200 m altitude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. U. parviflora
6a. Fruit 2 – 3 cm long (flowers not known). — West Papua, Arfak Mountains; at 850
m altitude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. U. arfakensis
b. Fruit 1–1.5(– 2) cm long (fruit not known in U. hippocrepicus). Flowers large, petals
(5 –)6 mm long or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Receptacle-tube inside and filaments glabrous; anthers not cordate at base. —
Whole of New Guinea, but not known from Vogelkop; at 1900 – 2900 m altitude .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. U. belensis
b. Receptacle-tube inside hairy (filaments glabrous); anthers deeply cordate at
base. — West Papua, Baliem Valley; at 2000 m altitude . . . . 4. U. hippocrepicus
1. Urceodiscus arfakensis W.J.de Wilde & Duyfjes,

Urceodiscus arfakensis W.J.de Wilde & Duyfjes, Blumea 51 (2006) 40. — Type: Sands MJS 6809
(holo K; iso L), West Papua, Arfak Mountains.
Climber to 3 m long, leafy stem 1–1.5 mm diam., plant sparsely stiff-hairy (hairs
less than 0.5 mm long), green on drying. Leaves: petiole c. 0.5 cm long, densely finely
stiff-hairy; blade unlobed, ovate-elliptic, 5 – 9 by 2 – 4 cm, both surfaces ± bullate,
scabrous by sparse stiff 0.5 mm long hairs, cystoliths distinct, veins scabrid-hairy,
base subtruncate or broadly rounded (hardly cordate), margin sparsely inconspicuously

dentate (teeth less than 1 mm long), apex (long) acute-acuminate, minutely mucronate.
Male and female flowers not known. Fruit ripening scarlet, ellipsoid, 2 – 3 by 1.5 – 2
cm; fruiting pedicel slender, 6 –7.5 cm long. Seeds numerous, whitish or pale brown,
ovoid, 4.5 – 5.5 by 3 – 3.5 by 2 mm, margin narrow.
Distribution — New Guinea (West Papua, Vogelkop (Arfak Mountains)); known
only from the type.
Habitat & Ecology — Montane forest; climbing near river in shade; at 850 m alti-
tude; fruiting in April.
2. Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes

Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 40, f. 10, 11.
— Melothria belensis Merr. & L.M.Perry, J. Arnold Arbor. 30 (1949) 57. — Type: Brass 11082
(holo A; iso BO, L), New Guinea (West Papua, Bele River).
Melothria cissybium M.Jacobs, Blumea 7 (1954) 617 (for the male flowering material only). — Type:
Womersley 5343 (sphalm. 5543) (holo L; iso LAE), Papua New Guinea, Western Highlands Prov-
ince, Al River.
Herbaceous climber to c. 4 m long, leafy stem 1(–1.5) mm diam., plant finely hairy,
glabrescent, green on drying. Leaves: petiole 0.5 – 2 cm long, (appressed-)hairy; blade
unlobed or towards the base lobed to 1/2 deep (blade then ± hastate), ovate to elliptic,
3.5 –14 by 2 – 5.5 cm, upper surface with stiff hairs and cystoliths, the veins more dense-
ly hairy, lower surface somewhat hairy or subglabrous, base subtruncate or broadly
shallowly (or deeply) cordate, margin sparsely finely or coarsely dentate (sometimes
shallowly undulate), apex long acute-acuminate, mucronate. Male inflorescences a 2 – 5
cm long peduncled few- to many-flowered condensed or zigzag raceme 0.5 – 5 cm long,
without or with co-axillary a single male or female flower. Male flowers: pedicel 2 – 20
mm long, minutely appressed-hairy; perianth 10 –15 mm long, expanded perianth 12 –
30 mm diam.; receptacle-tube urceolate-campanulate, sometimes faintly constricted in
the middle, 4 –7 by 3 – 5 mm, glabrous, throat glabrous; sepals 1– 2 mm long, glabrous;
petals free or up to 1/3 connate, yellow, (narrowly) elliptic, 6 – 9 by 4 – 5 mm, finely
hairy on both surfaces, obtuse or rounded; stamens inserted at c. 1/3 from the throat in
the receptacle-tube, filaments 4 – 5 mm long, straight, glabrous, anthers subcircular or
ovate(-elliptic) in outline, c. 1.5 mm diam., thecae c, 1 mm long, divergent, connective
broad, at apex truncate or broadly up to 1 mm produced, obtuse, hairy at both sides;
disc 1– 2(– 3) by 2 mm, entire or sometimes 3-parted by grooves and with 1– 3 small
nipples c. halfway. Female flowers solitary or co-axillary with a (later developing) male
inflorescence; pedicel 20 – 40 mm long; ovary ellipsoid, 2 – 4 mm long, glabrous, with
short or long neck 0.5 –1.5 mm long; corolla as in male flowers; style c. 4 mm long,
glabrous, stigma c. 4 mm diam., 3-lobed, the lobes half-patent, ± narrowly ovoid, c. 3
mm long, densely papillose; staminodes 1 mm long, glabrous; disc 0.5 mm high. Fruit
ripening bright shiny red, short-ellipsoid, 1– 2 cm long; fruiting pedicel 3 – 5 cm long.
Seeds numerous, ovoid, 4 – 5 by 3.5 – 4 by 3 mm, margin narrow.
Field-notes — Petals (orange-)yellow. Anthers orange, pollen yellow. Fruits edible.
Distribution — In the main range of New Guinea, except Vogelkop.
Habitat & Ecology — Edges of (Nothofagus) forest, scrub, secondary mixed (oak)

forest; climbing in tree fern; at 1900 – 2900 m altitude; flowering and fruiting mainly
April to December.
1a. Male raceme less than 0.5 cm long, with few subfascicled flowers; pedicels 2 – 3
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. var. conferta
b. Male raceme 1– 4(– 5) cm long, flowers more spaced; pedicels 5 – 20 mm long . . 2
2a. Male raceme c. 10-flowered; pedicels 5 –10(–15) mm long . . . . . . a. var. belensis
b. Male raceme lax, c. 5-flowered; pedicels (5 –)10 – 20 mm long . . . . . . c. var. laxa
a. var. belensis
Male raceme 1– 5 cm long, c. 10-flowered; pedicels 5 –10(–15) mm long; receptacle-
tube 4 – 5 mm long; disc 1–1.5 mm high. Female flowers and fruit as in the species.
— Fig. 92.
Distribution — In the main range of the whole of New Guinea, except Vogelkop.
b. var. conferta W.J.de Wilde & Duyfjes

Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes var. conferta W.J.de Wilde &
Duyfjes, Blumea 51 (2006) 34. — Type: Hoogland & Schodde 7537 (holo L; iso CANB), Papua
New Guinea, Western Highlands Province, Yobobos.
Male raceme less than 0.5 cm long, with 1– 5 clustered flowers; pedicels 1– 3 mm long;
receptacle-tube 4 – 5 mm long; disc 1–1.5 mm high. Female flowers not known. Fruit
as in the type-variety.
Distribution — New Guinea (Papua New Guinea, known from 2 collections: Morobe
Province (Schodde & Craven 4943) and Western Highlands Province (Hoogland &
Schodde 7537)).
c. var. laxa W.J.de Wilde & Duyfjes

Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes var. laxa W.J.de Wilde & Duyfjes,
Blumea 51 (2006) 43, f. 11b – g. — Type: Hoogland & Schodde 6798 (holo L; iso CANB), Papua
New Guinea, Western Highlands Province, Poio.
Male raceme lax, 2 – 5 cm long, 5 –10-flowered; pedicels 10 – 20 mm long; recepta-

cle-tube 5 –7 mm long; disc 1– 3 mm high. Female flowers not known. Fruit as in the
type-variety.
Distribution — New Guinea (Papua New Guinea: Western, Eastern and Southern
Highlands Provinces and Morobe Province).
3. Urceodiscus carrii W.J.de Wilde & Duyfjes

Urceodiscus carrii W.J.de Wilde & Duyfjes, Blumea 51 (2006) 44, pl. 1c, d. — Type: Carr 13647 (holo
BM; iso CANB, K, L, SING), East Papua New Guinea, Northern Province, Alola.
Small vine, leafy stem 1 mm diam., plant sparsely hairy, glabrescent, green on drying.
Leaves: petiole 1– 2 cm long, minutely appressed-hairy; blade unlobed or at base irregu-
larly 3 – 5-lobed to c. 1/3 deep, narrowly ovate, 7–14 by 3 – 8 cm, upper surface sparsely
stiff-hairy, with cystoliths, more densely hairy on veins, lower surface (sub)glabrous,
base shallowly cordate or subtruncate, margin sparsely short-dentate, apex long
acute-acuminate, mucronate. Male inflorescences in (5 –)10 – 20-flowered (non-zig-
zag) spike-like racemes of 0.5 – 2 cm long; peduncle 1–1.5 cm long. Male flowers:
pedicel 1– 3 mm long, minutely appressed-hairy; perianth c. 5 mm long, expanded
2 cm
2 mm
c
b
2 cm
1 mm
1 mm
e f g
Fig. 92. Urceodiscus belensis (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes var. belensis. a. Twig with
female flowers; b, c. female flowers; d. node with male inflorescence and one fruit; e. fruit surface,
enlarged; f, g. seed showing narrow margin (a – c: Borgmann 210; d – g: Kiprianus, Lawong & Gideon
LAE 69127).
perianth 5 – 6 mm diam.; receptacle-tube shallowly campanulate, 1–1.5 by 2(– 2.5) mm,

glabrous, throat glabrous; sepals 0.5 mm long; petals free, 2.5 – 3 mm long, rounded,
both surfaces gland-hairy; stamens inserted c. halfway the receptacle-tube, filaments
1.5 – 2 mm long, glabrous, straight, anthers subcircular in outline, 1 mm diam., thecae
0.5 mm long, situated lateral in the basal part, divergent, connective broad, at apex
broadly protruding, sparsely hairy; disc saucer-shaped, 0.5 by 2 mm. Female flowers
(solitary or) co-axillary with male raceme (which develops after the female flower);
pedicel 3 – 5 mm long; ovary ellipsoid, 3 – 3.5 mm long; corolla as in male flower; style
and stigma not investigated (see note). Fruit solitary, ripening glossy red, subglobose,
1–1.5 cm diam.; fruiting pedicel 3 – 6 cm long. Seeds numerous, pale brownish yellow,
ovoid, 4 – 4.5 by 2 – 3 by 2 – 2.5 mm, margin narrow.
Field-notes — Flowers cream, off-white or yellowish green.
Distribution — New Guinea (eastern Papua New Guinea).
Habitat & Ecology — Short-statue forest, rock faces of escarpment; at 1700 – 2500
m altitude; flowering and fruiting: July to December.
Note — The female flowers were not fully studied as Carr 13892 (BM) bears the
only female flower known.
4. Urceodiscus hippocrepicus W.J.de Wilde & Duyfjes

Urceodiscus hippocrepicus W.J.de Wilde & Duyfjes, Blumea 51 (2006) 44. — Type: Kostermans &
Soegeng 655 (holo L; iso A, BO, K), West Papua, Baliem Valley.
Small climber to 4? m long, leafy stem 1–1.5 mm diam., plant sparsely minutely
scabrous-hairy, brown-green on drying. Leaves: petiole 0.5 –1 cm long, minutely ap-
pressed-hairy; blade unlobed or hardly lobed, (narrowly) ovate, 6 –10 by 2 – 4.5 cm,
upper and lower surface ± bullate, scabrid by sparse minute stiff hairs and inconspicu-
ous cystoliths, veins more densely minutely appressed-hairy, base shallowly or deeply
cordate, often ± hastate, margin indistinctly sparsely dentate (teeth less than 0.5 mm
long), blade gradually narrowed towards the long acute-acuminate apex. Male inflo
rescences a 5- or 6-flowered short ± zigzag raceme c. 1 cm long; peduncle 2 – 3 cm
long, finely appressed-hairy. Male flowers: pedicel c. 10 mm long; perianth c. 10 mm
long, expanded perianth 10 –12 mm diam.; receptacle-tube 3 – 4 by 2 – 3 mm, sparsely
appressed-hairy, throat and inside hairy, hairs 0.5 mm long; sepals 1 mm long; petals
almost free, (narrowly) elliptic, 5 – 6 mm long, obtuse, densely hairy at both surfaces;
stamens inserted c. halfway in the receptacle-tube, filaments c. 2 mm long, glabrous,
anthers subelliptic in outline, 1.5 – 2 by c. 1.5 mm, apex rounded, base deeply cordate,
forming an 1 mm deep sinus, thecae 1.5(– 2) mm long, ± curved and nearly touching
at apex, extending downwards on the connective-lobes, connective ± narrow at apex
(where attached to the filament), densely hairy; disc thick-carnose, 1 by 2 mm, faintly
3-lobed. Female flowers and fruit not known.
Field-notes — Flowers yellow. Flowering in August.
Distribution — New Guinea (West Papua, Baliem Valley); known only from the type.
Habitat & Ecology — Wet places in low, open scrub; at 2000 m altitude.
Note — Because of the deeply cordate anther the two thecae are shaped like a horse-
shoe.
5. Urceodiscus parviflora W.J.de Wilde & Duyfjes

Urceodiscus parviflora W.J.de Wilde & Duyfjes, Blumea 51 (2006) 45. — Type: Van Royen & Sleumer
6002 (holo L; iso BO), West Papua, Cycloop Mountains.
Low climber; leafy stem c. 1 mm diam., plant sparsely hairy, largely glabrescent,
green on drying. Leaves: petiole 1–1.5 cm long, minutely hairy; blade unlobed, (ovate-
)narrowly elliptic, 5 – 9 by 2 – 3 cm, both surfaces sparsely hairy, hairs minute, stiff, cys-
toliths inconspicuous, veins more densely hairy, base subtruncate or shallowly broadly
cordate, margin (sparsely) repand-dentate, teeth 1– 2 mm long, apex acute-acuminate,
minutely mucronate. Male inflorescences arranged in a tiny and slender 10 –15-flowered
spike-like 0.5 –1 cm long raceme; peduncle c. 1 cm long, sparsely minutely appressed-
hairy. Male flowers: pedicel 1– 2 mm long, sparsely minutely appressed-hairy; perianth
c. 3 mm long, expanded perianth (as judged from well-advanced bud) c. 4 mm diam.;
receptacle-tube shallow, 0.5(–1) by 2 – 2.5 mm, throat sparsely minutely hairy; sepals
0.5(–1) mm long; petals free, 1.5(– 2) by 1 mm, obtuse, densely minutely hairy outside,
glabrous inside; stamens inserted about halfway in the receptacle-tube, filaments (im-
mature) 0.5 mm long, densely hairy in upper half, anthers ellipsoid, 1 by 0.7 mm, thecae
somewhat curved, 1 mm long, connective narrow, hairy, not produced; disc 0.2 – 0.5 by
c. 2 mm. Female flowers and fruit not known.
Field-notes — Leaves dark green above, greyish green beneath. Flowers pale orange-
yellow. Flowering in June.
Distribution — New Guinea (West Papua, Cycloop Mountains, path from Ifar to
Ormoe, at the camp-site); known only from the type.
Habitat & Ecology — In secondary regrowth; at 1220 m altitude.
6. Urceodiscus scabridula (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes,

Urceodiscus scabridula (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 45. — Mel
othria scabridula Merr. & L.M.Perry, J. Arnold Arbor. (1949) 56. — Type: Brass 10621 (holo A,
not seen; iso BO, L), West Papua, Lake Habbema.
Small (extensively scrambling) few meters long climber, leafy stem 1– 2 mm diam.,
plant densely or sparsely hairy, (partly) glabrescent, green-brown on drying. Leaves:
petiole 1– 2.5 cm long, short-hairy; blade (3- or) 5- (or 7–10)-lobed to 1/3 to nearly to
the base (then the segments narrowly elliptic, to 8 cm long), (narrowly) ovate in outline,
4 – 9 by 3 – 8 cm, upper surface with sparse stiff hairs especially on the veins, scabrous,
cystoliths present, lower surface (sub)glabrous, base shallowly broadly cordate, margin
sparsely coarsely serrate-dentate, teeth to 5 mm long, apex long-acuminate, mucronate.
Male inflorescences 2 – 5-flowered short (non-zigzag) 0.2 – 0.5 cm long racemes; pe-
duncle 1–1.5 cm long. Male flowers: pedicel 1– 4 mm long, minutely appressed-hairy;
perianth 5 – 6 mm long, expanded perianth 5 – 6 mm diam.; receptacle-tube 2 – 4 by
3 – 4 mm, glabrous, throat minutely hairy inside; sepals 0.5 –1 mm long, glabrous;
petals almost free or to c. 1.5 mm connate, (narrowly) elliptic, (3 –)4 – 5 by 1.5 – 2(– 3)
mm, subobtuse or rounded, papillose or (gland-)hairy outside, glabrous or papillose
inside; stamens inserted at c. 1/3 from the throat in the receptacle-tube, filaments c. 2
mm long, glabrous or (densely) hairy, hairs 0.3 – 0.5 mm long, at apex ± thickened and
1 mm
b
a
2 cm 1 mm 1 mm
d c
Fig. 93. Urceodiscus scabridula (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes. a. Habit with fruit and
immature male inflorescences; b. apex of male inflorescence; c. female flower, opened; d. seed (all:
Sands 7143).
c. 90° curved inwards, anthers dorsally attached to the filaments, ellipsoid, c. 1.5 by
1–1.5 mm, thecae vertical, straight, 1 mm long, connective narrow or broad with (few)
sparse minute hairs; disc a thick-carnose cupule, 0.5 –1.5(– 2) by 2(– 3) mm, margin
smooth or ± wavy. Female flowers solitary or co-axillary with male inflorescence; pedi-
cel 20 – 30 mm long; ovary subglobose-ellipsoid, c. 1.5 by 1 mm, neck 0.5 mm long,
perianth as in male flower; receptacle-tube c. 2 by 4 mm; style c. 2 mm long, at apex
3-armed, each arm c. 1.5 mm long, with elongated stigma-lobe c. 1.5 mm long, out-
curved, densely hairy; staminodes subulate, c. 1.5 mm long, hairy. Fruit ripening red,
2 mm
a b
Fig. 94. Urceodiscus scabridula (Merr. & L.M.Perry) W.J.de Wilde & Duyfjes. a, b. Male flower;
c. stamens (all: Willis FR 296).
blackish brown when dry, subglobose, 1– 2 cm diam., ± smooth or minutely pustulate;

fruiting pedicel 2.5 – 5 cm long. Seeds 10 – 20, pale brown, ovoid, 4 – 4.5 by 3 – 3.5 by
c. 2.5 mm, margin faint. — Fig. 93, 94.
Field-notes — Flowers yellow.
Distribution — New Guinea (south-western West Papua: near Lake Habbema; Val-
entijn Mountains; P.T. Freeport Indonesia Concession Area).
Habitat & Ecology — Mossy forest, on tree stumps; undergrowth in disturbed mon-
tane forest; forest clearings; at 2200 – 2800 m altitude; flowering and fruiting: August
to December.
Note — This species is remarkable by its filaments which are thickened at apex and
abruptly in-curved for c. 90°.
7. Urceodiscus viridis W.J.de Wilde & Duyfjes

Urceodiscus viridis W.J.de Wilde & Duyfjes, Blumea 51 (2006) 46, f. 12; pl. 1a. — Type: W. & M. Vink
BW 15396 (holo L; iso CANB, K), West Papua, Lake Ajamaru.
Small vine, leafy stem 1(–1.5) mm diam.; plant sparsely minutely hairy, glabrescent,
green on drying. Leaves: petiole 0.5 –1 cm long, partly appressed-hairy; blade unlobed
or at base with an odd lobe to 1/3 deep, (narrowly) ovate, 4 – 8 by (1.5 –)2 – 5 cm, upper
surface glabrous except for minute soft hairs on veins, cystoliths sparse, inconspicu-
ous, lower surface glabrous, base broadly rounded, subtruncate or shallowly cordate,
margin sparsely short serrate-dentate, teeth 1– 2(– 3) mm long, apex acute(-acuminate),
minutely mucronate. Male inflorescences 3 –7 in a short (non-zigzag) spike-like 0.5 –1
cm long raceme; peduncle 2 – 3 cm long. Male flowers: pedicel 1– 2 mm long, glabrous;
perianth 7–10 mm long, expanded perianth c. 10 mm diam.; receptacle-tube c. 5 by 2.5
mm, subglabrous, throat and upper half of tube minutely hairy inside; sepals c. 0.5 mm
long; petals free, 4 – 5 by 2.5 mm, subobtuse, minutely hairy outside, glabrous inside;
stamens inserted at c. 1/4 from the apex in the receptacle-tube, filaments c. 2 mm long,
straight, glabrous, anthers ± connivent, ellipsoid, c. 2 by 1 mm, thecae straight, c. 2 mm
2 mm
2 cm
1 mm
a c d
Fig. 95. Urceodiscus viridis W.J.de Wilde & Duyfjes. a. Twig with male inflorescences; b. detail of
male inflorescence, showing 4 buds; c, d. male flowers (all: W. & M. Vink BW 15396, type).
long, connective narrow, not produced; disc c. 1.5 by 2 mm. Female flowers and fruit
not known. — Fig. 95.
Field-notes — Rather common climber. Calyx green; corolla, connective and fila
ments light green; anthers yellow; flowering in March.
Distribution — New Guinea (West Papua, Vogelkop, Tanah Merah, west side of Lake
Ajamaru, 1°8' S, 132°13' E), known only from the type.
Habitat & Ecology — Young secondary forest on strongly humified limestone silt;
at 220 m altitude.
37. ZANONIA
Zanonia L. Sp. Pl. ed. 1 (1753) 1028; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 633, p.p.; Cogn. in
A.DC. & C.DC., Monogr. Phan. 3 (1881) 925, excl. Z. macrocarpa = Alsomitra (Blume) M.Roem.;
in Engl., Pflanzenr. 66, 4.275.I (1916) 27; Keraudren in Aubrév. & J.-F.Leroy, Fl. Cambodge, Laos
& Vietnam 15 (1975) 15, pl. 4; W.J.de Wilde, Blumea 52 (2007) 282; Fl. Thailand 9, 4 (2008) 540.
— Type species: Zanonia indica L.
Juppia Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 170. — Type species: Juppia borneensis
Merr.
Medium-sized liana to 15 m long, woody at base, leafy stem 3(– 5) mm diam., dio-
ecious. Probract absent. Tendrils (unbranched or) 2-branched at apex. Leaves: petiole
short, older twigs with raised leaf scars; blade simple, unlobed, (narrowly) ovate-el-
liptic, cystoliths absent, margin entire. Flowers small, creamy-white; buds globose;
perianth rotate; sepals 3 (but see note 1), concave, valvate in bud; petals 5, free, ± fleshy,
conduplicate-valvate with tips inflexed in bud; receptacle shallow. Male inflorescences
paniculate, many-flowered, ± pendulous; bracts small, linear, 1(– 2) mm long, glabrous
or hairy. Male flowers: pedicel short; receptacle (disc) broad, low; stamens 5, free, in-
serted on the disc, filaments short, anthers all 1-thecous, opening with apical transverse
slit; pistillode absent. Female flowers in (raceme-like) panicles; pedicel short or absent;
ovary clavate, imperfectly 3-locular, each locule with apically 2 pendulous ovules;
styles 3, short, horizontally two-horned, separate, inserted on slightly raised truncate
apex of ovary; staminodes absent or small (see note 2). Fruit rather large, capsular,
elongate-cylindrical, claviform, with truncate apex opening by 3 inward curving apical
valves. Seeds compressed, longitudinally winged all around the seed, faces smooth,
margin faint, edge entire.
Distribution — One species with two subspecies. South and NE India, South China,
Indochina, through Malesia east to New Guinea.
Notes — 1. The globose calyx in bud is membranous, and closed at apex, but mostly
the very apex shows 5 minute appendages proving that the calyx is morphologically
5-merous. However, at anthesis the globose bud splits into (2 or) 3 triangular ± concave
sepals. The five petals are thick-fleshy and drop off after anthesis.
2. In the female flowers of Giesen 76 (Zanonia indica subsp. orientalis var. paludosa)
the staminodes are distinctly stamen-like, each with a small anther, including an indication
of the apical slit at apex. It could not be verified whether they contain viable pollen.
1. Zanonia indica L.
Zanonia indica L., Syst. Nat. ed. 10, 2 (1759) 1292; Sp. Pl. ed. 2, 2 (1763) 1457; Miq., Fl. Ned. Ind. 1,
1 (1856) 658; C.B.Clarke in Hook.f., Fl. Brit. Ind. 2 (1879) 633; Cogn. in A.DC. & C.DC., Monogr.
Phan. 3. (1881) 926; Backer in Backer & Bakh.f., Fl. Java 1 (1964) 295; W.J.de Wilde & Duyfjes,
Blumea 52 (2007) 283. — Type: “Penar-valli mas” in Rheede, Hort. Malab. (1688) 8: 93 (‘39’),
t. 49 (lecto, designated by Chakravarty, Fasc. Fl. India 11 (1982) 126).
Distribution — South India, Sri Lanka, NE India, South China, Indochina, through

Malesia east to New Guinea; in Malesia: 1 subspecies.
Note — De Wilde & Duyfjes (Blumea 2007) distinguished in Zanonia indica two
subspecies, subsp. indica (South India and Sri Lanka) and subsp. orientalis W.J.de
Wilde & Duyfjes; only subsp. orientalis occurs in Malesia.
subsp. orientalis W.J.de Wilde & Duyfjes

Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes, Blumea 52 (2007) 284, f. 1, 2a – d; Fl.
Thailand 9, 4 (2008) 541, pl. 43, 1, 2. — Type: Phonsena 5192 (holo BKF; iso L), Thailand, Chon
Buri, Bo Thong, Khao Cha-ang On.
Zanonia indica L. var. angustifolia Cogn. in A.DC. & C. DC., Monogr. Phan. 3 (1881) 927. — Type:
Hooker f. & Thomson s.n. (holo K; iso P), East India, Chittagong.
Tinospora curtisii Ridl., J. Bot. 58 (1920) 148. — Type: Curtis 3464, male (holo K), Peninsular Ma-
laysia, Penang, Batu Feringhi, near the sea.
Alsomitra simplicifolia Merr., Philipp. J. Sci. 20 (1922) 470. — Type: Ramos & Edaño BS 37397 (holo
PNH†; iso BO, K, L, P), Philippines, Mindanao, Malangas.
Juppia borneensis Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 170. — Type: Ramos 1593 (holo
PNH†; iso K, L) Malaysia, Sabah, near Sandakan.
Liana 4 –15 m tall; leafy stem 3 – 5 mm diam., glabrous or hairy, often with lenticels.
Tendrils 5 –15 cm long, in juvenile plants at apices with irregularly shaped adhesive
pads to c. 5 mm diameter. Leaves: petiole 1.5 – 3 cm long; blade membranous or sub-
coriaceous, (broadly) ovate-elliptic, 8 –15 by 5 –12 cm, both surfaces glabrous or hairy,
base broadly rounded or cordate or sometimes somewhat hastate, apex (sub)obtuse,
minutely mucronate; 2 – 5-pinnately veined and one pair of basal veins, reticulation
distinct on lower surface. Male inflorescences glabrous or hairy, (10 –)15 – 60 cm long,
little or much branched (sometimes also branched from the base), lateral branches to
15 cm long, flowers often in bundles of up to 5, the bundles ± spaced, the larger in-
florescences often from the older wood. Male flowers: pedicels rather thick, 1– 3(– 5)
mm long, articulate at ± halfway; buds c. 2 mm diam.; corolla 5 –7 mm diam.; sepals
membranous, almost free, subtriangular, c. 2 mm long; petals (narrowly) ovate-ellip-
tic, 2.5 – 3.5 long, apex narrowed, subacute, adaxially papillose; stamens inserted on a
carnose disc, leaving the centre free, filaments 0.5 –1 mm long, fleshy, anthers trans-
versely ellipsoid, (0.5 –)1 mm long. Female inflorescences 5 – 40 cm long, raceme-like
or branched panicles, few- or many-flowered, flowers solitary (or 2). Female flowers:
sepals triangular, 2 – 4 mm long; petals (narrowly) ovate-elliptic, 3 – 8 mm long, suba-
cute, finely papillose-hairy; ovary subcylindrical-obconical, 5 –12 by 2 – 3 mm, gla-
brous or hairy; styles short, stigmas with 2 incurved horizontal horns, papillose. Fruit
pendulous, few to many per infructescence, variable in size, (4 –)5 –10 by 1.5 – 4.5(– 5)
cm, glabrous, glabrescent or hairy. Seeds 6, pale, elliptic, not ornamented, 15(– 20) by
8(–10) mm, winged all around, the wings leathery, elongated and rounded at both ends,
4 – 6(– 8) by 1.3 –1.5(– 2) cm.
Distribution — NE India, South China, Indochina, Thailand; in Malesia: Sumatra,
Peninsular Malaysia (Kedah, Perak, Penang, and Johor), Borneo (Kalimantan, Sarawak,
Sabah), Java, Philippines (Sulu Archipelago, Negros, and Mindanao), Sulawesi, Moluc-
cas (Aru Islands), New Guinea (West Papua, Papua New Guinea (Kaiser Wilhelmsland,
Morobe Province)); 2 varieties.
Habitat & Ecology — Forest edges, riversides, open forest on mountain slopes; (0 –)
20 – 800 m altitude; flowering and fruiting throughout the year.
Notes — 1. The size of the fruits shows a considerable variety in its length, vary-
ing from 6 – 9 cm, but strikingly small fruits, c. 4 by 2(– 2.5) cm, seem usual in New
Guinea.
2. Sterile material of Z. indica may resemble sterile Alsomitra macrocarpa. The
latter generally has longer petioles, and a characteristic short-hairy spot in the leaf
axil; also, the adhesive pads which may develop at the end of the tendril-branches are
larger and more elongate, to 1.3 cm long. Furthermore, the bark of twigs in Alsomitra
is not or but finely striate (coarsely striate in Zanonia), and without or with few small
lenticels.
e 1 cm j
a
2 cm
1 mm
1 mm
f i
2 cm
1 mm
2 mm g
c
Fig. 96. Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. pubescens Cogn. a. Node with
male inflorescence; b, c. male flower; d. ditto, seen from below showing 3-lobed calyx; e. node with
female inflorescence; f. female flower; g. apex of female flower, part of perianth removed showing
styles and stigmas; h, i. fruit and apex of fruit respectively; j. seed (a: Kerr 2048; b – d: Phonsena, De
Wilde & Duyfjes 5192 (from spirit); e – g: Ramos & Edaño BS 37397 (type of Alsomitra simplicifolia);
h – j: Geesink 8384).
2 mm
b
4 mm
4 mm
f c
4 mm
2 mm
5 mm
d
4 mm
g
a
Fig. 97. a – d. Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. pubescens Cogn.
a. Growing shoot apex; b. node of sterile shoot, note adhesive pads at apex of tendril; c. apex of female
inflorescence; d. female flower, perianth removed and ovary longitudinally opened to show position of
the ovules. — e, f. Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. paludosa W.J.de
Wilde & Duyfjes. e. Apex of female inflorescence; f. female flower. — g. Alsomitra macrocarpa
(Blume) M.Roem. Node with base of petiole and tendril with at apex adhesive pads (a: De Wilde &
Duyfjes 21759; b: Phonsena 3522; c. Korthals s.n. (33); d: De Wilde & Duyfjes 22125; e, f: Giesen 76
(type); g: McDonald BS 18711).
Fig. 98. Zanonia indica L. subsp. orientalis W.J.de

Wilde & Duyfjes var. paludosa W.J.de Wilde & Duyf
jes. Twig with infructescence (Anderson S 25559).
2 cm
1a. Ovary 8 –10 mm long; petals 4 – 5 mm long. Fruit 4 –10 cm long, pubescent, gla-
brescent or glabrous; pericarp finely wrinkled or irregularly pustulate . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. var. pubescens
b. Ovary c. 5 mm long; petals c. 3 mm long. Fruit c. 5 cm long, glabrous; pericarp
smooth or finely regularly pustulate . . . . . . . . . . . . . . . . . . . . . . . b. var. paludosa
a. var. pubescens Cogn.

Zanonia indica L. var. pubescens Cogn. in A.DC. & C. DC., Monogr. Phan. 3 (1881) 927. — Type:
Griffith 2521 (lecto K; iso P, designated by Chakravarty, Fasc. Fl. India 2 (1982) 126), India,
Khasia Hills.
Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. orientalis, nom. superfl. (De Wilde
& Duyfjes, Blumea 52 (2007) 288).
Climber 6 –15 m long, glabrous or hairy. Female flowers: ovary 8 –10(–12) mm

long; sepals 2 – 3 mm long; petals 3 – 5 mm long. Fruit 4.5 – 9(–10) cm long; pericarp
irregularly set with low wrinkles or pustules. — Fig. 8g, 96, 97a – d; Plate 20c, d.
Distribution — As the subspecies.
Note — Although the majority of the collections has (sub)glabrous leaves the name
of the variety, var. pubescens, had to be used because of taxonomic priority.
b. var. paludosa W.J.de Wilde & Duyfjes

Zanonia indica L. subsp. orientalis W.J.de Wilde & Duyfjes var. paludosa W.J.de Wilde & Duyfjes,
Blumea 52 (2007) 288, f. 2e, f, 3. — Type: Giesen 76 (holo L; iso BO, not seen), Kalimantan,
Negara River, Barito delta.
Climber 4 – 6 m tall, (sub)glabrous. Female flowers: ovary subglabrous, c. 5 mm long,

1.5 – 2 mm wide; sepals c. 2 mm long; petals c. 3 mm long. Fruit 4.5 – 5 cm long, 2 cm
wide, glabrous, smooth or very finely pustulate. Male flowers not known. — Fig. 97e, f,
98.
Distribution — Malesia: Borneo: Sarawak, Rejang delta; South Kalimantan, Barito
delta.
Habitat & Ecology — In riparian forest or transitional forest between mangrove and
peat swamp forest at sea level; flowering and fruiting: December & January.

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Flora Malesiana, Series I, Volume 19 (2010) 1– 333

and Paciﬁc); in Africa e.g., Momordica, Cucumis, Pilogyne, and Neoachmandra; in

HABITAT & ECOLOGY

Fevilleoideae Burnett, Outl. Bot. (1835) 756, 1092, 1129.

Dioecious. Tendrils distally 2-branched, spiralling below point of branching. Male

Monoecious or dioecious. Tendrils simple, or branched mostly at or below the middle

POLLINATION AND DISPERSAL

Flowers in Cucurbitaceae are unisexual, the plants either monoecious or dioecious.

The conspicuously coloured, berry-like, pulpy fruits of most Cucurbitaceae suggest

GERMINATION AND SEEDLING

INDUMENTUM, COLOUR OF PLANT ON DRYING

Cucurbitaceae differ from other scandent-climbing plant families by the presence of a

LEAVES AND LEAF GLANDS

INFLORESCENCES AND FLOWERS

Flowers are solitary or (few-)fascicled in the leaf axils, or arranged in an axillary

Bracts — The bracteate racemes of male flowers bear persistent or caducous bracts,

Receptacle-tube — The receptacle-tube is present in subfamily Cucurbitoideae. It is

Their shape in male flowers, whether entire or lobed, is taxonomically important in

Male flowers — The number of stamens is 3 or 5. In subfamily Fevilleoideae there are

Nectary & staminodes — A ring-shaped nectary, as well as staminodes are often

Fruit — In subfam. Cucurbitoideae it usually is a rather dry or juicy, indehiscent berry;

There is a large diversity in chromosome number within Cucurbitaceae. Schaefer &

Jeffrey (2001) gives a detailed worldwide account of cultivated Cucurbitaceae, while

Tribes Actinostemmateae, Fevilleeae, Gomphogyneae, Zanonieae (19 genera; Malesia:

Tribe Benincaseae (32 genera; Malesia: Anangia, Benincasa, Borneosicyos, Citrullus,

Tribe Bryonieae (Austrobryonia, Bryonia, Ecballium; Malesia: absent).

Tribe Coniandreae (21 genera; Malesia: Kedrostis).

Tribe Cucurbiteae (13 genera; Malesia: Cayaponia, Cucurbita).

Tribe Indofevilleeae (Indofevillea; Malesia: absent).

Tribe Momordiceae (Momordica; Malesia: 5 species).

Tribe Schizopeponeae (Biswarea, Edgaria, Herpetospermum, Schizopepon; Malesia:

Tribe Sicyeae (24 genera; Malesia: Cyclanthera, Gymnopetalum, Hodgsonia, Luffa,

Tribe Siraitieae (Microlagenaria, Siraitia; Malesia: Siraitia).

Earlier accounts on the taxonomic and evolutionary signiﬁcance of pollen characters in

KEYS TO THE GENERA

Note on the distinction of the subfamilies. — According to molecular phylogeny the

1 – GENERAL KEY FOR MALE FLOWERING MATERIAL

1a. Tendrils distally 2-branched. Inflorescences paniculate; bracts usually minute,

3a. Petals valvate. Anthers 3 – 4 mm long, 2 two-thecous, 1 one-thecous. Perianth

33a. Plant coarsely rough-hairy. Flowers subsessile or short-pedicelled (pedicel 2–10

2 – GENERAL KEY FOR FRUITING AND FEMALE FLOWERING MATERIAL

b. Fruit claviform-cylindric, bluntly triangular, 8 –10 cm wide. Seed toothed, with

16a. Fruit green or yellow, fenestrate, hardly beaked . . . . . . . . . . . . 34. Thladiantha

30a. Tendrils several-branched (2- or more-branched). Probract conspicuous. Petals

43a. Tendrils 2-branched. [Thecae plicate.] — East Java, naturalized . . . . . . . . . . . . .

1. Alsomitra macrocarpa (Blume) M.Roem.

Climber, 30 – 50 m long, stem at base 15 cm thick. Shoots grooved, 3 – 4 mm diam.,

1 or 2 times branched, 5 –15 cm long, peduncle (1–)2 cm long, sometimes with 1 or 2

Low suberect herb, tuberous(?); monoecious. Probract present in association with

1. Anangia macrosepala W.J.de Wilde & Duyfjes

Herb, several-stemmed, c. 30 cm high, subglabrous; stems suberect, much branched,

Fig. 4. Anangia macrosepala W.J.

1. Baijiania borneensis (Merr.) A.M.Lu & J.Q.Li

KEY TO THE VARIETIES

b. var. paludicola Duyfjes

1. Bayabusua clarkei (King) W.J.de Wilde

2 – 6 mm diameter. Leaves: petiole 0.5 – 3 cm long, 1– 2 mm wide; blade faintly laterally

Stout herbaceous annual climber; plant hairy; monoecious (sometimes flowers

1. Benincasa pruriens (Parkinson) W.J.de Wilde & Duyfjes

Annual herb, shoots 2 –7 m long, stem 3 – 5 mm diam., sparsely or densely grey or

Distribution — The cultivated form widespread in SE Asia and elsewhere.

KEY TO THE FORMS

b. forma hispida (Thunb.) W.J.de Wilde & Duyfjes

Herbaceous or (sub)woody perennial climbers, early glabrescent; dioecious. Pro

Liana, to 30 m long, leafy shoots 4 –7 mm diam., almost glabrous; dioecious. Pro