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SERIES I - SPERMATOPHYTA.
Flowering Plants
Vol. 10, part 1

Revisions

AceraCeade .... cence 4:.3, 592
Actinidiaceae s. str. ..... CS |
ASZGACEAG 165.0415. 2% ke WER 4: 267
Alismataceae .... 5: 317; 6: 915
Amaranthaceae
4: 69, 593; 6: 915
Anacardiacede...: 6... 6% 2! 8: 395
Ancistrocladaceae ....... 4: 8
Aponogetonaceae . 4: 11; 7: 213
AT ANACCAGA A shih 2 cota oe |
Aristolochiaceae ....... 10: 53
Balanophoraceae........ ie 763
Basellacere! .i. deni bana 5: 300
Batidacede hoe re ee 5: 414
Betulaceae:.. ..:. 5: 207; 6: 917
BISTIONIACEAE © via) e)oyeh aloes a 8: 114
BixaCEAG’S.4SEl, Maine ere ae 4; 239
Burmanniaceae...... 4: 13, 592
Burseraceae
522209; 5675062 9172 7-820
BitOMmACEAR A. eens 5: 118
BYDIGACEAG hisciet hoard PE A lsis)
CalhitrichaceaG 4.0022 a: 4: 251
Campanulaceae .... 6: 107, 928
Cannabinaceaes...,. ....-- 4: 223
Gapparidaceae a. sz. sock 6: 61
Caprifoliaceae
4: 175, 598; 6: 928
Cardiopteridaceae ....... fs» 93

Celastraceac... 6: 227, 389, 930

Centrolepidaceae........ 5: 421
Ceratophyllaceae........ AS Al
Chenopodiaceae

4: 99, 594; 6: 932
@lethrdcede... 5. mrten cer 7: 139
Cochlospermaceae....... 4: 61
Combretaceae

4: 533; 5: 564; 6: 932
Connaraceae .... 5: 495; 6: 933
Convolvulaceae.... 4: 388, 599;

5: 558; 627956; 7: $23

ME GTNACCAG . Sess.c\c. 2 cele oe 8: 85
Corynocarpaceae

4:5 262%,.5:) ‘S57
GGrassulacede ef eo oe 4: 197
Crypteroniaceae......... 8: 187
Syperacede.. .... «.. 1224355.9" 107
Watiscaceae®. 0. ok... bales 4: 382
Dichapetalaceae . 5: 305; 6: 941
Dilleniaceae ..... 4: 141; 7: 824
Dioscoreaceae .......... 4: 293

THE LuESTHER T. MERTZ LIBRARY

THE NEW YORK BOTANICAL GARDEN

TO REVISED FAMILIES

Dipsacateae.4 4 sieretie ao 4: 290
Dipterocarpaceae........ 9: 237
Droseraceae..... DA ig be pia i
Blatinacese::.. + is eee 4: 203
Epacridaceae*.... .s.7hs >. 6: 422
ENICACEAGH id nena t 6: 469, 943
Erythroxylaceae.:....... 5: 543
PaSaCene Vie eee ee ied 7: 265
FICMIGHEEAG 1) dr 55 tes nt oe 4: 267
Flacourtiaceae
Sols, SOI) OF 943) Ts 827
Flagellariaceae ...:...... 4: 245
Geraniaceae .........08- 6: 445
Gnetacedesc: . <2 4: 336; 6: 944
Gonystylaceaei. ones 4: 349
Goodeniaceae
5: 335, 567; 6: 949; 7: 827
Haemodoraceae......... Seo
Fialorapacedey. 2 a..nsi3e os Tt Zao
Hamamelidaceae ........ 5: 363
Hippocrateaceae ........ 6: 389
Hydrocaryaceae......... 4: 43
Hydrocharitaceae
5: 381; 6: 952
Hydrophyllaceae ........ 4: 207
IV PENIGACCAG# ater, ects rar ae
IGAGINAGEAE rece sat oe Tt
MiG acederas ne er ct cents, boss 82-77
Tuglandacede) ac. 36 als << 6: 143
NINGACGHE etch ae ont e o¥e ove 4: 210
Juncaginaceae .......... 42.557
Mabiatae: Hchiges icerse oc 8: 301
WGGACEAG Ty sirens ects 6 fHRTSE)
MeniACEAC). shellac ene 7: 219
Lentibulariaceae ........ 8: 275
Lilidcede-Dae bite ot arse eis 9: 189
Loganiaceae® .’.:.2..< 6: 293, 953
Lophopyxidaceae........ ae Oo
Malpighiaceae .......... Rel PS
Martyniaceae ........... 4: 216
Molluginaceae .......... 4: 267
Mornmeacedese <2 -7ac-452 4: 45
Myoporacéae <u... > 4: 265
WMiyniCaceae . wittn rosie cont: a5 °277
Najadacedern aceon a G2 157
Nyctaginaceae: 2 )0) 22: « 6: 450
INVSSACESE scoters camera 4: 29
Ochnacede sso. hese oe eR 7)
Olacaceae her nce to 10S el
Onasracede hice. see 8: 98
Opiliaceae ete 10: 31

a eee

Oxalidaceae-:\7.\...1; cae ae
Papaveraceaé ...> .sanene o:
Passifloraceae >. ..aaeure ee
Pedaliaceae.. . ...; {osm 4:
Pentaphragmataceae..... 4:
Pentaphylaceae ......... os
Philydraceae......7..seee 4:
Phytolaccaceae.......... 4:
Pittosporaceaé ....s28e% 5:
Plumbaginaceae......... 4:
Podostemaceae ... 4: 65; 6:
Polemoniaceae.......... 4:
Pontederiaceae: sass es 4:
Portulacaceaé yee a
Primulaceae’ ) yas 6:
Proteaceae ':< ase 5:
Punicaceaé «. .Gianevemee 4:
Restionaceae.. > estas eae 5
Rhizophoraceae
5: 429; 6: 965; 8:
Salicaceae\. ... (eure ae
Salvadoraceae Fazeeee eee 4:22
Sarcospermacede. sees fee
Saururaceae <2 sane 4:
Scyphostegiaceae
5: 2975.6:
Simaroubaceae ..... 6: 193,
Sonneratiaceae
4: 280, 513; 6:
Sparganiaceae .......... 4:
Sphenocleaceae ......... 4:
Stackhousiaceae......... -
Staphyleaceae =) 7240s 6:
Stylidiaceae.. ja2js0s eee 4:
Styracaceae’. .4..-.sieeeaees 4:
Symplocaceae =) ane 8:
Taccaceae .../1./.\4 sae ae
Thymelaeaceae
4: 349; 6: 1, 976; 7:
Trapaceaé': eae 4:
Trigoniaceae). eeee 4:
Triuridaceae:. 7 eee 10:
Turneraceaé: ..ieem ate 4:
Typhaceae. .... 2. aera .
Ulmaceae : {ace aes
Umbelliferae ...... 4: 113, 595;
5: 555; 6: 983; 7: 83
Valerianaceae . > ose. sees 4: 25
Violaceaé.. +... seams THe lye
Xyridaceae ........ 4: 366, °
Zygophyllaceae ......... t

7 10 OLACACEAE (H. Sleumer, Leyden)

Trees or erect, rarely scandent shrubs, sometimes hemi-, rarely autoparasitic.
Leaves spirally arranged, rarely distichous, simple, entire, often with parch-
ment-like and/or finely tuberculate surfaces, mostly penni-, rarely pli-nerved,
petioled, exstipulate, not rarely of a greyish-yellowish-olivaceous colour and
dull, especialiy in the dry state. /nflorescences axillary, rarely on old wood,
short racemes and panicles, or elongate spikes, often fascicles or glomerules,
these rarely reduced to a solitary flower. Flowers generally bisexual, rarely uni-
sexual (monoecious or andro-dioecious), generally actinomorphic, cyclic, 3—7-
merous, rarely heterostylous. Ca/yx small in anthesis, often very shortly 3—7-
lobed, -dentate, or -crenulate, the cup-like base free or adnate to the disk and/or
ovary to various degrees, afterwards sometimes accrescent, and then either free
from or connate with the fruit. Peta/s 3—7, free or connate below, valvate, ca-
ducous. Disk sometimes present, consisting of free glands, or cup-like, rarely
accrescent and then covering the fruit almost to the apex. Stamens 1—3-seriate,
hypogynous, 4—15 in number, epipetalous, or partly also episepalous, rarely in
part staminodial; anthers basi- or medifixed, with 2 thecae, or rarely with 1 the-
ca, dehiscing lengthwise. Ovary mostly superior, rarely semi-inferior when im-
mersed in the disk, or inferior when connate with the cup-like flower-axis
(Schoepfia), either 1-locular with 2—3 (—5, —7) ovules pendent from the apex
of a central free placenta (sometimes projecting into the stylar canal), or 3—5
(—7)-locular in the lower part only (rarely completely so), a single ovule hanging
then from the inner angle into each of the cells; ovules generally anatropous,
uni-, bi-, or ategmic; style, if any, conical, columnar or filiform, with a small,
sometimes 3—5-partite or -lobed, subsessile stigma. Fruit a drupe with a thin
and often fleshy, sometimes dehiscent or caducous exocarp, and a crustaceous
to woody endocarp, or concrescent with the cup-shaped floral axis, or with an
accrescent calyx or disk which then forms an external fleshy layer. Seed 1; testa
(if any) thin; endosperm abundant, starchy and/or oily, bearing the embryo at
its apex; cotyledons 2, 3, or 4.

Distribution. A pantropical family with about 27 genera and approximately 170 spp., pre-
dominantly in the tropics, a few in the subtropics.

In Malesia 9 genera with a total of 14 spp. Of these only Ochanostachys is strictly limited to
Malesia. Erythropalum, Harmandia and Scorodocarpus are Indo-Malesian. Some have a wider
Old World range, viz. Anacolosa (\ sp. in Central Africa, 2 spp. in Madagascar and 3 spp. in the
Pacific), Olax (also in Africa, Australia, and the Pacific), and Strombosia (also in Africa).
Schoepfia is Indo-Malesian, with c. 20 spp. also in tropical America. Ximenia is pantropical. The
genus Malania (limited to SW. China) is closely related to Scorodocarpus.

The Malesian representatives thus show a distinct alliance with those of SE. Asia, and a less
marked one with Australia and the Pacific (Anacolosa, Olax, Ximenia). Alliances are strong with
Africa in the genera Anacolosa, Olax, and Strombosia.

Ecology. Most Malesian Olacaceae occur in primary and secondary lowland (also littoral)
rain-forest. Olax and Ximenia are found mainly in drier vegetation types as teak forest, brush-
wood or beach vegetation. Ximenia sheds its leaves in the dry season.

The size of Malesian Olacaceae is mostly moderate to small, and the majority belongs to the
forest substage; they never become dominant,

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LEMBAGA BIOLOGI NASIONAL-LIPI

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Z FLORA MALESIANA [ser. I, vol. 10!

Parasitism. Several Olacaceae are known for their non host-specific parasitism, as is a common
feature in Santalaceae and Loranthaceae. Root haustoria have been found in Olax and Ximenia
in Asia, and also in Ptychopetalum and Schoepfia, both in America. The extent to which this
parasitism occurs in these and possibly in other genera is not known. For Malesia proper no data
on parasitism have been published but it can be expected for Olax scandens, Ximenia americana,
and maybe for Schoepfia fragrans. Cf. BARBER, Studies in root-parasitism. The haustoria of Olax
scandens. Mem. Dept. Agr. India, Bot. ser. 2 (4) (1907) 1—47; Kuyt, The biology of parasitic
flowering plants (1969) 65.

Dispersal. Little is known of the seed dispersal of Olacaceae; their fruits with fleshy pericarp
and a big seed point to a possible dispersal by animals, mainly monkeys and birds eating the fruits.
Fruits of the coastal Ximenia are able to float for some time in seawater (RIDLEY, Disp., 1930,
195, 265, 346).

Morphology. The family has an interesting morphology in that, though small in size, it ex-
hibits pro ratio a great pluriformity in important features as compared with many other families.
The habit may be erect or scandent, plants may be armed with spines or thorns (Ximenia, Olax)
or unarmed; in Erythropalum axillary tendrils occur (fig. 8), even rarely bifid. Of several genera
it has been proved that they are hemiparasitic.

Also in the flowers variability occurs in merousness, and stamens, which are usually epipeta-
lous, may also occur partly episepalous, rarely in part staminodial; stamens may be up to
3-seriate.

The ovary is superior, but may become through various ways of concrescence with disk and/or
receptacle inferior or lead to an inferior fruit.

In Erythropalum flowers are bisexual or andro-dioecious, in Ximenia flowers are bisexual or
rarely functionally unisexual, in Olax scandens and Schoepfia flowers are often heterostylous, in
Harmandia flowers are monoecious, in many others they are normally bisexual.

The ovary is either one-celled with a central placenta or the lower part is more-celled with ovules
pendent in these cells. Ovules may be bitegmic, unitegmic or even ategmic.

Phytochemically there is also variability: in Ochanostachys and Harmandia tissues contain cells
with milky juice, Scorodocarpus reeks of garlic, while Erythropalum has also a bad smell.

Leaf and wood anatomy and pollen morphology are also very diverse; see below.

This pluriformity is striking, because the family must be of ancient date, as can be derived from
the fact that it does not only range over the tropics of all continents but even three genera are trans-
Atlantic and one trans-Pacific.

Galls. Cf. DocTERS VAN LEEUWEN, Zoocecid. Neth. Ind. (1926) 175 (galls of Anacolosa frutes-
cens).

Embryology. Cf. AGARWAL, Phytomorphology 11 (1961) 269—272 (Strombosia); ibid. 13
(1963) 185—196 (Olax).

Phytochemistry. There is a more or less general tendency in Olacaceae to deposit oxalate
of lime in various parts, and silicic acid in leaves (not in wood). Seeds tend to be rich in oil. Trigly-
cerides with C-18 acetylenic acids such as santalbic (ximenyncic), isanic and isanolic acid occur
amply in seed oils (e.g. Ximenia americana), but are also present in roots, stems and leaves (Xime-
nia americana, Olax stricta), linking Olacaceae biochemically with Santalaceae and Opiliaceae.

The lack of knowledge about polyphenolic compounds in Olacaceae is astonishing. Tannins,
probably of mixed origin (mainly flavonoid type, but sometimes accompanied by galloyl tannins)
are abundantly present in the bark, roots or leaves of Ximenia americana, Anacolosa spp., Olax
spp., and others.

Prunasin-like (i.e. yielding HCN and benzaldehyde) cyanogenic glycosides are present in dif-
ferent parts of Ximenia spp. and Olax spp. Saponins seem to occur rather widely in Olacaceae.
Olacaceous sapogenins appear to be mainly triterpenoids; this character is shared with, among
others, Opiliaceae and Santalaceae.

Alcaloids are possibly present in some species of the palaeotropic genera Anacolosa, Olax, and
Strombosia.

1984] OLACACEAE (Sleumer) 3

Literature. R. HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 227; in Sleumer, Olacaceae,
Flora Neotropica, in press. — R. HEGNAUER.

Vegetative anatomy. For general accounts see SOLEREDER (1899, 1908), METCALFE &
CHALK (1950), REED (1955) and Baas et a/. (1982, with full references to older literature). Leaf
and wood anatomy of the Olacaceae are very diverse, but still support the concept of a natural
family. Stomatal type, secretory cavities, laticifers, silicified cells, nodal, petiole and midrib vas-
culature, idioblastic sclereids, type of vessel perforations, parenchyma distribution, ray type, and
fibre pitting all show distinct character states enabling a reconstruction of phylogenetic trends and
relationship patterns within the family (see Baas et a/., 1982; REED, 1955). Most Malesian genera
have their closest relatives in Africa and/or the New World. Ochanostachys is anatomically more
or less identical to Coula (Africa) and Minquartia (South America) and has both secretory cavities
and laticifers. Harmandia strongly resembles Aptandra from Africa and South America and
shares the occurrence of infrequent (reduced?) laticifers in the mesophyll. Anacolosa has its clos-
est relative in the neotropical genus Cathedra. Olax, Schoepfia, and Ximenia belong to a larger,
anatomically fairly homogeneous group including African and neotropical representatives. This
group also shares many characters with Anacolosa and Cathedra on the one hand, and with San-
talaceae and Loranthaceae on the other. Strombosia belongs to a well defined anatomical assem-
blage including Strombosiopsis and Diogoa from Africa and Tetrastylidium from South America.
This group is unrelated to Anacolosa, with which it has been placed in the same tribe Anacoloseae
in the past. Scorodocarpus shows remote affinities to Strombosia, but is closer anatomically to
the neotropical Brachynema. Erythropalum remains anatomically fairly isolated within the fami-
ly, but differences are insufficient to advocate a separate family. The geographical distribution
of the anatomical units (largely coinciding with the traditionally recognized tribes, with the ex-
ception of the Anacoloseae) is suggestive of considerable age and conservatism of the anatomical
character complexes. For a key to the genera based on leaf anatomy, see BAAS et al. (1982). A
detailed wood anatomical survey of the family is in preparation (L. VAN DEN OEVER, Blumea).

Literature: BAAS, VAN OOSTERHOUD & SCHOLTES, Allertonia 3 (1982) 155—210; METCALFE &
CHALK, Anatomy of the Dicotyledons I, Oxford (1950); REED, Mem. Soc. Brot. 10 (1955) 29—79;
SOLEREDER, Systematische Anatomie der Dicotyledonen & Erganzungsband, Stuttgart (1899 &
1908). — P. Baas.

Pollen morphology. General description. Olacaceae exhibit a variable pollen morphology.
In Olax and Ptychopetalum both intra- and interspecific, geographically based variation occurs.
In general Olacaceous pollen grains are single; size varies between 11 tm (Heisteria micrantha)
and 48 um (Olax benthamiana) and shape is basically subequiaxe, ranging from oblate to perob-
late in Anacolosa and Olax to subprolate in Diogoa and Tetrastylidium. Mostly grains are
isopolar, but subisopolar and heteropolar types occur also. This heteropolarity may be expressed
in sculpturing (Coula p.p., Ochanostachys), in apertures (Heisteria p.p., Strombosia p.p.), or in
shape and apertures (Aptandra, Harmandia, Ongokea, Schoepfia). In Coula, Heisteria and
Strombosia both isopolar and heteropolar types occur.

According to apertures, Olacaceae can be divided into four main groups, A, tricolpate with an
endoaperture which is not wider than the colpus or tricolporate with a rectangular or slightly ellip-
tical endoaperture at the equator, B, 3- (4-, 5-) stephanoporate and C, 6-diploporate. In Schoepfia
syncolpate ectoapertures are found at the proximal pole. The apertures are generally closed by
a granular-verrucate membrane.

The sculpture of the tectum varies between psilate (Erythropalum, Minquartia, Schoepfia), per-
forate (Scorodocarpus), reticulate (Diogoa, Strombosiopsis, Ximenia p.p.) and microechinulate
(Curupira, Octoknema). The sculpture may be different between apo- and mesocolpia. Chauno-
chiton has unique sculptured ridges. Infratectal structure is mostly granular but transitions to a
columellate structure are frequent. In Anacolosa and Chaunochiton distinct but always irregular
columellae are developed.

The footlayer is generally present, visible in the mesocolpia in Chaunochiton. This layer is often

4 FLORA MALESIANA [ser. I, vol. 10!

sculptured on the inner side of the apertural margin. The footlayer is especially thick when the
endexine is missing (Coula, Minquartia, Ochanostachys, Schoepfia p.p., Ximenia). The endexine
is mostly confined to the apertural areas, except in Anacolosa, Cathedra, Chaunochiton and Pha-
nerodiscus in which the endexine is thick and continuous, and in Erythropalum, Heisteria,
Schoepfia p.p., Scorodocarpus and Strombosia in which the endexine is very thin and continuous
in the mesocolpia.

The tricolpate isopolar pollen types of Group A, found in Coula, Curupira, Heisteria p.p., Ap-
tandropsis and Minquartia, and the tricolporate pollen types of Group A, found in Diogoa,
Strombosiopsis and Tetrastylidium are considered primitive.

Porate types in Aptandra, Brachynema, Dulacia, Harmandia, Olax p.p., Ongokea and Ptycho-
petalum p.p. as well as 6-diploporate grains characterizing Anacolosa, Cathedra, Phanerodiscus
and Ptychopetalum p.p. are probably derived.

The heteropolar-tetrahedral pollen grains of Schoepfia and the brevicolpate grains of Chauno-
chiton with its ectexinal ridges seem morphologically isolated and may also represent derived
types.

Intergeneric relationships. Couleae and Heisterieae p.p. (Heisteria, Aptandropsis) are pollen
morphologically related. The mutual affinities between Heisteria and Chaunochiton are weakly
expressed in aperture characters and by the loss of the tectum in the intercolpium, and this last
genus could be placed in a separate tribe of its own.

Anacolosae fall into two quite distinct generic groups, |, Diogoa, Scorodocarpus, Strombosia,
Strombosiopsis, Tetrastylidium and Il, Anacolosa, Cathedra and Phanerodiscus. Brachynema is
isolated.

Within Olaceae, the pollen of Ptychopetalum is very distinct from that of Olax and Dulacia.
It is distinct also from that of Anacolosa.

Pollen of Aptandreae (Aptandra, Harmandia and Ongokea) is uniform.

The morphology of Schoepfieae offers no clue to its affinities, but the ultrastructure of the ex-
ine draws them near to the Couleae.

Relationship of family. Olacaceae pollen shows resemblance to Opiliaceous and Santalaceous
pollen. Some similarity also exists between the pollen of the Olacaceae and Icacinaceae.

Fossil occurrences. Pollen of the Anacolosa type is known from the Maestrichtian onwards,
while O/ax type pollen has been recorded from the lower Eocene. The pollen of Ximenia has been
found in Quaternary sediment in East Africa.

Literature: R. BONNEFILLE, D. LOBREAU-CALLEN & G. RIOLLET, J. Biogeogr. 9 (1982) 469—486;
G. ERDTMAN, Pollen morphology and plant taxonomy, Angiosperms (1952) 295—297; S. FEUER,
Pollen morphology and evolution in the Santalales s. str. Thesis Univ. Mass. (1977), Amer. J.
Bot. 65 (1978) 759-763; D. LoBREAU-CALLEN, Adansonia sér. 2, 20 (1980) 29—89; Bot. Jahrb.
103 (1982) 371—412; Bull. Lab. Géol. Genéve (1983) in press; J. MULLER, Bot. Rev. 47 (1981) 84;
C. REED, Mem. Soc. Brot. 10 (1955) 29—79. — D. LOBREAU-CALLEN.

Chromosomes. Only in the allied Santalaceae species of a fair number of genera have been
examined; this yielded 2n = 20, 24, 30, 38, 40, and 72.

The two species of Olacaceae examined, one in Heisteria and one in Strombosia, yielded 2n =
38 and 40 respectively. The one species examined in Opiliaceae, viz. of Opilia, showed 2n = 20.
Although the evidence is small, it does support the affinity between the three families.

Literature: AN.A. FEpoRov (ed.), Chromosome numbers of flowering plants. Leningrad
(1969).

Taxonomy. The family as a whole is characterized by a free basal central placenta from which
a single ovule is pendent into each of the generally imperfect cells of the ovary, or in case of a
1-celled ovary, several ovules from the apex of such a free placenta. The ovules are bitegmic, or
more often unitegmic, or have — mainly in parasitic species — no integument at all. The ovary
is hypogynous in principle, but may become semi-hypogynous or even epigynous by concrescence
with the calyx, disk or flower-axis. The fruit is drupaceous, not rarely + included by the accres-
cent calyx or disk.

1984] OLACACEAE (Sleumer) 5

Olacaceae are regarded by ENGLER (Syllabus, 1924) to represent the most primitive family of
the Santalales with regard to the occurrence of hemiparasitism and the reduction in number of
the integuments. ENGLER has divided the family in the first edition of ENGLER & PRANTL, Nat.
Pfl. Fam. (III, 1, 1894; Nachtr. 1, 1897; Nachtr. 3, 1908) into 3 subfamilies with 6 tribes mainly
on the base of the presence or absence of integuments and the position of the micropyle on the
ovulum. These subdivisions have been maintained by the author in the second edition, 16b (1935).
However, the characters used by ENGLER to distinguish subfamilies were based on too scanty ob-
servations to prove the constancy needed for such high taxonomic rank as that of a subfamily.

Characters may prove more variable than assumed; for example AGARWAL (Phytomorphology
13, 1963, 185) found that in O/ax both unitegmic and ategmic species occur, which means that
more observations in embryology are needed.

The tribes distinguished by ENGLER are mainly based on the presence or absence of starch
and/or fatty constituents in the endosperm, and on the amount of fusion between the stamens.
These tribes are not well established as the mentioned chemical constituents are not fully known
yet in all members of the genera concerned. At the moment a subdivision of Olacaceae into natural
subfamilies and tribes is still open.

Uses. Scorodocarpus borneensis provides a deep red timber (Ku/im). The timber of other
genera (Anacolosa, Strombosia) is less important, usually of small size, and only locally used.
Ximenia americana has a hard yellowish wood similar to Sandal wood, and is used locally. Young
leaves of Strombosia javanica are eaten. Edible fruits are known of Scorodocarpus borneensis and
Ochanostachys amentacea. The kernel of Ximenia americana contains a strong purgative.

Note. The most important paper on Malesian Olacaceae has been written by VALETON (Crit.
Overzicht Olacineae; Inaug. Diss., Groningen, 1886). In a precursor I have given an account of
the genera and species of Asia, Malesia and adjacent areas (Blumea 26, 1980, 145—168). All
genera of this part of the world occur also in Malesia, with the exception of the monotypic genus
Malania which is endemic in SW. China.

KEY TO THE GENERA
based on flowering material

1. Leaves 3—S-plinerved, at base subpeltate. Climbing shrub; axillary tendrils often present
6. Erythropalum
1. Leaves penninerved (if + 3-plinerved a tree), never peltate. Tendrils absent.

Doncmens and staminodes present... 0) bi.iiac Slaie so las aera lead ie Oe eee Oe eae 1. Olax
2. Stamens present; staminodes absent.
3. Stamens fully fused into a tube. Flowers unisexual, plant monoecious ............... 2. Harmandia

3. Stamens not fused (sometimes partly adnate to the petals). Flowers generally bisexual.
4. Stamens 8 or 10, half of them epipetalous, the other half episepalous. Leaves usually mucronulate at

apex: Spines and/or thorns generally ‘present: |: v0). eee a ee ee 3. Ximenia
4. Stamens all epipetalous. Leaves not mucronulate at apex. Spines and/or thorns absent.
5. Flowers interruptedly arranged in elongate spikes...............-0..eeeeeeuee 4. Ochanostachys
5. Flowers arranged in short racemes, panicles, or mostly in fascicles.
6. Stamens 8 or 10, arranged in pairs before each petal ...................000. 5. Scorodocarpus
6. Stamens 4—7, solitary before each petal.
Wz Petals ienitively: free... 8 55 ic 3x iie iy vei cod:s vo vcs SpA bint IPR i dt ee 7. Strombosia

7. Petals fused in the lower half.
8. Calyx cupular, + truncate to very shortly 6-dentate. Petals thick-fleshy. Anthers apically penicil-
DEO dish dil SC AEOEV Mew Tae ees. Has Aca ae Cie eee 8. Anacolosa
8. Calyx indistinct, merely a rim with minute teeth. Flower supported by an epicalyx consisting of 3
concrescent bracts. Petals thin. Anthers not penicillate at apex...............4. 9. Schoepfia

6 FLORA MALESIANA [ser. I, vol. 10!

KEY TO THE GENERA
based on fruiting material

1. Leaves 3—5-plinerved, often subpeltate. Climber, often with axillary tendrils. Fruit long-stipitate narrowed
to the base. Pericanp Gebiscent. Inside meds) Seed) DING yar rei eis cies ee 6. Erythropalum
1. Leaves penninerved (if + triplinerved a tree). Fruit at most very short-stipitate, usually rounded or obtuse
at base. Seed whitish.
2. Calyx indistinct, merely with a rim. Fruit inferior, supported by a persistent epicalyx consisting of 3 con-
Crescerit' bracts. & ait ek Ae Be Se nee: shee at a AP Re 9. Schoepfia
2. Calyx distinct. No epicalyx.
3. Calyx much enlarged in fruit.
4. Enlarged calyx connate with the fruit only in its lower part, for the rest frill-like expanded, very large
2. Harmandia
4. Enlarged calyx enveloping the fruit for its entire or almost entire length................. 1. Olax
; 7. Strombosia
3. Calyx not accrescent in fruit.
5. Disk much accrescent, adnate to and almost entirely covering the fruit (which bears the persistent calyx

ALLS ID ASE) erento Serene etre tN scree a tel ore © Coe hone Gee teas chats Mielere mer ciate mera ee cete nts OMeee 8. Anacolosa
5. Disk, if present, not accrescent.
6Petiole veryadistinctlysswollenvdistalllyacmiar:- seco o oes a a cite tele eer ete on dele 5. Scorodocarpus

6. Petiole slightly or not thickened distally.
7. Leaves usually mucronulate at apex. Axillary spines and ramal thorns may be present 3. Ximenia
7 CAVES MOLMuUCKOnUlate al apexes SPINES Ol PhOnnSpalbSeitee precy orien 4. Ochanostachys

KEY TO THE GENERA
based on sterile material

1. Leaves usually mucronulate at apex, deciduous in the dry season. Branchlets usually with axillary spines
and/ombrachyblastsrenGinsamachOnmSpe spree cele ceca eet cenenetetc re tench sei aictetere aieteet ale tetera eteee 3. Ximenia
1. Leaves not mucronulate at apex, persistent. Branchlets usually without, in 1. Olax pr. p. sometimes with
ramal thorns.
2. Climbing shrubs
3. Leaves subpeltate, 3—5-plinerved. Branches often with spring-like lignescent tendrils 6. Erythropalum
3. Leaves not subpeltate, exclusively pinninerved. Tendrils absent .................... 1. Olax pr. p.
2. Erect shrubs or trees.
4. Leaves markedly distichous.

SRO DINES LOL ChOM Ss SOMeLIMeS PLESCMl Merrit arcs sachs ciacc Sates ate atrie irae 1. Olax pr. p.

See PINEsvOn LHOLNSsaADSENimee eee sek eet ee ee © © foc on slept Cuieetteleneve s aiseeds 2. Harmandia
4. Leaves indistinctly or not distichous.

6 Petiolerconspicuouslyathickenedidistally mer -taeitae kia ete sent -fob-t-t erections te tne ek 5. Scorodocarpus

6. Petiole hardly or not thickened distally.
7. Leaves usually showing scattered blackish points on both faces; nerves slightly impresien above
4. Ochanostachys
7. Leaves without such blackish points; nerves not properly impressed above.

8. Leaves usually with numerous fine pellucid points visible against strong light..... 7. Strombosia

8. Anacolosa

Sy Weavesinot pellucid=-punctate’ 4. - 2... ce oe oc I een ee vera teh <5 cele Sas 9. Schoepfia
1. OLAX

LINNE, Sp. Pl. (1753) 34; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 ((6s9)23i-
SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 24; Blumea 26 (1980) 154.
— Drebbelia Zou. Nat. Tijd. N. I. 14 (1857) 160; ENGL. in E. & P. Nat. Pfl.
Fam., Nachtr. 2 (1900) 18. — Fig. 1.

Trees, shrubs or subshrubs, sometimes climbing, occasionally armed with ra-

1984] OLACACEAE (Sleumer) a

mal thorns. Leaves spiral, sometimes (sub)distichous, penninerved. Flowers in
racemes, panicles or spikes, rarely solitary. Ca/yx cup-shaped, truncate or ob-
scurely dentate, small in anthesis, much accrescent in fruit. Peta/s 3, entire, or
all or in part bipartite and thus seemingly 6 (rarely 5), inserted on a conical disk,
free or connate in pairs. Stamens 8 (Mal.), partly adnate to the petals below, 3
of them fertile, and 5 staminodial (often bifid or bilobed, void of pollen); fila-
ments flat; anthers oblong. Ovary superior, 1-celled, or 3-celled in the lower
part only, with 3 anatropous uni- or ategmic ovules pending from the apex of
a free central short placenta; style short or elongate; stigma capitellate, 3-lobed.
Drupe oblongoid or ovoid to subglobular, included halfway or to almost the top
by the accrescent firmly membranous calyx, though not strictly connate with it;
pericarp parenchymatose; endocarp stony. Seed mostly 1; albumen copious,
containing oily substances.

Distr. About 40 (or less) spp. in the Old World tropics, subdivided by ENGLER into 4 African sections,
and a fifth sect. Triandrae ENGL. which comprises both African spp. and all those found in Asia, Malesia
(2 spp.), Australia, and the Pacific.

Ecol. Most species occur in drier vegetation types; a few are climbers; root-parasitism seems to be fre-
quent.

Morphology. Cf. FAGERLIND, Beobachtungen Uber die Kletterorgane bei Olax; Svensk Bot. Tidskr. 34
(1940) 26—34.

KEY TO THE SPECIES

1. Branchlets with a fine patent pubescence, usually armed with strong ramal thorns. Petals 7—9 mm, white,

not or hardly changing colour in the dry state. Drupe c. 1.5 cm long ................ 1. O. scandens
1. Branchlets more or less glabrous, unarmed. Petals 10—12 mm, becoming dark to blackish in the dry state.
OP 1G C0) 1) a ee Sareea Re me a ey arc ee Tae ae 2. O. imbricata

1. Olax scandens Roxs. P|. Corom. 2 (1798) 2, t. 102;
Mia. Fl. Ind. Bat. 1, 1 (1856) 785; Mast. Fl. Br. Ind.
1 (1875) 575; VALert. Crit. Overz. Olacin. (1886) 114;
Ript. Trans. Linn. Soc. London II, 3 (1893) 286;
Back. Fl. Bat. 1 (1907) 290; Voorl. (1908) 54;
Schoolfl. Java (1911) 222; Ripv. J. Str. Br. R. As.
Soc. 59 (1911) 84; Koorp. Exk. Fl. Java 2 (1912) 172;
Koorp.-Scuum. Syst. Verz. 1 (1912) 2; BEUMEE,
Flor.-anal. Onderz. Djatibosschen (1922) 113; Rip.
Fl. Mal. Pen. 1 (1922) 421; Heyne, Nutt. Pl. (1927)
592; Burk. Dict. (1935) 1578; Back. & BAKH./. FI.
Java 2 (1965) 64; SLeum. Blumea 26 (1980) 157. — O.
obtusa BL. Bijdr. (1825) 131; Mig. Fl. Ind. Bat. 1, 1
(1856) 785. — Drebbelia subarborescens Zou. Nat.
Tijd. N. 1. 14 (1857) 160. — Fig. 1.

Shrub with pendent branches, or generally scan-
dent, 2—20 m; stem 1—15 cm @; bark rather smooth,
grey; old branches with strong obtuse ramal straight
or slightly curved thorns. Branchlets often horizon-
tally spreading, patently puberulous or pubescent at
younger parts, glabrescent below, striate-wrinkled
longitudinally in dry specimens (not transversely
ridged!). Leaves almost distichous, ovate-elliptic-ob-
long, apex broadly attenuate to rounded, base slight-

ly inequilateral, attenuate to obtuse, not rarely round-
ed and plicate, thin-coriaceous, dark to yellowish
green, somewhat shining and glabrous above, initial-
ly short-pubescent at midrib beneath, glabrescent,
2—8 (—9.5) by (0.3—) 0.8—3.5 cm; nerves 5—8 pairs,
unequal-spreading, rather inconspicuous on both
faces as are the reticulations; petiole short-pubes-
cent, 5—7 (—10) mm. Racemes 1—3 per axil, simple
or branched, obliquely ascending, many-flowered,
densely short-hairy, bracteate at base, 0.5—3.5 cm;
bracteoles distichous, obtuse, keeled, pubescent, c. 2
mm, caducous; pedicels thickened at the very base,
glabrous, 1—1.5 mm. Ca/yx cup-shaped, truncate, ci-
liolate, 0.5—1 mm high, c. 1.5 mm @, much accres-
cent in fruit. Petals 3, of which 2 (rarely all) are split
about halfway, thus 5(—6) petals seemingly present,
linear-oblong, apex acute and incurved, glabrous,
white, scented, 7—9 by c, 1.5 mm. Stamens 3, in the
long-styled form reaching to the base of the sinus of
the petals, in the short-styled form reaching 2—2.5
mm beyond it; filaments free for a short part; anthers
oblong, c. 1.5 mm. Staminodes with very narrow
void, deeply bifid cells. Ovary ovoid, glabrous; style
either long (5~6 mm) or short (1.52.5 mm); stigma

8 FLORA MALESIANA

[ser. I, vol. 10!

Fig. 1. Olax scandens Roxs. In teak forest near Djombang, East Java (DE Voocp 893).

obscurely 3-lobed. Drupe broadly ovoid to subglo-
bose, covered for the lower 2/3 or more by the ac-
crescent firmly membranous calyx, apiculate, orange
to yellow (0.8—) 1.5 by (0.6—) 1 cm.

Distr. Widely distributed from Ceylon and tropi-
cal W. Himalayas through India, Burma, Indochina,
Thailand; in Malesia: Malay Peninsula, Java incl.
Kangean Is., Madura, Lesser Sunda Is. (Bali).

Ecol. Mostly in dry deciduous forest or scrub,
also in teak forest, not rarely on rocky ground, often
close to the sea (beach forest, dunes), rarely in light
rain-forest, up to c. 300 m.

Vern. Ganpi, M (Antjol), wangon, J, wuru
wuru, Md.

2. Olax imbricata Roxs. [Hort. Beng. (1814) 5, nom.
nud.] Fl. Ind. 1 (1820) 169; ed. Carey 1 (1832) 164;
DeEcNE, Nouv. Ann. Mus. Paris 3 (1834) 438; Herb.
Timor. Descr. (1835) 110; Zoti. & Mor. Syst. Verz.

(1846) 25; A. Gray, U.S. Expl. Exp. Bot. (1854) 305;
Mia. Fl. Ind. Bat. 1, 1 (1856) 785; Mast. Fl. Br. In-
dia 1 (1875) 575; F.-ViLL. Nov. App. (1880) 45; Vi-
DAL, Sin. Atlas (1883) 20, t. 30, f. A; Phan. Cuming.
Philip. (1885) 102; Rev. Pl. Vasc. Filip. (1886) 85;
VALET. Crit. Overz. Olacin. (1886) 115; CERON, Cat.
Manila (1892) 45; Kinc, J. As. Soc. Beng. 64, ii
(1895) 99, p.p.; Hocure. Bull. Inst. Bot. Btzg 11
(1904) 38; MerrR. Govt. Lab. Publ. Philip. 27 (1905)
32; Philip. J. Sc. 1 (1906) Suppl. 51; Back. FI. Bat.
1 (1907) 292; Voorl. (1908) 54; Merr. Philip. J. Sc.
3 (1908) Bot. 80; Back. Schoolfl. Java (1911) 222;
Merr. Fl. Manila (1912) 185; Koorp. Exk. Fl. Java
2 (1912) 171; MerRR. Sp. Blanc. (1918) 134; En. Born.
(1921) 242; Rip. Fl. Mal. Pen. 1 (1922) 421; MerRr.
En. Philip. 2 (1923) 116; SCHELLENB. Bot. Jahrb. 58
(1923) 158; Baku. Bull. Jard. Bot. Btzg III, 15 (1936)
49; Back. & BAKH. f. Fl. Java 2 (1965) 64; SLEUM.
Blumea 26 (1980) 156. — O. multiflora A. RICH. ex

1984]

Bari. Adansonia 3 (1862) 121. — Pseudaleia imbri-
cata (RoxB.) Hassk. ex VALET. Crit. Overz. Olacin.
(1886) 115, pr. syn. — Pseudaleia longistylis Hassk.
ex VALET. /.c., pr. syn. — O. semiinfera VALET. l.c.
116; Merr. En. Born. (1921) 242. — O. laxiflora
RIDL. Kew Bull. (1931) 34. — O. multiflora Rw . l.c.
— O. rosea RiDL. /.c..33; SincLair, Gard. Bull. Sing.
14 (1953) 31.

Shrub, usually climbing; branchlets unarmed, stri-
ate, somewhat pubescent initially, practically glab-
rous, dark red-brown when dry; lenticels pale.
Leaves ovate- to elliptic-oblong, apex subacuminate,
acute or blunt, base cuneate or rounded, (sub)coria-
ceous, shining above, glabrous on both faces, 4—15
(—18) by 2—7.5 cm, nerves 6—9 pairs, rather irregu-
larly curved-ascending, sometimes less in number
and more steeply ascending, slightly raised beneath;
petiole wrinkled, 5—10 mm. Racemes branched from
the base, many-flowered, 1—3 (—5) cm; floral bracts
ovate, concave, imbricate in 2 rows when young,
caducous, 2—3 mm. Calyx very small. Petals 3, lin-
ear-oblong, white or pinkish, 10O—12 mm. Stamens 3;
staminodia 5 or 6, bifid. Drupe subglobular, rarely

OLACACEAE (Sleumer) 9

oblongoid or obovoid, almost completely covered by
the thin accrescent orange calyx, 1.7—2.5 cm. Other-
wise as in O. scandens.

Distr. India, Ceylon, Burma, Andaman & Nico-
bar Is., Thailand, S. China (Hainan); in Malesia: Su-
matra, Malay Peninsula, Java incl. Madura, Borneo,
Celebes (incl. Kabaena & Buton Is.), Philippines
(incl. Sulu Arch.), Lesser Sunda Is. (Flores, Sumba-
wa, Alor, Timor), Moluccas (Tanimbar & Kei Is.),
New Guinea; also known from Formosa (Botel To-
bago), Micronesia (Palau), and the Solomon Is.

Ecol. In primary, also often in secondary forest,
dry brushwood, on coral limestone, but also occa-
sionally in mangrove or peat swamp, at low eleva-
tions, rarely up to 900 m.

Vern. Philippines: balagon, labnot, P. Bis., bi-
ton, malabagio, malabutong, Tag., ubet-ubet, Ilk.;
Lesser Sunda Is.: /eténg, Flores.

Excluded

Olax sumatrana Mia. Fl. Ind. Bat. Suppl. (1860)
342 = Cansjera scandens Roxs. (Opiliaceae).

2. HARMANDIA

PIERRE ex BAILL. Bull. Soc. Linn. Paris 2 (1889) 770; Batt. Hist. Pl. 11 (1892)
452; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 30; Blumea 26 (1980)
153. — Fig. 2.

Monoecious tree. Leaves distichous, penninerved, with infrequent laticifers.
Racemes corymbiform, short. Ca/yx in anthesis small, patelliform, shortly
4-dented, much accrescent afterwards to form a frill which includes the fruit be-
low. Petals 4in the o& ,6—8 inthe 9 , connate to an urceolate tube for the lower
3/4, free above in form of 4 lobes. Disk extra-staminal, annular, crenulate, thin,
finally disappearing. — o Flowers: Stamens 4, epipetalous; filaments fused to
a tube which bears the free anthers on top; connective thick. Ovary rudimen-
tary. — Q Flowers: Staminodial tube without anthers. Ovary pyramidal,
l-celled, with 2 (unitegmic or naked?) ovules pendent from the short basal pla-
centa; style short-conical; stigmas 3, sessile. Fruit drupaceous, concrescent with
the much enlarged calyx below; pericarp fleshy; endocarp thin-woody. Seed 1,
containing exclusively oil; embryo excentrically at apex of endosperm.

Distr. Monotypic. Indochina; in Malesia: Sumatra, Malay Peninsula, and Borneo. Fig. 3.
Ecol. Lowland forest.

1. Harmandia mekongensis Pierre ex Bait. Bull.
Soc. Linn. Paris 2 (1889) 770; Pierre, Fl. For. Coch.
(1892) t. 264; Gaon. FI. Gén. I.-C. 1 (1911) 818, f.
95; Suppl. (1948) 739. — H. kunstleri Kina, J. As.
Soc. Beng. 64, ii (1895) 100; Ripi. Fl. Mal. Pen. |
(1922) 421; Henpers. Gard. Bull. S. S. 4 (1928) 238;

Heyne, Nutt. Pl. (1927) 674 (‘Hernandia’) (cf. Kos-
TERM. Reinwardtia 2, 1953, 360); Descu, Mal. For.
Rec. 15, 2 (1954) 414; BALAN MENON, Mal. For. 24
(1961) 292 (wood); Wuirmore, Tree Fl. Malaya 2
(1973) 301, f. 1; Steum. Blumea 26 (1980) 153. —
Fig. 2.

10 FLORA MALESIANA

[ser. I, vol. 10!

Fig. 2. Harmandia mekongensis PIERRE. Mature
fruit with accrescent calyx, x2/3 (vAN BALGooy
2559):

Glabrous monoecious tree (6—)10—22(—40) m,
fluted towards the base; bark pale fawn, greenish or
whitish, flaky and corky; slash of inner bark whitish
to pale yellow, granular; wood pale yellow. Branch-
lets slender, striate, zig-zag, older parts with linear
lenticels. Leaves distichous, oblong or elliptic, some-
times lanceolate, short-acuminate, base cuneate to
obtuse or rounded, (sub)coriaceous or parchment-
like, shagreened from minute warts especially be-
neath, dull and brittle in the dry state, 5—7 (—9) by
2.5—4 cm, nerves (5—) 6—8 (—10) pairs, rather incon-
spicuous on both faces; petiole 3—4 mm. /nflorescen-
ces racemose-corymbiform, c. 5-flowered, 1—1.5
cm; bracts scaly, minute. Flowers small, green. Ca-
/yx cupular and low initially, hardly sinuate-dented,

very much accrescent in fruit. Petals connate below,
forming a thickish urceolate corolla, 2 mm. — o
Flowers: Stamens 4; filaments connate to a fleshy
tube, 1.5 mm; anthers cordate, 0.5 mm. Rudiment of
ovary generally present. — Q Flowers: Staminodial
tube without anthers. Ovary conical, tapering to a
short style; stigmas 3, sessile. Drupe ovoid-ellipsoid,
orange below, glaucous-green above, purple-black
with a waxy bloom when dry, 2.5 (—3) by 1.3 (—2)
cm, connate for c. 1 cm at the base with the enlarged
persisting calyx which is free and collar-like spread-
ing above, green turning yellow or pink-orange at
maturity, 5—8 (—11) cm across, (1—) 2—3 (—4) cm
high. Seed 1; pericarp fleshy, 0.5 mm; endocarp lig-
neous, 0.5 mm. ~

Distr. Indochina (Laos, Annam); in Malesia: Su-
matra (Atjeh: G. Leuser Nat. Res.; Palembang: Ra-
was), Malay Peninsula (Perak, Pahang, Trengganu,
Selangor, Negri Sembilan, Malacca), and Borneo
(Sabah: Keningau Distr.). Fig. 3.

Fig. 3. Range of Harmandia PIERRE ex BAILL.

Ecol. In primary lowland forest, hilly country,
up to 300 m, apparently rare. The fruits are eaten by
animals.

Vern. Mempudu tanah, M, kayu tadji, Palem-
bang.

3. XIMENIA

LINNE, Sp. Pl. ed. 1 (1753) 1193; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b
(1935) 22; Blumea 26 (1980) 166. — Fig. 4.

Shrubs or low trees; branches usually armed with axillary spines; brachyblasts
often ending in thorns. Leaves spiral, sometimes fascicled on brachyblasts, pen-

1984] OLACACEAE (Sleumer) 11

ninerved. /nflorescences axillary or at the end of brachyblasts, arranged in rath-
er few-flowered peduncled, sometimes umbel-like cymes, or in fascicles, rarely
solitary; bracts small. Flowers usually bisexual, rarely functionally unisexual.
Calyx small, cupular-expanded, 4 (—5)-dentate, persistent, hardly or not accres-
cent in fruit. Petals 4 (rarely 5), free, linear-oblong, finally revolute about half-
way, with a brush of hairs on the inner surface. Stamens 8 (rarely 10), free,
hypogynous, alternately epipetalous and episepalous; filaments filiform; an-
thers linear-oblong to subovate, basifixed, dehiscent lengthwise; connective
thick. Disk 0. Ovary sessile, superior, (3—) 4-locular; style slenderly columnar,
as long as ovary; stigma small, capitate; ovules anatropous, bitegmic, solitary
in each cell, pendulous from a free basal placenta. Fruit superior, drupaceous,
with a rather thin pulpy pericarp, and a crustaceous to woody endocarp. Seed
1; endosperm copious, containing oily substances; embryo very small.

Distr. 8 spp. inthe (sub)tropics, rather closely allied to each other, one of them (X. americana) pantropical

and -subtropical.

Ecol. In thickets along the sea-shore, or in dry forests, mainly at low elevations.

1. Ximenia americana LINNE, Sp. PI. (1735) 1193;
Decne, Herb. Timor. Descr. (1835) 111; Bi. Mus.
Bot. Lugd.-Bat. 1 (1850) 247; Mia. Fl. Ind. Bat. 1,
1 (1856) 786; Suppl. (1860) 136; Mast. Fl. Br. India
1 (1875) 574; Becc. Nuov. Giorn. Bot. Ital. 9 (1877)
278, t. 11, f. 1—11; F.-Vitt. Nov. App. (1880) 45; Bis-
SCHOP GREVELINK, PI. Ned. Ind. (1883) 221; HEMsL.
Rep. Challenger Bot. 1, 3 (1884) 132; VALET. Crit.
Overz. Olacin. (1886) 74, t. 2, 20—22; Wars. Bot.
Jahrb. 13 (1891) 299; Scuimp. Ind. Mal. Strandpfl.
(1891) 176; Kina, J. As. Soc. Beng. 64, ii (1895) 107;
VALET. in Koord. Minah. (1898) 391; Rip. J. Str.
Br. R. As. Soc. 32 (1900) 61; K. & V. Bijdr. Booms.
Java 5 (1900) 280; K.ScuH. & Laut. Fl. Schutzgeb.
(1901) 301; Merr. Philip. J. Sc. 1 (1906) Suppl. 190;
VateT. Pl. Pap. (1907) 8; Back. FI. Bat. 1 (1907)
288; Voorl. (1908) 53; Foxw. Philip. J. Sc. 4 (1909)
Bot. 450; Back. Schoolfl. Java (1911) 222; Gaan. FI.
Gén. I.-C. 1 (1911) 814; Koorp. Exk. Fl. Java 2
(1912) 172; RecHINGER, Bot. Ergebn. Wiss. Reise Sa-
lomon Ins. (1913) 107; Merr. Int. Rumph. (1917)
209; Brown, Minor Prod. Philip. For. 2 (1921) 274,
f. 23; Ript. Fl. Mal. Pen. 1 (1922) 424; Merr. En.
Philip. 2 (1923) 117; ScHELLENB. Bot. Jahrb. 58
(1923) 158; Craip, Fl. Siam. En. | (1926) 269; Hey-
NE, Nutt. Pl. (1927) 592; Wuire, J. Arn. Arb. 10
(1929) 211; Riot. Kew Bull. (1931) 33; Boosera,
Bot. Jahrb. 66 (1933) 13; Steum.in E. & P. Nat. Pfl.
Fam. ed. 2, 16b (1935) 23, f. 11; Corner, Wayside
Trees (1940) 728; Hoitu. & Lam, Blumea 5 (1942)
178; Quis. Medic. PI. Philip. (1951) 253; Back. &
Baku. f. Fl. Java 2 (1965) 64; Steum. Blumea 26
(1980) 166. — Vidara littorea Rumpu, Herb. Am-
boin. 2 (1741) 119, t. 37. — X. loranthifolia Span.
Linnaea 15 (1841) 177; Ic. Ined. t. 44. — Zizyphus lit-

torea TEysM. ex Hassk. Abh. Naturf. Ges. Halle 9
(1866) 176, nom. nud. — Fig. 4.

var. americana: DEFILipps, Bot. Soc. Broter. Il, 43
(1969) 195.

Glabrous sprawling or low-branching shrub or
tree, up to 10 m; bark greyish brownish. Branchlets
usually spiny, covered with red cork and roundish
lenticels. Leaves often closely arranged on short lat-
eral twigs, deciduous in the dry season, variable in
shape, size and texture, narrowly to broadly lanceo-
late, ovate, elliptic, obovate or sometimes suborbicu-
lar, generally obtuse on both ends, though apex gen-
erally minutely apiculate or mucronulate, and some-
times emarginate, (sub)coriaceous, yellowish green,
turning brownish blackish and becoming brittle in
drying, (2—) 2.5—5 (—8, —10) by (1—) 2—3 (—4, —6)
cm; nerves 3—5 (—7) pairs, rather faint; petiole 3—7
(—10) mm. /nflorescences axillary or near the ends of
short lateral branchlets (brachyblasts) in form of sub-
umbellate racemes or cymes, peduncled up to 1.5 cm,
3—9-flowered, up to 2.5 cm, pedicels ebracteolate,
3—7 (—12) mm. Flowers usually bisexual, white to
greenish, fragrant. Ca/yx cupular, subacutely 4(—5)-
toothed, ciliate, 0.5—1.5 mm. Petals 4 (5), linear-ob-
long, acute to rather obtuse, finally recurved for
about half their length, white-barbate inside, (5—)
8-10 (—12) by 1.5—2 mm. Stamens 8 (10); filaments
2.5—4 mm, sigmoid near apex; anthers 2—4 mm;
connective apiculate. Ovary ovoid-conical; style fili-
form, up to § mm. Drupe plum-like, subglobose to
ellipsoid, rarely ovoid, apiculate, yellow to orange or
scarlet, (1.7—) 2.5 (—3.5) by 1.5—3 cm; pericarp pul-
py, green; endocarp bony. Seed 1, 1.5—2.5 by 1.2—2
cm.

12 FLORA MALESIANA

[ser. I, vol. 10!

~

Fig. 4. Ximenia americana L. var. americana. South coast of New Guinea (photogr. C. KALKMAN).

Distr. Pantropical and -subtropical.

Ecol. In thickets immediately back of the beach
along sea-shore (Barringtonia formation), also in dry
savannah or forest, sometimes even in light rain-
forest, scattered, on stony or sandy ground; faculta-
tive root-parasite and auto-parasite.

Dispersal. The succulent pericarp is eaten by birds.
The kernel is light enough to float, and there is, in ad-
dition, a layer of air-bearing tissue beneath the shell
which allows the fruit to be water-borne for months
(Guppy, Plant seeds and currents W. Indies, 1917,

252; DEFiLipps, Webbia 30, 1976, 180).

Taxon. Ximenia americana comprises numerous
local forms of doubtful taxonomic significance.

Uses. Wood hard, close-grained, used as a sub-
stitute for white Sandal wood, because of its yellow-
ish brownish colour. The sour pulp of the fruit is eat-
en. The kernels are purgative, a fact already stated by
RUMPHIUS.

Vern. Bédara laut, bidari, pidaroh, M; Philip-
pines: bo-o, Samar I., pangungdn, Yak., paniungdn,
Sul., sulo-sulo, Bag.

4. OCHANOSTACHYS

Mast. Fl. Br. India 1 (1875) 576; ENGL. in E. & P. Nat. Pfl. Fam. Nachtr. 3
(1908) 99; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 12; Blumea 26
(1980) 153. — Petalinia Brecc. Malesia | (1883) 257. — Fig. 5.

Tree. Leaves spiral, penninerved; hairs, if any, of a dendritic type. Spikes
simple or sometimes 1|- or 2-branched, elongate, slender, the bisexual flowers in-
terruptedly solitary or arranged in groups of 2—4. Calyx small, cup-shaped,

1984] OLACACEAE (Sleumer) 13

owerbud, c. flower, two petals removed, both

Fig. 5. Ochanostachys amentacea Mast. a. Habit, x 3/5, b. fl
%8, d. fruit, x1 1/5 (a—¢ VAN BALGooy 2524, d FRI 13320).

14 FLORA MALESIANA

[ser. I, vol. 10!

4—5-toothed, not accrescent. Petals (3—) 4 (—5), free to almost the base. Sta-
mens generally 2, rarely 1 or 3 before each petal and adherent to its base; fila-
ments linear-subulate; anthers subglobular-didymous. Staminodes 0. Disk
hypogynous, fleshy, shallow, rather inconspicuous. Ovary superior, incom-
pletely (2—) 3 (—4)-celled below, 1-celled above; ovules bitegmic, each one pen-
dent from the top of a free basal placenta into the cell; style short, cylindric; stig-
ma minute, 3-lobed. Drupe subglobose; pericarp thin; endocarp woody. Seed
1, mainly containing starch, and very little fatty substances in the form of oil-
droplets; embryo very small in the apex of the endosperm.

Distr. Monotypic, endemic in Malesia: Sumatra, Banka, Malay Peninsula, and Borneo. Fig. 6.

Ecol. In lowland rain-forest.

1. Ochanostachys amentacea Mast. Fl. Br. India 1
(1875) 577; VALET. Crit. Overz. Olacin. (1886) 104;
BoerL. Handl. 1 (1890) 207; Kina, J. As. Soc. Beng.
64, ii (1895) 100; RipL. J. Str. Br. R. As. Soc. 33
(1900) 60; Hook. Ic. Pl. 27 (1901) t. 2689; Hocnr.
Bull. Inst. Bot. Btzg 22 (1905) 44; Foxw. Philip J.
Sc. 4 (1909) Bot. 449 (wood); WINKL. Bot. Jahrb. 49
(1913) 365; Merr. En. Born. (1921) 242; Rint. FI.
Mal. Pen. 1 (1922) 422, f. 42; S. Moore, J. Bot. 62
(1924) Suppl. 21; HEyNe, Nutt. Pl. (1927) 593;
Foxw. Mal. For. Rec. 3 (1927) 119, with plate; HEN-
DERS. Gard. Bull. S. S. 4 (1928) 238; Merr. Pl. Elm.
Born. (1929) 58; RipL. Kew Bull. (1931) 35, incl. var.
rufa STAPF ex RIDL.; DescH, Mal. For. Rec. 15, 2
(1954) 415, pl. 86, f. 1 (wood); BROWNE, For. Trees
Sarawak & Brunei (1955) 281; SrauFF. Vierteljahrs-
schr. Nat. Ges. Zurich 106 (1961) 414; Wyatt-SmITH
& Kocuum. Mal. For. Rec. 17 (1965) 309; SMyTHIEs,
Common Sarawak Trees (1965) 113; BURGEss, Tim-
bers of Sabah, Sabah For. Rec. 6 (1966) 420 (wood);
WHITMORE, Tree Fl. Malaya 2 (1973) 302, f. 2;
MEIER, Field Guide Trees W. Males. (1974) 222, f.
57; SLEUM. Blumea 26 (1980) 153. — Petalinia banca-
na Becc. Malesia 1 (1883) 258. — O. bancana
(BEcc.) VALET. Crit. Overz. Olacin. (1886) 104. —
Fig. 5.

Tree, (S—) 10—30 (rarely —50) m high; bole
straight, maybe fluted at base or shortly buttressed,
15—40 (rarely —80) cm g; bark grey-brown to brown-
red, shedding in thin irregular flakes to expose lighter
coloured patches so that the whole trunk is character-
istically mottled; slash of inner bark finely fibrous,
yellow-brownish, interspaced with blackish fibres
and discrete droplets of white latex. Branchlets glab-
rous or puberulous, rarely rufous-tomentellous-
scurfy at tips. Leaves ovate to elliptic or elliptic-
oblong, sometimes slightly inequilateral, apex
short-acuminate, tip blunt, base broadly cuneate to
rounded, subcoriaceous to coriaceous, usually glab-
rous, rarely rufous-tomentellous on the nerves be-
neath (Borneo), green and shining above, yellowish

green beneath when fresh, rather dull olivaceous-
brownish when dry, usually sparsely shallowly tuber-
cled on both faces, the tubercles (or slightly im-
pressed dots) in part blackish, (S—) 6—13 by (2.5—)
3—7 cm; nerves (4—) 5 (—6, —8) pairs, curved-
ascending, the upper ones inarching before the edge,
slightly though distinctly impressed above, much
raised beneath in dry specimens, transverse veins and
reticulation of veinlets rather inconspicuous; petiole
(1—) 1.5—2 (—3) cm, not or hardly thickened distally.
Spikes erect-ascending, (2—) 3—6 (—12) cm. Flowers
arranged interruptedly, either solitary or mostly 2 or
3 together in opposite clusters, all over subglabrous
or puberulous, rarely scurfy rufous-tomentellous
(Borneo); bracts minute, ovate, acute. Flowers green
to whitish-yellowish, subsessile, or pedicelled up to 1
mm. Calyx 4—5- toothed, | mm. Petals (3—) 4 (—S),
ovate to ovate-oblong, with a few coarse hairs inside,
2.5 by 1.5 mm. Filaments white-greenish; anthers
light brown. Ovary depressed-ovoid, lengthwise
striate, glabrous; style short-cylindric. Drupe supe-

Fig. 6. Range of the genera Ochanostachys Mast.
and Scorodocarpus BECC.

1984]

rior, subglobose, green turning yellow when fully
ripe, pendulous, (1.5—) 2—2.5 (—3) cm 9, on a slen-
der peduncle 2—3 mm; pericarp thin, exuding a milky
gum, often tubercled outside, getting loose finally;
endocarp woody, hardly 1 mm. Seed 1, subglobu-
lar.

Distr. Malesia: Sumatra, Banka, Malay Penin-
sula, Borneo. Fig. 6.

Ecol. Understorey tree in primary, also second-
ary lowland rain-forest, often in mixed Dipterocarp
forest, undulating country, hillsides and ridges, up to

OLACACEAE (Sleumer) 15

950 m, on loamy or sandy, rarely periodically inun-
dated ground, scattered, or locally frequent.

Uses. The hard and durable yellowish to purple-
brown wood is used for house constructions. The
fruit is said to be edible.

Vern. Sumatra: gaé, gai, goi, Karo, nahum, pé-
taling, pétikal, pimpin bulan, pitatar, Minangk.; ké-
map, Banka; Malaya: kétikal, kuning, mahun, mén-
tatai, Kedah, pétaling, the common Malay name;
Borneo: é€mpilang, énticol, éntikan, guru, pitotar,
santikal, Iban, tanggal, Dusan, kadasan, M.

5. SCORODOCARPUS

Becc. Nuov. Giorn. Bot. Ital. 9 (1877) 274, t. 11, f. 12—17; SLeuM. in E. & P.
Nat. Pfl. Fam. ed. 2, 16b (1935) 20; Blumea 26 (1980) 160. — Fig. 7.

Trees. Leaves spiral, penninerved. Flowers bisexual, in short racemes. Calyx
small, cupular, 4—S-crenate or -dentate, not enlarged in fruit. Petals 4 or 5,
hypogynous, narrow, coherent by their edges until full anthesis, brush-like
woolly inside. Stamens 8 or 10, adnate to the lower half of each petal in pairs,
the uppermost part of the filaments remaining free as are the linear-elongate an-
thers. Ovary superior, imperfectly 3—4-celled, with | (uni- or bitegmic?) ovule
pendent from the top of the almost free placenta into each cell; style elongate-
conical; stigma minutely 3—4-lobed. Drupe medium-sized, subglobular, with a
thin fleshy pericarp and a much thicker woody endocarp. Seed 1; albumen
fleshy, containing starch and tannin.

Distr. Monotypic. Peninsular Thailand; in Ma/esia: Sumatra, Lingga Is., Malay Peninsula, and Borneo.

Fig. 6.
Ecol. Lowland forest.

1. Scorodocarpus borneensis (BAILL.) Becc. Nuov.
Giorn. Bot. Ital. 9 (1877) 274, t. 11, f. 12-17; VALET.
Crit. Overz. Olacin. (1886) 89; BorrRL. Handl. 1
(1890) 205; Kina, J. As. Soc. Beng. 64, ii (1895) 108;
Rint. J. Str. Br. R. As. Soc. 33 (1900) 60; Hocur.
Bull. Inst. Bot. Btzg 22 (1905) 42; Foxw. Philip. J.
Sc. 4(1909) Bot. 449, pl. 22, f. 11 (wood); Merr. En.
Born. (1921) 242; Ript. Fl. Mal. Pen. 1 (1922) 424;
Burk. & Henpers. Gard. Bull. S. S. 3 (1925) 358;
Foxw. Mal. For. Rec. 3 (1927) 121 & plates; HEYNE,
Nutt. Pl. (1927) 593; Descu, Mal. For. Rec. 15, 2
(1954) 418, f. 2 (wood); Browne, For. Trees Sara-
wak & Brunei (1955) 280; SmytTHies, Common Sara-
wak Trees (1965) 113; Wyatr-SmitH & KocHum.
Mal. For. Rec. 17 (1965) 347; BurGess, Sabah For.
Rec. 6 (1966) 422 (wood); Wuitmore, Tree Fl. Ma-
laya 2 (1973) 303, f. 3; Mever, Field Guide Trees W.
Males. (1974) 224, f. 58; Rao, Mal. For. 38 (1975)
184, f. 1—5 (leaf anat.); Steum. Blumea 26 (1980)
160. — Ximenia borneensis Bait. Adansonia 11
(1874) 271. — Fig. 7.

A large tree, 10—40 (rarely —60) m, 20—60 (—80, or
more) cm @, all parts reeking of garlic or onion espe-
cially after rain and from cut or bruised parts; crown
dense; bole usually straight, notched, sometimes
with small buttresses; bark grey to dark brown, fis-
sured and thinly rectangularly flaky, inwards dark
red with coarse orange flecks; wood hard, slash gen-
erally yellow to orange-brown, rarely whitish.
Branchlets smooth and glabrous at tips, older parts
dark coloured with elongate lenticels. Leaves gener-
ally elliptic-, rarely lanceolate-oblong, apex rather
abruptly acuminate for 1—2 cm, base cuneate to
rounded, entire, subcoriaceous to coriaceous, shining
green above, paler beneath when fresh, dull olive-
green when dry, glabrous, often densely minutely tu-
bercled mainly on upper surface by spicular cells or
sclereids which show remarkably in dry leaves, 7—15
(-—22, —32) by 3—5 (—7, —12) cm; nerves 4—5 (—7)
pairs, distant, curved-ascending (one marginal rather
inconspicuous), flat above, much raised beneath as
are the midrib and the transverse veins, reticulations

FLORA MALESIANA

[ser. I, vol. 10!

v.&

Fig. 7. Scorodocarpus borneensis BEcc.a. Habit, x 2, b. flowerbud, c. flower, d. ditto, petals save one re-
moved, all x34, e. fruit, x '% (a—d FRI 4784, e FRI 17777).

rather faint; petiole swollen distally, 1—1.5 (—2) cm.
Racemes rusty- to greyish-puberulous; rachis 2 (—4)
cm; flowers along rachis either solitary or 2 or 3 ina
group; pedicels 1.5—2 mm. Calyx small with wavy
edge. Petals narrow-oblong, yellowish, pink or
usually creamy white, 8—10 (—15) by 2 mm, brushed-
woolly inside, finally reflexed. Anthers yellow, 3—4
mm. Ovary yellowish green, tapering to the thickish
white style. Drupe superior globose to rather pear-
shaped, with numerous vertical stripes or faint ribs in
the dry state, glabrous, green, (3—) 4—5 (—7.5) cm

@, on peduncle | by 0.5 cm; pericarp thin, fleshy; en-
docarp woody, 2—2.5 mm thick, wrinkled by an
outer layer of numerous vertical fibre-like stony
strands, and an inner one of compact stone cells.
Seed 1, subglobular.

Distr. Peninsular Thailand and Malesia: Suma-
tra, Lingga Is., Malay Peninsula, and Borneo. Fig. 6.

Ecol. In primary, or often disturbed or second-
ary lowland forest, on flat (sometimes seasonally
flooded) country or undulating hillsides, up to 600
(rarely —900) m, on sandy or clayey, rarely blackish

1984] OLACACEAE (Sleumer) 17

soil, scattered, but locally frequent.

Uses. A medium hardwood timber; wood of
rather fine texture and fairly durable, purple-brown,
used for constructions. The seeds are reported to be
edible; they have a taste of onion.

Vern. Generally known as kulim, M; locally
called bawang hutan, maju bawang, kisindah, mar-
sindu, sagad bérau, Murut, sédau, Kutei, sinoh, Iban
téradu, ungsunah, M.

6. ERYTHROPALUM

BL. Bijdr- (1826) 921; Fl. Jav. (1828) praefatio VII (‘Erythroropalum’); Hassk.
Cat. Hort. Bog. (1844) 191 (‘Erythropalla’); SLEuM. in E. & P. Nat. Pfl. Fam.
ed. 2, 20b (1940) 401, f. 121; Blumea 26 (1980) 151. — Monaria KorTu. ex Va-
LET. Crit. Overz. Olacin. (1886) 130, in syn. — Fig. 8.

Slender scandent shrub or liana, with axillary tendrils usually present. Leaves
spiral or subdistichous, slightly peltate, 3—S-plinerved, long-petioled. Flowers
very small, bisexual, or andro-dioecious, borne in loose, slender, peduncled and
repeatedly dichotomous many-flowered cymes; bracts minute. Ca/yx cupular,
with 4 or 5 short, broad, subimbricate teeth, the basal part accrescent and cover-
ing the fruit. Peta/s 5, ovate-triangular, coherent by their bases, recurved. Sta-
mens 5, inserted at the base of the petals, each provided there with 2 lateral
scales (or staminodes?); filaments very short; anthers ovate, introrse; connec-
tive thickish. Disk cup-shaped, 5-crenate. Ovary (rudimentary in the ©”) infe-
rior, tapering to a short conical style with a minutely 3-lobed stigma, (2—)
3-celled below, 1-celled above; placenta central, free, with 2 or 3 unitegmic
ovules pendent from its apex. Fruit drupaceous, crowned by the persistent calyx
lobes and remains of the disk, ellipsoid, stipitate-contracted downwards; peri-
carp thin-fleshy; endocarp crustaceous to woody, splitting into 3—6 segments.
Seed 1; embryo minute near the apex of the large albumen which contains oily
substances.

Distr. Monotypic. Widely spread in S. India, and from the E. Himalaya to Assam, Bengal, Burma and
the Andaman Is., in Indochina, Thailand, SW. China (incl. Hainan); in Malesia: Sumatra, Malay Peninsula,
Borneo, Java, Lesser Sunda Is. (Flores), N. Celebes (Minahasa), Philippines, Talaud Is.

Ecol. In forest or forest borders at low and medium altitudes.

1. Erythropalum scandens BL. Bijdr. (1826) 922; (1925) 358; Craip, Fl. Siam. En. | (1926) 271; BArRt-

Hassk. Pl. Jav. Rar. (1848) 193; Mig. Fl. Ind. Bat.
1, 1 (1856) 704; Mast. Fl. Br. India 1 (1875) 578; Va-
Let. Crit. Overz. Olacin. (1886) 130; Boerv. Hand.
1 (1890) 208; O.K. Rev. Gen. Pl. 1 (1891) 111;
Pierre, Fl. For. Coch. (1892) t. 269 f. A; Kina, J. As
Soc. Beng. 64, ii (1895) 130; Rint. J. Str. Br. R. As.
Soc. 33 (1900) 61; Hocur. Bull. Inst. Bot. Btzg 19
(1904) 39; ibid. 22 (1905) 102; Ann. Jard. Bot. Btzg
Suppl. 3, 2 (1910) 854, incl. var. abbreviatum
Hocur.; Back. Schoolfl. Java (1911) 223; Gaon. FI.
Gén. I.-C. 1 (1911) 822, f. 96; Koorp. Exk. Fl. Java
2 (1912) 172; Merr. Philip. J. Sc. 14 (1919) 242;
Rip. Fl. Mal. Pen. | (1922) 436; Merr. En. Philip.
2 (1923) 118; Burk. & Henpers. Gard. Bull. S. S. 3

LETT, Pap. Mich. Ac. Sc. 6 (1929) 49; Merr. PI. Elm.
Born. (1929) 58; Burk. Dict. (1935) 949; HoLtu. &
Lam, Blumea 5 (1942) 78; Gaan. Fl. Gén. I.-C.
Suppl. (1948) 741; Back. & Baku. /. Fl. Java 2
(1965) 65; Hartus. Fl. Batan |. (1966) 27; STEEN. Blu-
mea 15 (1967) 153; Steum. Blumea 26 (1980) 151. —
Modeccopsis vaga Grirr. Notulae 4 (1854) 633. — E.
vagum (Grirr.) Mast. Fl. Br. India | (1875) 578; Vi-
DAL, Phan. Cuming. Philip. (1885) 85; Ceron, Cat.
Pl. Herb. Manila (1892) 45. — E. grandifolium
Evmer, Leafl. Philip. Bot. 8 (1915) 2788. — Fig. 8.

Scandent shrub or liana, glabrous, 3—10 m; stem
flexible; tendrils often lignescent-thickened distally,
simple, or rarely bifid. Branches slender, elongate,

18 FLORA MALESIANA [ser. I, vol. 10!

Fig. 8. Erythropalum scandens Bu. a. Habit, x 2, b. flowerbud, x5, c. flower, in section, x 14, d. fruits,
one dehisced, « % (a—c KERR 20888, d VAN STEENIS 12722).

1984]

sparingly rebranched, the free ends dropping; bark
yellowish brownish, dotted with pale lenticels.
Leaves variable in shape and size, triangularly ovate
to ovate- or lanceolate-oblong, apex acuminate, tip
acute, base broadly attenuate-truncate, rarely sub-
cordate, mostly a little peltate, membranaceous to
firmly chartaceous, rarely coriaceous, deep green
above, glaucescent beneath when fresh, yellowish
greenish in the dry state, fetid when bruised, (S—)
6—16 (—20, —25) by 6—12 (—15) cm; basal nerves 1 or
2 pairs, widely divergent and ascendent, 3—6 upper
Pairs spreading, prominent beneath, reticulations
usually inconspicuous; petiole slightly thickened and
wrinkled on both ends, (2—) 3—5 (—10) cm. Jnflores-
cences peduncled, lax, very slender, repeatedly di-
chotomous and many-flowered cymes, up to 15 cm
long, these sometimes reduced to subsessile rather
few-flowered cymes or fascicles; pedicels filiform,
4—5 mm; bracts triangular-ovate, hardly 1 mm. Ca-
lyx cupular, 5-toothed, 1 (—1.5) mm. Petals 5,
ovate-triangular, glabrous, 1.5—2 mm. Stamens with
a tuft of hairs on either side; filaments very short; an-
thers ovate-cordate, c. 0.3 mm; connective thick.
Disk pentagonous, rather flat, fleshy, crenulate, 1.5
mm @, elevated in the centre to form the short conical
style with 3 small stigmas. Drupe pendulous, subglo-

OLACACEAE (Sleumer) 19

bose to ellipsoid or obovoid-pyriform, stipitate-at-
tenuate towards the base for 2—3 cm, crowned by the
persistent calyx lobes and the remains of the disk,
1.5—2 (—2.5) by 1.5—2.5 cm; pericarp thin-fleshy,
yellow to red, rarely whitish; endocarp crustaceous;
finally stellately splitting from top downwards into
3—6 reflexed segments, inside red. Seed indigo blue,
evil smelling.

Distr. S. India, E. Himalaya to Assam, Bengal,
Burma, the Andaman Is., Indochina, Thailand, SW.
China (incl. Hainan); in Malesia: Sumatra, Malay
Peninsula, Borneo, Java, Lesser Sunda Is. (Flores),
NE. Celebes (Minahasa), Philippines (incl. Sulu
Arch.), Talaud Is.

Mentioned to occur in the Kei Is. (S. Moluccas) by
WarBuURG(Bot. Jahrb. 13, 1891, 299) possibly due to
an erroneous identification. Sterile material can be
(and has been) confused with Cardiopteris molucca-
na and with Menispermaceae (e.g. Tinospora).

Ecol. Scattered in the substage of lowland and
submontane primary and secondary rain-forest or
forest borders, in mixed Dipterocarp forest, rarely
up to 2135 m.

Vern. Kulim akar, M, aroy uat bankong, S; Phi-
lippines: balingayo, saynat, Tag., barak-barak,
Mbo., pulipis, Sub.

7. STROMBOSIA

BL. Bijdr. (1826) 1154; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 21;
Blumea 26 (1980) 163. — Lavallea BatLL. Adansonia 2 (1862) 361. — Fig. 9.

Shrubs or trees; young twigs distinctly zig-zag. Leaves spiral, sometimes al-
most distichous, penninerved. Flowers bisexual, small, in shortly peduncled
cymes or in sessile fascicles. Calyx a shallow cup, 5-lobed to various depth,
showing a reddish brown prominent dot on tip of each lobe, subperigynous or
perigynous, finally epigynous, accrescent, adnate to the pericarp almost to the
top of the mature fruit. Petals (4—) 5, free, often reflexed at anthesis, hairy
within. Stamens 5 (sometimes 4), epipetalous; filaments flat, adnate to the pet-
als except for their uppermost part, and bearing numerous unicellular hairs; an-
thers didymous, dorsifixed. Disk hypogynous, prominent, (3—) 5-lobed. Ovary
initially superior, finally partly inferior, i.e. partly sunken into the receptacle,
almost entirely covered by the fleshy disk, 3—5 (—6)-celled below, 1-celled
above; placenta free, central, from which 3—5 (—6) anatropous unitegmic ovula
are pendent; style short to filiform-elongate; stigma subglobular, rather ob-
scurely 3—5 (—6)-lobed. Drupe crowned by the persistent calyx and style base;
pericarp (the outer part of which is formed by the accrescent calyx) thin-fleshy;
mesocarp crustaceous or woody. Seed 1, with a small embryo in the apex of the
fleshy albumen which contains oily substances and amorphous polysaccharides.

20 FLORA MALESIANA [ser. I, vole 0!

Fig. 9. Strombosia ceylanica GARDN.a. Habit, x 2, b. flower, x5, c. fruit, x “%.—S. javanica Bu. d. Inflor-
escence, X 1, e. fruit, x 4% (a—b Cult. Hort. Bog. III-G-122a, c DE WitpEc.s. 16551, d KOSTERMANS 10593,
e KOSTERMANS 9570).

1984]

OLACACEAE (Sleumer) 21

Distr. About 12 spp., c. 9 of which in tropical Africa, the rest in India (W. Peninsula) and Ceylon to Bur-
ma and Thailand; in Malesia: Sumatra, Malay Peninsula, Java, Borneo, the Philippines, and the N. Moluccas

(Morotai).
Ecol. Lowland forest.

KEY TO THE SPECIES

1. Cymes peduncled for 0.5—1 cm. Bracts and bracteoles caducous in early anthesis. Petals (6—) 8—10 mm.
Drupe obovoid-subturbinate, apex truncate, apiculate by the remaining hard style base. Leaves smooth on

upper surface, not pellucid-punctulate..........

1. S. javanica

1. Cymes or fascicles sessile. Bracts and bracteoles persisting into anthesis. Petals 2-5 mm. Drupe pyriform
when young, subglobose at full maturity, apex obtuse-rounded; remains of calyx and style base inconspi-

cuous

2. Leaves hardly or not pellucid-punctulate, their surfaces smooth. Petals 2mm....

2. S. philippinensis

2. Leaves generally distinctly pellucid-punctulate, their surfaces markedly parchment-like (shagreened) from
tiny surface wrinkles or bumps, often glaucous-green and dull in dry specimens. Petals (2.5—) 3—4 (—S)

1. Strombosia javanica BL. Bijdr. (1826) 1155;
Hassk. Cat. Hort. Bog. (1844) 232; Pl. Jav. Rar.
(1848) 238; Bi. Mus. Bot. Lugd.-Bat. 1 (1850) 251,
f. 47; Mig. Fl. Ind. Bat. 1, 1 (1856) 787; Mast. FI.
Br. India 1 (1875) 579; VALET. Crit. Overz. Olacin.
(1886) 86, pl. 1, 16 a—n, incl. var. sumatrana VALET.;
Kina, J. As. Soc. Beng. 64, ii (1895) 590; K. & V.
Bijdr. Booms. Java 5 (1900) 282; Hocnr. Bull. Inst.
Bot. Btzg 22 (1905) 43; K. & V. Atlas Booms. Java
1 (1913) t. 124; Merr. En. Born. (1921) 242; Rint.
Fl. Mal. Pen. 1 (1922) 425; Burk. & HENDERs. Gard.
Bull. S. S. 3 (1925) 358; HEYNE, Nutt. Pl. (1927) 594;
Foxw. Mal. For. Rec. 3 (1927) 125, 2 pl.; HENDERs.
Gard. Bull. S. S. 4 (1928) 238; Descu, Mal. For. Rec.
15, 2 (1954) 422, t. 87, f. 1 (wood); Browne, For.
Trees Sarawak & Brunei (1955) 282; Back. & BAKH.
f. Fl. Java 2 (1965) 65; Wyatt-SmitH & KocHum.
Mal. For. Rec. 17 (1965) 353; WHITMORE, Tree FI.
Malaya 2 (1973) 306, f. 4; SLEuM. Blumea 26 (1980)
164; pe VoGEL, Seed Dicot. (1980) 378, f. 141 (seed-
ling). — Fig. 9d—e.

Tree 10—25 (—40) m; trunk straight, often with
knobs, up to 70 (rarely —100) cm 9; crown dense;
bark grey to yellowish, shallowly irregularly fissured
or cracked; slash outer bark pink, fibrous, turning
pale brown. Branches slender. Leaves oblong to
elliptic- or ovate-oblong, apex shortly subacutely
acuminate, base obtuse to rounded, thick-membran-
aceous to subcoriaceous, deep to pale green when
fresh, grey to yellowish olivaceous or brownish in dry
specimens, smooth and shining above, practically
without pellucid points, (10O—) 12—18 (—24) by 4—8
cm; nerves 5—7 pairs (the lowest pair close to the
base), curved-ascending, flat above, raised beneath,
veins transverse, slightly prominent beneath; petiole
slightly swollen distally, 1.5—2 (—2.5) cm. Cymes
solitary or fascicled, 3—7-flowered, on puberulous
peduncle 5—10 mm, which bears a few basal very
small caducous bracts; pedicels glabrous, 3—5 mm,

3. S. ceylanica

ebracteate and ebracteolate already at anthesis.
Calyx patelliform, tube rather inconspicuous, 4—5-
angular, teeth obscure, c. 3 mm @. Petals ovate-
lanceolate, papillose-ciliolate, glabrous outside,
densely hairy inside, greenish white, reflexed at apex,
(6—) 8—10 by 2—3 mm. Fi/aments ciliate at the free
top; anthers ovate-oblong, 0.5 mm. Ovary deeply
5-furrowed lengthwise; style thick-columnar, as long
as Ovary; stigma obscurely 5-lobed. Drupe obovoid-
oblong to almost turbinate, apex truncate and a little
impressed, crowned by the remains of calyx, disk and
style, the latter forming a hard beak, green, 2—4 by
1.5—2.2 cm; pericarp thin, fleshy; endocarp woody,
0.5 mm.

Distr. Tenasserim to S. Thailand, in Malesia: Su-
matra (incl. Nias I.), Malay Peninsula, W. Java,
Borneo (incl. Natuna Is.). Fig. 10.

Ecol. Lowland rain-forest, also secondary for-
est, mixed Dipterocarp forest, undulating country,
up to c. 600 m, scattered though locally common.

Fig. 10. Range of Strombosia javanica Bt.

22 FLORA MALESIANA

[ser. I, vol. 10!

Uses. Young leaves are edible and have the taste
of katjang, /.e. various Leguminosae. Wood moder-
ately durable, hard and heavy, light yellowish
brown, locally used for house constructions and
cabinet work.

Vern. Kikatjang, kikojop, S, bayam badak, bé-
lian landak, dédali, éntelung, madang kalawar, sa-
nam sanam, M, leke-leke, Nias.

2. Strombosia philippinensis (BAILL.) RoLFE, J. Bot.
23 (1885) 211; VipaL, Phan. Cuming. (1885) 23, 102;
Rev. Pl. Vasc. Filip. (1886) 86; VALET. Crit. Overz.
Olacin. (1886) 87; CERON, Cat. Manila (1892) 45;
Foxw. Philip J. Sc. 2 (1907) Bot. 393 (wood); ibid.
4 (1909) Bot. 449, t. 22, f. 12 (wood); MERR. En. Phi-
lip. 2 (1923) 117; SLEUM. Blumea 26 (1980) 164. —
Lavallea philippinensis Batti. Adansonia 2 (1862)
361. — S. dubia VipALt, Sin. Atl. (1883) 20, t. 30, f.
D; Merr. Govt. Lab. Publ. Philip. 17 (1904) 15. —
S. minor ELMER ex MerR. En. Philip. 2 (1923) 117,
in obs. pr. syn.; ELMER, Leafl. Philip. Bot. 10 (1939)
3770, descr. angl. — S. elmeri SALvosa, Lexic. Phi-
lip. Trees, Bull. For. Prod. Res. Inst. Coll. Laguna
1 (1963) 125.

Tree 5—28 m; trunk up to 30 cm. Branchlets slen-
der, glabrous. Leaves subdistichous, ovate-oblong to
oblong, rarely lanceolate, apex shortly subacutely
acuminate, base cuneate to the petiole, thin-charta-
ceous, smooth above, finely wrinkled above, green
to brownish and rather dull in dry specimens, glab-
rous, hardly or not pellucid-punctulate, (6—) 8—12
(—16) by (3—) 4—6 (—7.5) cm; nerves 4—5 (—6) pairs
curved-ascending, slightly prominent beneath,
reticulations obscure; petiole slender, hardly thick-
ened distally, (I—) 1.5—2 cm. Flowers on axillary
very short multibracteolate subglobular axes, 5—8
per cluster; pedicels glabrous, c. 1 mm, with 1 or 2
persistent minute scaly bracteoles. Ca/yx cupular,
deeply 5-lobed, glabrous, lobes ovate, ciliate, 0.5
mm. Petals 5, linear-oblong, greenish-white, glab-
rous outside, hairy at apex inside, 2 by 0.5 mm. Sta-
mens 5; filaments fully adnate to the petals; anthers
ovate-oblong, 0.5 mm. Ovary superior, subglobose;
style filiform, 1—2 mm. Drupe subglobular, substipi-
tate-attenuate at base for 1—2 mm, apex obtuse, the
persistent style base short, c. 1 cm 9; pericarp thin-
fleshy, finely tubercled in the dry state as in S. ceyla-
nica; endocarp thin-woody.

Distr. Malesia: Philippines (Basilan, Catandua-
nes, Leyte, Luzon, Mindanao, Mindoro, Sibuyan),
N. Moluccas (Morotai, Halmahera).

Ecol. Forests at low and medium altitudes, local-
ly common.

Uses. Timber heavy, dull yellowish to pinkish,
used for house building.

Vern. Kamauydn, Tag., kamayudn, Tag.,
S.L.Bis., lardg, Ilk., lardk, Ibn., samayonan, Bik..,

sumayudn, Tag., tamahuyan, Bik., S.L.Bis., tama-
oydn, Neg., Tag., tamaudn, Tag., tamauydn, Ibn.,
Neg., Tag., tamayudn, tamayuéon, Bik., Tag.

3. Strombosia ceylanica GARDN. Calc. J. Nat. Hist.
6 (1845) 350; Mig. Fl. Ind. Bat. 1, 1 (1856) 787;
Mast. Fl. Br. India 1 (1875) 579; Vauer. Crit. Overz.
Olacin. (1886) 87; K. & V. Bijdr. Booms. Java 5
(1900) 284; Hocnr. Bull. Inst. Bot. Btzg 22 (1905)
43, incl. var. lucida (T. & B. ex VALET.) HOCHR., var.
membranacea (Bu.) Hocur. et var. sessilis HOCHR.;
Ann. Jard. Bot. Btzg Suppl. 3, 2 (1910) 854; Back.
Schoolfl. Java (1911) 223; Koorp. Exk. Fl. Java 2
(1912) 172; K. & V. Atlas Booms. Java 1 (1913) t.
125; Back. & Baku. f. Fl. Java 2 (1965) 64; SLEUM.
Blumea 26 (1980) 165. — Stemonurus? membrana-
ceus BL. Mus. Bot. Lugd. Bat. 1 (1850) 250. — S.
Javanica auct., non Bi.: THwaires, En. Pl. Ceyl.
(1858) 42. — Sphaerocarya leprosa Dauz. in Hook.
Kew J. Bot. 3 (1851) 34; Datz. & Giss. Bombay FI.
(1861) 223. — Lavallea ceylanica (GARDN.) BAILL.
Adansonia 2 (1862) 361. — Anacolosa maingayi
Mast. FI. Br. India | (1875) 580; VaLer. Crit. Overz.
Olacin. (1886) 93. — S. lucida T. & B. Cat. Hort.
Bog. (1866) 207, nom. nud.; ex VALET. Crit. Overz.
Olacin. (1886) 86, pl. 2, f. 18; K. & V. Bijdr. Booms.
Java 5 (1900) 286. — S. membranacea (BL.) VALET.
Crit. Overz. Olacin. (1886) 87; K. & V. Bijdr.
Booms. Java 5 (1900) 284. — S. multiflora KiNG, J.
As. Soc. Beng. 64, ii (1895) 102; Ript. Fl. Mal. Pen.
1 (1922) 425; WHiTMorE, Gard. Bull. Sing. 26 (1973)
285; Tree Fl. Malaya 2 (1973) 306. — S. rotundifolia
Kina, J. As. Soc. Beng. 64, ii (1895) 103; Rint. J.
Str. Br. R. As. Soc. 33 (1900) 60; Fl. Mal. Pen. 1
(1922) 425; Burk. & HENDERS. Gard. Bull. S. S. 3
(1925) 358; Foxw. Mal. For. Rec. 3 (1927) 123, fig.;
HENDERS. Gard. Bull. S. S. 4 (1928) 239; Descu,
Mal. For. Rec. 15, 2 (1954) 423, pl. 87, f. 2 (wood);
WYATT-SMITH & KocHum. Mal. For. Rec. 17 (1965)
352. — S. latifolia Starr, Kew Bull. (1906) 71;
Merr. En. Born. (1921) 242. —.S. rapaneoides S.
Moore, J. Bot. 62 (1924) Suppl. 22. — S. maingayi
(Mast.) WHITMORE, Gard. Bull. Sing. 26 (1973) 285;
Tree Fl. Malaya 2 (1973) 306; CorNer, Gard. Bull.
Sing., Suppl. 1 (1978) 207. — Fig. 9a—c.

Shrub or generally tree, 10—20 (—36) m; crown
compact; trunk straight, closely branched, occasion-
ally up to 1.2 m @ at base, sometimes buttressed; bark
grey to brown, peeling off in scroll-shaped patches.
Branches pendulous. Branchlets smooth, glabrous.
Leaves elliptic to ovate-oblong, sometimes ovate or
suborbicular, apex shortly acuminate, tip acute or
bluntish, rarely obtuse-rounded, base cuneate to ob-
tuse or rounded, slightly inequilateral, firmly mem-
branaceous when young, usually subcoriaceous,
rarely coriaceous in later stages, glabrous, somewhat
shining above when fresh, drying usually dull, green-

1984]

ish brown or glaucous-green, conspicuously parch-
ment-like (shagreened) from close wrinkled surface
and minutely pustular, + distinctly and densely pel-
lucid-punctulate, considerably varying in size even in
the same specimen, 8—15 (—25) by 3—7 (—8.5, —11)
cm; nerves (4—) 5—8 (rarely —12) pairs, curved-
ascendent, obsolete above, not much raised beneath,
transverse veins and reticulations rather inconspicu-
ous; petiole a little swollen distally, 4-10 (—17) mm.
Flowers from small woody warts, (1—) 3—6 (—15) per
fascicle; pedicels 1—2 mm; bracts and bracteoles sev-
eral, rounded, scale-like, reddish, minute. Calyx
5-lobed, lobes ovate, obtuse, ciliate. Peta/s 5, oblong
or elongately so, greenish white, glabrous outside,
hairy except the base inside, tips finally recurved
about halfway. Anthers ovate. Ovary semi-inferior
in a conical faintly 5-lobed disk; style filiform, (1—)
2—4mm. Drupe subsessile, pyriform when young, el-
lipsoid to subglobose with shortly attenuate base
when fully developed, apex with the remains of the
calyx obtuse-rounded, style base tiny, inconspi-
cuous, | .6—2 (—2.5) cm; pericarp thin-fleshy, pink to
purple, rugose or tuberculate; endocarp thin-woody.
Seed 1, c. 1.2 cm.

Distr. Ceylon and SW. India (Western Ghats
from Kanara southwards); in Malesia: Sumatra, Ma-
lay Peninsula, Anambas Is., West & Central Java,
Borneo. Fig. 11.

Ecol. In lowland forest and brushwood, mixed
Dipterocarp and even secondary forest, often close

OLACACEAE (Sleumer) 23

Fig. 11. Range of Strombosia ceylanica GARDN.

to the sea, scattered though locally common, on well-
drained flat land or lower slopes of ridges, up to c.
800 m.

Uses. Wood yellowish brown, hard, heavy, quite
durable, used for constructions.

Vern. Pétaling ayer, M; Borneo: bélian landak,
Iban, bungil, Sampit, kKambau, Dusun Kinabatan-
gan; Sumatra: damondjan, Eastcoast, médang huat,
Benkulen.

Note. Strombosia ceylanica is conceived here in
a broad sense including S. /atifolia, a variant with
coriaceous leaves and up to 12 pairs of nerves, which
in flower and fruit characters hardly differs from S.
ceylanica s. str.

8. ANACOLOSA

BL. Mus. Bot. Lugd. Bat. 1 (1850) 250, t. 46; SLEUM. in E. & P. Nat. Pfl. Fam.
ed. 2, 16 b (1935) 20; Blumea 26 (1980) 146. — Fig. 12-13.

Trees or erect shrubs (none scandent in Mal.). Leaves spiral, sometimes sub-
distichous, penninerved. Flowers in sessile (very rarely peduncled) cymes or fas-
cicles, often from bracteolate woody warts or short axes, rarely from trunk or
stem, bisexual. Ca/yx cupular, very shortly (S—) 6 (—7)-dentate, or subtruncate,
not enlarged after anthesis, subpersistent at base of the mature fruit. Petals (5—)
6 (—7) inserted on the margin of the cupular disk, fused in the lower part, fleshy,
concave below and including the stamens there, with a bearded keel above the
cavity. Stamens (5—)6(-—7); filaments short, flat; anthers broad-ovoid, the cells
distant and immersed in the thickened connective, the latter generally with a
long-hairy top. Disk hypogynous, adnate to the ovary, much accrescent in fruit,
6-denticulate or -furrowed. Ovary with its base or for a greater part immersed
in the disk, incompletely 2 (—3)-celled below, 1-celled above, with a central bas-
al placenta bearing 2 (—3) unitegmic ovules pendent from its apex; style short,
with a thickened or conical base; stigma very shortly lobed. Drupe included by
the enlarged disk almost to the top, and tipped by the remains of the style at the

24 FLORA MALESIANA [ser. I, vol. 10!

Fig. 12. Anacolosa frutescens (BL.) BL. a. Habit, x 1, b. flowerbud, c. flower, front part of calyx and some
petals removed, both x7, d. petal inside, e. anther, front side, e’. same from the back, all x 14, f. fruits,
x Y (a—e BACKER 22521, f SAN 70962).

1984]

OLACACEAE (Sleumer) 25

base subtended by the persistent calyx; pericarp thin-fleshy; endocarp thin-crus-
taceous. Seed 1; embryo minute, at the apex of the fleshy albumen which con-

tains starch and oil.

Distr. About 15 spp. in the Old World tropics, of which | sp. in Central Africa, 2 in Madagascar, the
rest in S. India (not in Ceylon) to Assam, Burma, Andaman & Nicobar Is., Indochina, Thailand, Malesia and
the Pacific; in Malesia 3 spp., one of which also in SE. Asia.

Ecol. Usually in lowland forest, rarely in montane rain-forest.

KEY TO THE SPECIES

Peeeeeetcom the stem 5... =<. sos dees bows

i dle ee By a oe Ue ees A ee 1. A. cauliflora

1. Flowers from foliate or defoliate axils of branchlets.
2. Flowers pedicelled for at least 2 mm. Fruit smooth in the dry state, finally yellow to orange

2. A. frutescens

2. Flowers subsessile (pedicels 0.5—1, rarely up to 2 mm). Fruit smooth or usually markedly tubercled in the

pommrate. fitially cherry red... 2. 2.500. oe eee

1. Anacolosa cauliflora SLEUM. Blumea 26 (1980)
148.

Treelet 3 m. branchlets glabrous. Leaves lanceo-
late, apex gradually long-attenuate, base cuneate to
almost rounded, glabrous, dull, brown and sub-
densely tubercled on both faces when dry, 20—26 by
4—5.5 cm, midrib impressed above, much prominent
beneath, nerves 4—5 pairs, curved-ascending and
looping before the edge, obscure above, raised be-
neath, reticulations obscure; petiole transversely fis-
sured, 5—7 by c. 2mm. /nflorescences from the stem,
glomerulate, each with numerous flowers on multi-
bracteolate axes 5—10 by 2—3 mm. Flowers white,
known only in bud. Ca/yx cupular, attenuate to-
wards the base to a kind of stipe, glabrous. Petals 6,
thickish, glabrous outside, barbate above the anthers
inside. Stamens 6; anthers barbate apically.

Distr. Malesia: NW. New Guinea (once found in
the Rouffaer R. area).

Ecol. In forest, at 250 m.

2. Anacolosa frutescens (BL.) BL. Mus. Bot. Lugd.
Bat. 1 (1850) 251, f. 46; Mig. Fl. Ind. Bat. 1, 1 (1856)
787; Baw. Adansonia 3 (1862) 118; VALET. Crit.
Overz. Olacin. (1886) 92; O.K. Rev. Gen. PI. 1 (1891)
111; K. & V. Bijdr. Booms. Java 5 (1900) 291; Ic.
Bog. 2 (1904) t. 136; Back. Schoolfl. Java (1911)
224; Koorp. Exk. Fl. Java 2 (1912) 172; K. & V. At-
las Booms. Java 1 (1913) t. 123; Ript. Kew Bull.
(1931) 34; Anperson, Gard. Bull. Sing. 20 (1963)
166; Back. & Baku. /. Fl. Java 2 (1965) 65; SLeuM.
Blumea 26 (1980) 150. — Stemonurus frutescens BL.
Bijdr. (1826) 649. — A. zollingeri Bait. Adansonia
3 (1862) 118, cf. Olacinea ignota, Z. & M. Syst. Verz.
(1846) 25. — A. heptandra MAinG. ex Mast. Fl. Br.
India 1 (1875) 581; Vater. Crit. Overz. Olacin.
(1886) 93; Kina, J. As. Soc. Beng. 64, ii (1895) 110;
Rint. Fl. Mal. Pen. 1 (1922) 425; Wuirmore, Tree

a ene pein meebo i bel ec ied Fe 3. A. papuana

Fl. Malaya 2 (1973) 300. — A. arborea K. & V. Bull.
Inst. Bot. Btzg 2 (1899) 9; Bijdr. Booms. Java 5
(1900) 288; Koorp. Nat. Tijd. N. 1. 60 (1901) 388;
Back. Schoolfl. Java (1911) 224; Koorp. Exk. FI.
Java 2 (1912) 172; K. & V. Atlas Booms. Java 1
(1913) t. 122. — A. luzoniensis MERR. Philip. J. Sc.
4 (1909) Bot. 253; Foxw. /.c. 449 (wood); BRown,
Minor Prod. Philip. For. 1 (1921) 270; WEsTER, Phi-
lip. Agr. Rev. 14 (1921) t. 35a; Bull. Agr. Philip. Is.
39 (1921) 272; Merr. En. Philip. 2 (1923) 117. — A.
celebica VALET. in Koord. Minah. (1898) 391;
Koorb.-SCHuo. Syst. Verz. 2, 3 (1914) 38. — Salacia
bartlettii RipL. Kew Bull. (1938) 239, cf. Dinc Hou,
Fl. Mal. I, 6 (1964) 420. — Fig. 12—13.

Erect shrub or tree, 5—25 (—30) m; trunk occasion-
ally up to to 1.4 m 9; outer bark smooth, flaking in
large thin pieces, grey to brown, inner one reddish.
Branchlets glabrous, grey-corticate below. Leaves
variable in shape and size, elliptic or elliptic-oblong
to lanceolate, apex broadly to narrowly acuminate,
tip blunt, base cuneate, slightly inequilateral, charta-
ceous to coriaceous, shining above when fresh,
brownish and rather dull in the dry state, then usually
with numerous tiny warts or tubercles on both faces,
or mainly so beneath, with numerous fine pellucid
points visible against strong light, (6.6—) 7—15 (—22)
by (3—) 4—6.5 (—9, —12) cm; nerves (4—) 5—6 pairs,
curved-ascending, the lowest pair close to the base,
raised beneath, reticulation of transverse veins and
veinlets lax, rather inconspicuous; petiole stoutish,
5—7 (—10) mm. Flowers on short warts or tubercles
(these rarely elongate to short scaly axes), (2—) S~15
per fascicle; bracts and bracteoles 0; pedicels glab-
rous or puberulous, 3—5 (—6) mm. Calyx cup-
shaped, shortly 5—7-lobed or subentire, glabrous or
pale rusty-puberulous, c. 3 mm diameter. Petals (S—)
6 (—7), ovate-lanceolate, connate about halfway,
thickish, glabrous or rarely puberulous outside, bar-

26 FLORA MALESIANA

[ser. I, vol. 10!

Fig. 13. Anacolosa frutescens (BL.) BL. Twig with fruits, + nat. size; Sarawak (Photogr. Dinc Hou 380).

bate in the upper free cristate and finally recurved
part inside, green-white, 2—3 (rarely —4) by 1—1.5
mm. Stamens 5—7; filaments flat; anthers barbate
apically. Ovary surrounded by the slightly
12-grooved disk; style short-conical; stigma minute,
shortly lobed. Drupe obovoid-ellipsoid to oblong-
oid, seated on the persistent non-accrescent calyx,
apex truncate and slightly 6-sulcate, the persistent
style base very short, at maturity yellow to orange,
(1.5—) 1.8—2 (—2.5) by (1.2—) 1.5—2 cm, on stout
pedicel 6—8 by 1.5 mm; pericarp thin-fleshy; endo-
carp thin-crustaceous. Seed 1; albumen copious.
Distr. Burma, Andaman & Nicobar Is., E. Thai-
land; in Malesia: Sumatra (Eastcoast, Palembang),
Malay Peninsula, West & Central Java, Borneo, NE.
Celebes (Minahasa), Moluccas (Sula Is.: Taliabu),
Philippines (Luzon, Mindoro, Panay, Masbate,

Fig. 14. Range of Anacolosa frutescens (BL.) BL.

Mindanao). Fig. 14.

Ecol. In lowland and submontane forest, mixed
Dipterocarp forest, along stream in kerangas forest,
sometimes in peat swamp forest (Borneo), also in
secondary forest, occasionally on limestone, scat-
tered though locally frequent, from sea-level up to
700 (rarely —1400) m.

Uses. Wood pale reddish brown, hard and
heavy, used for house posts, but said to be not dur-
able.

Vern. W. Java: kopi gunung, tangki leuweung,
S; Borneo: bélian landak, \ban; Philippines: a/u/oi,
gdlo, Tag., mataboto, S.L.Bis., yu-pa, Gad.

3. Anacolosa papuana SCHELLENB. Bot. Jahrb. 58
(1923) 157; SLEUM. Blumea 26 (1980) 148.

Small tree, occasionally up to 15 m; trunk up to 10
cm 9g; bark smooth, light brown, with shallow longi-
tudinal fissures. Branchlets glabrous, minutely lenti-
cellate. Leaves oblong to oblong-elliptic, apex short-
ly acuminate, tip acute or bluntish, base cuneate,
firmly membranaceous to subcoriaceous, dark green
above, lighter below when fresh, becoming dull and
brown in the dry state, glabrous, more or less densely
and finely tubercled and wrinkled on both faces
though mainly beneath in dry specimens, pellu-
cid-punctulate against strong light, variable in size
even in the same specimen, (8—) 10—26 by (3—) 4—7
(—10) cm; nerves 5—6 pairs, curved-ascending, raised
beneath only, reticulations obscure; petiole 0.5—1
(—1.3) cm. Flowers 2—8 per sessile cluster, the latter
on small woody scaly warts or tubercles, or on top of
cylindrical woody axes (2—3, rarely —6 mm); pedicels
0.5—1 (rarely —2) mm. Ca/yx cupular to almost pa-
telliform, entire, c. 3 mm g, attenuate at base to-
wards to pedicel. Petals (S—) 6 (—7), lanceolate,

1984]

acute, fused in the lower part, thickish, white or
cream, bearded at base of free part, glabrous below,
2—3 (rarely —4) by c. 1 mm. Stamens 5—7; filaments
flat, glabrous; anthers bearded distally. Disk cupu-
lar, flattish. Ovary ovoid, glabrous; style very short.
Drupe subglobular to slightly obovoid, the apex ob-
tuse-rounded, rarely subtruncate, the remains of the
style very short, smooth or usually shallowly tuber-
cled in the dry state, yellowish to orange initially, fi-

OLACACEAE (Sleumer)

27

—3.5) by 1.3—2 (—2.5) cm, on stoutish peduncle up
to 5 mm; pericarp thin-fleshy; endocarp woody,
smooth or warted, 0.5—1 mm. Seed 1.

Distr. Malesia: New Guinea, Solomon Is. (Bou-
gainville to San Cristobal).

Ecol. Understorey tree in lowland to mid-
mountain rain-forest, also advanced secondary for-
est, rarely in flood plain or swamp forest, from sea-
level up to c. 1675 m, locally common.

nally deep cherry-red, (1.5—) 1.8—2.5 (sometimes

9. SCHOEPFIA

SCHREB. Gen. (1789) 129; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1894) 233; Nach-
tr. 1 (1897) 145; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 30; STEEN.
Reinwardtia | (1952) 467; SteEuM. Blumea 26 (1980) 161. — Schoepfiopsis
Miers, J. Linn. Soc. Bot. 17 (1878) 75. — Fig. 15.

Trees or shrubs, often root-parasites. Leaves spiral, penninerved. Flowers bi-
sexual, fragrant, white or yellow, often heterostyled, in short racemes or cor-
ymbs; base of rachis with small imbricate scaly perulae (Mal.). Bract and brac-
teoles united at apex of pedicel into a small, acutely 3-lobed, persistent epicalyx.
Calyx inconspicuous, /.e. connate with the cup-shaped truncate flower-axis.
Petals 4—5 (—6), inserted on the edge of the flower axis, connate in the lower
1/2—2/3 to a tubular-campanulate corolla, free and revolute above, with a tuft
of hairs inside the tube behind each anther. Stamens 4—5 (—6), epipetalous, the
slender filaments adnate to the corolla tube for almost their full length, free at
apex; anthers free, 2-celled, attached below the middle. Disk epigynous, annu-
lar, fleshy. Upper half of the ovary superior, included by the disk, lower half
within the flower axis, 3-celled below, 1-celled above; style slender; stigma
3-lobed; placenta central, with 3 ategmic ovules pending from its apex. Fruit
drupaceous, subtended at base by the persistent epicalyx, crowned by the re-
mains of calyx, disk and corolla; epicarp (which originates from the somewhat
accrescent flower-axis) thin, fleshy; endocarp crustaceous to pergamaceous,
striate lengthwise. Seed 1; embryo very small at the apex of the fleshy albumen
which contains oily substances.

Distr. About 24 spp., of which c. 20 in (sub)tropical America, and c. 4 in SE. Asia, one of which in Male-
sia (N. Sumatra).
Ecol. In lowland to montane, even mossy forest.

1. Schoepfia fragrans WALL. in Roxas. FI. Ind. ed.
Wall. & Carey 2 (1824) 188; Mast. Fl. Br. India 1
(1875) 581, excl. GrirrirH t. 629; STEEN. Reinward-
tia 1 (1952) 470, f. 1; SLeum. Blumea 26 (1980) 162.
— Schoepfiopsis fragrans (WALL. in Rox.) Miers,
J. Linn. Soc. Bot. 17 (1878) 76. — Fig. 15.
Glabrous shrub or small tree, (1—) 3—5 (—12) m;
bark thick, corky, whitish grey or pale brown, with
fine horizontal fissures. Branchlets angular. Leaves

elliptic-oblong to lanceolate, apex acuminate, base
acute, subinequilateral, chartaceous to subcoriace-
ous (rarely coriaceous at higher altitudes), dark green
above, paler beneath when fresh, often tubercled,
5~9 (—12) by (1.2—) 3—5 (—6) cm; nerves 5—7 (—9)
pairs, inarching, inconspicuous above, a little raised
beneath; petiole slender, 4—6 (—7) mm. Racemes
solitary, 3~7 (~10)-flowered, 2.5—3 (—4) cm; ped-
uncle provided at base with several persistent small

28 FLORA MALESIANA

[ser. I, vol} 10!

Fig. 15. Schoepfia fragrans WALL. a. Habit, x1, b. flower, opened, supported by the 3 bracts at base, x2
(VAN STEENIS 9719).

perular bracts; pedicels slender, (5—) 8—10 (—12)
mm, distally with a cupule formed by 1 bract and 2
bracteoles, 0.5 mm. Calyx obconical, edge truncate,
adherent to the ovary and accrescent in fruit. Corolla
tubular, fleshy, whitish or pinkish to yellowish, or
even sulphur-yellow, with a scent of jasmine, tube
0.8—1 cm long, 4—5 mm @, grooved at base where it
is agglutinated (or connate) around the ovary; lobes
oblong, subacute, 4-5 mm, with a tuft of hairs at
their inner base above the insertion of the anthers.

Stamens (4) 5; filaments cohering to corolla tube; an-
thers ovate, subbilobed, free in the throat. Disk epi-
gynous, pulvinate. Ovary half-superior, turbinate,
5—6 mm long; style as long as or shorter than the co-
rolla tube; stigma generally 3-lobed. Drupe ellipsoid-
oblongoid, whitish or cream to yellow, (0.7—) 1—1.2
cm long, 0.7—0.8 mm @, crowned by the disk; peri-
carp succulent when ripe, 1—2 mm thick; endocarp
thin-crustaceous, striate lengthwise. Seed 1, white,
conforming with the endocarp.

1984] OLACACEAE (Sleumer) 29

Distr. Nepal, Bhutan, E. Bengal, Assam, Bur- Ecol. In primary montane forest or forest bor-
ma, SW. China, Thailand, Indochina; in Malesia: N. ders, also in mossy forest, (600—) 1400—2500
Sumatra (Atjeh: Gajo Lands). (—3000) m.

Excluded

Several genera formerly accommodated in a larger family concept ‘Olacineae’ are excluded and treated
under Icacinaceae (see Fl. Males. I, 7, 1971, 1—87) or Opiliaceae (this volume, pp. 31-52). The present list
is restricted to those genera and some species which have in the past been ascribed to Olacineae but which
belong to different families.

Bracea KinG, J. As. Soc. Beng. 64, ii (1898) 101 is according to StapF & KING, Ic. Pl. 7 (1901) t. 2690 =
Sarcosperma (Sarcospermataceae).

Ctenolophon Outv. Trans. Linn. Soc. 28 (1873) 516, t. 43 belongs according to WINKLER in E. & P. Nat.
Pfl. Fam. ed. 2, 19a (1931) 122 to Linaceae.

Erythropalum triandrum Quis. & MERR. Philip. J. Sc. 37 (1928) 143 = Alsomitra macrocarpa (BL.) ROEM.
(Cucurbitaceae), fide Kew.

Fissipetalum Merr. J. Str. Br. R. As. Soc. 85 (1922) 168 is according to Airy SHAW, Kew Bull. (1947) 22
= Ericybe Roxs. (Convolvulaceae).

Pteleocarpa Outv. Trans. Linn. Soc. 28 (1873) 515 has, with some doubt, been accommodated in Boragina-
ceae.

Strombosia? philippinensis [non (BAILL.) ROLFE] LAM& Ho tHuts, Blumea 5 (1942) 178, based on LAM3175
from Talaud Is., is identified by BAKHUIZEN f. & VAN STEENIS as Celtis paniculata (ENDL.) PLANCH. (U/ma-
cede).

Worcesterianthus MERR. Philip. J. Sc. 9 (1914) Bot. 288; ibid. 10 (1915) Bot. 270 is according to VAN STEE-
nis, Acta Bot. Neerl. 4 (1955) 478 = Microdesmis Hook. f. (Euphorbiaceae or Pandaceae respectively).

‘(aagwend eet) ole Se eee Se

\ \ ' A 7 .
Rua ne beet Me ALE ee) OP Acts een
quent) Fea & ORL WM wery Goo 2° oe ge? ee eorunent! thanily

4
+ ‘

ve

phaaice v3
fet

OPILIACEAE (P. Hiepko, Berlin)

Small evergreen trees, shrubs or lianas; two genera (Cansjera and Opilia) are
known to be root-parasites. Leaves distichous, simple, usually extremely var-
iable in form and size, entire, exstipulate, pinnately veined; dried leaves mostly
finely tubercled by cystoliths located in the mesophyll. /nflorescences axillary
or cauliflorous, panicle-like, racemose, umbellate (in Africa) or spicate; bracts
narrowly ovate or scale-like, in Opilia peltate, often early caducous. Flowers
small, (3—) 4—5) (—6)-merous, mainly bisexual, sometimes unisexual and plants
then dioecious (Gjellerupia, Melientha, and Agonandra) or gynodioecious
(Champereia). Perianth with valvate, free or sometimes partly united tepals (in
Q flowers of Gjellerupia wanting). Stamens as many as and opposite to the tep-
als (in Q flowers only small staminodes); anthers introrse, 2-celled, longitu-
dinally dehiscent. Disk intrastaminal, lobed (lobes alternating with the sta-
mens), annular, or cupular. Ovary superior, 1-celled; style short or none, stigma
entire or shallowly lobed. Ovule 1, pendulous from the apex of a central placen-
ta, anatropous, unitegmic and tenuinucellar. Fruit drupaceous, pericarp rather
thin, mesocarp + fleshy-juicy, endocarp woody or crustaceous. Seed large,
conform to the drupe, without testa; hilum basal, often in a funnel-shaped cavi-
ty. Embryo terete, embedded in rich, oily endosperm, nearly as long as the seed
or shorter, with 3—4 linear cotyledons, radicle often very short.

Distribution. There are 9 genera with about 30 spp., widespread in the tropics. Rhopalopilia
is restricted to Africa and Madagascar, Agonandra to South and Central America. In Malesia:
7 genera, 5 of these only known from the eastern Old World (1 endemic: Gjellerupia in New Gui-
nea); Opilia and Urobotrya occur also in tropical Africa.

Ecology. Some species of Opiliaceae occur as undergrowth in evergreen forest, primary and
secondary, e.g. Lepionurus sylvestris is in Java an indicator of everwet climate [cf. VAN STEENIS
in Back. & Bakh. f. Fl. Java 2, 1965, (70)]. Other species tolerate or prefer a more seasonal climate
and are constituents of deciduous forest. Cansjera and Opilia often occur in beach forest.

As to the altitude the species of Opiliaceae are usually found at low and medium altitude below
1000 m, only some species ascend with several collections up to 1600 or even 2000 m (Lepionurus
in Sumatra).

Some species are mainly growing on sandy soil, others are more often found on limestone.

Habit. Most species are small trees of about 3—8 m, sometimes gregariously growing (Urobo-
trya spp.) or tiny shrubs (Lepionurus often less than 1 m high); only Champereia and Melientha
can attain sizes of more than 10 m. The lianas (Cansjera and Opilia) are climbing up to 30 m,
but they are often recorded as erect shrubs, too. The stem of a young Cansjera rheedii is growing
in an inclined position and the branches are spreading (cf. HiepKo& Weser, Willdenowia 8, 1978,
354, f. 1); if there is no tree for climbing up it becomes an erect shrub.

Pollination. The flowers of all species of this family possess nectar-secreting disks; some flow-
ers are fragrant. They are evidently entomophilous. The inflorescences of Champereia manillana
are often visited by ants.

Dispersal. The fruits of Opiliaceae are drupaceous and vary in size from less than | cm long
(Champereia) to 4cm (Melientha). For Champereia it is reported that the fruits are eaten by birds.

Galls. Galled fruits (or flowers?), in form and size like peas, have been observed in some speci-
mens of Opilia amentacea (in N. Borneo, Philippines, and New Guinea).

Parasitism. Root-parasitism has been proved to occur in Cansjera leptostachya, C. rheedii and
Opilia amentacea (see under these species). According to HiepKo& Weser (/.c.), Cansjera rheedii

(31)

32 FLORA MALESIANA [ser. I, vol. 10!

forms 4 morphologically different types of haustoria. The largest type of haustoria is formed in
the root hair zone and may grow up to | cm or more in diameter. Selfparasitism is common in
this species, and it was shown that it is also mycotrophic (cf. WEBER, Naturwissenschaften 64,
1977, 640 f.).

Morphology. The panicle-like inflorescences of Champereia and Melientha are irregularly
branched (fig. 2, 5), the flowers are pedicelled or sessile. In the racemose inflorescences of some
genera the pedicelled flowers are arranged in ternate groups (in the axil of each bract) along the
rachis, showing that these inflorescences are not genuine racemes but more complex types of in-
florescences. Occurrence of bracteoles in some species of Urobotrya stresses this opinion.

The perianth of Opiliaceae was often described as composed of a ‘minute and inconspicuous’
calyx and + free petals. Other authors use these terms only for the description of Opilia, whereas
the flowers of the other genera are called monochlamydeous (e.g. BACK. & BAKH. f. Fl. Java 2,
1965, 66). Since the flowers of Opilia show solely a slightly cupuliform torus which hardly can
be called calyx, I use the terms perianth and tepals for the entire family.

Anatomy. Wood. Most Opiliaceae are wood anatomically rather homogeneous. The light
coloured wood shows often faint to distinct growth rings. The vessels are predominantly solitary,
but up to 40—80% in radial multiples and tangential groups in Lepionurus and Urobotrya lati-
squama. The shrubby species show small vessels, the lianas have vessel diameters up to 200 um.
Fibres are often thick-walled, with minutely bordered pits in Gjellerupia and Lepionurus, and
bordered pits up to 4—6 um in the other genera. The rays are 2—6-seriate mingled with few
uniseriates; ray cells are weakly procumbent and square/upright, except in Agonandra, Opilia,
Champereia, and Melientha, where the rays are composed of clearly procumbent cells. With the
exception of Agonandra, all genera show cystoliths of calcium carbonate in enlarged ray cells.
The presence or absence of stalks and the size and shape of the cystoliths seem to have diagnostic
value. Parenchyma strands are apotracheal, diffuse or diffuse-inaggregate.

Leaves. Cystoliths occur in all representatives, commonly in pairs of clusters in enlarged meso-
phyll cells or ray cells of the vascular tissue. Size, shape and refractive properties seem to be taxo-
nomically relevant. Within the family, uniseriate and branched hairs (the last mentioned in species
of Opilia, Rhopalopilia and Cansjera) are found. They may cover leaf surfaces, or may be re-
stricted to midrib and veins, or are lacking. Stomata are paracytic with two to several subsidiary
cells. Some variation in differentiation of the mesophyll is found: either a homogeneous tissue
of cubic cells, or two layers of palisade parenchyma and spongy parenchyma, with or without a
hypodermis.

Both wood and leaf characters point to a very close relationship between Champereia and Me-
lientha. Of the other Malesian taxa, Gjellerupia, Lepionurus, and Urobotrya are very similar.
Opilia and Cansjera, as well as the African Rhopalopilia and the neotropical Agonandra, seem
to have a slightly isolated position.

Literature: Descu, Mal. For. Rec. 152 (1954) 431; EpeLHorr, Bot. Jahrb. 8 (1887) 100—153;
KOEK-NOORMAN & VAN RIJCKEVORSEL, Willdenowia 13 (1983) 147—174; METCALFE & CHALK,
Anat. Dicot. Oxford (1950) 379—381; REED, Mem. Soc. Brot. 10 (1955) 29—79; SOLEREDER, Syst.
Anat. Dicot. Stuttgart (1899) 227—237, 829—830; ibid. (1908) 81—83. — J. KoEK-NOORMAN.

Embryology. Detailed embryological investigations on members of this family are very rare.
Only in Cansjera rheedii (Swamy, 1960) and Opilia amentacea (SHAMANNA, 1955; Swamy & Rao,
1963) the male and female gametophyte have been studied. In the tetrasporangiate anther a glan-
dular tapetum with 2—4-nucleate cells is developed. The pollen grains are 2-celled when shed.

The anatropous ovule is unitegmic and has a much reduced nucellus. The nucellar tissue and
the integument collapse in later stages of ovular development (therefore the ovules often have
been described as ategmic). The chalazal megaspore of the linear tetrad develops into a Polygo-
num type embryo sac. A chalazal caecum grows down into the solid part of the gynoecium and
so the embryo sac becomes U-shaped at maturity. The endosperm is cellular. Its chalazal chamber
grows towards the base of the ovary forming a l-nucleate haustorium which in Opilia amentacea

1984] OPILIACEAE (Hiepko) 33

reaches the pedicel of the flower. In Cansjera rheedii the haustorium is branched and secondary
haustoria are developed. The embryogeny has not yet been studied.

Literature: Davis, Embryol. Angiosp. (1966) 193 f.; FAGERLIND, Svensk Bot. Tidskr. 42 (1948)
195—229; SHAMANNA, Curr. Sci. 24 (1955) 165—167; Swamy, Phytomorphology 10 (1960)
397—409; Swamy & Rao, ibid. 13 (1963) 423-428.

Palynology. The first detailed palynological survey of Opiliaceae (together with the other
families of Olacales) was given by REED(1955). The results of my own studies during the last years
have not yet been published.

The pollen grains of Opiliaceae (in Mal.) are always simple, spherical or suboblate (Gjellerupia,
Lepionurus), and mostly distinctly tricolporate. The ectoaperture is a furrow bordered by a
smooth margin (Urobotrya and Opilia). In Lepionurus the ectoapertures are very short and dis-
tinct. If the pollen is reticulate, the meshes are interrupted by the colpi (Champereia, Gjellerupia,
Melientha, and Urobotrya) or closed by a murus (Lepionurus).

The apertural membrane is granulate only on the endoaperture in Cansjera, Champereia, and
Gjellerupia, but on the whole surface of the furrow in Opilia and Urobotrya.

The ectexine consists of tectum, infratectal layer and foot-layer. The tectum varies in thickness
and may be smooth, perforate, echinulate (A gonandra) or reticulate, sometimes with crested muri
(Champereia, Lepionurus and Melientha), while in some genera the meshes are not closed (e.g.
Gjellerupia). The infratectal layer is columellate, except in Cansjera in which it is granular. The
foot-layer is often sculptured on the inner side of the apertural margin. Endexine is generally pres-
ent.

The pollen of Opiliaceae is similar to the pollen of Olacaceae (LOBREAU-CALLEN, 1980) where
we also find many suboblate grains. It also resembles pollen of /cacinaceae. The pollen grains of
Cansjera are similar to those of certain genera of Santalaceae (Scleropyrum, Pyrularia, etc.) and
to those of Octoknemaceae. These families have an endosculptate foot-layer and an ornamented
apertural membrane (except many /cacinaceae).

Literature: ERDTMAN, Pollen morphology and plant taxonomy, Angiosperms (1952) 298—299;
LoBREAU-CALLEN, Adansonia sér. 2, 20 (1980) 29—89; REED, Mem. Soc. Brot. 10 (1955) 29-79.
— D. LOBREAU-CALLEN.

Chromosomes. Chromosome numbers of 3 spp. have been recorded, two of these species
occur in Malesia: Lepionurus sylvestris (n = 10) and Opilia amentacea (= O. celtidifolia, 2n =
20, African material counted). The neotropical Agonandra racemosa has the same chromosome
number (n = 10).

Literature: KHosLA, Nucleus 21 (1978) 211—218; MANGENoT & MANGENOT, Bull. Jard. Bot.
Brux. 28 (1958) 323; SEAvEy, Taxon 24 (1975) 671.

Phytochemistry. HEGNAUER (1969) stressed the paucity of phytochemical information on
Opiliaceae. The most interesting feature then known was the presence of acetylenic fatty acids in
the lipids of roots, stem and leaves of Cansjera leptostachya Bru. This connects Opiliaceae bio-
chemically with Olacaceae and Santalaceae. In the meantime phytochemical screenings of some
medicinally used African Opiliaceous plants demonstrated the presence of saponins in two Mada-
gascan species of Rhopalopilia (DEBRAY c.s., 1971), in Rhopalopilia pallens PiERRE (BOUQUET,
1970) and in Opilia celtidifolia (GUILL. & PERR.) ENDL. (HAERDI, 1964, /.c. sub ‘Uses’; BOUQUET
& Depray, 1974; SHIHATAC.s., 1977). Alkaloid-like substances were also detected in the Madagas-
can Rhopalopilia spp., in Rhopalopilia pallens and in Opilia celtidifolia, but confirmation of the
presence of true alkaloids by isolation and characterization is still lacking. Saponins of the bark
of Opilia celtidifolia have triterpenic sapogenins, i.e. oleanolic acid and hederagenin, according
to SHIHATAC.S. (1977). This is in line with the saponins of African Olacaceae which have recently
been shown to have mainly oleanolic acid and hederagenin as sapogenins.

Literature: Bouquet, Plantes méd. Congo-Brazzaville, Thése (Pharm.), Univ. Paris (1970) 37;
Bovouet & Desray, Plantes méd. Céte d’Ivoire, Trav. Doc. O.R.S.T.O.M. no. 32, Paris (1974)
133; Despray c.s. Contr. inventaire plantes méd. Madagascar, Trav. Doc. O.R.S.T.O.M. no. 8,

34 FLORA MALESIANA [ser. I, vol. 10!

Paris (1971) 31; HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 248—249; SHrHatac.s. Plan-
ta Medica 31 (1977) 60—67. — R. HEGNAUER.

Taxonomy. Before VALETON (1886) established the Opiliaceae as a distinct family the genera
of this group have been placed by different authors in several other families. BENTHAM & HOOKER
(1862) e.g. treated the tribe Opilieae (Lepionurus, Cansjera, Opilia, and Agonandra) as a part of
their Olacineae, whereas Champereia was a member of their Santalaceae (B. & H., 1883). In the
treatment of BAILLON (1892) the Opilieae (including Opilia, Lepionurus, Champereia, Melientha,
Agonandra, and Cansjera) made part of the family Loranthaceae, which included also the Ola-
ceae, Santaleae and several other groups today mostly considered to form distinct families. ENG-
LER (1889) treated Champereia also as a genus of Santalaceae; the other genera of our Opiliaceae
were placed in two different tribes of the family Olacaceae, namely the Opilieae (Opilia, Cansjera,
and Lepionurus) and Agonandreae (Agonandra).

In 1897 ENGLER accepted the family Opiliaceae as established by VALETON (/.c.) and transfered
Champereia according to the treatment of VALETON from Santalaceae to the tribe Opilieae of this
family. In the classification of SLEUMER (1935) the same two tribes are set down: Opilieae and
Agonandreae. The second tribe is composed of the genera Gye/lerupia and Agonandra. Since Gjel-
lerupia is, with respect to morphological, anatomical, and palynological characters, obviously
more closely allied to Urobotrya and Lepionurus it has also to be included in the Opilieae.

Most present-day authors consider Opiliaceae in our circumscription as a distinct family placed
along with Olacaceae and Santalaceae in the order Santalales (or Olacales). THORNE (1981) very
recently included this family in the rank of a subfamily in his Olacaceae.

Literature: BAILLON, Hist. Pl. 11 (1892) 456—458; BENTHAM & HOOKER, Gen. PI. | (1862) 349;
ibid. 3 (1883) 231; ENGLERin E. & P. Nat. Pfl. Fam. 3, 1 (1889) 214, 240—241; ibid. Nachtr. 1
(1897) 143; SLEUMERin E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 33—41, 339; THoRNE in Young
& Seigler, Phytochemistry and angiosperm phylogeny (1981); VALETON, Crit. Overz. Olacin.
(1886) 136-161.

Uses. Young leaves and inflorescences (incl. young fruits) of Champereia manillana and Me-
lientha suavis are frequently used as a vegetable. The fruits (juicy mesocarp) of some species are
also eaten locally and occasionally: Cansjera leptostachya (Northern Australia), Champereia ma-
nillana (in many parts of the range), Melientha suavis (in Thailand), and Opilia spp. (in Northern
Australia and different parts of Africa).

Some species are used in local folk medicine (pounded or as a decoction): Champereia manilla-
na (leaves and roots applied for ulcers, rheumatism, headache, and stomachache); Lepionurus
sylvestris (roots or whole plant applied for fever or headache); Opilia amentacea (roots and/or
leaves are in Africa applied for fever, headache, or intestinal parasites; in W. Africa the plant is
said to have a purgative, diuretic, and abortive action); Urobotrya siamensis (in Thailand used
for a medicine against intestinal parasites, in large amount a deadly poison).

The wood of Melientha suavis is often used for charcoal in Thailand.

Literature: BURKILL, Dict. Econ. Prod. Mal. Pen. (1935) 526, 1353; DRUET & CoMEAU, Ann.
Univ. Abidjan, sér. C, 14 (1978) 57-67; DuNntopc.s. N. Territ. Bot. Bull. 1 (1976) 59; HAERDI,
Acta Tropica, Suppl. 8 (1964) 109; H1ePpKo, Willdenowia 9 (1979) 13—56; IRVINE, Woody plants
of Ghana (1961) 474; Wors.Ley, Acta Ethnographica 10 (1961) 153-190.

KEY TO THE GENERA

1. Inflorescence a panicle or panicle-like, in axils of leaves, often also on older branches and on the main
trunk.

2. Shrub or small tree, polygamous (gynodioecious). Flowers distinctly pedicelled. Drupe ellipsoid, 8—16

TYVIND SLOT 0. wns sice acts tar ait IED its Re serch Ss one cea ep SS PALS (cB asa 3) a oceans Ces 1. Champereia

2. Small tree. Flowers unisexual, sessile, the female flowers often with a very short pedicel, elongated in

fruiting state. Drupercllipsoid. 2-5-—4 cml lOMP cree ora cree Ge nie sie cine) costs ciers ote sient 2. Melientha

1984] O PILIACEAE (Hiepko) 535)

1. Inflorescence a raceme or spike, in axils of leaves, rarely on older branches or on the trunk.
3. Flowers in racemes, mostly 3 per bract. Bracts broadly ovate to ovate or cordate, caducous before anthe-
sis. Tepals free or united at the base only (rarely female flowers without perianth).
4. Shrub or small tree. Rachis of raceme glabrous or puberulous, pedicels glabrous. Bracts basally attached,
not peltate. Drupe usually less than 1.5 cm long.
5. Flowers bisexual. Raceme at least 2.5 cm, mostly longer. Drupe ellipsoid.
6. Tepals free, recurved. Stamens exceeding the perianth. Disk annular. Drupe 8-16 mm
3. Urobotrya
6. Tepals united at base, tube cupular, lobes spreading. Stamens not exceeding the perianth. Disk cupular
with irregularly lobed margin. Drupe 9—16 mm long, resting on the thickened disk 4. Lepionurus
5. Plant dioecious. Raceme 1—2 cm long. Female flowers without perianth, male flowers with free tepals.
emir = OLDICWIAT occ ss ss ceeisin SC Ra oie nie elas sala a3 Siareie Oo eis) otek OER er 5. Gjellerupia
4. Liana, sometimes an erect shrub. Rachis of raceme and pedicels densely covered with brownish hairs.
Braerpeitate. Drupe ellipsoid,-1'.5—3 cm long... .. /+ ... .....).tocs eterna ners e 6. Opilia
3. Flowers in spikes, each flower in the axil of a small (c. 1 mm long) persistent bract. Tepals united, tube
meee wopes sorter than thestube -..... 0... seu dee os © « «: ote eden bers eRe R eaten esate is =) 7. Cansjera

1. CHAMPEREIA

GriFF. Calc. J. Nat. Hist. 4 (1843) 237; Flora 27, 2 (1844) 436; Not. Pl. As. 4
(1854) 362; Ic. Pl. As. 4 (1854) t. 537 (‘Champereya’); B. & H. Gen. Pl. 3 (1883)
231; VALET. Crit. Overz. Olacin. (1886) 150; ENGL. in E. & P. Nat. Pfl. Fam.
3, 1 (1889) 214 (sub Santalaceae); BoERL. Handl. | (1890) 210; Baty. Hist. Pl.
11 (1892) 457 (sub Loranthaceae); ENGL. in E. & P. Nat. Pfl. Fam. Nachtr. 1
(1897) 143 (sub Opiliaceae); SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
37; HiepKo, Willdenowia 9 (1979) 14. — Malulucban Buco, Fl. Filip. (1837)
188, nom. illeg. — Opilia sect. Opiliastrum BatLL. Adansonia 3 (1862) 123. —
Govantesia LLANos, Rev. Progr. Cienc. 15 (1865) 191. — Nallogia BAIL. Bull.
Soc. Linn. Paris 2 (1892) 985; Hist. Pl. 11 (1892) 478. — Fig. 1, 2, 5b.

Shrubs or small trees; branchlets glabrous. Leaves coriaceous-fleshy. Plants
polygamous with 9 or Q flowers in panicles. Panicles axillary, often also on
older branches or on the main trunk. /nflorescences with 9 flowers widely
branched; Q inflorescences more dense with stout branches, rachises sometimes
finely puberulous. Flowers 5- (sometimes 4- or 6-)merous, with pedicels, solitary
or fascicled along the branches of the inflorescence; bracts minute, fugacious.
— 9 Flowers: tepals reflexed; filaments filiform; ovary small, conical, half im-
mersed in the fleshy, annular disk; stigma sessile. — 9 Flowers: tepals adjacent
to the ovary; stamens rudimentary; disk lobed. Drupe shortly pedicelled, ellips-
oid; pericarp thin, 0.8—1.2 mm thick, mesocarp fleshy, endocarp woody. Em-
bryo nearly as long as the seed, radicle small, with 3 long cotyledons.

Distr. One variable species: Andamans, Burma, Thailand, Vietnam, Taiwan, and Malesia. Fig. 3.
Ecol. Open evergreen forest and dry monsoon forest, from lowland up to c, 1600 m, mostly below 900 m.

1. Champereia manillana (BL.) Mere. Philip. J. Sc. 580, f. 862 (‘Cansjera leptostachya’), KANEHIRA,
7 (1912) Bot. 233; Fl. Manila (1912) 185; Philip. J. Form. Trees, ed. 2 (1936) 176, f. 128; Merr. J. Arn.
Sc. 11 (1916) Bot. 268; Sp. Blanc. (1918) 133; En. = Arb. 19 (1938) 25; Corner, Ways. Trees (1940) 514,
Philip. 2 (1923) 116; Koorp. Exk. Fl. Java 4 (1925) _ f. 173; Descu, Mal. For. Rec, 152 (1954) 431; Liv,

36 FLORA MALESIANA

[ser. I, vol. 10!

ya ne
Na ETI UNTEN Oe
, y

pata Lalueahal eid Ail

a)
ara

Fig. 1. Champereia manillana (BL.) MERR. a—b. 9 Flower, c. Q flower, d. Q flower, two tepals removed
(a—b GEESINK & HiEPKO 7823, c—d HiepKo 364). After HreEpKo, 1979.

Illust. Lign. Pl. Taiwan 2 (1962) 808; L1, Woody FI.
Taiwan (1963) 142, f. 50 (sub Santalaceae); Back. &
Baku. f. Fl. Java 2 (1965) 67; Hatus. Mem. Fac. Ag-
ric. Kagoshima Univ. 5, 3 (1966)27; HUANG, Taiwania
14 (1968) 229 (fig. of pollen); Li, Fl. Taiwan 2 (1976)
233, f.279; HieEpKo, Willdenowia 9 (1979) 16. — Cans-
Jera manillana Bu. Mus. Bot. Lugd. Bat. 1 (1851) 246.
— Opilia manillana Baw... Adansonia 3 (1862) 124. —
Opilia cumingiana Batt. /.c. — Govantesia maluluc-
ban Lianos, Rev. Progr. Cienc. 15 (1865) 191. —
Champereya gnetocarpa Kurz, J. Bot. 13 (1875) 325;
J. As. Soc. Beng. 45, ii (1876) 123. — Champereya
griffithiana PLANCH. ex Kurz, J. As. Soc. Beng. 44,
ii (1875) 154; Hook. f. Fl. Br. India 5 (1886) 236
(‘Champereia’ sub Santalaceae); GAMBLE, J. As. Soc.
Beng. 75, ii (1912) 277; Parxis. For. Fl. Andam.
(1923) 231. — C. griffithii PLANCH. ex Kurz, For. Fl.
Burma 2 (1877) 330 (nom. illeg.); Via, Sin. Atlas

(1883) t. 81, f. D; Phan. Cuming. (1885) 141; Rev. Pl.
Vasc. Filip. (1886) 232; Rm . Fl. Mal. Pen. 3 (1924)
172; BurK. Dict. (1935) 520. — Nallogia gaudichau-
diana Batt. Bull. Soc. Linn. Paris 2 (1892) 985. — C.
gaudichaudiana ( BaiLL.) TiEGH. Bull. Soc. Bot. Fr. 41
(1894) 65. — C. cumingiana (Batt.) MERR. Philip. J.
Sc. 1 (1906) Suppl. 50; Iro, Illust. Form. Pl. (1927) t.
523. — C. platyphylla MeErR. Philip. J. Sc. 11 (1916)
Bot. 177; En. Philip. 2 (1923) 116. — C. oblongifolia
Merk. Philip. J. Sc. 11 (1916) Bot. 177; En. Philip. 2
(1923) 116. — C. lanceolata MERR. Un. Cal. Publ.
Bot. 15 (1929) 57. — Fig. 1, 2, 5b.

Small tree, mostly 4—8 m, sometimes up to 20 m,
or shrub; stem 5—12 (—35) cm a; bark smooth, pale.
Slash wood white to cream. Leaves glabrous, ovate,
oblong, or lanceolate, (4.5—) 6—18 (—25) by (1.5—)
2—8 (—11) cm; apex slightly acuminate or acute; base
shortly attenuate to attenuate, rarely rounded; mid-

OPILIACEAE (Hiepko) 37

Fig. 2. Champereia manillana ( Bu.) MERR. Left: twig with inflorescences with 9° flowers (RAHMAT SI TOROES
3297); right: young infructescence developed from a Q inflorescence (H1EPKo 364). After HiEpKo, 1979.

rib above prominulous; nerves 5—7 (—8) pairs; mid-
rib and nerves prominent beneath; petiole 3—5 (—8)
mm. Panicles solitary or in groups of 2—4; main
rachis up to 20 cm long; bracts ovate, acute, 0.5—1
mm long. — 9 Flowers: pedicels 2—5 mm, thicken-
ed upwards; tepals yellowish green, 1—1.5 mm,
oblong, acute; stamens as long as the tepals, anthers
yellow, oval, 0.3 mm long; disk green, annual,
crenulate; ovary green, 0.5 mm long. — 9 Flowers
green; pedicels c. 0.5 mm long; tepals c. 0.5 mm,
acute; staminodes minute, 0.2 mm long, scaly; disk-
lobes smaller than staminodes; ovary cylindric to
ovoid, c. 0.5 mm long; stigma sessile, cushion-
shaped. Drupe orange-red, (8—) 10—12 (—15) by 7-9
mm; pedicels c. 1.5—2 (—4) mm.

Distr. Andamans and SE. Asia to Taiwan,
throughout Malesia to NW. New Guinea. Fig. 3.

Ecol. In open evergreen forest, primary and sec-
ondary, and in dry monsoon forest. Mostly at low
and medium altitudes, from sea level up to 700 m,
sometimes to 900 m (Malay Peninsula) or even 1600
m (N. Borneo). Fi. fr. Jan.—Dec., only in the north-
ern part of the area of distribution (e.g. Luzon) more
concentrated: f/. Dec.—April, /r. Jan.—May.

Inflorescences are frequently visited by ants.
Fruits eaten by birds.

Uses. Young leaves and young fruits are eaten as

vegetables; according to BurkILL (Dict. 1935, 520)
and many labels; BuRKILL (/.c.) and many collectors
recorded the fruits to be eaten in Thailand, Malaya,
the Kangean Is., Flores, N. Borneo, and the Philip-
pines (Luzon, Palawan). Leaves and roots are
pounded to make a poultice for ulcers, and the boiled
root is used for rheumatism in Malaya (BuRKILL,
l.c.). Mindanao: leaves pounded and applied for
headache and stomachache (fide FRAKE in sched.).

Vern. Malay Peninsula: be/kan (sakai), chempe-
rai, chimpri, chipreh, poko kuching-kuching, shar-
ing some of these names with Lepionurus sylvestris;
Sumatra (Simalur): futup-mateh; Flores: sasang, sui;
Philippines (Merr. En. Philip. 2, 1923, 116):
garimo, liongliong, luingluing, malakabuan, mala-
lukban, malardyap, marispdris, Tag., ichikamanok,
Tagb., panalaydpin, \\k., panalaydpon, Sbl., sulan-
manok, Sub., talaminuk, \v.; the main name in Lu-
zon: malulukban; Palawan: duro-manok, laniti;
Mindanao: gelenjup, getipun; Celebes: borongbeni-
si, kajuwatu; Talaud Is.: amaloana, aramalu; Am-
boina: sayor garing.

Notes. The species is extremely variable in vege-
tative characters, especially in form and size of
leaves. The greatest variation is found in N. Borneo
and the Philippines. Specimens from Luzon (and
Taiwan) often have relatively small leaves; the largest

38 FLORA MALESIANA

[ser. I, vol. 10!

Fig. 3. Range of Champereia manillana (Bl.) MEeRR. After HrepKo, 1979.

leaves are found in Samar and N. Borneo. But leaves
of similar extreme sizes occur in other localities, too,
and the number of main side-nerves is rather con-
stant. Therefore it is impossible to accept the species
described and named by MERRILL.

The flowers are rather uniform. The ovary of the

® flowers obviously develops rarely into a fruit. The
variation of the size of the fruits is considerable, but
the extreme forms are irregularly scattered over the
whole area of distribution, e.g. relatively large fruits
occur at the western (Andamans) and at the north-
eastern border (Samar) of this area.

2. MELIENTHA

PIERRE, Bull. Soc. Linn. Paris 1 (1888) 762; Baty. Hist. Pl. 11 (1892) 457;
ENGL. in E. & P. Nat. Pfl. Fam. Nachtr. 1 (1897) 143; SLeuM. in E. & P. Nat.
Pfl. Fam. ed. 2, 16b (1935) 36; H1EPKo, Willdenowia 9 (1979) 23. — Fig. 4, 5a.

Small trees; branchlets glabrous. Leaves glabrous, coriaceous-fleshy, in dry
state hard and brittle. Plants dioecious. F/owers in panicle-like branched inflor-
escences; rachises minutely papillate to puberulous. /nflorescences mostly on
the main trunk but also on branches and even in the axils of the uppermost
leaves. Flowers 4- or 5-merous. — © Flowers sessile, solitary or in groups of
3—5 (mainly at the end of the rachises) in the axil of a minute bract. Tepals re-
flexed. Filaments very short, attached to the base of the tepals; anthers relatively

1984]

3 mm

O PILIACEAE (Hiepko) 39

mm

Fig. 4. Melientha suavis PIERRE spp. suavis. a. Part of a 9 inflorescence, b. 9 flower, two tepals removed,
c. LS ofa o& flower-bud, d. o flower, the stamens removed, e. o' flower (a—c PuT666, d—e MAXWELL75-70).
After HrepKo, 1979.

large. Disk lobes fleshy, as large as the rudimentary ovary. — Q Flowers soli-
tary per bract, sometimes in groups of 3—4, with very short pedicels. Tepals ad-
jacent to the ovary; the small staminodes alternating with broad disk lobes.
Drupe pedicelled, ellipsoid to slightly ovoid or obovoid; pericarp thin, 1.5—2
mm thick, mesocarp fleshy-juicy, endocarp woody. Embryo nearly as long as
the seed, with small radicle (2 mm) and 3—4 long, narrow cotyledons.

Distr. Monotypic, SE. Asia (Indochina, Thailand) and West Malesia: Malaya, Borneo, and the Philip-

pines. Fig. 6.

Ecol. Primary, mostly deciduous forest, from the lowland up to 1500 m.

1. Melientha suavis Pierre, Bull. Soc. Linn. Paris 1
(1888) 763; Fl. Coch. fasc. 17 (1892) t. 264B; Gac-
NEP. Fl. Gén. I.-C. 1 (1911) 802, f. 89; ibid. Suppl.
1 (1948) 731; HiepKo, Willdenowia 9 (1979) 23. —
M. acuminata Mere. Philip. J. Sc. 29 (1926) 477. —
Fig. 4, Sa.

Small tree up to 13 m. Leaves lanceolate, elliptic to
ovate (or rarely obovate), (4—) 6—12 (—16) by 2.5—5
(—7) cm; apex obtuse- or retuse-mucronulate, some-

times acute to acuminate; base cuneate-attenuate;
nerves 5—6 (—8) pairs; hardly prominulous on both
sides; petiole 1-5 mm. /nflorescences often in
groups on swellings at the trunk or solitary on
branches and in axils of leaves; main rachis up to 15
cm, in fruiting state up to 20 cm; bracts ovate, acute,
c. 0.5 mm long. Flowers: see under the subspecies.
Drupe yellow, 2.3—4 by 1.5—2 cm; pedicels 3—7
mm.

40 FLORA MALESIANA

[ser. I, vol. 10!

{ | af as
\

Fig. 5. Melientha suavis PIERRE ssp. suavis. a. In-
fructescences. — Champereia manillana (Bu.) MERR.
b. Infructescence developed from an inflorescence
with 9 flowers (@ MAXWELL 75-452, b HarpIAL &
SAMSURI 226). After HiEPKO, 1979.

KEY TO THE SUBSPECIES

1. Drupe ellipsoid or slightly ovoid, 2.3—3 cm long
a. ssp. suavis

1. Drupe slightly obovoid, 3.5—4 cm long
b. ssp. macrocarpa

a. ssp. suavis.

Small tree up to 11 m; bark grey, smooth or fis-
sured; wood white.Leaves: see under the species; pet-
iole 1—2 mm. Main rachis of the infructescence up to
5 mm 9. — o Flowers: tepals greenish, c. 1.5 mm,
oblong, acute; anthers yellow, almost sessile, oval,
1—1.5 mm long. Disk lobes and rudiment of ovary
thick, irregularly angular, c. 0.5 mm long. — Q
Flowers green; pedicels less than 0.5 mm; tepals c. 1
mm, acute; staminodes shorter than 0.5 mm; disk
lobes as long as the staminodes, but much broader.
Ovary globose, c. 1 mm, stigma sessile. Drupe ellip-
soid or + ovoid, 2.3—3 by 1.5—1.7 cm; in herb. usu-
ally yellowish brown; pedicels 3—5 mm.

Distr. Thailand, Laos, Vietnam, Cambodia; in
Malesia: Malay Peninsula and Philippines (Minda-
nao). Fig. 6.

Ecol. In deciduous forest, locally common, rare-
ly in dry evergreen forest. From sea level (in beach
forest) up to 600 m. F/. Dec.—March; fr. April—July.
Flowers strongly fragrant.

Uses. Young shoots and inflorescences are eaten
after boiling as a vegetable (notes of many collec-
tors). Fruits edible.

Vern. Philippines: malatado, Mindanao.

Note. Melientha suavis ssp. suavis varies consid-
erably in leaf characters. The apex of the leaves is of-
ten obtuse-mucronulate but more or less acuminate
leaves are to be found at several points of the range
of the subspecies.

b. ssp. macrocarpa HieEpPKo, Willdenowia 9 (1979)
28.

Small tree up to 13 m, girth of the stem up to 45
cm; bark smooth, grey; wood white. Leaves lanceo-
late, elliptic or slightly obovate, 8—15 by 2.3—4.5 cm;
apex shortly acuminate; base cuneate-attenuate; pet-
iole 2—5 mm long. Flowers not seen (according to
KEP 80403 the inflorescences are attached to the
stem and the flowers are ‘apetalous, 4 green sepals,
4 stamens’). Main rachis of the infructescence up to
7 mm g. Drupe ellipsoid to + obovoid, 3.5—4 by 2
cm, in herb. dark brown; pedicels 7 mm long.

Distr. Malesia: N. Borneo (Mt Kinabalu: Kota
Belud). Fig. 6.

Ecol. In primary forest up to 1500 m, on black
rocky soil. Fr. July and Aug.

Vern. TJangal.

Note. The fruits of this subspecies differ consid-
erably in form and size, the structure of the pericarp
and of the seed corresponds to that of M. suavis ssp.
suavis.

Fig. 6. Range of the genus Melientha PIERRE: M. sua-
vis PIERRE ssp. suavis (dots), M. suavis PIERRE ssp.
macrocarpa HieEPKO (triangles). After H1EPKo, 1979.

1984] OPILIACEAE (Hiepko) 41

3. UROBOTRYA

Stapf, J. Linn. Soc. London 37 (1905) 89; HrieEpKo, Ber. Deut. Bot. Ges. 84
(1972) 662; Willdenowia 9 (1979) 29. — Opilia subg. Urobotrya (STAPF) ENGL.
Bot. Jahrb. 43 (1909) 171; SteuM. in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
38. — Fig. 7.

Bb cise,
imm

a

Fig. 7. Urobotrya parviflora HiePKo. a. Flower-bud, b. bract (with 3 buds), c. flower, one stamen cut off,

d. LS of pistil and disk, e. flower, tepals and stamens removed, f. pistil with disk, after flowering, g. fruit,

h. LS of seed (a, c, e Nep1738, holotype, b SAN 35987, d KOSTERMANS 21116, f KOSTERMANS 21069, g AMDJAH
370, A Woop 1273). After HiepKo, 1979,

42 FLORA MALESIANA [ser. I, vol. 10!

Section Lepionuroides

Hrerko, Willdenowia 6 (1972) 471; Nat. Hist. Bull. Siam Soc. 27 (1978) 121;
Willdenowia 9 (1979) 29.

Shrubs or small trees, twigs glabrous or puberulous. Leaves glabrous or mid-
rib hairy, thinly-coriaceous. Flowers bisexual, in racemes, usually three pedi-
celled flowers per bract; rachis of inflorescence slender, glabrous or puberulous.
Bracts broad, green, with hyaline ciliate margin, densely imbricate, caducous
before anthesis, only some basal (smaller) bracts persistent. Flowers 3- or 4 (—5)
-merous. Tepals free, oblong, acute. Stamens exceeding the perianth. Disk an-
nular, fleshy. Ovary conical to cylindric; stigma sessile. Drupe ellipsoid, meso-
carp thinly-fleshy; embryo with 3 long cotyledons.

Distr. The range of this section: 5 spp. in Thailand, S. Burma, Laos, S. China, Vietnam; in Malesia: 2
spp. in Borneo and Flores. Fig. 8.

Ecol. Evergreen (in Thailand rarely deciduous) forest, from the lowland up to c. 550 (—1000) m.

Uses. Leaves and/or fruits of U. siamensis HtepKOin Thailand locally used as a medicine against intestinal
parasites, in large amounts a deadly poison.

Taxon. The genus was originally restricted to western tropical Africa (Urobotrya sect. Urobotrya with
2 spp.). This section is characterized by much longer racemes with small, narrowly triangular bracts.

Two species from Indochina were originally described under Lepionurus. The young inflorescences are in-
deed very similar to those of that genus, but the structure of the flowers differs considerably (cf. H1EPKo, Ber.
Deut. Bot. Ges. 84, 1972, 661—663). Anatomical and palynological data support the opinion that Urobotrya
and Lepionurus are closely allied mutually and to Gjellerupia.

KEY LO) THE SPECIES

1. Racemes 4.5—5.5 cm long, rachis glabrous. Tepals mostly 3. Drupe 14-16 by 7 mm 1. U. floresensis
1. Racemes 8—12 cm long, rachis densely puberulous. Tepals mostly 4. Drupe 13 by 8.5 mm
2. U. parviflora

1. Urobotrya floresensis HiEPpKOo, Willdenowia 9
(1979) 32.

Small treelet, up to 3 m. Twigs puberulous. Leaves
glabrous, only the midrib underneath with short
hairs; ovate to elliptic, (5—) 8—12 (—16) by (2-)
3—4.5 (—6.5) cm; apex shortly acuminate, base
rounded to cuneate; nerves 7—8 on each side of the
midrib; petiole 1-2 mm. J/nflorescences axillary,
solitary; rachis (3—) 4.5—5.5 cm, glabrous. Bracts
broadly ovate, apiculate, 2.5—3 by 2.5—3 mm.
Flowers 3 per bract, without bracteoles; pedicels 1.5
mm. Tepals 3, rarely 4, c. 1 mm. Stamens white, fila-
ment c. | mm. Disk cup-shaped, up to nearly half the
length of the ovary. Ovary cylindric to conical, 0.7
mm long; stigma tripartite. Drupe 14—16 by 7 mm;
pedicel 3 mm.

Distr. Malesia: Lesser Sunda Is. (W. Flores:
Manggarai), 5 collections. Fig. 8.

Fig. 8. Range of Urobotrya STAPF sect. Lepionuro-

Ecol. From the lowland up to 800 m, according
to ScHMuTz (in sched.) gregarious (like in U. siamen-
SiS).

Vern. Sasang manuk (cf. Opilia amentacea).

Note. This species is distinguished by several

ides HierKo: U. floresensis H1EPKO (rhomb), U. /a-

tisquama (GAGNEP.) HIEPKO (triangles), U. /ongipes

(GAGNEP.) HtepKo (star), U. parviflora HiEPKO

(squares), U. siamensis HiepKo (dots). After HiEp-
Ko, 1979.

1984]

characters partly typical for some other species of the
genus: puberulous twigs, relatively small flowers,
and large fruits (like U. parviflora), glabrous and
comparatively short rachis (like U. siamensis). A uni-
que character of U. floresensis is the trimerous flow-
er, not only with three tepals and stamens, but also
showing a tripartite stigma.

2. Urobotrya parviflora HiepKo, Willdenowia 6
(1972) 474; ibid. 9 (1979) 34. — Cansjera sp.? MERR.
Un. Cal. Publ. Bot. 15 (1929) 57. — Fig. 7.
Shrub, 1—5 m, twigs puberulous. Leaves glabrous,
but midrib pilose on both sides, elliptic to broadly
ovate or lanceolate, (6—) 8—13 (—17) by (1—) 2.5—5
(—7) cm; apex shortly acuminate, base rounded or
cuneate; midrib prominent and rounded, nerves less
prominent beneath, 6—8 pairs; petiole 1—3 (—5) mm
long. /nflorescences axillary, usually solitary, rarely

OPILIACEAE (Hiepko) 43

in twos; rachis 8—12 cm long, densely puberulous.
Bracts broadly ovate, acuminate, 3—4 by 4 mn, fine-
ly hairy on both sides. Flowers 3 per bract, without
bracteoles; pedicels 1—1.5 mm. Tepals (3—) 4 (—5),
whitish, c. 1 mm. Stamens white, filaments c. 1 mm,
anthers elliptic, c. 0.5 mm long. Disk annular, low.
Ovary conical, c. 0.5 mm long. Drupe slightly apicu-
late, red, mesocarp juicy, 13 by 8.5 mm; pedicels up
to 2.5 mm.

Distr. Malesia: Borneo (Brunei, Sabah, N. &
NE. Kalimantan). Fig. 8.

Ecol. In primary and secondary evergreen forest,
from sea level up to 540 m. Fi. fr. Jan.—Dec.

Note. Form and size of the leaves are extremely
variable; besides rather broadly ovate leaves narrow-
ly lanceolate leaves are found (KOSTERMANS 21116:
12 by 1 cm). Inflorescences, flowers and fruits are
fairly uniform.

4. LEPIONURUS

BL. Bijdr. (1826) 1148; ENDL. Gen. PI. 2 (1840) 1041; B. & H. Gen. Pl. 1 (1862)
349; VALET. Crit. Overz. Olacin. (1886) 151; ENGL. in E. & P. Nat. Pfl. Fam.
3, 1 (1889) 241 (sub Olacaceae); BAIL. Hist. Pl. 11 (1892) 456 (sub Lorantha-
ceae); SLEUM.in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 35; HtepKo, Willdeno-
wia 9 (1979) 38. — Leptonium GrirF. Calc. J. Nat. Hist. 4 (1843) 236; Flora 27,
2 (1844) 435. — Opilia sect. Lepionurus (Bu.) BaAILL. Adansonia 3 (1862) 124.
— Fig. 9, 10.

Shrubs, erect or straggling, usually glabrous, sometimes young twigs with
short hairs. Leaves glabrous, thinly coriaceous. Flowers bisexual, in axillary ra-
cemes, three flowers per bract; rachis of inflorescence slender, glabrous. Bracts
broad scaly, pale green, with hyaline, shortly ciliate margin, densely imbricate,
caducous before anthesis (lowermost bracts smaller, sterile and persisting).
Flowers (3—) 4 (—5)-merous. Perianth united, deeply lobed. Stamens not ex-
ceeding the perianth, filaments flattened. Disk cupular, with irregularly lobed
margin. Ovary ovoid-conical; stigma + sessile, entire or shallowly 4-lobed.
Drupe ellipsoid to somewhat ovoid or obovoid; pericarp thin, mesocarp juicy,
endocarp crustaceous. Embryo nearly as long as the seed, radicle small, with
3—4 long, linear cotyledons.

Distr. Monotypic. SE. Asia (Nepal to Vietnam) and W. Malesia: Sumatra, Malaya, Java, Borneo. Fig. 11.
Ecol. Undergrowth in evergreen forest, from the lowland up to 1250 (—2000) m.

1. Lepionurus sylvestris BL. Bijdr. (1826) 1148; Mio.
Fl. Ind. Bat. 1, 1 (1856) 784; Kurz, For. Fl. Burma
2 (1877) 330 (sub Santalaceae); Vater. Crit. Overz.
Olacin. (1886) 153, incl. var. lanceolata VALET.;
Boert. Handi. | (1890) 210; Kina, J. As. Soc. Beng.
64, ii (1895) 593; Branovis, Indian Trees (1906) 150;
Gaonep. Fl, Gén. I.-C. 1 (1911) 806; Koorp. Exk.

Fl. Java 2 (1912) 170; ibid. 4 (1925) 581, f. 863; Ript.
Fl. Mal. Pen. 3 (1924) 172; Burk. Dict. (1935) 1353;
KANJILAL ¢c.S. Fl. Assam 1 (1936) 250; Corner,
Ways. Trees (1940) 515; Back. & Baku. /. Fl. Java
2 (1965) 67; HiepKo, Willdenowia 9 (1979) 29, — L.
Javanicus G, Don, Gen. Syst. 2 (1832) 16, nom, illeg.
— Leptonium oblongifolium Grier. Cale. J. Nat.

44 FLORA MALESIANA

[ser. I, vol. 10!

ry } F te tS
| 3 A § sain 4
| | 4 ai ye
/ pS i } (
m i ; 2mm
ae |
a 7

Fig. 9. Lepionurus sylvestris BL. a. Two flowers of a
triad, one just opened, b. bract, c. LS of flower (N1-
COLSON 3072). After HiEpKo, 1979.

Hist. 4 (1843) 237; Flora 27, 2 (1844) 435; Not. Pl.
As. 4 (1854) 368; Ic. Pl. As. 4 (1854) pl. 536. — Opi-
lia acuminata WALL. [Cat. (1829) 243, n. 7206, nom.
nud.| ex BatLt. Adansonia 3 (1862) 124. — L. ob-
longifolius (GRiFF.) Mast. Fl. Br. India 1 (1875) 583;
Koorp. Exk. Fl. Java 2 (1912) 170; Ript. Fl. Mal.
Pen. 3 (1924) 173, incl. var. angustifolius RiDL. —
Fig. 9, 10.

Shrub, usually less than 2 m, rarely up to 6 m.
Leaves extremely variable in shape, (5.5—) 10—16
(—25) by (1.5—) 3—7 (—9) cm, ratio 2—4 (—10), widest

Fig. 10. Lepionurus sylvestris BL. Young inflorescen-
ces (GEESINK, HIEPKO & PHENGKLAI 7567). Photogr.
Hrepko, Nov. 1974.

above, at, or below the middle: obovate, oblong,
lanceolate or ovate; apex acutely acuminate, base
shortly attenuate or attenuate; (S—) 8—10 (—13) pairs
of nerves, midrib and side-nerves often prominent
beneath; petiole 1—5 (—8) mm. Racemes 1—8 (—17)
per axil; rachis erect, drooping or pendulous, 2—5 cm
(in fruit up to6cm). Bracts broadly ovate, acuminate
or apiculate, 4—5 (—7.5) by 3—5 (—8) mm. Flowers 3
per bract, on a tubercle, without bracteoles; pedicels
1—2 mm. Tepals united, tube 0.5 mm long, resting on
the cupular hypanthium. Perianth yellowish, 2—4.5
mm across; segments patent, ovate, acute. Stamens
inserted below the margin of the disk, as long as the
perianth tube; anthers oval, 0.5 mm long. Pistil c. 1
mm long. Drupe resting on the thickened disk,
orange-red, 9-16 by 6—10 mm; pedicel 2—2.5 mm,
thick, seemingly longer through the enlarged tubercle
on the thickened rachis. Embryo nearly as long as the
seed, radicle about half as long as the cotyledons.

Distr. Nepal, Sikkim, Assam, Burma, S. China,
Thailand, and S. Vietnam; in Malesia: Sumatra, Ma-
lay Peninsula (Common), W. Java (common), rarely
in Central Java, Borneo (Sarawak, Sabah, Kaliman-
tan). Fig. 11.

Ecol. Usually in evergreen forest, locally com-
mon undergrowth, from sea level up to 1250 m, rare-
ly up to 2000 m (Sumatra). Fi. fr. Jan—Dec.

Razi(Lloydia 20, 1958, 238) mentioned Lepionu-
rus in his list of phanerogamic parasites, but he does
not give any evidence as proof of this statement.

Uses. In Peninsular Thailand the roots are local-
ly used for a medicine against fever. In Pahang (Ma-
laya) a poultice of the plant or of the root is applied
for headache (BuRKILL, 1935).

Fig. 11. Range of Lepionurus sylvestris Bi. After
Hiepxko, 1979.

1984] OPILIACEAE (Hiepko) 45

Notes. As already pointed out by VALETON(Crit. variable. Whereas in the greater part of the range
Overz. Olacin. 1886, 152) the leaves of L. sylvestris 1—8 racemes are found, one third of the specimens

are extraordinarily variable in shape and size. Ex- from Assam show in part more than 10 racemes per
tremely narrow leaves (ratio about 10) are especially —_axil.

striking, but such forms occur sporadically in all The size of the flowers and the differentiation of
parts of the range of the species next to plants with the rim of the disk are variable as well. Since this
a more common leaf shape (Burma, Thailand, Mala- variability is quite irregular it is impossible to distin-
ya, Sumatra). guish varieties.

The number of inflorescences per axil is also very

5. GJELLERUPIA

Laut. Nova Guinea 8 (1912) 817, t. 149; SLEUM. in E. & P. Nat. Pfl. Fam. ed.
2, 16b (1935) 40; STEEN. Nova Guinea, Bot. 12 (1963) 192; HiEpKo, Willdeno-
wia 9 (1979) 36. — Fig. 12.

Shrubs or small trees, twigs puberulous. Leaves glabrous, but midrib hairy
above, coriaceous. Plants dioecious. F/owers in racemes, | —3 pedicelled flowers
per bract; rachis of inflorescence slender, glabrous, rarely with some scattered
hairs. Bracts broadly cordate, green, with hyaling ciliate margin, densely imbri-
cate, caducous before anthesis, only some basal (smaller) bracts persistent. —
o Flowers (3—) 4 (—5)-merous. Tepals free, oblong, acute, reflexed. Stamens

v
/

2cm

Fig. 12. Gjellerupia papuana Laut. a. Twig with infructescence, b. Q flower, c. bract froma o inflorescence,
d. & flower, one tepal and one stamen removed (a DarsysHine & HOOGLAND 8232, b GseLLERUP 170, syntype,
¢ KOSTERMANS & SOEGENG 266, d KosTERMANS & SOEGENG 390). After HiepKo, 1979,

46 FLORA MALESIANA [ser. I, vol. 10!

exceeding the perianth. Disk annular, fleshy. Pistil rudimentary. — 9 Flowers
without perianth and stamens. Disk annular, thinly-fleshy. Ovary + conical;
stigma sessile. Drupe almost globular, mesocarp juicy, endocarp thinly crusta-
ceous; embryo nearly as long as the seed, with 3—4 long, linear cotyledons.

Distr. Monotypic. Malesia: New Guinea.

Ecol. Undergrowth in high evergreen forest, often on limestone ridges, from the lowland up to 200 m.

Note. Gjellerupia was reduced by Hatusma (Bot. Mag. Tokyo 65, 1952, 110) to Lepionurus sylvestris
BL., but VAN STEENIS (/.c.) pointed out that the observations and conclusions of HATUSIMA are erroneous and
that Gje/lerupia is a distinct genus. It is closely allied to Urobotrya. The male flowers of Gjellerupia show
a striking similarity with those of e.g. Urobotrya siamensis. Furthermore the pollen type (LOBREAU-CALLEN,

pers. comm.) and the placentation are the same in both genera.

1. Gjellerupia papuana Laut. Nova Guinea 8 (1912)
817, t. 149; SCHELLENB. Bot. Jahrb. 58 (1923) 157;
HiePko, Willdenowia 9 (1979) 37. — Fig. 12.
Shrub or small tree up to 6 m, with few horizontal
branches. Bark light grey, smooth. Wood hard,
straw coloured. Leaves ovate to narrowly lanceolate,
5—15 (—17) by 1.5—4 (—5.5) cm; apex acute to acumi-
nate, base attenuate to rounded; lateral nerves 8—15
pairs, midrib prominent beneath; petiole 1—4 mm.
Inflorescences axillary, usually solitary, rarely 2 or 3
together; rachis 1—2 cm long (in fruiting state up to
2.5cm). Bracts 2—3 by 2—3.5 mm. — o& Flowers 1—3

mm long. Disk undulate. Rudimentary pistil cylin-
dric, up toc. 1 mm long, spindly. — 9 Flowers 1—3
per bract. Tepals and stamens 0. Disk 0.5 mm. Ovary
conical, | mm long. Drupe red, 10—12 mm g; pedicel
5—7 mm long, often bent.

Distr. Malesia: New Guinea (Geelvink Bay, Ja-
yapura, West Sepik & Sepik Distr.).

Ecol. Locally common as undergrowth in high
evergreen forest, often on limestone ridges; from sea
level up to 200 m. Fi. fr. Jan.—Dec.

Vern. Maroa, Orne lang.

Note. The species is rather variable in form and

size of the leaves; flowers and fruits are fairly uni-
form.

per bract; pedicels 1.5—4 mm long. Tepals 1.5—2 mm
long. Stamens 1.5—2.5 mm; anthers subcordate, 0.3

6. OPILIA

Roxp. Pl: Corom: 2°(1802) 31, t. 158; R.&.S..Syst.. Veg. 5 (1819) 2732p
Gen. PI. 2 (1840) 1041; B. & H. Gen. PI. 1 (1862) 350; BariL. Adansonia 3 (1862)
123; Bru. Fl. Austr. 1 (1863) 394; Oriv. Fl. Trop. Afr. 1 (1868) 352; VALET.
Crit. Overz. Olacin. (1886) 153; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 240
(sub Olacaceae); BAILL. Hist. Pl. 11 (1892) 456; ENGL. in E. & P. Nat. Pfl. Fam.
Nachtr. 1 (1897) 143; SLEuM.in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 38 (incl.
Urobotrya); Lucas, Fl. Trop. E. Afr., Opil. (1968) 1; H1repKo, Willdenowia 12
(1982) 161. — Groutia GuILL. & PERR. Fl. Seneg. Tent. (1831) 100, t. 22. — Te-
tanosia RicH. ex M. ROEMER, Syn. Hesper. 1 (1846) 23. — Pentitdis Zipp. ex
Bui. Mus. Bot. Lugd. Bat. 1 (1851) 246, pro syn.

Lianas, sometimes erect shrubs, root parasites; young branchlets glabrous or
tomentose to puberulous. Leaves coriaceous. Flowers bisexual, in axillary ra-
cemes, three per bract. Rachis of racemes and pedicels densely covered with
brownish or yellowish hairs. Bracts peltate, broadly ovate, densely imbricate,
caducous before anthesis. Tepals free, 5—4, recurved. Stamens exceeding the
perianth. Disk lobed, with 5—4 thick and fleshy, irregularly toothed lobes alter-
nating with the stamens. Ovary cylindric to ellipsoid, stigma sessile. Drupe ellip-
soid, puberulous, mesocarp fleshy, endocarp thin, woody. Embryo nearly as
long as the seed, radicle extremely small (c. 0.5 mm), with 3 cotyledons.

1984]

OPILIACEAE (Hiepko) 47

Distr. Intropical Africa 2 spp., O. amentacea also from India through Burma, Thailand, Indochina, and

Malesia to the Solomon Is. and N. Australia. Fig. 13.

Ecol. In dry deciduous forest, often in beach forest, from the lowland up to 600 (—1200) m.

1. Opilia amentacea Roxs. Pl. Corom. 2 (1802) 31,
t. 158; Fl. Ind. ed. Carey 2 (1832) 87; Mia. Fl. Ind.
Bat. 1, 1 (1856) 784; Bru. Fl. Austr. 1 (1863) 394;
Outv. Fl. Trop. Afr. 1 (1868) 352; Mast. Fl. Br. In-
dia 1 (1875) 583; Kurz, For. Fl. Burma 1 (1877) 238;
Vial, Sin. Atlas (1883) t. 30, f. B; Rev. Pl. Vasc.
Filip. (1886) 86; VALET. Crit. Overz. Olacin. (1886)
154; Boer. Handl. | (1890) 212; Wars. Bot. Jahrb.
13 (1891) 300; Trim. Fl. Ceyl. 1 (1893) 258; K. Scu.
& Laut. Fl. Schutzgeb. (1900) 301; BRaANopis, Indian
Trees (1906) 150; Merr. Philip. J. Sc. 1 (1906)
Suppl. 50; THONNER, Blitenpfl. Afr. (1908) t. 36;
GAGNEP. Fl. Gén. I.-C. 1 (1911) 804; Koorp. Exk.
Fl. Java 2 (1912) 170; Merr. Fl. Manila (1912) 184;
GAMBLE, Fl. Pres. Madras | (1915) 192; Ewart &
Davies, Fl. N. Territ. (1917) 90; Merr. En. Philip.
2 (1923) 115; Harngs, Bot. Bihar Orissa 1 (1925) 190;
Koorp. Exk. Fl. Java 4 (1925) 579, f. 861; Back. &
Baku. f. Fl. Java 2 (1965) 66; HreEpKo, Willdenowia
12 (1982) 162, f. 1-4. — Groutia celtidifolia GUILL. &
Perr. Fl. Seneg. Tent. (1831) 100, t. 22. — Ximenia
(?) olacioides W. & A. Prod. (1834) 89. — O. celtidi-
folia (GuitLt. & PERR.) ENDL. ex WALP. Rep. Bot.

“.@

Syst. 1 (1842) 377; Keay, Fl. W. Trop. Afr. ed. 2, 1,2
(1958) 651; Garcia, Fl. Zambes. 2 (1963) 336; Lucas,
Fl. Trop. E. Afr., Opil. (1968) 2. — Tetanosia olacioi-
des (W. & A.) M. RoEMER, Syn. Hesper. | (1846) 23.
— O. pentitdis BL. Mus. Bot. Lugd. Bat. 1 (1851) 246;
Mia. Fl. Ind. Bat. 1, 1 (1856) 784; VALET. Crit. Overz.
Olacin. (1886) 155. — O. javanica Mia. Fl. Ind. Bat.
1, 1 (1856) 784. — O. tomentella (OLIv.) ENGL.
Pflanzenw. Ost-Afr. C (1895) 168; Garcia, Fl. Zam-
bes. 2 (1963) 338. — O. thorelii GAGNEP. Not. Syst. |
(1910) 206; Fl. Gén. I.-C. 1 (1911) 804, f. 90. — O. fra-
grans ELMER, Leafl. Philip. Bot. 5 (1912) 1824.
Liana up to 30 m or erect shrub; bark smooth or
fissured, pale to dark grey, branches glabrous or
glabrescent. Leaves mostly glabrous; ovate, oblong,
or lanceolate, S—14 (—16) by 2—5 cm; apex acumi-
nate, acute, or obtuse; base attenuate, sometimes
rounded; midrib prominent beneath; nerves (6—)
7—9 (11) pairs; petiole 3—7 (—10) mm. Racemes
1—S in the axil of one leaf, 1.5—3.5 cm long when
flowering; bracts 2-3 mm @, with ciliate margin.
Pedicels of 1.5—2 mm. Tepals yellowish green, ob-
long with a short inflexed top, shortly pubescent out-

+

it of

re%..

fot
cS

Fig. 13. Range of Opilia amentacea Roxs. in the eastern Old World.

48 FLORA MALESIANA

side, c. 1.5 mm long. Filaments filiform, 1.5 mm; an-
thers oval, 0.3 mm long. Disk lobes subclavate,
green, c. 0.5 mm long. Ovary c. 1 mm long. Drupe
orange-yellow, 1.5—3 by 1.25—1.75 cm; pedicels
thickened upwards, 5—7 mm.

Distr. Tropical Africa, and from India and Sri
Lanka through Burma, Thailand, Indochina, and
Malesia to the Solomon Is. and Australia. Fig. 13.

Ecol. Indry deciduous forests or thickets (in New
Guinea in light rain-forest), often on seashore or
along streams; on limestone, sandstone, or volcanic
tuff; from sea Jevel up to 600 m (in New Guinea up
to 1000 m). Fl. fr. Jan.—Dec. Flowers sweet scented.

The root-parasitism was studied by BARBER (Proc.
Cambridge Phil. Soc. 14, 1907, 246—256).

Pea-shaped galled flowers have been observed in

[ser. I, vol. 10!

some specimens from the Philippines, N. Borneo,
and from New Guinea.

Uses. The fleshy mesocarp of the fruit is edible,
but it is only reported from different parts of Africa
and N. Australia that the fruits are eaten.

Vern. Philippines: agaroiroi, P. Bis., aratig,
campenaya, toolongan, Yagb.; Komodo: Janda;
Flores: sasang manuk (cf. Urobotrya floresensis).

Note. Like the other members of Opiliaceae this
species is extremely variable in vegetative characters.
Form, size, and texture of the leaves vary consider-
ably as well as the measurements of the fruits, but the
geographical distribution of these differences is irre-
gular. The differences in the indumentum of the
young twigs are also irregularly distributed. Inflores-
cences and flowers are uniform.

7. CANSJERA

Juss. Gen. (1789) 448, nom. cons.; ENDL. Gen. PI. 1 (1837) 331; MEISN. in DC.
Prod. 14 (1857) 519; B. & H. Gen. Pl. 1 (1862) 349; Bait_. Adansonia 3 (1862)

Fig. 14. Cansjera rheedii J.F. GMELIN. a. Flower with bract, without indumentum, b. disk scale, adaxial view,

c. pistil and disk, one scale removed. — C. parvifolia Kurz. d. Pistil with disk scales, e. flower with bract,

without indumentum. — C. /eptostachya Bru. f. Flower with bract, g. pistil with disk scales (a—c GEESINK
& Hiepko 7831, d—e HELFER s.n., f—g NGF 30718). After HrepKo, 1979.

1984] OPILIACEAE (Hiepko) 49

124; Bru. Fl. Austr. 1 (1863) 191; VALET. Crit. Overz. Olacin. (1886) 156; ENGL.
in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 241; S-eum.in E. & P. Nat. Pfl. Fam. ed.
2, 16b (1935) 36; HtEPKo, Willdenowia 9 (1979) 43. — Tsjeru-caniram RHEEDE
[Hort. Mal. 7 (1688) 3, t. 2] ex ADANs. Fam. PI. 2 (1763) 80; PFEIFFER, Nom.
2 (1874) 1501 (‘Tsjerucanirum’); O.K. Rev. Gen. Pl. 1 (1891) 112 (‘Tsje-
rucaniram’); BAILL. Hist. Pl. 11 (1892) 458 (sub Loranthaceae). — Fig. 14, 16,
7.

Lianas or erect shrubs, root parasites; branches often conspicuously zigzag;
twigs densely covered with mostly upcurved hairs. Leaves herbaceous to thinly-
fleshy or coriaceous, glabrous or hairy. Flowers bisexual, in axillary spikes, each
flower in the axil of a small persisting bract. Rachis of spike and bracts densely
hairy, perianth pilose. Tepa/s united; the urceolate or campanulate perianth
with 4, small, recurved lobes, exceptionally 5-lobed. Stamens not exceeding the
perianth tube; filaments filiform. Disk scales alternating with the stamens.
Ovary ovoid to cylindric; style short, not or hardly exceeding the perianth tube;
stigma capitate, + 4-lobed. Drupe + ellipsoid, sessile on the lacerated perianth;
1 or 2 drupes per infructescence; mesocarp fleshy-juicy, endocarp thin, brittle.
Embryo much shorter than the seed, with 3—4 cotyledons.

Distr. 3 spp., from India and Sri Lanka to S. China, Malesia, and N. Australia. Fig. 15.
Ecol. In evergreen and deciduous forest, from the lowland up to 1000 m, in S. China up to 1400 m.

e
3

ans 3

a ‘ =_ a nd .
ee | <a
ss

-

Fig. 15. Range of the genus Cansjera Juss.: C. leptostachya Bru. (squares), C. parvifolia Kurz (stars), C.
rheedii J.F. Gmewin (dots). After HierKo, 1979.

50 FLORA MALESIANA

[ser. I, vol. 10!

KEY TO THE SPECIES

1. Leaves ovate to broadly lanceolate (or elliptic), + acuminate. Perianth tube (2—) 2.5—3.5 mm long

1. C. rheedii

1. Leaves lanceolate, long-narrowed. Perianth tube c. 1.5 mm long................. 2. C. leptostachya

1. Cansjera rheedii J.F. GMELIN, Syst. Nat. 2 (1791)
280 (‘Cansiera’); Meisn. Denkschr. Kon.-Bayer.
Bot. Ges. Regensburg 3 (1841) 290; WiaGurt, Ic. 5
(1852) t. 1861; Bru. Fl. Hongk. (1861) 296; BRANDIS,
For. Fl. India (1874) 75; Mast. Fl. Br. India 1 (1875)
582; Kurz, J. As. Soc. Beng. 45, ii (1876) 123; For.
Fl. Burma | (1877) 237; VALET. Crit. Overz. Olacin.
(1886) 158; Trim. Fl. Ceyl. 1 (1893) 259; Kina, J. As.
Soc. Beng. 64, ii (1895) 592; BRANpis, Indian Trees
(1906) 149, f. 69; GaGnep. Fl. Gén. I.-C. 1 (1911)
809; MerR. Philip. J. Sc. 7 (1912) Bot. 265; GAMBLE,
Fl. Pres. Madras (1915) 193; MerR. En. Born. (1921)
242; En. Philip. 2 (1923) 115; Parkins. For. Fl. An-
dam. (1923) 125; Hartnes, Bot. Bihar Orissa 1 (2)
(1925) 191; Merr. Lingn. Sc. J. 5 (1927) 70; CuuN
& CHANG, Fl. Hainan. 2 (1965) 458, f. 522; PATEL,
For. Fl. Melghat (1968) 70; HiEPKo, Willdenowia 9
(1979) 45. — C. malabarica LAmK. (incl. var. B) En-
cycl. 3 (1792) 433, nom. illeg.; Tabl. Encycl. 2 (1792)
429, pl. 289. — C. scandens Roxs. Pl. Corom. 2
(1799) 2, t. 103 (‘Cansiera’); Fl. Ind. ed. Carey 1
(1832) 441. — Daphne polystachya WILLD. Sp. Pl. 2
(1799) 420, nom. illeg. — Daphne monostachya
WIiLLp. /.c. — C. lanceolata BtH. London J. Bot. 1
(1842) 491; M. Roemer, Syn. Hesper. | (1846) 16. —
C. zizyphifolia GrirF. Calc. J. Nat. Hist. 4 (1843)
236 (‘Cansiera zyziphifolia’); Flora 27 (1844) 435;
Not. Pl. As. 4 (1854) 360; Ic. Pl. As. 4 (1854) pl. 537;
Kurz, For. Fl. Burma 1 (1877) 237; Rip. Fl. Mal.
Pen. 3 (1924) 172; Srnciatr, Gard. Bull. Sing. 14
(1953) 35. — C. polystachya (WILLD.) M. ROEMER,
Syn. Hesper. 1 (1846) 144. — C. monostachya
(WitLp.) M. Roemer, /.c. 16. — Olax sumatrana
Mia. FI. Ind. Bat. Suppl. (1861) 342. — Fig. 14a—c,
16.

Liana, climbing up to 8 (—11) m, with hanging
branches, or erect shrub with spiny stem. Leaves co-
riaceous and brittle in dry state, glabrous, ovate to
lanceolate or elliptic, (3—) S—9 (—13) by 1.5—4 (5)
cm; apex + acuminate; base shortly attenuate to at-
tenuate, rarely rounded; midrib and nerves some-
times prominent beneath, nerves 5—7 pairs; petiole
3-5 mm, densely hairy. Spikes 1—3 (—S) in the axil
of one leaf, 1.3—2.5 (—4) cm long when flowering;
bracts ovate to triangular, acute, 1 mm long. Per-
ianth urceolate, greenish yellow, tube (2—) 2.5—3
mm long, lobes recurved, 0.5 mm. Filaments c. 2
mm; anthers broadly oval, reaching as far as the
throat of the perianth tube. Disk scales slightly
fleshy, ovate, acute, irregularly toothed, c. 0.75 mm
long. Ovary + cylindric, c. 1mm long; stylec. 1 mm,

Fig. 16. Cansjera rheedii J.F. GMELIN. Inflorescence
and fruit (GEESINK & HiepKo 7831). Photogr. HIEp-
KO, Dec. 1974.

long persistent; stigma 4-lobed. Drupe orange,
10—13 (—15) by 7—9 (—12) mm. Seed with deeply
sunken basal hilum; embryo about 1/3 as long as the
seed.

Distr. From Nepal, India, and Sri Lanka to S.
China and western Malesia: Sumatra (East Coast,
Palembang), Malaya (mostly southern half), Borneo
(NW. Kalimantan, Sarawak, Sabah), Philippines
(Mindoro, Cebu, Sulu Is.). Fig. 15.

Ecol. In deciduous and evergreen forest, often in
beach forest, from sea level up to 1000 m, in S. China
up to 1400 m. Often on sandy soil. F/. Jan.—Dec.

Root parasite; roots and haustoria have been stud-
ied recently (cf. WeBER, Naturwissenschaften 64,
1977, 640, fig.; H1epkKo & WEBER, Willdenowia 8,
1978, 351-362; WeBeErR, Beitr. Biol. Pfl. 53, 1978,
371—410; WeEBER & HILDENBRAND, Ber. Deut. Bot.
Ges. 91, 1978, 231—242).

Vern. Malay Peninsula: buah champerei; Sanda-
kan: tomou.

1984]

Notes. Stem of young shrubs growing in an in-
clined position, branches spreading. The spines be-
come teat-like through secondary growth (cf.
HierkKo & WEBER, Willdenowia 8, 1978, 356, f. 3).

Form, size, and venation of the leaves are variable.
Often a pair of arcuate side-nerves shortly above the
base are nearly as strong as the midrib; such 3-nerved
forms have been named C. zizyphifolia. The flowers
are fairly uniform, but the fruits vary in size and
form (sometimes more globular).

2. Cansjera leptostachya Bru. London J. Bot. 2
(1843) 231; M. Roemer, Syn. Hesper. | (1846) 16;
Melsn. in DC. Prod. 14 (1857) 519; Bru. Fl. Austr.
1 (1863) 394; HemsL. Bot. Chall. 1, 3 (1885) 235; Va-
LET. Crit. Overz. Olacin. (1886) 159; Wars. Bot.
Jahrb. 13 (1891) 299; K. Scu. & Laut. Fl. Schutzgeb.
(1900) 301; Vater. Bull. Dép. Agric. Ind. Néerl. 10
(1907) 8; SCHELLENB. Bot. Jahrb. 58 (1923) 156;
Back. & BAKH./f. Fl. Java 2 (1965) 67; HiePKo, Will-
denowia 9 (1979) 49. — C. timorensis DECNE, Voy.
Venus, Bot. (1864) 12 (‘Candjera’); Atl. (1846) pl. 8;
Forses, Wand. (1885) 502. — Fig. 14f—g, 17.

Liana, up to 6 m, branches hanging, or erect
shrub; young twigs puberulous, often soon becoming
glabrous. Leaves herbaceous in dry state, glabrous,
ovate-lanceolate, long-narrowed, 4—9 (—11) by (1—)
1.5—4 cm; apex acute or + acuminate; base at-
tenuate to shortly attenuate; midrib and main lateral
nerves rarely somewhat prominent beneath, (6—)
7—9 pairs of mostly inconspicuous nerves; petiole
(2—) 4—6 mn, hairy. Spikes 1—4 (—5) in the axil of
one leaf, (1—) 2—3 cm long when flowering; bracts
lanceolate, 0.5—1 mm long. Perianth urceolate,
greenish yellow or white, tube c. 1.5 mm long, lobes
recurved. Stamens as long as the perianth tube. Disk
scales slightly fleshy, oblong, apex 3-toothed, c. 0.5
mm long. Ovary ovoid, c. 1 mm long; style c. 0.5 mm
long; stigma shallowly 4-lobed. Drupe orange-red,
ellipsoid to nearly globular, 11—15 by 9-13 mm.

Distr. Northern Australia; in Malesia: New
Guinea (incl. Bismarcks), Moluccas (Key and Sula
Is.), Lesser Sunda Is. (Sumba, Alor, Timor), and E.
Java (Surabaya). Fig. 15.

VALETON (/.c. 1886, 159) mentioned ‘Nova Zeelan-
dia’, but this is obviously an error for ‘Nova Gui-
nea’.

Ecol. In evergreen forest or in semi-deciduous
thickets, often climbing on the edge of woods; from
sea level up to 700 m; on calcareous rocks (Java) or
on sandy soil. According to STAUFFER (in sched.)
parasitic on Leguminosae and Sapindaceae. Fl.
Jan.—Dec. Flowers with sweet scent.

Uses. According to DuNLopc.s. (N. Territ. Bot.
Bull. 1, 1976, 59) the fruits are edible.

Vern. Lesser Sunda Is.: kema raberi, Sumba,

OPILIACEAE (Hiepko) Sy |

Fig. 17. Cansjera leptostachya Bru. With the flower-

ing pendent branches. Ifar near Hollandia (Jayapu-

ra), W. New Guinea (VAN RoyYEN & SLEUMER 6174;
photogr. SLEUMER, July 1961).

kape bila, Alor; Moluccas: mé6 menumpang, Sula
Is.

Note. Cansjera leptostachya is undoubtedly
closely allied to C. rheedii. But since it differs from
this species in several floral and vegetative characters
(inflorescences more lax, perianth tube clearly shor-
ter; leaves smaller, lanceolate, and more herbaceous;
spines never reported) | prefer to maintain C. lepto-
stachya as a distinct species.

52 FLORA MALESIANA [ser. I, vol. 10!

Excluded

Cansjera grossularioides Buco, FI. Filip. (1837) 73 (‘Cansiera’) = Antidesma ghaesembilla GAERTN. (Eu-
phorbiaceae).

Cansjera pentandra Buco, l.c. = Antidesma pentandrum (Buco) MerR. (Euphorbiaceae).

Cansjera rheedii Buco, l.c., non J.F. GMELIN = Antidesma pentandrum (Buco) MeErR. (Euphorbiaceae).

Champereia perrottetiana BatLL. [Adansonia 3 (1862) 125] is doubtless a Scleropyrum sp. and probably
rightly regarded as Scleropyrum pentandrum (DENNST.) MABBERLEY [= S. wallichianum (W. & A.) ARN.],
Santalaceae; cf. BAILLON, Hist. Pl. 11 (1892) 467; Hook. f. Fl. Br. India 5 (1886) 235.

Lepionurus pubescens Ript. Trans. Linn. Soc. Bot. 9, 1 (1916) 27 = Scleropyrum aurantiacum (LAUvT. &
K. ScH.) PILGER (Santalaceae).

ARISTOLOCHIACEAE (Ding Hou, Leyden)

Perennial herbs, more commonly woody at the base, undershrubs or shrubs,
erect, scrambling or scandent, sometimes high lianas. Rhizome not rarely tuber-
ous. Branches often slightly swollen and jointed at nodes. Hairs simple, uni- or
multicellular, short ones often with a hooked apex. Leaves simple, spiral or al-
ternate, petioled (without an abscission zone), exstipulate; midrib usually prom-
inent beneath, elevated or flat above; nervation commonly palmate, or pinnate,
nerves often obliquely extending towards the margin. Flowers bisexual, actino-
morphic or zygomorphic, solitary, fasciculate, or in axillary or cauligerous,
racemose, paniculate or cymose inflorescences, usually only one or two flowers
open at a time; bracts present and often persistent; pedicel often hardly distinct
from the ovary. Calyx petaloid, gamosepalous, 3- (or 6-) lobed or 1-lipped;
lobes valvate or induplicate. Peta/s (in Mal.) absent. Disk (?) 0, rarely present
(e.g. a few Thottea spp.). Stamens 6 (4 or 5 in some extra-Mal. Aristolochia
spp.) or 6—c. 36 (—46), in 1 whorl or in 2 (3 or 4) whorls (Thottea); filaments
free or slightly mutually united at the base, and/or almost completely adnate to
the style column to form a gynostemium; anthers free (7hottea) or dorsally unit-
ed with the style column (Aristolochia), each consisting of 2 thecae with 4 pollen
sacs, extrorse, rarely introrse (extra-Mal. spp.), dehiscing longitudinally. Ovary
inferior (rarely half-inferior in extra-Mal. genera), 4—6-carpellate, 4—6-celled,
syncarpous (or + apocarpous in extra-Mal. Saruma); placentae parietal (dis-
tinct when young, then intruding and connivent axially, thus often seemingly ax-
ile); ovules usually many, anatropous, in | or 2 vertical rows in each locule of
the ovary, horizontal or pendulous; style-column 3—many-lobed, sometimes
some of the lobes redivided; stigmas or stigmatic tissue apical, lateral, or on the
surface of style lobes. Fruits capsular or siliquiform (follicular or cocci in extra-
Mal. genera), 4—6-celled; dehiscing apically towards the base (basipetal, e.g.
Thottea) or basally towards the apex (acropetal, e.g. most Aristolochia); septici-
dal, rarely septifragal (some extra-Mal. Aristolochia) or bursting irregularly
(extra-Mal. Asarum); rarely indehiscent (W. African Pararistolochia). Seeds
many in each locule (1-seeded in extra-Mal. Euglypha), often coated with re-
mains of placental tissue (membranous when dry), horizontal or pendulous,
variously shaped; ovate, deltoid or triangular, flat, convex-concave, or longitu-
dinally curved, or oblong (and triangular in cross-section), rugose, finely verru-
cose, or smooth, immarginate (Thottea; Aristolochia, p.p.) or winged (Aristo-
lochia, p.p.); albumen fleshy, copious; embryo minute, cotyledons two, dis-
tinct.

Distribution. There are 7 genera, Aristolochia worldwide, Asarum over the northern hemi-
sphere, Thottea in continental Southeast Asia and Malesia, Pararisto/ochia in tropical Africa, and
3 monotypic genera, viz. Saruma in China, Holostylis and Euglypha in South America. As to
number of species, Aristolochia is by far the largest with some 300 spp., largely concentrated in
the New World, especially in Central and South America, in Malesia with 28 spp.; Asarum (incl.
Hexastylis and Heterotropa) with possibly some 70 spp. in northern temperate regions, Thottea
with 26 spp., of which 22 in Malesia, and Pararistolochia with 12 spp. in West Africa.

(53)

54 FLORA MALESIANA [ser. I, vol. 10!

Ecology. In Malesia Aristolochiaceae occur mostly locally, often sporadic, exceptions being
Thottea tomentosa which may be a locally common undershrub and Aristolochia tagala which
is often a common slender twiner in thickets. Usually the species are confined to the primary for-
est, from the lowland to montane stations, in various forest types, dryland and swampy forest,
on limestone, in secondary forest and bamboo groves, only a few species ascending to 1500—2250
m altitude.

Aristolochiaceae as host plants for butterflies. Certain groups of Papilionidae are bound to
Aristolochiaceae as a host plant and this is true of Malesian Aristolochia and Thottea. EHRLICH
& RAVEN (1964) have made a survey and found that in the family Papilionidae, the swallow tail
butterflies having 3 subfamilies, the holarctic and oriental subfamily Parnassiinae with 5 genera,
feed only on Aristolochiaceae. In the tropical worldwide subfamily Papilioninae with 3 tribes, the
tribe Troidini is almost confined to Aristolochiaceae as host plant. The bond between the butter-
flies and their host may be different, some are monophagous, others are oligophagous (feeding
on a few species) and still others are polyphagous.

Obviously there is a choice, coinciding with the taxonomy of the butterflies and the phytochem-
istry of the host plants. EHRLICH & RAVEN use the term ‘co-evolution’ in this respect, but it should
be pointed out that in this case the benefit is only for the butterflies (/.e. their larvae); they do
not serve in pollination; in proper co-evolution both parties are interdependent.

It is found by entomologists that the female butterflies are attracted by the scent of the plant
to lay their eggs. The evolving caterpillars feed on the host and in the herbarium one may find
traces of this, in the way of leaf perforations or erose leaf margins; the larvae also feed on new
shoots and buds. Pupae are generally found near the base of the stem of the host, and that is in
some very large rain-forest lianas far away from the foliage of the host.

Several entomologists have published on the relations of Aristolochiaceae and butterflies in
Malesia, e.g. STRAATMAN (on N. Sumatra, SE. New Guinea, Queensland, and the Solomons), IGaA-
RASHI (On the Philippines and New Guinea), while HAUGUM listed them from the Papuan region.
I gave a summary (1983) and JAcoss a review (1982).

Literature: EHRLICH& RAVEN, Evolution 18 (1964) 586—608; Haucum, The Lepidoptera group
of 1968, Newsletter 2 (1981) 171—184; Dinc Hou, Blumea 29 (1983) 223—249; IGARASHI, Food
plants of Papilionidae (1979); JAcoss, Fl. Males. Bull. 35 (1982) 3747—3749; STRAATMAN &
NIEUWENHUIS, Tijdschr. Entom. 104 (1961) 31—41; J. Lep. Soc. 16 (1962) 99-103; ibid. 23 (1969)
69-76; ibid. 25 (1971) 58-64.

Pollination. Already two centuries ago SPRENGEL Suggested insect pollination in Aristolo-
chia and a century ago HILDEBRAND found the flowers proterogynous and concluded to cross-
pollination. As a matter of fact the flowers represent a beautiful trap with a ‘slide zone’ on the
limb above the tube which is inside usually provided with retrorse hairs preventing insects to leave
during anthesis. They are trapped in the utricle which provides them with nectar and usually also
other food substance of glands. BAKER C.s. (1973) added that also stigmatic secretions containing
amino-acids would add to the nutritional potential in the utricle. The insects, mostly flies, some-
times also ants, are attracted to the flowers by the putrescent odour, sometimes an offensive smell
of decaying meat, emitted during anthesis by the flower or its stalk, and this occurs also in other
genera of the family. PETcH (1924) found that some species are visited by only one kind of fly,
but in other species he found up to 13 different kinds; the two native Ceylon species were visited
by one kind of fly only. In some intricate-built flowers of South American species insects are guid-
ed to the sexual organs by a window-pane in the utricle. After the flower withers, and the hairs
have lost turgescence, the insects can crawl out, loaded with pollen and can visit another flower,
leading to cross-pollination.

This is only a generalization, as it appears from the very large study by PETCH (1924) that there
is a great variability among the species: mostly flowers open at daybreak or shortly before and
wither after 24 hours, but there are species which show a second-day revival; some have no food
bodies; in some species the tube is wide and flies can easily escape; in other species the tube has

1984] ARISTOLOCHIACEAE (Ding Hou) 55

no hinged hairs. For that reason one cannot give a single answer to whether cross-pollination is
necessary for the setting of the fruit in all species.

Burck (1890, 1892) made extensive experiments, including bagging flowers efc., on three exotic
species in the Botanic Gardens at Bogor (viz. Aristolochia barbata, A. elegans and A. ornithoce-
phala = A. brasiliensis) and concluded that they are autogamous. PETCH (1924) studied in detail
some dozen species at Peradeniya in Ceylon and concluded that, ‘although Aristolochias are
adapted for cross-fertilization, some species can be self-fertilised. It is evident that all grades of
self-fertility or self-sterility may be expected within the genus.’

Observations on pollination in Thottea are very scant; its flowers are regular and open and do
not offer a complicated structure as in Aristolochia. They emit also a putrid smell, are mostly
dark-coloured and their flower is also proterogynous, as BACKER (1918) observed in Apama to-
mentosa at Bogor. He stated that this species propagates very well vegetatively by stooling and
that very few fruits are produced, both in cultivation and in the field in bamboo groves at Depok.
BACKER observed flies visiting the flowers; he hypothesized that cross-pollination might be pos-
sible during the transition period from the female to the male stage. As a matter of fact I found
(1981) the styles or style-lobes (with their stigmas or stigmatic surfaces) reflexed or twisted at an-
thesis, facilitating contact with pollen grains, which I found germinated in flowers of Thottea tri-
serialis. Self-pollination and fertilisation may hence also occur in Thottea.

For Asarum reports also vary and both self-pollination and cross-pollination by flies or fungus
gnats seem to occur (VOGEL, 1978).

Literature: BACKER, Trop. Natuur 7 (1918) 177-183, 4 fig. (on Apama); ibid. 8 (1919)
133—138, 150—155, 161—168, fig. 5-15; H.G. BAKER c.s. in Brantjes (ed.), Pollination and dis-
persal (1973) 47—60; Burck, Ann. Jard. Bot. Btzg 8 (1890) 149-157, t. 23; Bot. Zeit. 50 (1892)
121-129, 137—144, t. 3; CAMMERLOHER, Oest. Bot. Z. 72 (1923) 180—198; t. 5—6; Dinc Hou, Blu-
mea 27 (1981) 314; ibid. 29 (1983) 223—249; LEEMAN, Bull. Soc. Bot. Geneve 19 (1927) 149-159,
fig. 98-107; K.L. Lu, Syst. Bot. 7 (1982) 150—157, t. 1-3 (both on Asarum); PETCH, Ann. R.
Bot. Gard. Perad. 8 (1924) 1—108, t. 1-5; Prerrer, Ann. Mo. Bot. Gard. 53 (1966) 119-120; S.
VocEL, Flora 167 (1978) 329—366, fig. 1—12.

Morphology. Habit. In Malesia there are two main habit types: 1) perennial herbs which are
often woody at the base; they are either a) erect undershrubs or shrub-like, up to 3 m high, some-
times slightly higher as in most species of Thottea and some of Aristolochia (e.g. A. humilis, A.
macgregorii, A. sericea, etc.) or b) spreading, scrambling or twining up to several metres high,
as in Thottea corymbosa and some species of Aristolochia (e.g., A. glaucifolia, A. jackii, A. linne-
mannii, A. minutiflora, etc.); and 2) woody twiners or high lianas from a few metres up to c. 50
m high, with an old stem up to 2 (—4) cm @ (most species of Aristolochia).

In absence of field data on the habit (erect or climbing) sterile specimens can hardly be identified
to the genus (Aristolochia or Thottea). Sterile specimens of erect plants can be discriminated if
they are sufficiently ample; see the paragraph ‘leaf architecture’ under Thottea.

As to the direction of twining, I do not know whether it is constant for the species of Aristolo-
chia. | observed that plants of A. tagala, A. ringens and A. foveolata germinated from seeds and,
growing in my office, appear to have no definite direction to twist and may go either right or left
as stated by MENNINGER (Flowering vines of the world, 1970, 91—99, phot. 42—45).

Lianas of Artistolochia twining on high trees bear leaves often at the top and flowers and/or
fruits at the lower part of the stem. Occasionally only ‘leafless’ fertile herbarium specimens were
available because the leaves were difficult to locate or to collect.

Roots and rootstocks. The roots, sometimes also root-like tubers, of (some) Aristolochia are
fleshy, sometimes with bitter taste, and of various shapes (e.g. globose, ovoid, cylindric,
fusiform, turnip-shaped, efc.), which are characteristic for some species. They have sometimes
been collected, recorded and used for species delimitation (cf. Davis& Kuan, Notes R. Bot. Gard.
Edinb. 23, 1961, 515—546; Liana, Acta Phytotax. Sinica 13, 1975, 10-28; Cuow & Hwan, /.c.
108—109). From Malesia nothing is known about root structure.

56 FLORA MALESIANA [ser. I, vol. 10!

Fig. 1. Scanning electron micrographs showing features of leaf undersurfaces. — a—c. Thottea: a. T. corym-

bosa (GriFF.) DING Hou, b. T. dependens (PLANCH.) KLoTzscu, c. T. muluensis DING Hou. — d—f. Aristolo-

chia: d. A. macgregorii MERR., e. A. gaudichaudii DUCHARTRE, f. A. tagala CHaAmisso. All x 270 (a & c hairs

and mostly curved or hooked thickenings, b hair and ring- or loop-shaped papillae, d hooked hair and stomata

with striae, e glabrous surface and scattered stomata with raised rim, f scattered hooked hairs and stomata

with raised rim) (@ CARRICK 1489, b SIDEK BIN KI1AH 295, c ARGENT c.S. 760, d BARTLETT 15090, e BW 11439,
J WEBER 1074).

1984] ARISTOLOCHIACEAE (Ding Hou) 57

The recumbent rootstocks or rhizomes of Thottea and some Aristolochia species develop off-
shoots or runners, which sprawl on the ground or produce erect stems. When the motherplant
dies, these stems become free and grow on as separate individual plants, a method of vegetative
propagation.

The stems of woody vines of Aristolochia are mostly terete, or sometimes slightly flattened (fig.
16), and are up to 4cm or more in diameter. The bark of the old stems is corky and is often longitu-
dinally fissured or prominently ridged or sometimes rather smooth.

On a cross-section one can observe, by using a handlens, conspicuous anatomical features of
the Aristolochiaceae: the vascular bundles are arranged in a ring and widely separated from one
another by the broad medullary rays. On the cross-section of a rather flattened stem, where the
cambium is more active towards two opposite directions, the vascular bundles elongate according-
ly and the whole section appears like the numeral ‘8’ (cf. METCALFE& CHALK, Anatomy of Dicoty-
ledons 2, 1965, 1114, 1117, f. 237, 268; Poncy, Adansonia 17, 1978, 466, 476, f. 1).

The ‘8’-shaped appearance of the cross-section of the stem has been used as one of the generic
characters for separating the tropical African Pararistolochia from Aristolochia (with circular
stem) (cf. Poncy, /.c.). In Malesia old stems of Aristolochia decandra and A. coadunata are some-
times also flattened. Fig. 16.

Leaves. Leaves of Aristolochiaceae can provide useful characters especially for identification
of sterile collections. Fig. 1, 8. However, in some species, they are heteromorphic or very variable
in shape, size, texture, efc.; they vary sometimes also between those of fertile and vegetative
branches, apical and lower parts of a (high) woody vine, juvenile and adult stages, efc. (e.g. in
Aristolochia dielsiana, A. tagala, A. zollingeriana; Thottea tomentosa).

The leaves of Malesian Aristolochia vary in size; the largest known to me occur in A. dielsiana
measuring up to,37 by 23 cm; according to R. STRAATMAN they can reach to 100 by 70 cm.

The leaves are usually distinctly petioled. The petiole is often more than 2 cm long, sometimes
up to 13 cm; it is very short only in a few species, e.g., Aristolochia macgregorii (c. 3 mm), A.
sericea (2—5 mm). In Thottea petioles are usually short.

The leaf does not possess an abscission zone either on the petiole or at its base. The old or dried
leaf just hangs on the plant for some time and then breaks irregularly from the petiole, leaving
no scar on the stem. This is very characteristic for the species of this family.

The undersurface of the leaf has interesting sculpture features or ornamentation, e.g. hair types
or density of hairs, cuticular thickenings or markings, protuberances of epidermal cells, efc.,
which are useful as diagnostic characters, especially for identification of steriie collections (cf.
Blumea 27, 1981, 310—311, f. 5S—33). Fig. 1. For example, Thottea dependens has papillae form-
ing rings or curves (fig. 1b), 7. mu/uensis, T. pennilobata, and a few others show crescent, curved
or hooked thickenings (fig. lc), Aristolochia macgregorii has stomata with extended striae of
thickenings (fig. 1d). Such characters can easily be examined under a normal binocular with a
magnification up to about x 60; sometimes they can even be observed with a handlens.

Also the venation types are often characteristic; the main ones are illustrated in fig. 8.

Series of axillary buds. In some species of Thottea and Aristolochia sometimes 2 or 3 (—5) buds
occur in a leaf axil, especially in the terminal one. These buds may develop into flowering and/or
vegetative branches, e.g. in Thottea corymbosa, Aristolochia sericea, A. gaudichaudii, etc. (cf.
Scumipt in E. & P. Nat. Pfl. Fam. ed. 2, 16b, 1935, 210—211, f. 106; DeLaiGuE, Soc. Bot. Fr.,
Mém. 1971, 167—177, f. 1—6).

Flowers. The flower in Aristolochiaceae is probably essentially provided with a calyx and a co-
rolla, but the latter is almost always suppressed. It is still present in the monotypic Chinese genus
Saruma which is assumed to be the most primitive of the family. It is also found as 3 rudimentary,
subulate segments in Asarum canadense, as a relict feature.

Flowers are very important for species delimitation. Unfortunately, for many Malesian species
flowering material is scanty in the herbarium. Some tropical species of Aristolochia have rather
large flowers, the largest being the neotropical A. grandiflora Sw., with a limb up to 50 cm wide

58 FLORA MALESIANA [ser. I, vol. 10!

and a total flower length up to 3 m, a serious competitor of Rafflesia which is mostly held as the
largest flower in the world. In contrast with this, Aristolochia flowers have often a thin, delicate
texture difficult to handle in dried material.

The flowers in Aristolochia open only one or two, or a few at a time. The flowering duration
is often very short, one to a few days. They are sometimes deformed after pressing and drying.
The flowers of Aristolochia deliquesce sometimes rapidly; they also fall and decay quickly follow-
ing pollination and fertilization (cf. PFEIFER, Ann. Mo. Bot. Gard. 53, 1966, 119).

The flowers are bisexual; they emerge terminally or laterally in the axils of leaves or bracts,
and/or cauligerous; they are solitary, fasciculate, or arranged in cymes, racemes or panicles. The
flowering branches or rachides are sometimes with spacious internodes (e.g., Aristolochia jackii,
A. schlechteri) or strongly reduced with internodes hardly visible (e.g., A. crassinervia, A.
sericea). The flowers are pedicelled. There is often hardly any external distinction visible between
the pedicel and the ovary; they have been treated here as one unit.

The perianth or calyx is 3-lobed and actinomorphic in Thottea. In Aristolochia it is rather spe-
cialized and usually zygomorphic; it consists of three (sometimes not sharply separated) parts:
utricle, tube and limb. Between the perianth and the ovary, there is often a constriction or articula-
tion, sometimes with a lobed rim where the perianth breaks off from the fruit.

The utricle is the basal inflated part of the perianth. It is often globose, subglobose, ellipsoid,
ovoid or obovoid. On the inner surface of the utricle, there are usually two symmetrically placed
glandular, usually ellipsoid swellings at the apical part. They are food bodies, composed of dense
glandular hairs, serving for imprisoned insects (cf. PETcH, Ann. R. Bot. Gard. Perad. 8, 1924,
28). Sometimes there are two small bosses or depressions shown on the outer surface correspond-
ing to the position of the food bodies inside (cf. Curtis’ Bot. Mag. t. 7429). Some Malesian spe-
cies have six such glandular food bodies (e.g., Aristolochia foveolata, A. papillifolia). The distal
end of the utricle is gradually or abruptly narrowed into a cylindric tube which may be straight
or curved. The base of the ‘tube’, specially in some extra-Malesian species, slightly elongates and
projects into the utricular cavity; the flange-like part inside the cavity has been called syrinx (cf.
Preirer, Ann. Mo. Bot. Gard. 53, 1966, 116, f. 1). The tube gradually or abruptly and slightly
enlarges its size at the apical part and merges with the expanded limb. For the diameter of the
tube, only the cylindric, middle part has been taken.

The limb is 1-lipped (in many species), sometimes distinctly 3-lobed (e.g., Aristolochia decan-
dra, A. momandul), occasionally rim-like and obscurely 3-lobed (A. coadunata), or rarely 6-lobed
(A. schlechteri).

The colour of the perianth appears sometimes to vary with the developing stage of the flower,
as recorded in field notes. It is characteristic in some species. Unfortunately, I could not use it
in keys, because it has only erratically been recorded in field notes.

Perianth of Aristolochia. As mentioned above, the perianth of Aristolochia should be regarded
as homologous with a calyx and of course be homologous with the perianth in other genera of
the family (e.g. Asarum, Thottea). In several species it is also 3-lobed, but in many others it is
entire. Some authors have, however, a different opinion about its morphological derivation.

Lorcu (Evolution 13, 1959, 415—416, f. 1) observed a shoot of Aristolochia maurorum bearing
a series of teratological leaves and proposed a new interpretation of the perianth of this genus.
He stated that ‘the perianth is the metamorphosed first leaf of a lateral branch’ and ‘. . . agrees
in form with an involute normal foliage leaf.’

HAGERUuP (Bull. Res. Counc. Israel 10, sect. D, 1961, 348—351, f. 1—14) studied both the vena-
tion and the development of the leaf and the perianth of Aristolochia (especially A. elegans). He
concluded that ‘The perianth is not compounded of several united leaves but consists of only a
single leaf (like the spathe of the Araceae).’

GuEbes(Flora, ser. B, 158, 1968, 167—179, f. 1-5) and TionG CHur1 HuonG(Morph. and taxon.
studies on some Aristolochiaceous plants in Singapore, 1979/80, 43—45, not published) made
comparative, morphological studies on the vegetative leaf and the perianth of Aristolochia (e.g.,

1984] ARISTOLOCHIACEAE (Ding Hou) 59

A. clematitis, A. grandiflora, A. peltata). Their results confirmed the interpretation and findings
of LorcH and HAGERUP.

It should be remarked that the Aristolochia species, studied morphologically and anatomically
by the three authors all possess a 1-lipped perianth. Their thesis should be tested for species in
which the limb is rim-like or obscurely 3-lobed (e.g. A. coadunata, A. griffithii), or distinctly
3-lobed (e.g. the tropical West African species of Pararistolochia; and A. decandra, A. moman-
dul), and the 6-lobed species A. schlechteri.

Stamens and styles. The number and arrangement of the stamens in Thottea show an interesting
series of reduction. Fig. 4—7. The stamens in this genus range from 36 (—46) (e.g. T. grandiflora)
to as few as 6 (e.g. T. tomentosa); they are from free and arranged in 4 series (7. parviflora),
through partly free and in 3 (7. ¢riserialis) or 2 series (most of the species), to united with the style
column and just in | series (several species, e.g. 7. corymbosa).

In Aristolochia the stamens are adnate to the style column to form a gynostemium. All Malesian
species have 6 stamens, except A. decandra which has 10. Fig. 15. Each anther consists of two
thecae with four microsporangia (pollen sacs) (cf. JoHRI & BHATNAGAR, Phytomorphology 5,
1955, 124—125, f. 8, 44—47; Narr& NARAYANAN, Lloydia 24, 1961, 199—200, f. 1—3). The thecae
of astamen are in some Aristolochia species (e.g. A. jackii) separated from each other by a rather
broad connective.

The styles appear to be free in Thottea parviflora. They are united with the stamens into a short
column (gynostemium) in all other species of Thottea and Aristolochia. The style column may
be discoid or obtuse at apex and then divides, or sometimes redivides, into a number of slender
or finger-like lobes. The number of styles or style lobes varies in species of Thottea from c. 20
(e.g. T. macrophylla) to only 2 or 3 (e.g. T. paucifida; fig. 5). In Malesian Aristolochia the style
has 6 lobes (except 3 in A. coadunata and 10 in A. decandra).

The lobes of style column (or gynostemium) are glabrous (often sticky when fresh) or some-
times (densely) hairy (covered with hooked and/or straight hairs or papillae). In Thottea, they
are erect or spreading when young and often reflexed or irregularly twisted at anthesis (cf. DING
Hou, Blumea 27, 1981, 311—314, f. 38-50).

In Aristolochia, changes occur in the structure and shape of the style lobes at anthesis. When
young, they are distinctly separate from one another. At first the style lobes may be rather thin
with longitudinally reflexed margins and their basal parts covering the apices of the unopened an-
thers. At anthesis, the style lobes slightly swell, flatten, and become erect and adherent; their api-
cal parts bend inward, and the anthers become exposed. The lobes form then almost a funnel;
their apical parts and inner surfaces have a rather thick layer of slime (cf. BACKER, Trop. Natuur
8, 1919, 134—136, f. 1—4; Preirer, Tax. rev. pentand. sp. Aristolochia, 1970, 8—9, f. 1).

In herbarium specimens of both Aristolochia and Thottea | observed that the style lobes are
sometimes covered with pollen grains which even may have germinated. These lobes certainly pos-
sess stigmatic surface. However, some botanists assume that the lobes are not true stigmas, and
that the connectives of the anthers have assumed stigmatic functions (cf. BurcK, Ann. Jard. Bot.
Btzg 8, 1890, 151—153, f. 4-8; Wiruis, Dict. Fl. Pl. Ferns, rev. by Airy SHAW, 8th ed., 1973,
92). This idea seems a bit far-fetched, as for example Aristolochia coadunata has only 3 style
lobes, but the usual 6 stamens. Also in Thottea the number of stigmatic lobes does not correspond
with the number of stamens; in 7. tomentosa, with 6 stamens the number of lobes is 3 or 4.

Ovary and placentation. The ovary is inferior (but half inferior in extra-Malesian monotypic
Saruma and some species of Asarum). It is linear, cylindric or fusiform, and is 4- to 6-carpellate
and syncarpous (apocarpous in Saruma).

The placentas are parietal when young and gradually become imperfectly 4—6-celled. Whenever
I dissected a flower, I observed that the placentation appears to be axile. The pseudo-axile appear-
ance is due to intrusion and fusion of the placental partitions in later stages (cf. LEEMANN, Bull.
Soc. Bot. Genéve 19, 1927, 140-146, f. 82-92; Jour1& BHATNAGAR, Phytomorphology 5, 1955,
123-124, f. 1-7).

60 FLORA MALESIANA [ser. I, vol. 10!

Fig. 2. Scanning electron micrographs of seeds. — a—d. Aristolochia: A. transtillifera DING Hou, a. undersur-
face, showing testa with finely granulate thickenings, x 12, b. upper surface, showing the funicle with laterally
dilated extension flattened against the seed, x 15; A. philippinensis WarB., c. undersurface, showing testa
with rather coarse, wart-like thickenings, x 15, d. upper surface, showing the funicle with laterally dilated
extension covering the seed, x 15. — e—g. Thottea: T. macrantha (BoOERL.) DinG Hou, e. seed with coating
tissue partially peeled off, x 20, f. surface view of the testa, with periclinal walls peeled off, showing a bundle
of fibrous thickenings in each cell lumen, x 700; 7. reniloba Dinc Hou, g. seed with tuberculate testa, coated
with dried, membranous tissue, x 21 (a & b SAN 19008, c & d PNH 10592, e & f LORzING 12434, g DE WILDE
& DE WILDE-DUYFIES 18829).

1984] ARISTOLOCHIACEAE (Ding Hou) 61

The ovules are anatropous and bitegmic (cf. JoHRI& BHATNAGAR, /.c. 128-132, f. 48—54; Narr
& NARAYANAN, Lloydia 24, 1961, 200—201, f. 11—15). They are usually numerous and are
horizontally or pendulously superposed in one or two series in each locule of the ovary.

Fruits and seeds. Fruits and seeds are very characteristic for the Aristolochiaceae. The fruits
are usually capsular (e.g. Aristolochia; fig. 17) or siliquiform (Thottea; fig. 4) (but follicular in
the extra-Malesian Saruma and cocci in Euglypha). They are 4—6-loculed, usually dehiscent, sep-
ticidal, acropetal (and the opened, hanging fruit basket-like as characteristic in Aristolochia; fig.
17), or basipetal. They are indehiscent in the tropical African Pararistolochia and possibly also
in the New Guinean Aristolochia dielsiana (see there). They are usually glabrous or rarely hairy
(Thottea) (cf. ScumipT in E. & P. Nat. Pfl. Fam. ed. 2, 16b, 1935, 220—222, f. 116—117).

The size of the fruits is very variable: in Malesian representatives: the length ranges from c. 1
em (e.g. Aristolochia sericea) to 20 (—38) cm (e.g. A. dielsiana; Thottea tricornis) and the width
from c. 0.5 cm (Thottea) to c. 4 cm (Aristolochia).

The fruit wall is often slightly lignified (but strongly lignified in Pararistolochia and some Aris-
tolochia species). The valves of the capsules in Aristolochia can sometimes easily be separated in
epi-, meso- and endocarp.

The seeds are usually numerous, horizontally or pendulously superposed, and immersed in the
spongy cellular tissue in each locule of the capsules (but only one seed developed in extra-Mal.
Euglypha). They may be divided into two main types according to their general appearance: 1)
compressed and flat (Aristolochia, Asarum and Holostylis) and 2) oblong, fusiform, or broadly
ovoid, obscurely or distinctly triangular (Thottea). Fig. 2. However, the flat seeds sometimes may
be longitudinally slightly or strongly concave (e.g. Aristolochia singalangensis; Thottea curvise-
men and Thottea sp.).

The seeds are not winged or with a rim-like or marginal wing (Aristolochia). They are often
slightly or prominently transversely corrugate or rugose (Thottea) and are smooth or warty on
the testa (Aristolochia). The irregular surface of the testa in Aristolochia is due to unequal divi-
sions and outgrowth of the epidermal cells (cf. JoHR1& BHATNAGAR, Phytomorphology 5, 1955,
133, f. 90-91).

In Aristolochia the seeds in many species have an almost unique feature in that the large funicle
is rather fleshy, thick, dilated laterally, flattened against the upper surface of the seed and general-
ly larger than it (JoHRI & BHATNAGAR, /.c.; CORNER, Seeds Dicot. 1, 1976, 73-74; ibid. 2, 1976,
f. 27—29). This fleshy funicle is equivalent to an elaiosome and is important in seed dispersal. In
the dry state it becomes almost membranous and usually covers the seed (fig. 2b, d).

In Thottea, after the coating membranous tissue is removed, the testa cells appear as reticula-
tions or papillae; each of these cells has a strong thickening projecting into the cell lumen. If the
soft tissue of the testa has been removed or brushed off, one can easily observe the two layers
of crossed fibres of the tegument (cf. SOLEREDER, Bot. Jahrb. 10, 1889, 504—507, t. 13, f. 19-21;
Jouri& BHATNAGAR, Phytomorphology 5, 1955, 133-137, f. 83-95; Corner, Seeds Dicot. /.c.;
Dinc Hou, Blumea 27, 1981, 314—315, f. 51—69). Fig. 2e—g.

According to CorNER (/.c.) the attachment of the integument along the course of the raphe and
the development of two layers of crossed fibres in the tegument forming the mechanical layer of
the seed coat are the chief characters of the seeds in Aristolochiaceae. He also stated that the tegu-
ment of Caricaceae seems strikingly similar to the one of this family in having the same set of
crossed fibres.

The endosperm of the seeds in Aristolochiaceae is copious and fleshy. The embryo is minute
with two distinct cotyledons and is enclosed in the endosperm close to the hilum (cf. Jonri &
BHATNAGAR, /.c.).

Seed germination and seedlings. The seed germination of some species of Aristolochia and Asa-
rum (s./.) has been reported as epigeal, with the cotyledons spreading above the ground.

In Aristolochia, during germination, the radicle protrudes through the hilum or near it or
through the testa. The cotyledons are rather fleshy, suborbicular or broad-ovate, with simple

62 FLORA MALESIANA [ser. I, vol. 10!

venation (midrib with a few lateral nerves or veins). The first two leaves are opposite; they develop
from almost the same plane as the cotyledons and are at right angles with them. The foliage leaves,
following the first pair mentioned above, are scattered (cf. LUBBock, Contribution to our know-
ledge of seedlings 2, 1892, 444—446, f. 624; TlionG CHu1 Huona, Morph. and taxon. studies on
some Aristolochiaceous pl. in Singapore, 1980, 27—28, pl. 4 & 5, f. 4 & 5, not published).

Seed dispersal. The winged seed of some Aristolochia species may help in dispersal. More im-
portant seems the elaiosome (fleshy funicle) which is probably attractive to ants.

Anatomy (for oil cells, silicified cells and crystals see under Phytochemistry). METCALFE &
CHALK (1950) provided a general survey of the vegetative anatomy of the family: hairs simple uni-
cellular or uniseriate and/or with a hooked terminal cell with silicified tip (‘bracket hairs’). Sto-
mata usually anomocytic. Stems typically with broad medullary rays. Secondary phloem occa-
sionally with stone cells but devoid of fibres. Wood with very wide vessels in climbers, but rather
narrow ones in erect species. Vessels with simple perforations and coarse pits. Fibres with bor-
dered pits (especially conspicuous in Aristolochia). Parenchyma paratracheal, often scanty. Rays
mostly wide and forming broad interfascicular bands, but narrow (up to 3-seriate) in some species
of Apama (= Thottea), heterocellular. GUEDEs (1968) described the petiole anatomy of some
Aristolochia species; ALEYKUTTY & INAMDAR (1980) provided detailed accounts of hair types in
the family; Pup (1983) reported on the diverse ontogeny of the stomatal complex in species of
Aristolochia and on the predominance of paracytic stomata in Aristolochia leuconeura.

Vegetative anatomy is in agreement with the view that Aristolochiaceae are related to the
Magnoliales.

Literature: ALEYKUTTY & INAMDAR, Fedde, Rep. 91 (1980) 95—108; GUEDEs, Flora, Jena 158B
(1968) 167-179; MeTCALFE & CHALK, Anatomy of the Dicotyledons II, Oxford (1950); PHP,
Curr. Sci. 52 (1983) 223—224. — P. Baas.

Palynology. The pollen of Aristolochiaceae varies in size between 27 um in Saruma henryi
and 73 um in Aristolochia grandiflora and is generally spherical-suboblate or ellipsoidal. Two
main types can be recognized, the first of which is restricted to Saruma. This genus has monocol-
pate, reticulate pollen which is rather primitive and similar to the basic type found in Chlorantha-
ceae, Annonaceae and in many monocotyledons. The second, more derived type found in the re-
maining genera is characterized both by a variable aperture configuration and exine structure.

In Aristolochia the pollen grains are inaperturate, indistinctly monocolpate or periporate and
the exine may be psilate, scabrate, echinate or areolate. Pararistolochia has an indistinctly out-
lined distal aperture and differs from the preceding genus mainly in its rugulate-areolate exine.
The pollen of Asarum is inaperturate or irregularly pericolpate-periporate and this variation may
even occur in a single species (Asarum virginicum). The exine has separated verrucae.

Thottea pollen is generally inaperturate, but indistinctly monocolpate or periporate grains have
been reported also. The exine is verrucate-areolate. In T. paucifida the areolae are hardly devel-
oped, while in 7. dependens, T. dinghoui, T. macrantha, T. tomentosa and T. tricornis the areo-
lae are widely spaced, thin-walled and not centrally supported by columellae. The intervening ex-
ine here is tectate-columellate. Densely spaced areolae are found in 7. grandiflora and T. parvi-
flora and in these species the columellae are reduced to the margins of thin-walled areolae. In the
former species the areolae are perforated by rather large holes. T. corymbosa is deviating in the
larger, rather densely spaced areolae which are finely perforated and supported by rather densely
spaced columellae.

With the exception of Saruma, the genera of Aristolochiaceae cannot be separated on pollen
morphological characters, although some species may be distinct. The comparatively primitive
Saruma pollen type indicates that the taxonomic relations of the family are with Magnoliales.

Literature: G. ERDTMAN, Pollen morphology and plant taxonomy, Angiosperms (1952) 61—62;
D. LoBREAU-CALLEN, Adansonia 17 (1978) 470—472; J.W. WALKER, Amer. J. Bot. 61 (1974)
1112—1137; Linn. Soc. Symp. Series 1 (1976) 251—308. — J. MULLER.

Phytochemistry. The chemical characters of Aristolochiaceae have been summarized and

1984] ARISTOLOCHIACEAE (Ding Hou) 63

discussed from a taxonomic point of view by HEGNAUER (1960, 1964) and a comprehensive phyto-
chemical review of the family was given by MUNAVALLI & VIEL (1969).

Members of the family tend to deposit SiO, and calcium oxalate in their tissues. Heavy silicifi-
cation of cell walls (hairs, epidermis, mesophyll) and cell lumina (silica bodies of various shapes)
is especially frequent in the tropical members of the three genera in Malesia. Calcium oxalate oc-
curs in the form of prismatic and needle-shaped crystals which are accompanied or replaced in
species of Aristolochia by druses.

All members of the family possess oil cells producing appreciable amounts of essential oil of
taxon-specific composition. These idioblasts occur in roots, rhizomes, leaves and flowers. De-
pending on taxa and chemodemes monoterpenes, sesquiterpenes or (and) phenylpropanoids are
the main constituents of these essential oils.

The nitrophenanthrenes called aristolochic acids and debilic acid and the biogenetically related
phenanthrenoid aristolactams occur practically everywhere in Aristolochia and have been traced
in species of Thottea and Asarum.

Consideration of the chemistry leads to the conclusion that the affinity of Aristolochiaceae is
closest with Annonaceae as suggested formerly by VON WETTSTEIN. The most convincing evidence
comes from the co-occurrence of heavy silification, essential oil in idioblasts and benzylisoquino-
line alkaloids and their degradation products. Both families should be included in Polycarpicae
(compare, e.g., Magnoliiflorae, DAHLGREN, 1980).

Literature: DAHLGREN, Bot. J. Linn. Soc. 80 (1980) 91—124; HEGNAUER, Pharmazie 15 (1960)
634—642; Chemotaxonomie der Pflanzen 3 (1964) 184—199, 639; MUNAVALLI & VIEL, Ann.
Pharm. Franc. 27 (1969) 449—464, 519-533, 601—614. — Editor’s extract from a large report of
R. HEGNAUER.

Chromosomes. In Aristolochiaceae, chromosome data have been reported for about 90 spe-
cies of mainly the two (large) genera, viz. Aristolochia and Asarum (s.1., incl. Heterotropa and
Hexastylis) and only one species of Apama (= Thottea).

In Aristolochia the somatic chromosomes have been reported as 2n = 8, 10, 12, 14, 24, 26, 28,
32. The number in this genus is, with some deviations, rather uniform: 2n= 14 (in most of the
tropical species) and 2n = 28 (in most of the temperate zones) (cf. GREGORY, 1956). There is one
widely distributed species, occurring also in Malesia, A. tagala, having 2n= 14; | examined the
material of this species from Celebes and New Guinea and obtained the same number. The other
numbers occur very unfrequently: 2n = 12 (or 24) four times, 2n =8, 10, 16, 32 each once, mostly
for extra-tropical species.

In the extra-Malesian genus Asarum (s./.) the chromosomes of many species have been reported
mostly with 2n = 24 (for Asiatic species) and 2n = 26. There are only a few species with 2n = 36,
40, or 48.

For the genus Thottea, there is only one species of Apama (= Thottea) from India being known
with 2n = 26 (cf. Feporov, 1969).

There is still no chromosome information known for the three monotypic genera, viz. Eugly-
pha, Holostylis and Saruma.,

Literature: DARLINGTON & Wy te, Chromosome Atlas ed. 2, 1955, 29; A.A. Feporov (ed.),
Chromosome numbers of flowering plants, 1969, 58-59; M.P. Grecory, Amer. J. Bot. 43, 1956,
110-112, tab. 1 & 2, fig. 1-154; R.J. Moore (ed.), Index to plant chromosome numbers, Regn.
Veget. 90, 1973, 162—163; ibid. 91, 1974, 31; ibid. 96, 1977, 26-27; Tanaka, Bot. Mag. Tokyo
49, 1935, 709-746, f. 1-43.

Taxonomy. Though certainly natural, the family is rather heterogeneous: small creeping or
erect herbs and large woody lianas, flowers regular or zygomorphic, stamens 6 to many, perianth
simple or double, efc. Several genera have outstanding structures: Asarum has a leathery capsule
bursting irregularly, Aristolochia has a bent, zygomorphic complicated flower, the curious South
American genus Euglypha has also a utricle but not a bent flower and besides has a fruit consisting
of 6 one-seeded cocci attached to a sort of columella, the South American genus Holostylis is like

64 FLORA MALESIANA [ser. I, vol. 10!

Aristolochia in flower, but its flower is also straight and does not possess a utricle, Thottea has
up to 4 whorls of stamens (up to 36—46), and the Chinese genus Saruma has a double perianth
and 6 halfway free follicles.

Whether the West African genus Pararistolochia can be maintained is liable to doubt. KEAy
(Fl. W. Trop. Afr. ed. 2, 1, 1, 1954, 77) distinguished it from Aristolochia by: ‘Fruit indehiscent,
elongated, strongly ribbed, cucumber-like’, but these characters seem also to occur in the New
Guinean A. dielsii ScHmipT (see p. 105). — Editor.

Affinities. In the past many suggestions have been made and there is unanimity that most char-
acters point to the assemblage of primitive families in the Dicotyledones, especially through those
of the genus Saruma. Since WAGNER’S research (Oest. Bot. Z. 57, 1907, 265—271) the general
opinion prevails that among the living plants the closest affinity is with Annonaceae in the general
Magnoliales concept.

Uses. Some American Aristolochia species are cultivated for their (rather large) beautiful
flowers as ornamentals, e.g. A. brasiliensis MART. & Zucc., A. elegans MART. & Zucc., A. gigan-
tea Mart. & Zucc., A. grandiflora Sw., A. ringens VAHL, etc.

In Malesia some indigenous Aristolochia and Thottea species are locally cultivated as food
plants for the larvae of the beautiful (swallowtail) butterflies, for commercial purposes.

Some members of the Aristolochiaceae have been used for drugs, medicine, or medicinal prod-
ucts, especially in the Far East and Southeast Asia. According to published records, such plants
or their derivatives have been applied to remedy snake bites, stomach-ache, dysentery, rheumatic
affections, colds, headache, toothache, or to reduce swellings and high blood pressure, efc. Aris-
tolochic acid has been reported possessing the capacity to reduce growth of certain types of cancer
in mice. For medicinal uses of Malesian plants see the records under the species concerned. For
further details one should consult the following literature.

Literature: Brown, Minor Prod. Philip. For. 3 (1921) 183; BurxkiL1, Dict. (1935) 188—189,
239-240, 2156-2157; CHow & Hwang, Acta Phytotax. Sinica 13 (1975) 108—109; HEyYNE, Nutt.
Pl. (1927) 596—597; Lianc, Acta Phytotax. Sinica 13 (1975) 10—28; PERRY, Medic. Pl. E. & SE.
Asia (1980) 45—48; PrerFer, Ann. Mo. Bot. Gard. 53 (1966) 121; QuisuMBING, Medic. PI. Philip.
(1951) 254—256; ScHmipT in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 225—226.

Notes for collectors. For proper identification flowers are essential. In many cases sterile
material is insufficient. Field notes should include information on flower colour and its variation
with age of the flowers. As flowers are often delicate in texture, it is advisable to insert some dry
material inside the flower before drying, e.g. wool, dry moss, or thin paper, which facilitates later
examination in the herbarium. Colour photographs and flowers in liquid are desirable.

If possible, roots should be collected; nothing is known about them in Malesian species.

In several species fruits are not yet known; attention should be given to their development to
maturity and release of seeds.

KEY, 20) THE GENERA

1. Flowers actinomorphic. Perianth narrowed at the base and gradually, slightly widened towards the apex,
always 3-lobed. Stamens 6—c. 36 (—46), in 1 or 2, rarely 3 or 4, whorls. Capsules usually siliquiform, elon-
gated, + 4-angular, 5—10 mm wide, 4-celled. Seeds oblong, ellipsoid, or broadly ovoid, in cross-section
usually 3-angular. Stems usually erect, bearing small, bract-like leaves in the lower half, then one small
leaivetollowedepyanonmal foliage leaves a: = tac ctiee ie dace 4c: oa wks oclces wveie cle s.« ote acheenene 1. Thottea

1. Flowers zygomorphic. Perianth curved, consisting of 3 parts: the inflated basal utricle, the narrowed and
neck-like tube, and the elongated, enlarged, or expanded, often 1-lipped, sometimes 3 (—6)-lobed limb. Sta-
mens 6 (10 in A. decandra), always in 1 whorl. Capsules not siliquiform, often 6-angular or -ridged, (10—)
15—40 mm wide, 6-celled. Seeds ovate, deltoid, or triangular, often flat. Stems usually scandent, also in
ELECSPECles || DEAN CAOM ya t Ol AG GREAVES waite cr te ges ach chs. cyan a She im -lSSel fun cko oud cusp oeuouaroin Ay 2. Aristolochia

1984] ARISTOLOCHIACEAE (Ding Hou) 65

1. THOTTEA

ROTTBOELL, Nye Samling Kongel. Danske Vidensk. Selsk. Skr. 2 (1783) 529;
KiotzscH, Monatsb. Akad. Berlin (1859) 588; DUCHARTRE in DC. Prod. 15, 1
(1864) 428; ScHmipTin E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 232; Dinc Hou,
Blumea 27 (1981) 303. — [A/pam RHEEDE, Hort. Malab. 6 (1686) 51, t. 28.] —
Apama LAMK, Encycl. Meth. Bot. 1 (1783) 91; Tabl. Encycl. Méth. (1823) t.
640; ScHMIDT, fide supra. — Bragantia Lour. Fl. Coch. (1790) 528; ed. Willd.
(1793) 645, non VANDELLI (1771). — Ceramium BL. Bijdr. (1826-27) 1134, non
ROTH (1797), nec ADANSON (1763). — Munnickia Bi. ex RcuB. Consp. (1828)
85. — Vanhallia SCHULT. in R. & S. Syst. 7 (1829) xviii & 166. — Trimeriza
LINDL. Bot. Reg. (1832) sub t. 1543, in note. — Asiphonia GrirrF. Trans. Linn.
Soc. 19 (1845) 333. — Lobbia PLancu. in Hook. Lond. J. Bot. 6 (1847) 144.
— Strakaea PReEsL, Epim. Bot. (1851) 221. — Fig. 1-8.

Herbs, woody at the base, or undershrubs, rarely shrubs, single or tufted,
simple or (sparsely) branched, erect, sometimes rhizomatous or scrambling.
Stems bearing small, bract-like leaves in the lower 2/3—1/2, then one smaller
leaf, followed by normal foliage leaves; (young) branches or branchlets some-
times zigzag. Leaves entire; petiole grooved above. Flowers actinomorphic,
axillary or subradical, solitary or a few on short branches, in spicate or
racemose, cymose or corymbose, or cincinnal, usually few-flowered inflores-
cences. Bract usually opposite to the flower. Flower buds (not including the
ovary) often distinctly triangular in top view. Perianth broad-campanulate, ur-
ceolate, bowl- or cup-shaped, 3-lobed; lobes valvate, caducous. Disk (?) 0, rare-
ly cupular, adnate to the perianth tube with the apical part free and ring-like
(e.g. T. tomentosa). Stamens 6—c. 36 (—46) in 1| or 2, rarely 3 or 4 whorls, free
or adnate to the style column. Ovary 4-angular, 4-celled; style (2—) 5—20-lobed,
lobes linear or linear-lanceolate. Capsules usually siliquiform, elongate, vari-
able in length, 5—10 mm wide, + 4-angular, sometimes cruciform in cross-sec-
tion, dehiscing apically towards the base, or splitting from the central part to-
wards both ends. Seeds oblong, ellipsoid, or broadly ovoid, usually 3-angular
in cross-section, rarely boat-shaped, often coated with remains of the placenta;
testa crustaceous or hard, usually (transversally) rugose, or deeply furrowed,
rarely rather smooth or sparse granular.

Distr. Indo-Malesia (c. 26 spp.): India (4 spp.), Sri Lanka (1), Bangladesh (1), Burma (3), Thailand (4),
Vietnam (2), China (Hainan, |), and Malesia (22): Sumatra, Malay Peninsula, Java, Borneo, Philippines, and
Celebes.

Ecol. Often growing sporadically, occasionally locally abundant, in shady places in tropical lowland for-
est, rarely up to c. 1200 m.

Notes. Thottea species possess a distinctive leaf architecture: the lower half or two-thirds of the stem car-
ries many (8—12) small, scale- or bract-like, alternate reduced leaves, followed by a single small leaf, which
is in turn followed by normal foliage leaves. This was observed by VAN STEENISin Hortus Bogoriense on speci-
mens of 7. borneensis and T. macrantha and found to be a constant feature in all herbarium specimens with
a complete stem.

The occurrence of these three leaf types in this sequence on a single stem has proved useful to recognize

66 FLORA MALESIANA [ser. I, vol. 10!

sterile specimens from some erect species of Aristolochia (e.g. A. philippinensis and A. sericea) which have
a similar habit, but the stems of which carry only the normal foliage leaf type.

SYNOPSIS OF SPOTTING CHARACTERS
Species are indicated by their numbers

Stem bearing 1—5 (mostly 2 or 3) leaves at the apical part: 9, 20.

Leaf with 3 prominent nerves, reaching often to the apex, connected with almost parallel and transverse veins:
19.

Leaves with lateral nerves pinnately arranged more or less at regular intervals; venation on the lower surface
prominently and closely reticulate: 11, 13.

Leaves villous or densely tomentose beneath and hairs covering almost the whole surface: 7, 8, 20 (young
leaves).

Leaf base distinctly cordate, the sinus rather narrow and the auricles or basal lobes often overlapping: 4 (p.p.),
5), @

Leaves distinctly papillate beneath; papillae forming rings or curves: 14.

Flowers with funnel-shaped perianth, up to c. 12.5 cm long, the largest in this genus: 4.

Flowers with folded perianth more or less round in outline in side view, c. 7 cm @, base cordate: 3.

Flowers with stamens arranged in 4 whorls: 1.

Flowers with stamens arranged in 3 whorls: 2.

Flowers with 6 stamens in | whorl; style lobes covered with (often hooked) hairs at the upper part: 20.

Flowers with 6 stamens in | whorl; anthers with connectives distinctly protruding 0.5—1 mm beyond them: 22.

Style lobes 2 or rarely 3, glabrous: 9.

Seeds boat-shaped, rather smooth and only sparsely granulate on both surfaces: 7.

KEY TO THE SPECIES

1. Stamens arranged in 4 whorls (shown distinctly in flower buds) or appearing scattered (in open flowers)
1. T. parviflora
1. Stamens arranged in 1 to 3 whorls.
DE Stamens anraneedunesnwhOnisieewnches cree tanto: Seach ras le ls ahve auhe nie aaa aula ene 2. T. triserialis
2. Stamens arranged in | or 2 whorls.
3. Stamens arranged in 2 whorls.
4. Perianth base prominently cordate, in side view with 2 distinct auricles. Perianth of the (mature) flower,
folded in side view, rounded in outline, c. 7 cm @; sinus 1—1.5 cm deep. (Leaves unknown)
3. T. straatmanii
4. Perianth base obtuse or rounded (in side view).
5. Leaf base distinctly cordate, the sinus rather narrow and the auricles or basal lobes often overlapping.
6: Blowers large? perianth up toc, 12cm! long’ when mature 2.- 325.2 --4-s.4¢ 4. 4. T. grandiflora
6. Flowers smaller; perianth less than 3 cm long when mature.
7. Perianth c. 23 mm long, deeply 3-lobed; lobes triangular, inner surface densely covered with papillae
and glandular hairs. Stamens with papillate filaments. Style lobes 18—20.... 5. T. macrophylla
7. Perianth 8—10 mm long, shallowly or obscurely lobed; lobes nearly semi-orbicular or triangular, in-
ner surface densely covered with (mainly hooked) hairs. Stamens with glabrous filaments. Style c.
DRG Des meme test reerets. Mkt e ne ae an tats dtl bd Beles ie a hin an die Sole Paleo ee 6. T. robusta
5. Leaf base obtuse, rounded, or cuneate.
8. Leaves with hairs densely covering the whole lower surface, so the latter usually hidden; hairs bent at
right angles near the base and parallel to the surface.
9. Hairs on the lower leaf surface appearing very thin in dry state and sticking together (resembling a
layer of gelatine). Perianth lobes with the margin not reflexed at anthesis. Seeds flattened, boat-

Shaped wrathen Smoothies cael meres titre ere crete ties eo aoc SR e aes SRE OS 7. T. curvisemen
9. Hairs on the lower leaf surface appearing thicker and free from one another. Perianth lobes with the
margin reflexed at anthesis. Seeds + ellipsoid, triangular, strongly rugose .... 8. T. borneensis

8. Leaves with hairs rather loosely or sparsely covering the lower surface, so the latter always visible;
hairs irregularly spreading, curved or twisted.
105 Blowers large; perianth up! tore: 12vemblong, when open ..2.=5....-22-o-+- +. 4. T. grandiflora
10. Flowers much smaller; perianth at most 4.5 cm long.

1984]

ARISTOLOCHIACEAE (Ding Hou) 67

11. Stamens 3 in the upper whorl, 9 in the lower. Style 2- or 3-lobed ........... 9. T. paucifida
11. Stamens 6—15 (—18) in the upper whorl, 9—15 (—24) in the lower. Style (4—) 5—19-lobed.
12. Perianth shallowly or obscurely lobed, sometimes + entire.
13. Leaf venation loosely reticulated, veins or veinlets often parallel to one another. Inflorescences
at the upper part of stem, in the axils of foliage leaves. Fruits twisted, 15—25 cm long, densely

RAINY rns. So ts Oia us Tee eras

AE Se erase con ob eae oe 10. T. tricornis

13. Leaf venation closely reticulated. Inflorescences near the base of stem, in the axils of bract-like,
reduced leaves. Fruits straight, less than 10 cm long, sparsely hairy or almost glabrous

11. T. beccarii

12. Perianth distinctly lobed, often divided to c. half or more of its length.

14. Perianth tube cylindric (c. 1.5 cm long)

WP ES ee ie AG Bo Shyais ¢. 12. T. rhizantha

14. Perianth tube campanulate, short cupular, obscure, or 0.
15. Leaves with distinctly pinnate, often rather evenly spaced nerves; venation closely reticulate

(almost tesselate), prominent beneath

SP es See as acnéan: 13. T. philippinensis

15. Leaves with | or 2 pairs of basal nerves and some lateral ones from the midrib, not evenly
spaced; venation loosely reticulate, slightly elevated beneath.
16. Leaves with the inner pair of basal nerves emerging from the base. Inflorescences at the upper
part of stem, in the axils of foliage leaves.
17. Perianth campanulate, 15—25 mm long; contracted at the lower 1/3 and then erecto-patent,
glabrous oltside: lobes trianeilardlat. «2... «0-1-1. oe eet ete 14. T. dependens
17. Perianth short-cupular, 10—13 mm long, sparsely puberulous outside; lobes reniform, the

margin reflexed at anthesis........

2 aR each Vee, Sn ee 15. T. reniloba

16. Leaves with the inner pair of basal nerves emerging near the base or a few mm from it. Inflor-
escences at the basal part of stem or near the ground, in the axils of bracts or reduced leaves

(almost cauligerous).

18. Perianth c. 15 mm long, lobed to c. 2/3 of the length............. 16. T. pennilobata
18. Perianth 25—45 mm long, lobed up to c. half of the length.
19. Perianth lobes semi-orbicular, 10 by 15 mm, at apex subrounded or slightly apiculate

19. Perianth lobes triangular, 20—25 by c. 20 mm, at apex acuminate ...

3. Stamens arranged in | whorl.

17. T. celebica
18. T. muluensis

20. Leaves with the inner pair of basal nerves extending to the apex, joined by predominantly rather close,

transverse, parallel veins. Inflorescences corymbose or paniculate............. 19. T. corymbosa

20. Leaves with the inner basal pair of nerves extending usually to c. half, rarely more, of the length,
joined by rather loose, transverse and reticulate veins. Inflorescences usually spicate or racemose.

21. Plant bearing 1—5 (often 2 or 3) foliage leaves. Leaves densely tomentose or villous beneath especially

when young. Inflorescences near the base of stem. Style lobes densely covered with (hooked) hairs at

RIETHENCAU DAIL: o.214 caiiae. 0 # CS ee ae > eel

jo a:b -a-e dud REPS GE RER ORE aE TS aon eres 20. T. tomentosa

21. Plant bearing many foliage leaves. Leaves pubescent or puberulous beneath. Inflorescences at the

upper part of stem. Style lobes glabrous.

22. Perianth lobes broadly rounded, c. 12 by 4mm, emarginate. Stamens 9—12; connective not protrud-

Iie Beyoud the AutMers 6 shee western emis was

ies s\ ccs e-wrone 0:0. el so wee 21. T. macrantha

22. Perianth lobes ovate to lanceolate, 8-15 by 5—8 mm, apex acute, acuminate, or obtuse. Stamens 6;
connective protruding c. 0.5 mm beyond the anthers .......... 0000000 eeeee 22. T. sumatrana

1. Thottea parviflora Riptey, J. Str. Br. R. As. Soc.
n. 57 (1911; nec 1910) 89; Fl. Mal. Pen. 3 (1924) 17;
Scumiptin E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
232; Burk. Dict. 2 (1935) 2157; Dinc Hou, Blumea
27 (1981) 305, f. 38-40, 70D. — Fig. 7n—p.

Erect shrub, up to 2 m high. Branches subterete or
obscurely angular, c. 7 mm @, pubescent. Leaves
chartaceous, ovate, obovate, broad-elliptic, elliptic,
or lanceolate, 10—26 by 4.5—9 cm; apex acuminate;
base rounded, cuneate, or attenuate; sparsely puber-
ulous or almost glabrous above, puberulous beneath;
basal nerves 2 or 3 pairs, emerging 2—5 mm from the

midrib above the base, similar to the lateral nerves,
ascending upward to 2/3 or more of the blade, the
outer | or 2 pairs much weaker and shorter; lateral
nerves 6—9 pairs, elevated beneath, flat but distinct,
or slightly elevated above; veins + parallel and scala-
riform, sometimes + transverse at the basal part be-
tween the inner pair of basal nerves, connected with
parallel or reticulate veinlets, elevated beneath, faint
above; petiole 3—7 mm, puberulous. /nflorescences
axillary, often in the axils of leaf scars, usually simple
and spiciform, up to 1.5 cm long; bracts lanceolate,
1—4 mm long, puberulous on both surfaces. Pedicel

68 FLORA MALESIANA [ser. I, vol. 10!

Fig. 3. Thottea grandiflora RottTs., x 1/3. Possesses the largest flowers in the genus. Singapore (Photogr.
CORNER).

1984]

and ovary very short, c. 6mm long, densely puberul-
ous. Perianth white, whitish green with a pink basal
patch inside, pale pink or pink, pale purplish or vio-
let, discoid, 2-4 mm long, c. 6 mm g, with longitu-
dinal and loosely reticulate veins, loosely puberulous
outside and papillate inside, glabrescent; lobes
semi-orbicular, 1.5—3 by 3.5—4.5 mm. Stamens
(15—) 20—22, in 4 whorls (can easily be observed in
young buds); filaments 0.4—1 mm, short-hairy; an-
thers oblong, c. 0.7 mm long. Sty/e almost branched
from the base, c. 1.7 mm, lobes usually 4 or 5, glab-
rous. Capsules slender, up to 9 cm long, acute or
pointed at both ends, obscurely 4-angular, slightly
twisted, loosely puberulous. Seeds ellipsoid, 3—3.5
by 1.5 mm, triangular, irregularly and transversely
corrugated.

Distr. Peninsular Thailand; in Malesia: Malay
Peninsula (Kedah, also Langkawi, Perak, Kelantan,
Pahang, Selangor).

Ecol. In lowland forest, occasionally in swampy
forest, up to 150 m; in Thailand occasionally found
also on granitic rock in the forest, up to 700—1055 m.
Fl. March—July, fr. March—August.

Uses. Rootstock is eaten with rice for remedy of
coughs.

Vern. Chudok, Pahang.

2. Thottea triserialis DinG Hou, Blumea 27 (1981)
330, f. 35, 41, 42.

Shrublet of 120 cm high. Branches terete, c. 5mm
@, pubescent. Leaves chartaceous, elliptic, broad-el-
liptic, slightly obovate, or ovate, 23—34 by 11-23
cm; apex acuminate or acute; base rounded or obtuse
in outline but cordate (sinus narrow, 0.5—0.75 cm
deep, auricles overlapping or touching each other);
glabrous above, sparsely pubescent beneath; basal
nerves 2—3 pairs, starting almost from the insertion
of the petiole, the inner one ascending upward to
1/2—2/3 of the blade, similar to the lateral ones, the
outer | or 2 short and weak; lateral nerves 7 or 8
pairs, prominently elevated below, often flat but dis-
tinct above; veins scalariform, connected with loose-
ly reticulate or straight veinlets, elevated beneath,
obscure above; petiole very short or obscure, some-
times up to c. 5 mm, pubescent. /nflorescences in the
axils of foliage leaves or their scars, spiciform, soli-
tary or fasciculate, 3—5 cm long, pubescent; bracts
lanceolate, elliptic, or oblanceolate, 3-15 mm long,
pubescent. Pedicel and ovary 18—20 mm long, dense-
ly pubescent. Perianth pink or pinkish brown, cupu-
lar, 10-15 mm long, slightly contracted at the lower
1/3—1/2, almost orbicular in outline when open
(10—15 mm @), veins loosely reticulate, sparsely pu-
berulous on both surfaces; tube short, terete, c. 5mm
long; lobes semi-orbicular, 5—10 by 11-15 mm, apex
acute or cuspidate. Stamens in 3 whorls: upper row
5—8, middle 7—12, lower 12—16; filaments hairy,

ARISTOLOCHIACEAE (Ding Hou) 69

0.5—1 mm; anthers oblong, c. 1.25 mm long. Style
column c. 2 mm long, lobes 11—20, 1.5—2 mm, glab-
rous. Capsule (very young) siliquiform, 12 cm long,
pubescent. Seeds flat (?).

Distr. Malesia: Borneo (Sarawak: Lundu Distr.;
G. Pueh). Twice collected.

Ecol. In primary lowland dipterocarp forest, on
gentle ridge slope, 600—1080 m. Fi. fr. April.

Note. Vegetatively not distinguishable from 7.
macrophylla and T. robusta. The inner surface of the
perianth is glabrous; it is densely hairy in the other
two species (glandular hairs in the former, mainly
hooked-hairy ones in the latter). The stamens are ar-
ranged in 3 whorls, a unique character in the genus.

3. Thottea straatmanii DinGc Hou, Blumea 28 (1983)
352)'f.-6:

Plant c. 2.5 m high. Leaves not preserved, (from
a sketch) ovate-oblong, c. 30 cm long. Inflorescence
cauligerous, spiciform, 2 cm long, puberulous, inter-
nodes 7—10 mm long; bracts leafy, ovate, 4.5—13 by
2.5—8 mm, puberulous on both surfaces. Pedicel and
ovary c. 17 mm long, puberulous. Perianth when
folded in side view + orbicular in outline, c. 7 cm 9,
cordate at base, sinus c. 12 mm deep, distinctly bi-
auriculate; perianth deeply lobed, lobes + orbicular,
c. 7 cm @, puberulous outside, loosely glandular
hairy inside, veins rather loosely reticulate. Stamens
in 2 whorls: upper row c. 18, lower c. 24; filaments
glabrous, very short or 0; anthers oblong, 1.5—2 mm
long. Style column c. 3 mm long, lobes c. 12, c. 1.5
mm. Capsule siliquiform, slightly curved, 15—21 cm
long, slightly 4-angular, narrowed at both ends, pu-
berulous. Seeds broad-ellipsoid, triangular, 3.5 by 2
mm, transverse-rugose, deeply grooved.

Distr. Malesia: NE. Sumatra (East Coast: Laut
Tador).

Ecol. Growing very locally in wet shady places in
open forest, at c. 100 m.

Notes. In flower size the second largest in the ge-
nus (perianth c. 7 cm in diam.), next to 7. grandiflo-
ra (12.5 cm).

Closely related to 7. reniloba with which it shares
the spaced bracts and distinct internodes, the deeply
lobed perianth, 2-whorled stamens, pubescent cap-
sules and deeply grooved seeds, but different by lar-
ger, + orbicular perianth lobes, the higher number
of stamens (upper whorl c. 18, lower c. 24) and the
transverse-rugose seeds.

The forest of the type locality is now destroyed.

4. Thottea grandiflora Rorrsoett, Nye Samling
Kongel. Danske Vidensk. Selsk. Skr. 2 (1783) 529, t.
2; BENN. & Brown, PI. Jav. Rar. | (1838) 45; Grier.
Trans. Linn. Soc. 19 (1845) 325, t. 36; Ann. Sc. Nat.
Bot. 7 (1847) 328; Notul. 4 (1854) 346; Ic. Pl. Asiat.
(1854) t. 530 & 531; Mig. Fl. Ind. Bat. 1, 1 (1858)

70 FLORA MALESIANA [ser. I, vol. 10!

\\

Ae 77

>

Se 3 Se SS ae

ara:

4

\
|

1

. 1
\N
\
i

|

I

!
etl
N
\

!

oe

Fig. 4. Thottea robusta STEEN. a. Habit, nat. size, b. young inflorescence, c. open flower, both x 2, d. gyno-
stemium, x7, e. dehisced fruit, nat. size (VAN STEENIS 1270).

1984]

ARISTOLOCHIACEAE (Ding Hou) 71

1068; KLotzscu, Monatsb. Akad. Berl. (1859) 5 & 9,
t. 1 f. 3; DUCHARTRE in DC. Prod. 15, 1 (1864) 428;
Hook. f. Fl. Br. India 5 (1886) 74; SOLEREDER, Bot.
Jahrb. 10 (1889) 429 & 478; in E. & P. Nat. Pfl. Fam.
3, 1 (1889) 272; Ript. J. Str. Br. R. As. Soc. n. 33
(1900) 127; Kinc& Gams te, J. As. Soc. Beng. 75, ii
(1912) 27; Rip. Fl. Mal. Pen. 3 (1924) 16; HEYNE,
Nutt. Pl. (1927) 596; Burk. Dict. 2 (1935) 2156;
ScHmipT in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
232; HEND. Mal. Wild Fl. (1951) 424, f. 383; DinG
Hou, Blumea 27 (1981) 308, f. 8—10, 317, f. 62. —
Fig. 3.

Erect shrub, up to 2 m high. Branches terete, c. 1
cm @, villous. Leaves coriaceous, obovate, elliptic,
ovate-oblong, or lanceolate, (1S—) 20—30 (—45) by
9—10 (—25) cm; apex acute, short-acuminate, some-
times cuspidate; base obtuse, sometimes subcordate,
rarely cuneate; villous, glabrescent or almost glab-
rous above, hispid-pubescent beneath; basal nerves 2
or 3 pairs, the inner one obliquely extending upward
to c. half the blade, the outer one weaker and shor-
ter, running along the margin; lateral nerves 10—12
pairs; all nerves prominent below, slightly elevated
above; veins + parallel or reticulate, elevated below,
distinct above; petiole 0.5—1.5 cm, villous. /nflores-
cences usually at the lower part of the stem in the ax-
ils of (fallen) leaves, simple or sparsely branched,
sometimes branched near the base and seemingly fas-
cicled, spiciform or racemiform, 1—7 cm long, vil-
lous; bracts lanceolate or elliptic, 1—3 cm long, vil-
lous on both surfaces. Pedicel and ovary up to 4 cm
long, villous. Perianth deep claret-coloured and pur-
ple mottled, funnel-shaped, up toc. 12.5 cm long and
as broad at the mouth (largest flower in this genus),
with distinct and reticulate veins; pubescent without
especially on the venation, puberulous inside, usually
glabrescent; tube about half the length of the peri-
anth; lobes triangular or suborbicular, 5—6 by 6—7
cm, acute or rounded at the apex. Stamens in 2
whorls: upper row c. 15 (—18), lower c. 15 (—24); fila-
ments glabrous, very short; anthers oblong, c.
1.5—2 mm long. Sty/e column short; lobes 8—19, c.
2.5 mm. Capsules slender, 10—15 cm long, straight
or twisted, 4-angular, pubescent. Seeds ellipsoid,
3—4 mm long, trigonous, acute at both ends, rugose-
tubercled.

Distr. Peninsular Burma (Moulmein); in Male-
sia: Malay Peninsula (Perak, Trengganu, Pahang,
Negri Sembilan, Malacca, Johore, Singapore).

Ecol. In lowland forest, up to 600 m. Fi. fr. al-
most all year round.

Vern. Grobo, Malacca; sel-wohl, Pahang.

5. Thottea macrophylla Becc. Nuov. Giorn. Bot.
Ital. 2 (1870) 5, t. 1: f. 1-6; Sreen. Bull. Jard. Bot.
Btzg II, 12 (1932) 205; Scumiptin E. & P. Nat. Pfl.
Fam. ed. 2, 16b (1935) 232; Dino Hou, Blumea 27

(1981) 312, f. 34, 330.

Erect shrub. Branches terete, c. 5 mm @, pubes-
cent. Leaves coriaceous, elliptic-obovate, or obo-
vate, 23—35 by 14—20 cm; apex acuminate; base ob-
tuse or rounded in outline, shallow-cordate, sinus
narrow, with auricles slightly overlapping each
other; glabrous above, pubescent beneath; basal
nerves 3 pairs, the inner pair similar to lateral ones,
extending upward to c. 2/3 of the blade, outer 2
much weaker and shorter, close to the margin; lateral
nerves c. 7 pairs, prominent below, distinct or slight-
ly elevated above; veins crossbar-like, parallel, scala-
riform, veinlets transverse or reticulate, elevated and
prominent beneath, distinct or obscure above; peti-
ole 0.5—1 cm long, pubescent. /nflorescences in the
axils of foliage leaves, solitary or sometimes 2, spici-
form, c. 1.5 cm long, densely pubescent; bracts el-
liptic, obovate or oblanceolate, 3.5—9 by 1.5—3 mm,
densely puberulous on both surfaces. Pedicel and
ovary 17—20 mm long, densely puberulous. Perianth
campanulate, c. 23 mm long, up toc. 45 mm g, deep-
ly 3-lobed, outer surface with distinctly pubescent re-
ticulations, less hairy between the veins, inner sur-
face densely covered with papillae and glandular
hairs (appearing carpet-like); tube very short; lobes
triangular, c. 18 by 24 mm, short-acuminate. Sta-
mens in 2 whorls: upper row 10 or 11, lower 16 or 17;
filaments papillate, 0.5—1 mm; anthers oblong, c.
1.5 mm long. Style column c. 3 mm long; lobes
18—20, c. 3 mm long, glabrous. Capsules unknown.

Distr. Malesia: Borneo (Sarawak: Mt Matang).
Twice collected.

Ecol. In forest, c. 750 m. Fi. April & July.

6. Thottea robusta STEEN. Bull. Jard. Bot. Btzg III,
12 (1932) 20S, f. 11; ScHmipT in E. & P. Nat. Pf.
Fam. ed. 2, 16b (1935) 232; Dinc Hou, Blumea 27
(1981) 330, f. 36, 37, 57-60. — Fig. 4.

Erect shrub, 3—4 m high. Branches terete, c. 1 cm
@, villous on young parts, glabrescent. Leaves char-
taceous, variable in shape and size, obovate, sub-
rhomboidal, ovate-oblong to oblong-lanceolate,
17.5—40 by 7.5—25 cm; apex acute or acuminate;
base broadly rounded in outline and distinctly cor-
date, the auricles overlapping each other; glabrous
above, pubescent or villous beneath, especially on
the midrib and venation; basal nerves 2 or 3 pairs,
emerging flabellately from the base, the inner one as-
cending upward to + halfway, similar to the lateral
nerves in thickness and appearance, the outer | or 2
much weaker and shorter and close to the margin;
lateral nerves 6—8 pairs, prominent beneath, distinct
or slightly elevated above; veins transverse, parallel
and scalariform, connected with crossbar-like or reti-
culate veinlets, elevated below, distinct or obscure
above; petiole stout, 0.5—1.5 cm, densely pubescent.
Inflorescence in the axils of foliage leaves, solitary or

2 FLORA MALESIANA [ser. I, vol. 10!

Fig. 5. Thottea paucifida Dinc Hou. a. Habit, x 2, b. inflorescence, x 2, c—d. flowers, x 442, e—g. gynoste-
mia, with 2- or 3-fid style, x 9 (BRooKE 10009). Courtesy of Blumea.

1984]

ARISTOLOCHIACEAE (Ding Hou) 73

in fascicles of 2 or 3, sometimes sparsely shortly
branched, spiciform, 0.5—3.5 cm long, densely vil-
lous; bracts ovate to spathulate, crowded, 5—9 mm
long, densely pubescent or villous. Pedicel and ovary
short, c. 5 mm, densely pubescent or villous. Peri-
anth pale wine-red, shallowly broad-campanulate,
8—10 mm long, c. 17 mm 9; outer surface with
strongly prominent, villous reticulations, less hairy
between the veins; inner surface densely covered with
short (mainly hooked) hairs; tube 3—5 mm long,
shallowly or obscurely lobed, lobes nearly semi-orbi-
cular or triangular, c. 5 by 10 mm. Stamens in 2
whorls: upper row 7 or 8, lower c. 14; filaments gla-
brous, 0—0.7 mm; anthers oblong, 0.7—1 mm long.
Style column short, c. 1.5 mm long, lobes c. 12, gla-
brous. Capsules narrow spindle-shaped, straight or
falcate, not or only slightly twisted at the top, 8—11.5
cm long, villous. Seeds ovoid, c. 4 by 2 mn, trigo-
nous, tuberculate.

Distr. Malesia: Natuna Islands (NW off Borneo)
(Bunguran: E. slope of G. Ranai); once collected.

Ecol. Primary forest, along stream, c. 250 m. Fi.
fr. April.

Vern. Kaju ribal, M.

7. Thottea curvisemen DinG Hou, Blumea 27 (1981)
320, f. 32, 33, 64-66.

Erect shrub, 1—1.30 m high. Branches subterete,
4—6 mm @, pubescent. Leaves chartaceous, broad-
elliptic or elliptic, 18.5—26 by 9—18 cm; apex acumi-
nate; base rotund; glabrous above, sericeous be-
neath; basal nerves 2 (rarely 3) pairs, the inner one
ascending obliquely upward to halfway, the outer |
or 2 weak and short, close to the margin; lateral
nerves 6—9 pairs; veins distinctly crossbar-like;
nerves and veins prominent beneath, visible rarely
distinct above; petiole c. 1 cm, pubescent. /nflores-
cences axillary, in axils of bracts or reduced leaves,
spiciform, c. 3 cm long, puberulous; bracts elliptic or
slightly obovate, 2.5—3 mm long, both surfaces pu-
berulous. Pedicel and ovary 7—12 mm long, puberu-
lous. Perianth bright purple, short-cupular, c. 12.5
mm long, with distinct, longitudinal veins, sparsely
puberulous outside, densely glandular hairy inside;
tube short-cupular, c. 9.5 mm long; lobes arcuate, c.
3 by 10 mm, obscurely cuspidate at the apex. Sta-
mens in 2 whorls: upper one 8—10, lower 11—14; fila-
ments glabrous, 1—1.5 mm; anthers oblong, 1—1.5
mm long. Style column c. 2 mm long; lobes 6~12, ra-
diate, c. 1.5 mm long. Capsule (only one seen) nar-
row fusiform, 4.5 by 0.6. cm, straight, distinctly 4-an-
gular, almost glabrous. Seeds flattened, boat-
shaped, broad-ellipsoid in side view, c. 2—2.5 mm
long, + obtuse or truncate at both ends (depending
on the position in the fruit), rather smooth with only
sparse granules on both surfaces.

Distr. Malesia: Borneo: Sarawak (Kapit: Bukit

Raya, Pelagus), once collected.

Ecol. In lowland dipterocarp forest, 240 m, on
slopes of steep ridges. Fi. fr. August.

Note. Allied to 7. borneensis.

8. Thottea borneensis VALET. Ic. Bog. (1908) t. 261;
ScumipT in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
232, f. 120P; Dinc Hou, Blumea 27 (1981) 321, f.
29-31.

Erect shrub up to 2 m high. Branches terete, up to
2cm 9, glabrous. Leaves chartaceous, elliptic, ovate-
oblong, or obovate, 17—27 (—30) by 6.5—14 (—19)
cm; apex acute, acuminate, sometimes cuspidate;
base obtuse or cuneate; glabrous above, densely seri-
ceous beneath; basal nerves 2 pairs, the inner one as
prominent as the lateral ones, extending upward to
2/3 or more of the blade, the outer one very weak
and short; lateral nerves 5—8 pairs, prominent be-
neath, distinct above; veins and veinlets transverse or
slightly curved, + scalariform, or loosely reticulate,
slightly elevated below, visible or obscure above; pet-
iole 1—1.5 cm, slightly pubescent. /nflorescences ax-
illary, simple or sparsely branched near base, 4—7 cm
long, with one or a few flowers at the apical part, pu-
bescent; bracts many, ovate or lanceolate, 1-3 mm
long, densely puberulous outside, sparsely puberu-
lous or glabrous inside. Pedicel and ovary 7—12 mm
long, densely puberulous. Perianth cupular, 8—12
mm long, c. 15 mm, dark purple; densely pubescent
outside, sparsely puberulous inside; tube 4-6 mm
long; lobes semi-orbicular or broadly ovate, 4.5—6
by 4.5—9 mm, erect, at anthesis the marginal part re-
flexed and the base biauriculate. Stamens in 2
whorls: upper row 10—13, lower 14—17; filaments
glabrous, c. 0.5 mm; anthers oblong, c. 1 mm long.
Style column obscure; lobes 9, radiately ascending,
c. 1 mm long. Capsules siliquiform, pendulous,
twisted, 9 cm long. Seeds + ellipsoid, c. 5 by 2mm,
strongly rugulose.

Distr. Malesia: Sumatra (West Coast: Padang)
and Borneo (Landak; Kapuas: Mt Biang). Cultivated
in the Hort. Bogor., from plants collected by TEys-
MANNin Borneo, under the numbers XI-B-XIII. 76 &
134.

Ecol. Fil. Sept.—Nov. No other field ecological
data recorded.

Note. See the note under 7. reniloba.

9. Thottea paucifida Dino Hou, Blumea 27 (1981)
324, f. 23, 24, 71. — Fig. 5.

Undershrub, creeping below and rooting, then as-
cending, up to 30 cm high. Branches subterete, c. 3
mm @, puberulous, glabrescent. Leaves chartaceous,
4.5—8 by 2.5—4.5 cm; apex acute or obtuse; base ro-
tund or obscurely cordate; sparsely pubescent on
both surfaces; basal nerves | or 2 pairs, usually thin-
ner than the lateral ones, ascending upward to about

74 FLORA MALESIANA

[ser. I, vol. 10!

halfway; lateral nerves 4—6 pairs, slightly elevated
below, distinct or faint above; veins crossbar-like
and loosely reticulate, distinct or faint below, ob-
scure above; veinlets obscure; petiole 2—3 mm, pu-
bescent. /nflorescences axillary, in the axils of fallen
leaves near the basal part of the stem, spiciform, sim-
ple or once branched, 1—3 cm long, puberulous;
bracts linear, 2—5 mm long, densely puberulous on
both surfaces. Pedicel and ovary 4.5—5 mm long,
densely puberulous. Perianth cream colour, cupular,
6 mm long, with loose reticulations; tube 3 mm long,
densely papillate inside; lobes subrotund or triangu-
lar, 3 by 3.5 mm, sparsely puberulous inside, apicu-
late at the apex. Stamens in 2 whorls: upper row 3,
lower 6; filaments glabrous, 0—1 mm; anthers ob-
long, c. 0.5 mm long. Sty/e column 1.25 mm long;
lobes 2 or 3, c. 0.5 mm. Capsules unknown.

Distr. Malesia: Borneo (Sarawak: Div. 5, La-
was); once collected.

Ecol. Onthe banks of a stream through stands of
rubber and other trees. F/. May.

Note. Allied to 7. tomentosa, but leaves lax-pu-
bescent underneath, perianth without annular ridge
in the apical part of the tube, 9 stamens in 2 whorls,
and 2 (or 3) glabrous style lobes.

10. Thottea tricornis MatNGay ex Hook. /f. Fl. Br.
India 5 (1886) 74; SoLEREDER, Bot. Jahrb. 10 (1889)
430 & 506; KING & GamBLE, J. As. Soc. Beng. 75, ii
(1912) 29; Ripi. Fl. Mal. Pen. 3 (1924) 16; ScumiptT
inE. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 232; DinG
Hou, Blumea 27 (1981) 318, f. 2, 17, 18, 43, 44, 61,
70B. — Fig. 7f-i.

Erect shrub, up to 2 m high. Branches terete or
slightly angular, c. 1 cm @, pubescent. Leaves charta-
ceous or subcoriaceous, elliptic, lanceolate, obovate-
oblong, or oblanceolate, 20—30 by 8.5—16 cm; gla-
brous above, pubescent beneath; apex acute or acu-
minate; base cuneate or rounded; basal nerves 2
pairs, emerging from the base, the inner one ascend-
ing upward reaching to c. 2/3 or higher, similar to
the lateral ones, the outer one weak and short; lateral
nerves 6—8 pairs, prominently elevated beneath,
slightly elevated above; veins transverse, scalari-
form, connected by crossbar-like or reticulate vein-
lets, elevated beneath, often faint above; petiole
5—17 mm, pubescent. /nflorescences in the axils of
foliage leaves, racemi- or paniculiform, up to 5 cm
long, pubescent; bracts ovate, lanceolate, or linear,
3—6 mm long, puberulous. Pedicel and ovary 1.25—2
cm, densely pubescent or velvety. Perianth magenta
or violet, campanulate, slightly contracted at the
lower part, 1.5—2 cm long, suborbicular or six-angu-
lar in outline, 2.5—3.5 cm g, densely pubescent out-
side, inner surface densely covered with glandular
hairs and appearing mat-like; obscurely lobed, lobes
triangular, 5—10 by 16—30 mm, apex acute or mu-

cronate. Stamens in 2 whorls, upper row 6—10, lower
10—14; filaments glabrous, c. 1 mm; anthers oblong,
c. 1 mm long. Style column c. 2 mm long; lobes
5—13, c. 1.5 mm, glabrous. Capsules slender, rather
long, 15—25 cm long, slightly curved or twisted,
4-angular, densely pubescent or velvety. Seeds ob-
long, 4—5 by 2 mm, trigonous, coarsely granulate.

Distr. Peninsular Thailand (Chawng), and Male-
sia: Malay Peninsula (Perak, Pahang, Selangor, Ma-
lacca).

Ecol. Undergrowth in forest, 300—600 m. FY.
Feb.—May, fr. Feb.—Aug.

Vern. Melada, Selangor; telinga kelawar, Pa-
hang.

11. Thottea beccarii Dinc Hou, Blumea 27 (1981)
SHIBIs te AB, A5, Sil, S77.

Erect shrublet, 75—120 cm high. Branches + te-
rete, 0.5—0.7 cm g, lower part rather straight, upper
part slightly zigzag, slightly pubescent, glabrescent.
Leaves firmly chartaceous, elliptic, rarely lanceolate
or oblanceolate, 20—35 by 7.5—15 cm; apex short-
acuminate; base cuneate; sparsely pubescent above
when young, glabrescent, often almost glabrous
when old, pubescent beneath; nerves 6—9 pairs, usu-
ally pinnate, usually the basal pair for 0.5—0.7 cm
united with the midrib, ascending up to 1/2—2/3 of
the blade, sometimes one weak, short pair starting
from the very base and extending along the margin;
veins and veinlets closely reticulate, or + crossbar-
like; both nerves and veins prominent below, obscure
above; petiole very short, 5—7 mm, slightly pubes-
cent. Inflorescences near the basal part of stem, | or
2 in an axil of ascale-like leaf, c. 5cm long, spiciform,
slightly puberulous; bracts lanceolate, linear, or ob-
lanceolate, 5—6 mm long, rarely 2-lobed, sparsely pu-
berulous on both surfaces. Pedicel and ovary 15—17.5
mm long, sparsely puberulous. Perianth cupular,
12—17.5 mm long, 20—25 mm g, obscurely lobed;
tubec. 10 mm long; lobes arcuate, 3—5 by 1S—20 mm,
slightly acute at the apex. Stamens in 2 whorls: upper
row 7—10, lower 13 or 14; filaments hairy (?) or gla-
brous, 0.3—0.6 mm; anthers oblong, c. 1 mm long.
Style column c. 2 mm long; lobes c. 10, erect, c. 1 mm
long. Capsules siliquiform, 6.5—9 cm _ long,
4-angular, straight, narrowed and pointed at both
ends, sparsely hairy or almost glabrous. Seeds broad-
ellipsoid or subglobose, obscurely triangular on cross-
section, 2 by 1.5—1.3 mm, prominently rugose, with
transverse bars and tubercles, deeply furrowed.

Distr. Malesia: Sumatra (Padang and Asahan), 4
collections.

Ecol. At c. 360 m. Fi. fr. August.

Vern. Kaju pinggu batu, Asahan.

Notes. Allied to 7. borneensis but differing by
the closely reticulate venation and scattered hairs; in
T. borneensis the venation is scalariform or loosely

1984]

ARISTOLOCHIACEAE (Ding Hou) 75

reticulate and hairs are densely matted beneath.

Also allied to T. tricornis by the closely reticulate
venation and inflorescences near the base of the
stem, in the axils of bract-like reduced leaves, and
further by the fruit and seed.

12. Thottea rhizantha Becc. Nuov. Giorn. Bot. Ital.
2 (1870) 6, t. 1: 7-10; ScHmipTin E. & P. Nat. Pf.
Fam. ed. 2, 16b (1935) 232, f. 120K; Dinc Hou, Blu-
mea 27 (1981) 311, f. 45—47.

Erect shrub or treelet, up to c. 1.25 m high.
Branches terete, c. 1 cm 9, densely tomentose or vil-
lous, sometimes glabrescent. Leaves subcoriaceous,
elliptic, oblanceolate, or oblong, 21—42.5 by
9.5—16.5 cm, apex acuminate; base cuneate or ob-
tuse; glabrous above, tomentose beneath especially
on the midrib, nerves and veins; basal nerves 2 pairs,
emerging from the base, the inner one extending up-
ward to about halfway, similar in thickness and ap-
pearance to the lateral ones, occasionally with some
secondary nerves, Outer one very weak, short and
close to the margin; lateral nerves 8—10 pairs, elevat-
ed and prominent beneath, distinct or flat above;
veins usually transverse and scalariform, connected
with crossbar-like or loosely reticulate veinlets, ele-
vated beneath, often faint or invisible above; the pet-
iole c. 5 mm, densely pubescent. /nflorescences near
the base of the stem, in the axils of bracts, few-
branched, paniculiform or racemiform, internodes
spacious, villous; bracts ovate, c. 10 mm long, vil-
lous. Pedicel and ovary 10—15 mm, pubescent. Peri-
anth outside faintly violet tinged, inside violet at the
base, white at the top, or red with white, funnel-
shaped, 3—3.5 cm long, c. 3.5 cm @, with distinct
longitudinal and loosely reticulate veins, pubescent
outside, glandularly hairy inside; tube cylindric, c.
1.5 cm long; lobes suborbicular, 1.5—2 by 1.2—2 cm,
apical part rounded, acute, or rarely apiculate. Sta-
mens in 2 whorls: upper row 6—8, lower 13-15; fila-
ments glabrous, 0—0.5 mm; anthers oblong, c. 1.5
mm long. Style column 2.5 mm long, lobes 5—7, c.
1.5 mm, glabrous. Capsules unknown.

Distr. Malesia: Sumatra (Djambi: Sg. Lesing
near Pauh); Borneo (Sarawak: Bellaga near Bintulu;
Kapit Distr.).

Ecol. In primary hill forest on sandstone substra-
tum, mixed dipterocarp forest on ridge, or on ridge
in old secondary forest, up to 500 m.

Uses. Roots boiled in water is taken to cure go-
norrhoea in Sarawak.

Vern. Sumatra: mai-mai, Sg. Lesing; Sarawak:
keh, Punan lang.

13. Thottea philippinensis Quis. Philip. J. Sc. 41
(1930) 322, t. 2; Dinc Hou, Blumea 27 (1981) 306, f.
1; ibid. 29 (1983) 242. — Fig. 8a.

Erect undershrub up to c. 70 cm high. Branches

subterete, c. 0.5 cm 9, pubescent. Leaves thin-coria-
ceous, lanceolate to narrow-lanceolate, oblong-ellip-
tic or elliptic-lanceolate, 16—26 (—39) by 4—9 (—13)
em; apex acuminate; base rounded or cuneate; gla-
brous above, pubescent beneath; nerves pinnate, ba-
sal pair weak and short, close to the margin, up to
1/4—1/3 of the blade; lateral nerves 8—13 pairs, ele-
vated and prominent beneath, slender above; veins
and veinlets closely reticulate, prominent beneath,
rather faint above; petiole c. 5mm, pubescent. /nflo-
rescences at the basal part of the stem, in the axils of
reduced leaves, spiciform, up to 5 cm long, pubes-
cent. Bracts elliptic, 2—5 mm long, puberulous on
both surfaces. Pedicel and ovary 8—13 mm, densely
puberulous. Perianth light bluish purple, blue and
pink, or dark red outside and whitish inside, cam-
panulate, 16—22 mm long, 10—20 mm 9, with loosely
reticulate veins, puberulous outside, glandular hairy
inside; tube 10—15 mm long; lobes triangular or
semi-orbicular, 6—7 by 11—16 mm, apex acute or
apiculate (the apical part usually incurved and the
apex seemingly obtuse). Stamens in 2 whorls: upper
row 8—10, lower 12—14; filaments glabrous, 1—1.5
mm; anthers oblong, 1.3 mm long. St#y/e column 5—7
mm; lobes 4—6, glabrous. Capsule fusiform (only
open, empty valves seen), c. 3 cm long, pubescent.
Seeds not seen.

Distr. Malesia: Philippines (Mindanao: Lanao
Prov.) and Borneo: Sarawak (4th Div., Lambir Nat.
Park). Twice collected.

Ecol. In dipterocarp forest and on sheltered
sandstone cliff, 150—450 m. Fi. fr. March, Sept.

Vern. Taguibunon, Lanao.

Note. See the note under 7. celebica.

14. Thottea dependens (PLANCH.) KLorzscH, Mo-
natsbl. Akad. Berl. (1859) 589; DucHARTRE in DC.
Prod. 15, 1 (1864) 428; Hook. f. Fl. Br. India 5
(1886) 74; SOLEREDER, Bot. Jahrb. 10 (1889) 429; in
E. & P. Nat. Pfl. Fam. 3, 1 (1889) 272; Riptey, J.
Str. Br. As. Soc. n. 33 (1900) 127; Kinc & GAMBLE,
J. As. Soc. Beng. 75, ii (1912) 28; GresHorr, Meded.
Lands Pl. Tuin 29 (1930) 132; Burk. & HAniFr,
Gard. Bull. S. S. 6 (1930) 240; ScumipT in E. & P.
Nat. Pfl. Fam. ed. 2, 16b (1935) 232, f. 120 L—O;
Burk. Dict. 2 (1935) 2156; Dinc Hou, Blumea 27
(1981) 311, f. 5-7. — Lobbia dependens PLANCH. in
Hook. Lond. J. Bot. 6 (1847) 144, t. 3; Mig. Fl. Ind.
Bat. 1, 1 (1858) 1068. — Fig. 1b.

Erect shrub, up to 2.5 m high. Branches subterete,
c. 0.5 cm @, glabrous. Leaves chartaceous, elliptic,
obovate, rarely ovate, 12—29 by 5—15 cm; apex acu-
minate; base cuneate or acute; glabrous above,
sparsely puberulous or pubescent, sometimes seem-
ingly glabrous beneath; basal nerves 2 pairs, the in-
ner one similar to lateral nerves, extending upward to
about halfway, the outer one weak, short, close to

76 FLORA MALESIANA [ser. I, vole!

Nea
ND

uD

Fig. 6. Thottea penitilobata Dinc How. a. Habit, x ¥2, 6. basal part of flowering stem, x 4, c. inflorescence,
nat. size, d—e. flowers, x2, f. gynostemium, x 5. — T. muluensis DING Hou. g. Inflorescence on stem, x 4,
h. flower, nat. size, i. gynostemium, x 5 (a—f ARGENT c.s. 691, g—i ARGENT C.s. 760). Courtesy of Blumea.

1984]

the margin; lateral nerves 6—9 pairs, elevated and
prominent below, less so above; veins transverse or
+ parallel and reticulate, slightly elevated on both
surfaces; petiole S—10 mm, glabrous. /nflorescences
at the basal part of the stem, in the axils of leaves or
fallen leaves, or cauliflorous, simple or sparsely
branched, up to 4 (—7) cm long, pubescent; bracts
lanceolate, oblanceolate or elliptic, 6—9 mm long,
densely puberulous on both surfaces. Pedicel and
ovary 7—15 mm long, pubescent. Perianth pale yel-
low with center and margin streaked with claret-
colour, brown, deep reddish pink, dark purple out-
side and pink inside, or purple; campanulate, con-
tracted at the lower 1/3 and then erecto-patent,
15—25 mm long, c. 25 mm 9g, glabrous on both sur-
faces; tube urceolate, c. 10 mm long; lobes triangu-
lar, 6—10(—15) by 10—15 (—25) mm, acute, each with
longitudinal (c. 7) and reticulate veins. Stamens in 2
whorls: upper row 7—10, lower 13—16; filaments gla-
brous, |—2 mm; anthers oblong, c. 1.5 mm long.
Style column c. 4 mm long; lobes (4—) 6—9, spread-
ing, c. 3 mm long. Capsules slender, 5—10 cm long,
4-angular, straight or slightly twisted at the apical
part. Seeds ellipsoid, trigonous, 3—4 mm long, acute
at both ends, rugose-tubercled.

Distr. Malesia: Malay Peninsula (Perak, Din-
dings, Trengganu, Pahang, Selangor, Penang, Sin-
gapore).

Ecol. In forest, up to c. 500 m. F/. March—Oct.,
fr. Jan.—Nov.

Uses. Leaves used as medicine for cutaneous dis-
ease.

Vern. Tlinga berwang, Perak.

Note. See also the note under the Bornean 7. mu-
luensis.

15. Thottea reniloba DinG Hou, Blumea 27 (1981)
326, f. 19, 20, 53. — Fig. 2g.

Erect subshrub, up to 2.5 m high. Branches +
curved, terete or slightly compressed, 0.5—1 cm @,
pubescent. Leaves chartaceous, oblanceolate, lance-
olate, or elliptic, 1S—29 by 5.5—14 cm; apex acumi-
nate; base cuneate or obtuse; glabrous, sometimes
slightly pubescent on the midrib above, loosely pu-
bescent beneath; basal nerves 2 pairs, the inner one
starting near the base, ascending up to 2/3 or higher,
similar to the lateral ones in thickness and appear-
ance, the outer one weak and short, close to the mar-
gin; lateral nerves 5—8 pairs, occasionally with shor-
ter ones between them, elevated beneath, faint
above; veins transverse, scalariform, connected with
crossbar-like or loosely reticulate veinlets, slightly
elevated below, flat or obscure above; petiole 4—10
mm, pubescent. /nflorescences in the axiles of foliage
leaves, simple or sparsely branched, up to 8.5 cm
long, spici- or racemiform, internodes spacious, pu-
berulous; bracts oblanceolate to spathulate, or nar-

ARISTOLOCHIACEAE (Ding Hou) 77

row-elliptic, 3—7 mm long, puberulous on both sur-
faces. Pedicel and ovary rather long, 14—30 mm, pu-
berulous. Perianth dirty purplish red, or dark
brown, short-cupular, 10O—13 mm long, 15-20 mmo,
veins rather loosely reticulate, sparsely puberulous
outside, slightly (glandular) hairy inside; tube 5—6
mm long; lobes reniform, 5—7 by 11—20 mm, margin
reflexed at anthesis. Stamens in 2 whorls: upper row
9-12, lower 9—14; filaments glabrous, very short;
anthers oblong, c. 1 mm long. Sty/e column short, c.
1.5 mm long; lobes 8 or 9, c. 2 mm, glabrous. Cap-
sules (rather young) pendent, siliquiform, twisted,
narrowed at both ends, puberulous. Seeds ellipsoid,
c. 2.5 by 1.5 mm, tubercled, deeply furrowed.

Distr. Malesia: Northern Sumatra (Atjeh: Gu-
nung Leuser Nature Reserve; Tapanuli: Div. Pa-
dang; East Coast: Asahan and Upper Bila).

Ecol. In lowland forest, at base of steep sand-
stone rock or over basalt rock, up to 125 m. Fi. April
—Sept., fr. July.

Note. The flowers remind of 7. borneensis; dif-
fers from that species by loosely pubescent (not
densely sericeous) leaf undersurface, venation at
base reticulate (not transverse), rachis with spaced
bracts and long internodes, perianth without a dis-
tinct annular fold or ridge inside the mouth of the
tube, and reniform lobes.

16. Thottea penitilobata Dinc Hou, Blumea 27
(1981) 324, f. 14-16, 72A. — Fig. 6a—f.

Erect shrub, c. 1 m high. Branches terete, c. 0.5 cm
9, slightly puberulous, glabrescent. Leaves chartace-
ous, oblanceolate or elliptic, (12—) 16—22 by 5—9.5
cm; apex acuminate; base cuneate or slightly obtuse;
glabrous above, sparsely puberulous beneath; basal
nerves 2 pairs, the inner one similar to the lateral
nerves, emerging near the base and obliquely ascend-
ing halfway, the outer one very weak, short, and
close to the margin; lateral nerves 4—7 pairs, elevated
beneath, distinct above; veins transverse or scalari-
form, connected by transverse or reticulated veinlets,
slightly elevated beneath, visible or rather obscure
above; petiole 0.5—0.7 cm, subterete, sparsely puber-
ulous. /nflorescences near the basal part of the stem,
in axils of reduced leaves, simple or rarely with short
branches, spiciform, 3—6.5 cm long, puberulous;
bracts ovate or lanceolate, 2—7 mm long, puberulous
on both surfaces. Pedicel and ovary c. 6 mm long,
sparsely puberulous. Perianth c. 15 mm long, deeply
lobed, sparsely puberulous outside, glabrescent,
densely (glandular) papillate on the inner surface ex-
cept glabrous at the apical and marginal parts of the
lobe; veins invisible outside, obscurely reticulate near
marginal parts of lobes; tube very short; lobes broad-
ovate or suborbiculate, 10-12 by c. 11 mm, acute,
short-acuminate, or obtuse. Stamens in 2 whorls: up-
per row 10, lower c, 14; filaments glabrous, very

78 FLORA MALESIANA

[ser. I, vol. 10!

short; anthers oblong, c. 1.25 mm long. Sty/e column
short; lobes (6—) 10—14, 1.5—2 mm, glabrous. Cap-
sules unknown.

Distr. Malesia: Borneo (Sarawak: 4th Div., Gu-
nong Mulu Nat. Park; Kalimantan: Kalteng Prov.);
twice collected.

Ecol. On bank at riverside in lowland rain-forest
and in primary dipterocarp forest, 40—150 m. Fi.
Nov., Jan.

Vern. Kayu manis, Kalimantan.

Note. Closely allied to T. muluensis, from which
it cannot be separated in sterile state, but quite differ-
ent in flower structure: perianth 1.5 cm long, deeply
lobed, with broad or suborbicular lobes; 10—12 style
lobes; in 7. muluensis the perianth is 3.7—4.5 cm
long, only lobed halfway, with triangular lobes; 7
style lobes.

17. Thottea celebica DiInG Hou, Blumea 27 (1981)
318, f. 21, 22, 70A. — Fig. 7a—e.

Erect undershrub, up to 70 cm high. Branches te-
rete, 5-7 mm @g, pubescent. Leaves chartaceous to
coriaceous, 22.5—32 by 6.5—9 cm; apex acuminate;
base obtuse; glabrous above, sparsely puberulous be-
neath; basal nerves 2 pairs, the inner one branching
from the midrib c. 4 mm above the base and extend-
ing upward to about halfway, the outer one much
weaker and shorter and close to the margin; lateral
nerves c. 10 pairs; veins transversal or reticulate;
nerves and veins elevated and prominent beneath,
less so above; petiole short, c. 0.5 cm, puberulous.
Inflorescences at the base of the stem, simple, spici-
form, 5.5 cm long, puberulous; bracts elliptic, 4—8
mm long, sometimes 2-lobed, puberulous on both
surfaces. Pedicel and ovary c. 12.5 mm long, puberu-
lous. Perianth dark purplish red, campanulate, c. 25
mm long, c. 30 mm 9g, with several longitudinal veins
distinct outside, obscure inside, puberulous, glabres-
cent outside, glandular hairy inside, especially at the
lower 1/3; tube cupular, the lower half contracted
and cylindric (c. 6 mm long); lobes semi-orbicular, c.
10 by 15 mm, subrotund or slightly apiculate at the
apex. Stamens in 2 whorls: upper row 10—12, lower
12—15; filaments glabrous, 1—2 mm; anthers 1—1.7
mm long. Sty/e column c. 1 mm long; lobes c. 12,
erect, c. 2mm long. Capsules (very young) slender,
4-angular, twisted, c. 4 cm long, sparsely puberu-
lous.

Distr. Malesia: Central Celebes (Lambarese,
NE. Palopo); once collected.

Ecol. Open shady places in forest at low altitude.
Fl. & very young fr. July.

Note. The only species so far known from Cele-
bes. Allied to 7. philippinensis, which has a closely
reticulate prominent venation underneath the leaves,
a slightly shorter perianth not contracted in the lower
third, and 4—6 style lobes against c. 12 in T. celebica.

18. Thottea muluensis DING Hou, Blumea 27 (1981)
322, f. 72B. — Fig. 1c, 6g—i.

Erect shrub, c. 1 m high. Branches terete, c. 6 mm
@, glabrous. Leaves chartaceous, elliptic, lanceolate,
or oblanceolate, 15—29 by 5—12 cm; apex acute to
acuminate; base cuneate; glabrous above, sparsely
puberulous beneath; basal nerves one pair, emerging
slightly above the base and ascending to about half-
way; lateral nerves 6—10 pairs; nerves elevated and
prominent below, flat and distinct above; veins
transverse, scalariform, connected with loosely reti-
culated veinlets, elevated below, visible or obscure
above; petiole c. 1 cm, sparsely puberulous. /nflores-
cences at the basal part of the stem or near the
ground, usually in the axils of bracts, single or
sparsely branched near the base, racemiform, 12—14
cm long, puberulous; bracts lanceolate, S—10 mm
long, puberulous on both surfaces. Pedicel and
ovary 9-11 mm long, puberulous. Perianth dark
purplish or dark purplish maroon, campanulate,
37—45 mm long, with longitudinal and reticulate
veins, sparsely puberulous on both surfaces; tube
17—20 mm long; lobes triangular, 20—25 by c. 20
mm, acuminate. Stamens in 2 whorls: upper row 9,
lower 12; filaments glabrous, very short, c. 1.3 mm;
anthers oblong, 0.7—1 mm long. Style column c. 3
mm long; lobes 7, c. 1.5 mm, glabrous. Capsules un-
known.

Distr. Malesia: Borneo (Sarawak: Gunong Mulu
Nat. Park).

Ecol. Somewhat open position in lowland forest,
c. 35 m. Fl, Oct.—Nov.

Uses. Said to be used for birth control by Punan
people.

Note. Allied to 7. dependens which has, how-
ever, the undersurface of the leaf covered by papillae
forming rings or loops and a glabrous perianth. See
also note under 7. penitilobata.

19. Thottea corymbosa (GriFF.) Dinc Hou, Blumea
27 (1981) 320, f. 4. — Bragantia corymbosa GriFF.
Trans. Linn. Soc. 19 (1845) 335; Ann. Sc. Nat. Bot.
7 (1847) 340; Mia. Fl. Ind. Bat. 1, 1 (1858) 1068;
Kxiotzscu, Monatsb. Akad. Berl. (1859) 591, t. 1, f.
4; DucHARTREin DC. Prod. 15, 1 (1864) 429; Hook.
f. Fl. Br. India 5 (1886) 73; SOLEREDER, Bot. Jahrb.
10 (1889) 431. — Asiphonia piperiformis GRIFF.
Trans. Linn. Soc. 19 (1845) 333; Ann. Sc. Nat. Bot.
7 (1847) 338; Notul. 4 (1854) 344; Icon. Pl. Asiat. 4
(1854) t. 528, f. 1. — Strakaea melastomaefolia
PresL, Epim. Bot. (1851) 221. — Asiphonia sp.
GrirF. Notul. 4 (1854) 346; Icon. Pl. Asiat. 4 (1854)
t. 528, f. 2. — Bragantia melastomaefolia DUCHAR-
TREin DC. Prod. 15, 1 (1864) 429; WarBuRG, Pflan-
zenwelt 1 (1913) 521. — Apama corymbosa (GRiFF.)
WILLD. ex SOLEREDERin E. & P. Nat. Pfl. Fam. 3, 1
(1889) 272; O.K. Rev. Gen. Pl. 1 (1891) 563; Kinc&

1984]

GaMBLE, J. As. Soc. Beng. 75, ii (1912) 25; Moore,
J. Bot. 63 (1925) 83; HEyNeg, Nutt. Pl. (1927) 596;
Burk. Dict. | (1935) 188; ScHmipT in E. & P. Nat.
Pfl. Fam. ed. 2, 16b (1935) 233; HEND. Mal. Wild FI.
(1951) 420, f. 382A—C. — Fig. 1a, 8d.

Shrub, spreading, sometimes scrambling, up to 5
m high. Branches terete, up to 2 cm 9, densely puber-
ulous. Leaves chartaceous or subcoriaceous, ovate to
lanceolate, elliptic or elliptic-oblong, sometimes ob-
ovate, 6.5—17.5 by 2.5—8.5 cm; apex cuspidate or
acuminate; base cuneate, rounded or obtuse; upper
surface glabrous except scattered puberulous on the
midrib, nerves and veins, lower surface puberulous;
basal nerves 2 pairs, the inner one as prominent as
the midrib (the leaf appearing as 3-ribbed), ascend-
ing upward and reaching often to the apical part of
the blade, the outer one much weaker, close to the
margin, shorter than the inner one; lateral nerves
0—3 pairs, when present usually at the upper half of
the blade; veins transverse, parallel (+ perpendicular
to the midrib), joined by crossbar-like or loosely reti-
culate veinlets; nerves and veins elevated and promi-
nent beneath, slightly elevated and rather fine above;
petiole subsessile to c. 8 mm, puberulous. /nflores-
cences terminal and/or axillary in the upper leaf-
axils, few- and lax-branched, paniculiform or corym-
bose, up to 10 cm long, puberulous; bracts subulate
or linear, up to 8 mm long, puberulous. Pedicel and
ovary 7-20 mm, puberulous. Perianth yellow,
greenish or cream coloured outside, pale lilac inside,
3—3.5 mm long, when spreading c. 7 mm @, densely
puberulous outside, glabrous inside, veins invisible,
deeply 3-lobed; lobes broad-ovate or suborbicular,
2.5—3 by 2.5—3.5 mm, apex acute. Stamens in 1
whorl, 7—10, rarely more; filaments 0 or obscure; an-
thers oblong, c. 1 mm long, covered with short,
hooked hairs. Sty/e column obscure, lobes 4, gla-
brous. Capsules slender, long, up to 38 cm long,
4-angular, slightly twisted, puberulous. Seeds ovoid,
trigonous, 4—6 by 2.5—3.5 mm, rugose.

Distr. Malesia: Widely distributed but scattered
in Sumatra and the Malay Peninsula (incl. Penang);
Borneo (Kalimantan: Bukit Kasian), once collected.

Ecol. In forest, sometimes on the edge of forest,
occasionally in shaded forest in limestone zone, from
the lowland up to 1050 m. F/. fr. all the year round.

Uses. Pounded leaves are put inside the hollow of
the tooth to remedy toothache. The central part of
the roots is chewed along with betel-nut as a diuretic,
if needed, during confinement (BURKILL, /.c.).

Vern. Sumatra: (andor) lasi, bandar puluh, Asa-
han, kadudu rimbu, Djambi, subie siang, Riouw
(Kuala Belilas). Malay Peninsula: akar chambai olar,
a. julong bukit, a. serai, andor lasi, bunga changi u-
lar, ékor pelandok, jangat, lerkor, mahjar pahit, tin-
jau biuti, M.

ARISTOLOCHIACEAE (Ding Hou) 79

20. Thottea tomentosa (BL.) Dinc Hou, Blumea 27
(1981) 328, f. 48, 49. — Ceramium tomentosum BL.
Bijdr. (1826-27) 1135. — Bragantia tomentosa BL.
En. Fl. Jav. (1827) 82; BENN. in Benn. & Brown, PI.
Jav. Rar. 1 (1838) 43, t. 11; Grirr. Trans. Linn. Soc.
19 (1845) 336; LinpL. Veg. Kingd. (1846) 794; GRIFF.
Ann. Sc. Nat. Bot. 7 (1847) 340; ZoLL. Syst. Verz. 2
(1854) 118; Mig. Fl. Ind. Bat. 1, 1 (1858) 1068; Du-
CHARTRE in DC. Prod. 15, 1 (1864) 431; Hook.f. FI.
Br. India 5 (1886) 73, incl. var. lanuginosa Hoox./.;
CLARKE, J. Linn. Soc. Bot. 25 (1889) 61; SOLEREDER,
Bot. Jahrb. 10 (1889) 431; Riot. J. Str. Br. R. As.
Soc. n. 57 (1910) 89; ibid. n. 59 (1911) 161. — Vanhal-
lia tomentosa J.A. & J.H. SCHULTEs, Syst. Veg. 7
(1829) 166. — Bragantia blumii LINDL. Bot. Reg. 18
(1832) sub t. 1543, in note; Veg. Kingd. (1846) 793,
f. 526. — Cyclodiscus tomentosus KLoTzscH, Mo-
natsb. Akad. Berl. (1859) 592. — Apama tomentosa
ENGL. ex SOLEREDER in E. & P. Nat. Pfl. Fam. 3, 1
(1889) 272; O. K. Rev. Gen. Pl. 1 (1891) 563; KinG
& GAMBLE, J. As. Soc. Beng. 75, 11 (1912) 25, incl.
var. lanuginosa (Hook. f.) K. & G. /.c. 26; BACK.
Trop. Natuur 7 (1918) 179, f. 1—4; ibid. 8 (1919) 164;
BeuMEE, ibid. 8 (1919) 15; Rip. Fl. Mal. Pen. 3
(1924) 15; Moore, J. Bot. 63 (1925) Suppl. 83;
Koorp. Exk. Fl. Java 4 (Atlas) (1926) 589, f. 871;
Heyng, Nutt. Pl. (1927) 596; STEEN. Bull. Jard. Bot.
Btzg III, 12 (1932) 204; Burk. Dict. 1 (1935) 189;
ScumipT in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
234, f. 120OA—G; KANJILALC.s. Fl. Assam 4 (1940) 30;
HEND. Mal. Wild FI. (1951) 423, f. 382E; Back. &
BakH. f. Fl. Java 1 (1963) 162. — Bragantia affinis
PLANCH. ex ROLFE, Kew Bull. (1913) 265; MERR. En.
Philip. 2 (1923) 120. — Bragantia brevipes MERR.
Philip. J. Sc. 17 (1920) 248; En. Philip. 2 (1923) 120.
— Apama affinis WeissE, Ber. Deut. Bot. Ges. 45
(1927) 235; ScHmipTin E. & P. Nat. Pfl. Fam. ed. 2,
16b (1935) 234. — Apama brevipes WeIssE, Ber.
Deut. Bot. Ges. 45 (1927) 235; ScHmipT in E. & P.
Nat. Pfl. Fam. ed. 2, 16b (1935) 234.

Subwoody herb, from creeping base erect to
erecto-patent, 10—35 cm; stems one to several or
sometimes many together, usually simple. Branches
rather slender, c. 5mm @, bearing 1—5 (often 2 or 3)
foliage leaves at the apical part, furrowed or slightly
angular, tomentose. Leaves chartaceous or subcoria-
ceous, variable in shape and size even on one plant,
ovate, elliptic or broadly elliptic, elliptic-oblong, ob-
ovate or oblanceolate, rarely suborbiculate, (4—)
7-18 (—24) by (1.7—) 3-15 (—17) cm; apex acute,
acuminate, or obtuse; base obtuse, rounded, some-
times subcordate or cordate; glabrous above, densely
pubescent, tomentose or villous beneath especially
when young, sometimes glabrescent; basal nerves 2,
rarely 3 pairs, starting from the base and ascending
upward to more than halfway, similar to the lateral
nerves, the outer | or 2 rather faint and short, close

FLORA MALESIANA [ser. I, vol. 10!

PEER

i
tlhe
le

s

=
ae
oS
ssi

Le
Gril
<oN

x\ PD
uN

Fig. 7. Thottea celebica Dinc Hov.a. Habit, x %, b. inflorescence, c. flower, nat. size, d. gynostemium, x5,
e. young infructescence, x 2. — T. tricornis MAINGAY. f—g. Flowers, nat. size, A. indument of inner surface
of perianth, x 10, i. gynostemium, x5. — T. sumatrana (MeErRR.) Dinc Hou. j—k. Flowers, nat. size, /. gyno-
stemium, x5, m. 3-lobed style, x 5. — T. parviflora Riwi. n. Flower, nat. size, o. ditto, perianth removed,
x 5, p. floral diagram showing 4 whorls of stamens (a—e STRAATMANS./1., f-i VAN Batcooy 2627, j—m Kos-
TERMANS 284, nm VAN BEUSEKOM & PHENGKLAI 694, 0—p GEESINK & SANTISUK 5101). Courtesy of Blumea.

1984]

ARISTOLOCHIACEAE (Ding Hou) 81

to the margin; lateral nerves 4—9 pairs, joined by
rather closely reticulate and loosely crossbar-like
veins and veinlets, elevated beneath, distinct some-
times obscure above; petiole S—15 mm, pubescent.
Inflorescences near the base of the branches, some-
times hidden under fallen leaves, in the axils of bracts
or not well developed (small) leaves, often simple and
spiciform, up toc. 12 cm long, pubescent; bracts lan-
ceolate to linear, up to c. 10 mm long, pubescent.
Pedicel and ovary 8—22 mm long, densely pubescent.
Perianth pale yellow, yellow with purple, purple,
pale red, or red (colour changing with age of the
flower), urceolate-campanulate, 6—12.5 mm long,
12—16mm 9g, with longitudinal and loosely reticulate
veins, pubescent outside, glabrous inside; tube 3—5
mm long, with a thin ‘disk’ adnate to the inner side,
slightly protruding above the tube (c. 0.5 mm) like a
narrow rim; lobes broadly ovate, suborbicular, or
subreniform, 3—7.5 by 4—8 mm. Stamens 6, in 1
whorl; filaments glabrous, 1—1.5 mm; anthers ob-
long, 1.5—2 mm long, connective slightly produced
beyond the anthers. Sty/e column c. 2 mm long; lobes
3 (or 4), 1.5—2 mm, hairy often at the apical part.
Capsule slender, 3.5—5 (—15), often obscurely
4-angular, pubescent, glabrescent. Seeds oblong, tri-
gonous, c. 4 by 2 mm, rugose.

Distr. India (Assam: Manipur & S. Andaman
Is.), Bangladesh (Sylhet), Burma (Moulmein), South
Vietnam (Bien Hoa), Peninsular Thailand; in Male-
sia: Malay Peninsula (throughout), Sumatra, West &
Central Java, and Philippines (Jayabas, Alabat I.,
Panay, Mindanao), not yet found in Borneo. Culti-
vated in Hort. Bog. n. XI-B-XIII-138.

Ecol. Inshady, moist places in forest, sometimes
in bamboo or teak forest, occasionally in secondary
forest, rarely on limestone, locally sometimes com-
mon, from the lowland up to 1200 m. Fi. fr. often all
the year round.

Uses. In Malaya the plant is used for poulticing
skin-complaints and boils (along with ///ligera). In Ja-
va the stems and leaves may be pounded and the juice
swallowed for coughs. The roots and leaves are used
as a diuretic during confinement. In W. Java also
used against snake-bites (BURKILL /.c.; HEYNE,/.c.).

Vern. Malay Peninsula: kaneb, kemed, seréng-
kong, M. Java: singa dapur, s. depa, J; kaliwaro,
singa depa, S.

Notes. T. tomentosa is the widest ranging species
of the genus. It is the only species in Java.

Fruits are surprisingly rare and hitherto accepted
as indehiscent. However, in a Thailand collection it
had split with 4 valves.

The species is allied to the Indian 7. siliquosa
(Lamk) Dinc Hou and the Bornean 7. paucifida (see
note under the latter); 7. siliquosa has 9 stamens in
3 groups, the anthers dorsally and the style lobes
densely hooked-hairy.

21. Thottea macrantha (BoERL.) DinG Hou, Blumea

27 (1981) 321. — Bragantia macrantha Boerrt.
Handl. 3 (1900) 64; Vater. Icon. Bog. 3 (1908) sub
t. 260, emend. — Apama macrantha WEIssE, Ber.

Deut. Bot. Ges. 45 (1927) 234, in obs.; STEEN. Bull.
Jard. Bot. Btzg III, 12 (1932) 204; Scumipr in E. &
P. Nat. Pfl. Fam. ed. 2, 16b (1935) 234, f. 120H—J.
— T. hirsuta Rw ey, J. Mal. Br. R. As. Soc. 1 (1923)
87. — Fig. 2e-f.

Herb woody at base, or shrub-like, up to 4m high.
Branches terete, S—~7 mm @, pubescent or hirsute.
Leaves chartaceous to subcoriaceous, obovate to ob-
lanceolate, elliptic-oblong, or lanceolate, (13—)
25—36 (—41) by (S—) 8—12.5 (—21) cm; apex short-
acuminate or acuminate, sometimes apiculate, base
cuneate; glabrous above, pubescent beneath; basal
nerves 2 or 3 pairs, the inner one starting from the
base or c. 5 mm above it, ascending upward and
reaching to more than halfway, similar to the lateral
nerves, the outer | or 2 much weaker and shorter,
close to the margin; lateral nerves 7—10 pairs, elevat-
ed and prominent beneath, slightly elevated or flat
above; veins transverse, scalariform, some loosely
reticulate, joined by weaker crossbar-like or loosely
reticulate veinlets, slightly elevated beneath, visible
or obscure above; petiole 5—8 (—15) mm, hirsute. /n-
florescences axillary, 1—3, often in axils of foliage
leaves, simple or sparsely branched, spiciform or ra-
cemiform, rarely paniculiform, up to 7 cm long, pu-
bescent; bracts oblanceolate, those at the lower part
shorter, pubescent outside, glabrous inside. Pedicel
and ovary 15—18 mm, pubescent. Perianth reddish
or dark brown and puberulous outside, white and
glandular-hairy inside, campanulate, c. 20 mm long
and wide; tube c. 8 mm long; lobes broadly rounded,
c. 12 by 4 mm, emarginate, reflexed. Stamens in 1
whorl, 9—12, patent or reflexed; filaments c. 3.5 mm,
glabrous; anthers oblong, 1.2—2 mm long. Style col-
umn short, lobes 9-12, c. 2mm, glabrous. Capsules
erect, elongate, up to 11 cm long, 4-angular, pubes-
cent, glabrescent. Seeds ellipsoid, c. 3 by 1.5 mm,
transverse-rugose.

Distr. Malesia: Northern Sumatra (Atjeh, Sibo-
langit, Taram, Ketambe, Deli, Asahan). Cultivated
in Hort. Bog. sub n. XI-B-XIII-133.

Ecol. In primary and young forest, mostly on
slopes, sometimes in secondary forest, rare, some-
times locally abundant, in lowland up to 450 m. F/.
Sr. Feb.—Sept.

Vern. Ambolas tombak, Asahan.

Note. Ruipiey described 7. hirsuta to have 2
whorls of stamens, but the single flower of the type
I examined has the stamens in one whorl.

22. Thottea sumatrana (Merr.) Dinc Hou, Blumea
27 (1981) 328, f. 70C. — Apama sumatrana MERR.
Pap. Mich. Ac. Sc. 23 (1937) 178. — Fig. 7j-m.

82 FLORA MALESIANA

[ser. I, vol. 10!

a sees Rah
Gee Ise py
E e

Be, “s eate Ee

eae

‘v,
weer
?

ats
Rew &

eth,
‘s
sae Sgt
YaCrrE
camp EN

Fig. 8. Samples illustrating leaf venation patterns: a & c densely reticulate, b loosely reticulate, d trabeculate.

— a. Thottea philippinensis Quis., X 2/3 (FB 30249). — b. Aristolochia momandul Scumipt, x 3/5 (NGF

27848). — c. A. crassinervia ScumipT, x 4/5 (NGF 45343). — d. Thottea corymbosa (GrirF.) DinG Hou,
x 2/3 (Hou 732). (a undersurface of not cleared leaf, b—d cleared leaves).

1984]

ARISTOLOCHIACEAE (Ding Hou) 83

Herb woody at base, shrub-like, up to 1 m high.
Branches sulcate or irregularly angular, c. 5 mm @,
sparsely pubescent. Leaves chartaceous, elliptic, lan-
ceolate, obovate, rarely oblanceolate, 10-21 by
4.5—7.5 cm; apex acute, acuminate, or short-acumi-
nate; base obtuse, rotund, or cuneate; glabrous
above, pubescent beneath; basal nerves 2 or 3 pairs,
the inner one ascending upward to about halfway,
similar to the lateral nerves, the outer 1 or 2 much
weaker and shorter, close to the margin; lateral
nerves 8—14 pairs, joined by transverse, subparallel,
scalariform or sometimes loosely reticulate veins and
veinlets, elevated beneath, often obscure above, peti-
ole 3—5 (—10) mm, pubescent. /nflorescences ax-
illary, usually in the axils of foliage leaves, spiciform
or racemiform, solitary or rarely 2 in an axil, pubes-
cent; bracts crowded, lanceolate, elliptic, oblanceo-
late or spathulate, 3—10 mm long, pubescent outside,
glabrous inside. Pedicel and ovary 5—10 mm long,
pubescent. Perianth dirty yellowish white, inside
blackish red, or dark-red, campanulate, or funnel-
shaped, 14—20 mm long, c. 12 mm @, veins loosely
reticulate, sparsely puberulous outside, glabrous in-
side; tube c. 5 mm long; lobes ovate to lanceolate,
8—15 by 5—8 mn, apex acute, acuminate, or obtuse.
Stamens 6, in 1 whorl; filaments glabrous, c. 1 mm;

anthers oblong, c. 1 mm long (excl. the protrudin
connective). Sty/e column c. 2 mm long, lobes 3, 1—
mm, glabrous. Capsules slender, up to 16 cm long,
cylindric or slightly angular, twisted, sparsely pubes-
cent. Seeds ovoid, trigonous, c. 4.5 by 2.5 mm, trans-
versely rugose.

Distr. Peninsular Burma (Ta Pe) and Peninsular
Thailand (Pattani and near Neckey); in Malesia:
Northern Sumatra (Tapanuli: Padang Lawas; East
Coast: Laut Tador & Gedong Biara Estates) and N.
Malay Peninsula (Perlis and Kedah).

Ecol. In lowland forest, by stream, on sandy
loam soil, up to c. 150 m. Fil. fr. April—August.

Note. Sterile specimens are difficult to identify.
In flower easily recognized by a deeply lobed peri-
anth, 6 stamens in one whorl, anthers with a protrud-
ing connective, and 3 style-lobes.

8
5

Insufficiently known

Thottea sp. DinG Hou, Blumea 27 (1981) 329.

In Kalimantan (Sg. Dingei) JAHERI (nm. 790, BO)
collected an as yet undescribed species. It resembles
T. tomentosa, but the single immature flower pos-
sesses 2 whorls of stamens. Better material is needed
for a proper description.

2. ARISTOLOCHIA

LINNE, Sp. Pl. (1753) 960; KLotzscH, Monatsb. Akad. Berlin (1859) 593; Du-
CHARTREin DC. Prod. 15, | (1864) 432; ScHmipTin E. & P. Nat. Pfl. Fam. ed.
2, 16b (1935) 235; HOEHNE, FI. Bras. 15 (1942) 23; PFerrer, Ann. Mo. Bot. Gard.
53 (1966) 122; Tax. Rev. Pentam. Sp. Aristolochia (1970) 15; Dinc Hou, Blumea
29 (1983) 223. — Fig. 1, 2, 8-18.

Herbaceous perennials, undershrubs or shrubs, usually scandent, scrambling,
twining, or climbing, sometimes high lianas, often with prostrate or tuberous rhi-
zomes or rootstocks; vegetative parts upon breaking or only the flowers often
with a bad, mostly putrid smell. (Old) woody stems mostly with a thick-corky
and fissured bark, showing broad, medullary rays on cross-section. Leaves en-
tire, sometimes 3-lobed (irregularly up to 7-lobed in extra-Malesian spp. ). Petiole
grooved above. Flowers usually zygomorphic, rarely actinomorphic, solitary,
fasciculate, or in inflorescences (cymose, racemose, spicate, paniculate), axillary
or cauligerous. Perianth straight, curved, or S-shaped, inflated at the basal part
(utricle), then contracted or narrowed above in an often cylindric or fun-
nel-shaped part (tube), gradually elongated, enlarged and expanded into the
1-lipped or 3 (—6)-lobed limb, the utricle often inside provided with 2 (—6) glan-
dular bodies. Stamens mostly 6 (4, 5, or more in extra-Malesian spp.) (10 in the
Malesian A. decandra), adnate to the style column in a gynostemium. Ovary ob-
long or elongate, slightly 6-angular and 6-celled (5-celled in extra-Malesian spp. );
style column mostly 6-lobed (5-lobed in extra-Malesian spp.). Fruits capsular,

84 FLORA MALESIANA [ser. I, vol. 10!

6-celled (5-celled in extra-Malesian spp.), dehiscent (dehiscing usually acropetal-
ly, rarely basipetally: A. singalangensis) or indehiscent (A. die/siana ?, and extra-
Malesian spp.). Seeds ovate, deltoid, or triangular, often winged, flat, convex-
concave, or slightly longitudinally curved; testa crustaceous or hard, finely ver-
rucose or smooth; funicle often fleshy, thickened, usually covering the whole
seed, usually persistent on the seed as an elaiosome, membranous when dry.

Distr. About 400 species, widely distributed mainly throughout the tropics and subtropics and some in the
warm temperate regions, also in Australia; throughout Malesia: 28 species.

Ecol. Usually scattered, only rarely locally abundant, in primary forests, sometimes in (old) secondary for-
ests or in thickets, occasionally occurring in beach or swampy forest, limestone regions, or submontane to mon-
tane forest; mostly at low and medium altitudes, sometimes found higher, above 1500 m, up to 2250 m (in
Papua New Guinea). ;

For the relation with butterflies, pollination in relation to flower morphology, seed dispersal, efc., I refer
to the general chapters.

Morph. See the general chapters, also for chromosomes, phytochemistry, anatomy and palynology.

Uses. See the general chapters.

Notes. Inthe key and descriptions the pedicel and ovary are taken as a whole, like under Thottea, because
these organs merge imperceptibly.

For the diameter of the floral tube, only the cylindric middle part is used for size.

In general flowering material is necessary for correct identification.

In Malesia a number of exotic species is cultivated; a dozen of these are entered in the key by BACKER & BaK-
HUIZEN f., Flora of Java 1 (1963) 162—164.

For convenience of identification, local keys for islands have been added to the general key.

GENERAL KEY TO THE SPECIES
based on fertile specimens

1. Leaves deeply 3-lobed, in shape reminding of the letter “W’.
2. Leaf base shallowly concave, subcordate, or almost truncate, rarely cuneate. Inflorescences with small,

not amplexicaul bracts c. 1.5 mm long. Seeds distinctly winged ...................-44- 1. A. jackii
2. Leaf base cuneate. Inflorescences with conspicuous, amplexicaul bracts 7-10 (—15) mm long. Seeds not
WIVES en G o-aisbu 6 ceo eecha.G Guo 0-9 bb £0, olouEO CERES 66 Bao emo crerNeIS coin G aicid Gctomrors euepsryats cca 2. A. curtisii

1. Leaves never deeply 3-lobed as above, mostly entire.
3. Leaves villous or densely tomentose underneath, often concealing the surface (especially when young).
4. Perianth limb distinctly 3-lobed; lobes triangular, 3 by 4.5—6 cm, obtuse. (Fruit dehiscing from the apex
downward ssseedsistronely CONVEX-CONGAVE)) ees ai-l i-th sell -atele tele teri eee or 3. A. singalangensis
4. Perianth limb rim-like, the limb 0.5—1 cm wide, obscurely 3-lobed. (Fruit unknown) 10. A. coadunata
3. Leaves shortly, minutely, or sparsely hairy on the undersurface (surface always exposed), or glabrous on
both surfaces.

5. Leaves glabrous on both surfaces.

6. Leaf base subtruncate, slightly concave, obtuse, or rounded, sometimes slightly cuneate. (Seeds not

winged).
7. Plants erect, up toc. 1 m high. Leaves with pinnately arranged lateral nerves. Petiole less than 1 (—2)
Gin ona, Pater Woijojeals op ga sonasroaccoobagcuopabacgodaqoongododgobr 18. A. philippinensis

7. Twiners or stout climbers, up to 15 m high. Leaves palmately nerved. Petiole 3—12 cm long.
8. Leaves distinctly papillate beneath; 5-nerved, joined by reticulate veins. Flowers 1-lipped
13. A. papillifolia
8. Leaves smooth beneath; 3- or 5-nerved, joined by + transverse veins. Flowers unknown
14. A. transtillifera
6. Leaf base subcordate or cordate (usually adult leaves), or subtruncate (especially when young in A. gau-
dichaudii).
9. Perianth puberulous outside; limb narrow-elliptic, 5-6 cm long.................. 22. A. leytensis
9. Perianth glabrous outside; limb obovate to oblanceolate, 1.7—2.5 cm long.
10. Perianth tube 8—12 mm long; limb obovate or oblanceolate, 17—20 mm long. Fruits oblong 4—6
byes stein SCedsnvilgedes meme nee rey ucts ches troteleuruersielcl aie set leet ten -rarette <p -t=vell 27. A. gaudichaudii

1984] ARISTOLOCHIACEAE (Ding Hou) 85

10. Perianth tube 16—19 mm long; limb obovate-oblong, 20—25 mm long. Fruits unknown 8. A. klossii
5. Leaves shortly, minutely, or sparsely hairy on the undersurface.
11. Plants erect, up to c. | m high.
12. Petiole (20—) 30—40 mm long. Seeds verrucose only on the marginal part beneath, smooth above
19. A. humilis
12. Petiole at most 10 mm long. Seeds verrucose on both surfaces.
13. Leaf base obtuse or rounded, subacute or cuneate. Perianth with reflexed tube 17. A. samarensis
13. Leaf base distinctly cordate, sometimes slightly cordate, very rarely associated with some obtuse
ones. Perianth straight, sometimes slightly curved.
14. Leaf base deeply cordate, auricles often overlapping or surrounding the stem 20. A. macgregorii
14. Leaf base slightly cordate, rarely associated with some obtuse ones, auricles obscure or 0
21. A. sericea
11. Twiners or lianas, much taller, up to many metres high.
15. Perianth 6- or 3-lobed.
PERIATNIG ODER Sear LTS ees Se eee 24. A. schlechteri
16. Perianth 3-lobed.
17. Perianth 19—21 cm long. Stamens 10. Style-column 10-lobed. Leaves suborbicular or broad-ovate,
deeply cordate; palmately/S=sion7-nenved’ =.22)J2)2. hisses lo ateedalevteew ave 16. A. decandra
17. Perianth less than 9 cm long. Stamens 6. Style column 6-lobed.
18. Flower buds or perianth lobes with the apical 20—30 mm strongly contracted or narrowed and
often tail-like when dry (easily broken).
19. Perianth lobes suborbicular, c. 10 mm g, with an apical tail-like part c. 20 mm long
11. A. momandul
19. Perianth lobes triangular to narrow-triangular, 35—5S0 by 12—14 mm at the bases (apical 25—30

munvoften tail-like when dry) ust seas ee SR cee een eer 25. A. dielsiana
18. Flower buds or perianth lobes without a tail-like apical part as above. Perianth lobes triangular,
5—6) by/8—10) ml, apex: ODtuUSEsOrsTelUSe is -ictae oe eet eel cenevatee es tensa 26. A. engleriana

15. Perianth 1-lipped.
20. Leaf base often rounded or obtuse, not cordate.

21. Inner pair of basal nerves reaching nearly the leaf apex joined by 10—14 pairs of loose, transverse
or slightly curved cross-veins. Perianth without a stipe; limb ovate-oblong or lanceolate, 6—9 by
BSE emai civics SERS. Deere Ree ES as IS. RELI: Se oer 15. A. naviculilimba

21. Leaves with | pair of basal nerves reaching upward to about halfway; lateral nerves 4 or 5 pairs,
obliquely ascending to the margin. Perianth with a distinct stipe of 5 mm; limb oblong, much smal-
eG) AF! by0.7 sense Reha, SRE LA OE SOL UL Se A ee een 12. A. rumphii

20. Leaf base cordate or subcordate.
22. Leaf veins and veinlets closely reticulate or foveolate-reticulate, prominent beneath. Cf. fig. 8c.
23. Perianth with a distinct stipe of 3-5 mm. Seeds winged. Leaves variable in shape and size
5. A. zollingeriana
23. Perianth without a stipe. Seeds not winged.
24. Perianth limb ovate-oblong, 30 by 14—16 mm. Seeds triangular, c. 5 by 4 mm, verrucose on both

SUT FACES SIs Se SR 6 Va RRS FEC UIE eS AS ee ate ee 4. A. foveolata
24. Perianth limb linear, 20 by 4 mm. Seeds triangular or deltoid, 7 by 6-7 mm, rather smooth or
obscurely muriculate:on both surfaces: 2.2.1. 6.2% 0... GER ee 23. A. crassinervia

22. Leaf veins and veinlets loosely reticulate, distinct or obscure beneath. Cf. fig. 8b.
25. Leaves triangular or deltoid. (Leaves palmately 5-nerved. Perianth with a distinct stipe of c. 2.5
msti: Seeds winged) ix. situ: 2s the Saas WS ee Pee Oa ie etoteni at toe 28. A. linnemannii
25. Leaves often ovate, ovate-oblong, or lanceolate.
26. Leaves with 3—5 pairs of lateral nerves from the midrib, obliquely ascending. Perianth with a
distinct stipe.

27. Perianth limb 20—30 (—40) mm long. Seeds winged ...........060 cece eens 9. A. tagala
27. Perianth limb 50-60 mm long (ripe seeds unknown) ........6.60 6000s e es 22. A. leytensis

26. Leaves without lateral nerves from the midrib. Perianth without a stipe.
28. Basal nerves 2 (or 3 pairs), the inner pair ascending upward to c. 2/3 of the blade. Flowers
small: utricle 3~6 mm long, tube 2.5—5 mm long, limb 11-12 mm long . 6. A. minutiflora
28. Basal nerves | pair, ascending upward to the apex. Flowers larger: utricle 10-12 mm long, tube
5-11 mm long, limb 15—25 mm IONG.........0. severe eeecreeeevevenes 7. A. glaucifolia

86 FLORA MALESIANA [ser. I, vol. 10!

KEY TO THE SPECIES
Sumatra, Malay Peninsula and neighbouring islands

1. Leaves deeply 3-lobed, in shape often reminding of the letter ‘W’.
2. Leaf base shallowly concave, subcordate, or almost truncate, rarely cuneate. Inflorescences with small,

not amplexicaul bracts (c. 1.5 mm long). Seeds distinctly winged...................... 1. A. jackii
2. Leaf base cuneate. Inflorescences with conspicuous amplexicaul bracts (7—10, rarely up to 15 mm long).
SES GSHIOL SWAT eC epee ae ier Sete Hi Shave PN cr MEO, TREE ea caPet ernie ET tI POT? eae 2. A. curtisii

1. Leaves entire.
3. Leaves villous or densely tomentose underneath, often concealing the surface (especially when young).
4. Perianth limb distinctly 3-lobed; lobes triangular, 3 by 4.5—6 cm, obtuse. (Fruit dehiscing from the apex
CONMIMITE SITs Sioa hy Coy, conNeeE KS) cnasnadedacoousconouguosude aan ss 3. A. singalangensis
4. Perianth limb rim-like, the limb 0.5—1 cm wide, obscurely 3-lobed. (Fruit unknown) 10. A coadunata
3. Leaves shortly, minutely, or sparsely hairy underneath (surface always exposed), or glabrous.
5. Leaf veins and veinlets closely reticulate or foveolate-reticulate, prominent beneath.
Op, RAM EValiol WA NO WE BSUS. Sreeaky avo Wl oe oo ononenodoaascecocucogenenonodo: 4. A. foveolata
6. Perianth with a distinct stipe of 3—5 mm: Seeds winged’ ..................... 5. A. zollingeriana
5. Leaf veins and veinlets loosely reticulate, distinct or obscure beneath.
7. Perianth without a stipe. Seeds not winged. (Leaves thin-chartaceous).

8. Leaves palmately 5 (—7)-nerved, papillate beneath) iaeceshie sinker. es shee 6. A. minutiflora
8. Leaves 3-curvinerved, with irregular (finely reticulate, wax) thickenings and scattered black dots (se-
cretonyscells))ibeneatliaeeyk chord tek Mieuders eeccets Sotele anata Hicnec tached aaMGLanA eda AAR 7. A. glaucifolia

7. Perianth with a distinct stipe. Seeds winged. (Leaves chartaceous).
9. Leaf with the two basal lobes widely separate from each other (sinus 7—9 cm wide at the base). Perianth

limb obovate-oblong, 10—14 mm wide; apex slightly retuse or mucronate........... 8. A. klossii
9. Leaf with the two basal lobes close to each other or often connivent at the base. Perianth limb lanceo-
late to narrowly lanceolate; 6—8 mmiwide; apex acute!) sa. e2 a ee le rs ele pene 9. A tagala

KEY TO THE SPECIES
Java, Lesser Sunda Is., Moluccas, Celebes and neighbouring islands

1. Leaves deeply 3-lobed, in shape often reminding of the letter ‘W’...................... 1. A. jackii

1. Leaves entire.

2. Leaves villous or densely tomentose on the undersurface. Perianth with the tube bent backward and con-
tacting closely laterally with the utricle; limb rim-like, obscurely 3-lobed.......... 10. A. coadunata

2. Leaves shortly or minutely hairy beneath. Perianth straight or slightly curved; limb distinctly 3-lobed or
1-lipped.

3. Flower buds with a distinct, long tail-like apex c. 20 mm long (easily broken when dry). Limb distinctly
3-lobed. Capsules oblong or ellipsoid, 5.5—9 by 3.5—4 cm, strongly 6-ridged. (Seeds not winged, rather
SMO OMMONMb OLS unfaces)) ck ctl cic erase vic. sss See cece lies Re Chon ane eee 11. A. momandul

3. Flower buds without a long tail-like apex. Limb 1-lipped. Capsules short-cylindric, subglobose, slightly
pyriform, or oblong, 2—4 by 1.5—3 cm, often slightly 6-ridged.

4. Leaf base obtuse, sometimes slightly cuneate, rarely truncate. Seeds not winged ... 12. A. rumphii
4. Leaf base cordate. Seeds winged.

5. Leaf veins and veinlets closely reticulate, densely covered with minute hairs beneath (examining with
a handlens or under the dissecting microscope). Perianth lobes obovate-oblong, 25—30 by 10 mm,
longitudinally reflexed at anthesis. Seeds 4—5S by 4 mm (incl. the c. 0.6 mm broad wing)

5. A. zollingeriana

5. Leaf veins and veinlets loosely reticulate, sparsely shortly hairy or (nearly) glabrous beneath. Perianth
lobes lanceolate to narrowly lanceolate, 20—30 (—40) by 6—8 mm, not longitudinally reflexed at anthe-
sis. Seeds 8.5—10 mm long and wide (incl. the c. 2 mm broad wing) ............... 9. A. tagala

KEY TO THE SPECIES
Borneo

1. Leaves deeply 3-lobed, in shape often resembling the letter ‘W’.....................08. 1. A. jackii
1. Leaves entire.

1984] ARISTOLOCHIACEAE (Ding Hou) 87

2. Leaf base not cordate, but obtuse or rounded, shallowly concave, subtruncate, or slightly cuneate.
3. Leaves glabrous beneath.

4. Leaves distinctly papillate beneath (handlens or binocular), 5-nerved, joined by reticulate and trans-

“CET RE WET eae PSR pacts Oe ee Ci Oe Oe eI ie A ee 13. A. papillifolia
4. Leaves not papillate beneath, 3 (—S)-nerved, joined by transverse veins ....... 14. A. transtillifera
3. Leaves minutely hairy beneath especially on the venation ................... 15. A. naviculilimba

2. Leaf base distinctly cordate.

5. Petiole 10—13 cm long. Flowers large: utricle 5—6 cm long, tube c. 5 cm long, limb 4—9 cm long, 3-lobed.
CP TUETO TL ESS Seated Goya sce Ae tes OF OH Oa IS CER eR AA CRS OMLT Chis RA otic 16. A. decandra

5. Petiole much shorter. Flowers much smaller. Perianth 1-lipped. Stamens 6.
6. Leaf veins and veinlets closely foveolate-reticulate, prominent beneath. Utricle with 6 (or 2) glandular
ere IO SCCUS TOL WITRER) © nya eisetetae <5) ban coda ein wie in a wate ech Tee 4. A. foveolata
6. Leaf veins and veinlets loosely reticulate or transverse, distinct or rather faint beneath. Utricle with 2

glandular bodies inside.

7. Leaf undersurface papillate (under the binocular). Perianth without a stipe; utricle 3—6 mm long, tube

2.5—5 mm long, limb 11—12 mm long. Seeds not winged .................... 6. A. minutiflora
7. Leaf undersurface not papillate. Perianth with a distinct stipe; utricle 3—9 mm long, tube 5—10 (—15)
mm long, limb 20—30 (—40) mm long. Seeds distinctly winged .................... 9. A. tagala
KEY TO THE SPECIES
Philippines
1. Leaves deeply 3-lobed, in shape often resembling the letter ‘W’....................000- 1. A. jackii

1. Leaves entire, very rarely remotely minutely toothed (A. philippinensis).
2. Leaf base obtuse or rounded, subacute, or cuneate. (Plants erect, up to c. 1 m high).
3. Perianth with tube and limb bent backward and parallel to the utricle. Utricle 35 mm long, tube 10-15 mm
long, limb 45 mm long. Seeds verrucose on both surfaces. Petiole 6-10 mm long 17. A. samarensis
3. Perianth straight. Utricle 6-7 mm long, tube S—15 mm long, limb 18—25 mm long.
4. Leaves glabrous on both surfaces. Petiole 4—8 (—20) mm long. Seeds verrucose beneath and marginal

CUS! CE DNs Fe a lon ee a ee MEAS err PR ALES 0 18. A. philippinensis
4. Leaves glabrous above, sparsely short-hairy beneath. Petiole (20—) 30—40 mm long. Seeds verrucose
only on the marginal part beneath, smooth above ..................:ece eee eeees 19. A. humilis

2. Leaf base distinctly cordate, sometimes slightly or shallowly cordate, very rarely associated with some ob-
tuse ones (A. sericea).
5. Erect plants up to | m high. Petiole 2-10 mm long.
6. Leaf base deeply cordate; auricles often overlapping or surrounding the stem .. 20. A. macgregorii
6. Leaf base slightly or shallowly cordate, rarely associated with some obtuse ones; auricles obscure or 0
21. A. sericea
5. Twiners or climbers. Petiole 20—70 mm long.
7. Leaf veins and veinlets foveolate-reticulate, prominent beneath. Utricle with 6 (or 2) glandular bodies
panide.Seeds not withsed® 2517 8 5 eh eal. Sas 5. A PERS SE a oe a ee eae 4. A. foveolata
7. Leaf veins and veinlets often loosely, rarely closely, reticulate, or transverse, distinct or rather faint be-
neath. Utricle with 2 glandular bodies inside. Seeds distinctly winged (not known in A. /eyfensis).
8. Leaf veins and veinlets (closely reticulate) densely covered with minute hairs beneath (handlens or bi-
nocular). Seeds (proper) verrucose on both surfaces.......... 0000 eee e ee eees 5. A. zollingeriana
8. Leaf veins and veinlets (loosely reticulate) sparsely short-hairy or (nearly) glabrous beneath.
9. Perianth tube 20 mm long, limb 50-60 mm long. Mature seeds not known ..... 22. A. leytensis
9. Perianth tube 5—10 (—15) mm long, limb 20-30 (—40) mm long. Seeds verrucose beneath, less so
BV io Se SO WIG IRS SGN Abate SMe teak td eke a cig etn eae ae 9. A. tagala

KEY TO THE SPECIES
New Guinea and neighbouring islands

1. Leaves deeply 3-lobed, in shape often reminding of the letter ‘W'.. 0.0... cece cee eee es 1. A. jackii

1. Leaves entire.

2. Leaf veins closely reticulate, prominent beneath. Fruits cylindric or oblong, 2.5—4.5 by 2—3 cm, minutely
granular. Leaves cordate, auricles usually much overlapping; apex acuminate .... 23. A. crassinervia

88 FLORA MALESIANA

[ser. I, vol. 10!

2. Leaf veins loosely reticulate and some crossbar-like, often slightly elevated beneath.

3. Perianth limb 6-lobed. (Leaf base cordate).....

3. Perianth limb 3-lobed or 1-lipped.

Sti oh eee. Qe eee 24. A. schlechteri

4. Perianth without a stipe; limb 3-lobed. Seeds not winged.
5. Flower buds or perianth lobes with the apical 20—30 mm strongly contracted or narrowed and often

tail-like when dry (easily broken).

6. Perianth lobes suborbicular, c. 10 mm @, with an apical tail-like apex c. 20 mm long

11. A. momandul

6. Perianth lobes triangular to narrow-triangular, 35—50 by 12—14 mm at the base, apical 25-30 mm

oftenttal-hkesvhent diverse meee

4. 2a aba its fee 25. A. dielsiana

5. Flower buds or perianth lobes without a tail-like apical part. Perianth lobes triangular, 5—6 by 8—10

mm, apex OLHWS OF WKS 55 0o50000n00nnn 04

» sava.racp Joanne We rh.a) RAS, Mena SR oe 26. A. engleriana

4. Perianth with a distinct stipe; limb 1-lipped. Seeds distinctly winged.
7. Leaves glabrous. Perianth limb obovate or oblanceolate, 17—20 by 8—12 mm. Seeds (incl. wing) trans-
verse-oblong, 6—11.5 by 12—16 mm, smooth on both surfaces.............. 27. A. gaudichaudii
7. Leaves minutely hairy or sparsely short-hairy beneath. Seeds (incl. wing) triangular, 6-10 by 5—10

mm, usually verrucose on both surfaces.

8. Leaves triangular or deltoid in outline, minutely hairy beneath. Inflorescences 2—3 (—15) mm long;
internodes condensed, obscure or invisible; bracts minute, c. 1 mm long. Perianth lobes oblong, c.

2 OumimelOnsiers 5 Ace een se ee:

AM: DY Rae Sa oe ere 28. A. linnemannii

8. Leaves variable in shape and size, ovate, ovate-oblong, rarely suborbicular, sparsely shortly hairy be-
neath. Inflorescences 20—60 mm long; internodes spacious, distinct; bracts up to 10 mm long. Peri-

anth lobes lanceolate, 20—30 (—40) mm long

1. Aristolochia jackii StEUD. Nom. Bot. ed. 2, 1
(1840) 132; Mia. Sum. (1860) 150; Dinc Hou, Blu-
mea 29 (1983) 230. — A. hastata Jack, Malay Misc.
2, 7 (1822) 6 [reimpr. in Hook. J. Bot. 1 (1834) 362;
GriFF. Calc. J. Nat. Hist. 4 (1843) 358; TRUBNER,
Oriental Series II, 2 (1887) 249], nom illeg., non
H.B.K. (1817), nec NuTTALL (1818); KLotzscu, Mo-
natsber. Akad. Berl. (1859) 597, Jack sphalm. as
JACQUIN, excl. ZOLLINGER 2744; DUCHARTRE in DC.
Prod. 15, 1 (1864) 482; Merr. J. Arn. Arb. 33 (1952)
217. — A. ungulifolia Masters, J. Linn. Soc. Bot.
14 (1875) 494; Gard. Chron. n.s. 14 (1880) 116, f. 28;
Hook. f. in Curtis’ Bot. Mag. 121 (1895) t. 7424;
RIpDLey, J. Str. Br. R. As. Soc. n. 33 (1900) 126; KinG
& GAMBLE, J. As. Soc. Beng. 75, ii (1912) 30;
ScHMIpT, Bot. Jahrb. 58 (1923) 488; RipLey, FI.
Mal. Pen. 3 (1924) 18; ScumipT in E. & P. Nat. Pf.
Fam. ed. 2, 16b (1935) 241; HeNb. Mal. Nat. J. 6
(1951) 422, f. 381C. — A. tripartita Back. Trop. Na-
tuur 8 (1919) 161 & 165, f. 14; Bull. Jard. Bot. Btzg
III, 2 (1920) 322; ScumipTin E. & P. Nat. Pfl. Fam.
ed. 2, 16b (1935) 241; Back. & Baku. f. Fl. Java 1
(1963) 162. — Fig. 9.

Undershrub, spreading or twining, up to 10 m
high. Branches. obscurely angular, 3—5 mm a, gla-
brous. Leaves chartaceous or subcoriaceous, broadly
or transversely ovate or suborbicular in outline,
11—23 by 15—24 cm, deeply 3-lobed, in shape often
reminding of the letter ‘W’, glabrous; base emargi-
nate, subcordate, or almost truncate, rarely cuneate;
midlobe usually obovate- or ovate-oblong, 5.5—20
by 6.5—10 cm, apex obtuse or acute, rarely short acu-
minate; lateral lobes oblong, falcate or semilunar,

Sarniiaeies Sieieel oGsiis i eee 9. A. tagala

curved upward, sometimes spreading almost hori-
zontally, 4—15 by 2.5—6 cm, rounded or obtuse at
the apex; midrib with 2 or 3 pairs of lateral nerves;
basal nerves 2, each once or twice branched (the leaf
seemingly 5—7-nerved); nerves elevated and promi-
nent beneath, rather less so above; veins loosely
transverse or reticulate, slightly elevated on both sur-
faces; petiole 2—7 cm, glabrous. /nflorescences in ax-
ils of foliage leaves, spiciform or racemiform, up to
c. 7 (—25) cm long, internodes distinct; bracts rather
loose, ovate, c. 1.5 mm long, glabrous beneath,
short-hairy above. Pedicels and ovary 1.5—3 cm, gla-
brous. Perianth purple-brown or -red, or purple,
slightly curved, 7.5—11 cm long, glabrous outside;
utricle ellipsoid or obovoid, 2—3.5 cm long, with a
stipe of 6—7 mm, inside hairy, with 2 ellipsoid, glan-
dular bodies; tube 2—3.5 cm long, hairy inside; limb
1-lipped, oblong or ovate-oblong, elliptic, or spathu-
late, erect or reclined, 3.5—4.5 by 2—2.5 cm, tomen-
tose or villous on the upper surface and mouth of the
tube. Stamens 6; anthers ellipsoid-oblong, c. 1.5 mm
long. Style column 5—7 mm long, 6-lobed; lobes con-
ical, with a prominent annular ring at their base.
Capsules oblong, 5—6 by 2.5 cm, 6-angular, gla-
brous. Seeds triangular-orbicular, 4—5 by 5—7 mm
(excl. wing), lower surface smooth except a few warts
at the apical and basal ends, upper surface slightly
ridged at the centre; marginate, the wing 3—5 mm
wide.

Distr. Malesia: N. Sumatra (Medan, Sibolangit,
Asahan, Natal), Malay Peninsula (Pahang, Jarak I.
in Malacca Strait, Singapore), Java (Preanger, Nusa
Kambangan I., Mt Wilis), Borneo: Sabah (Labuan,

1984]

ARISTOLOCHIACEAE (Ding Hou) 89

Fig. 9. Aristolochia jackii SteuD. NW. Kalimantan,
near Njarumkop (Photogr. Father A. ELsENER, HS2,
Oct. 1964).

Tuaran), NW. Kalimantan (Njarumkop), SW. Phi-
lippines (N. Palawan), New Guinea (?) (Sepik and
Madang Distr.). Cultivated in Hort. Bog. sub n. XV-
D-46.

Ecol. In forest, sometimes in swampy forest be-
hind the sea coast, from sea level up to 1200 m. Fi.
Feb., June, Nov., fr. Jan., Feb., April, Sept., Oct.

Vern. Sabah: fawayagon, Tuaran; New Guinea:
bagup, Sepik.

Notes. Closely related to A. curtisii, sharing
deeply digitately 3-lobed leaves, spaced flowers and
bracts, and a 1-lipped perianth, but easily distin-
guished from it by a truncate, emarginate or subcor-
date leaf base, much smaller bracts (c. 1.5 mm), a
longer perianth (7.5—11 cm), and winged seeds.

Professor JUMALON (Cebu-city, Philippines), who
raised plants from seeds from Palawan, recorded
that leaves can attain 50 by 30 cm, or even more.

2. Aristolochia curtisii Kinc, Ann. Bot. Gard. Calc.
5 (1896) 161, t. 195; Gamaie, Kew Bull. (1910) 78;

KING & GAMBLE, J. As. Soc. Beng. 75, ii (1912) 32;
RIDLey, Fl. Mal. Pen. 3 (1924) 18; Dinc Hou, Blu-
mea 29 (1983) 227.

Climber up to 5 m high. Branches obscurely sul-
cate or slightly angular, c. 3 mm @. Leaves charta-
ceous, deeply 3-lobed (broadly hastately 3-lobed
when young), 10—23 by 10—30 cm; base cuneate; gla-
brous on both surfaces; middle lobe oblanceolate,
8—18 by 2—6.5 cm, acuminate; lateral lobes spathu-
late, slightly incurved, 6.5—14 by 2.5—5.5 cm, round-
ed at the apex; nervation pedately flabellate,
3-nerved at the base; outer nerves at first along the
margin, then giving off: a) 2 interior nerves to the
central lobe and each of them ascending upwards to
the apex, and b) often 2 exterior nerves for each of
the outer lobes, respectively, extending to the apex;
midrib and nerves elevated and prominent beneath,
rather less so above; veins loosely transverse and reti-
culate, distinct beneath, rather faint above; petiole
5—10 cm, subterete. /nflorescences in the axils of fo-
liage leaves, 1—3, spiciform, up to 6.5 cm long, gla-
brous, internodes distinct; bracts amplexicaul, con-
spicuous, reddish, ovate to lanceolate, 7—10 (—15)
mm long, glabrous. Pedicel and ovary 6—8 mm long,
glabrous. Perianth blue and crimson, straight, with
obscure venation, glabrous; utricle ellipsoid, 20 by 8
mm; tube cylindric, 10 by 1 mm; limb 1-lipped, lin-
ear, 20 by | mm. Stamens 6, filaments with very
small anthers. Style column very short, 6-lobed.
Capsules oblong, 3—4 by 1.5 cm, obscurely 6-ribbed,
obtuse. Seeds broad-ovate, not winged, c. Sby 4mm,
granular on both surfaces, funicle spindle-shaped, c.
4.5 by 2 mm.

Distr. Peninsular Thailand (Khaw Pok Hill,
Khsoon) and in Malesia: Malay Peninsula (Penang).

Ecol. In dense forest, 1S0—450 m. Fi. March, fr.
June—August.

Note. Closely allied to A. jackii STEUD.

3. Aristolochia singalangensis KorTH. ex DinG Hou,
Blumea 29 (1983) 224, f. 1, 2a, 3c & d, 7a.

Liana up to 20 m high. Stems terete or slightly flat-
tened, 1—1.5 (rarely more) cm @; branchlets tomen-
tose or villous, glabrescent. Leaves subcoriaceous,
suborbicular, broadly ovate, sometimes ovate, rarely
ovate-oblong, (14—) 24—33 by (6—) 11—24 cm; apex
acuminate or shortly so; base cordate, sinus 1.5 (—3)
cm deep; upper surface pubescent on the midrib and
nerves; undersurface villous or densely tomentose,
glabrescent; basal nerves one pair reaching upward
to halfway or higher; lateral nerves pinnate, 4 or 5
pairs, prominent beneath, distinct, sometimes de-
veins crossbar-like or reticulate,
slightly elevated with distinct areoles beneath, rather
l4cm,c.
5 mm @, villous or tomentose, glabrescent. /n/lores-

pressed above;
obscure above; petiole stout, terete, (3—) 6

cences cauligerous, solitary, racemiform, axis c. 4cm

90 FLORA MALESIANA [ser. I, vol. 10!

Fig. i0. Aristolochia singalangensis KorTH. ex D1nG Hou. Photograph of a coloured drawing in L made after
the living plant; leaves x 1/3, twig with flowers and fruit x 2.

1984]

long, internodes spacious, tomentose or densely pu-
bescent; bracts small, triangular, c. 3 mm long,
densely pubescent. Pedicel and ovary up to 7 cm
long, tomentose or pubescent. Perianth pale yellow-
ish green, pubescent outside; utricle cylindric, c. 7 by
1—1.5 cm, the apical 1—1.5 cm strongly bent back-
ward; tube cylindric, 4 by 0.6—1.2 cm, closely paral-
lel to the utricle; limb deeply 3-lobed, lobes triangu-
lar, 3 by 4.5—6 cm, apex obtuse. Stamens and style
column unknown. Capsules elongate-oblong, 14—15
by 2.5—3 cm, 6-ridged, dehiscing from the apex
downward, tomentose or pubescent. Seeds convex
on the lower side and ovoid-like in side view or seen
from beneath, not winged, c. 8 by 5 mm, deeply con-
cave on the upper side with a prominent, central,
longitudinal, septum-like funiculus, testa smooth on
both surfaces.

Distr. Malesia: Sumatra (Atjeh, near Pematang
Siantar, Mt Singalang, Palembang).

Ecol. At edge of grassy marshland in the forest,
along a trail in depleted forest, and on flat forest
ridge, (c. 350—) 750—1700 m.

Taxon. A. singalangensis is closely allied to the
Himalayan A. griffithii DUCHARTRE and the Chinese
A. kwangsiensis LiANG with which it shares all essen-
tial structural characters of the leaves, flowers, fruit
and seed. It can be distinguished from them by the
different size of the flora parts and the deeply
3-lobed perianth, of which the lobes are pale yellow-
ish green on the inner surface (not pinkish purple or
purple).

Sterile specimens may resemble Phyfocrene species
(Ucacinaceae), but can readily be distinguished by the
absence of an abscission zone in the petiole, hence
leaving no scar after withering.

Style column and stamens are as yet unknown.

4. Aristolochia foveolata Merr. Philip. J. Sc. 13
(1918) Bot. 280; En. Philip. 2 (1923) 119; IGARASHI,
Food PI. Papilionidae (1979) t. 26 (fig. on the lower
right) & 27, as Aristolochia sp. 2; Liu & Lat, Quart.
J. Taiwan Mus. 33 (1980) 247; Dinc Hou, Blumea 29
(1983) 227. — A. kaoi Liu & Lat, Fl. Taiwan 2 (1976)
573, t. 411; Hsu(ed.), The Rare & Threatened Plants
of Taiwan (1980) 45, col. phot.

Twiner up to 10 (—40) m high. Old stems terete,
1.5—2 cm @, bark corky, longitudinally fissured or
ridged. Branches terete, 2—6 mm @, striate, glabrous.
Leaves chartaceous to coriaceous, ovate to lanceo-
late in outline, sometimes broad-ovate, rarely sub-
orbicular, 7—18 (—24) by (3—) 4—8 (—21) cm; apex
acuminate, rarely cuspidate; base cordate, sinus
1—2.5 cm deep, sometimes auriculate with auricles
overlapping, rarely shallowly cordate, concave, or
subtruncate (especially when young); glabrous
above, densely puberulous beneath; nervation pal-
mate, appearing as 5 (—7)-nerved; inner pair of

ARISTOLOCHIACEAE (Ding Hou) 91

nerves nearly reaching the apex; outer pair much
shorter, branched at the base with | or 2 branches ex-
tending to the margin or auricles; nerves prominent
beneath, distinct above, joined by closely foveo-
late-reticulate and crossbar-like veins and veinlets;
veins and veinlets slightly elevated and prominent be-
neath, distinct or rather faint above; petiole 2—4
(—7.5) cm, glabrous. /nflorescences in axils of leaves
or cauligerous, often with very short branches, inter-
nodes hardly visible and flowers almost fasciculate;
bracts lanceolate, 4-5 mm long, minutely hairy on
both surfaces. Pedicel and ovary 28—40 mm long,
slightly twisted, glabrous. Perianth maroon or pur-
ple-brown, at first straight or horizontal then curved,
veins faint, glabrous outside, with scattered, glandu-
lar hairs inside; utricle subglobose, c. 7 mm @, not
stiped, with 6 (or 2) glandular, ellipsoid bodies (c. 2
by 1 mm) (6, rarely 2, depressions shown on the outer
surface); tube 10—14 by 2.5 mm; limb 1-lipped,
ovate-oblong, 30 by 14—16 mm. Gynostemium c. 2.5
mm long. Stamens 6; anthers oblong, c. 1 mm long.
Style 6-lobed, lobes triangular, 1.5 mm long, with an
annular ring at the base. Capsules cylindric or obo-
void, not angular or ridged, 2.5—4 by 1.5 cm, gla-
brous (minutely granulate examined with hand lens).
Seeds triangular, c. 5 by 4mm, not winged, verrucose
on both surfaces; funicle broadened, membranous,
and covering the upper surface.

Distr. China (Taiwan); in Malesia: NE. Suma-
tra, Malay Peninsula (Trengganu), Borneo (Sabah:
Mt Kinabalu, Sandakan; Sarawak: Upper Rejang R.
and Kuching; Kalimantan: Landak R.), Philippines
(Catanduanes and Palawan). Fig. 11.

Ecol. In primary, sometimes in secondary forest,
often at low and medium altitudes, sometimes found
at 1500-2100 m. Fi. May—Aug., Dec., fr. May,
Oct:, Dec:

Taxon. The species resembles A. faga/a in leaf

Fig. 11. Localities of Aristolochia foveolata MERR.

92 FLORA MALESIANA

[ser. I, vol. 10!

shape (usually with cordate base) and the 1-lipped
perianth but is easily to distinguish from that species
by the palmate nervation of the leaves, the undersur-
face being distinctly foveolate-reticulate and densely
puberulous, the perianth which is not strongly con-
tracted and stipe-like at base, and the immarginate
seeds.

Notes. The leaves are rather variable in shape
and size (ovate to lanceolate, rarely suborbicular,
and I = 7/3, 94/5, 12/9, 18/11, 24/21 cm) and leaf
base (deeply cordate with auricles sometimes over-
lapping to cordate or subtruncate). This variability
may well occur in one specimen, as was observed in
the field and in specimens which I cultivated.

The species is often collected by entomologists as
the larvae of the butterfly Trogonoptera brookiana
feed on it. Near Berastagi (NE. Sumatra) it is even lo-
cally cultivated for this purpose.

5. Aristolochia zollingeriana Mia. Fl. Ind. Bat. 1, |
(1858) 1066; DucHaARTRE in DC. Prod. 15, 1 (1864)
482; Back. Trop. Natuur 8 (1919) 162, f. 15, 165;
Koorp. Exk. Fl. Java 4 (Atlas) (1926) 591, f. 873;
SCHMIDT in E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935)
241; Back. & BAKH. f/f. Fl. Java 1 (1963) 162; DinG
Hou, Blumea 29 (1983) 232, f. 2c, 3a & b. — A. has-
tata (non Jack, sphalm. as JACQUIN) KLotzscH, Mo-
natsb. Akad. Berl. (1859) 597, quoad ZOLLINGER
2744. — A. ramosii MerRR. Philip. J. Sc. 29 (1926)
478. — A. kankauensis Sasaki, Trans. Nat. Hist.
Soc. Form. 21 (1931) 251; Liu & Lat, Fl. Taiwan 2
(1979) 572; Hsu(ed.), The Rare & Threatened Plants
of Taiwan (1980) 45, col. phot. — A. roxburghiana
ssp. kankauensis (SASAKI) KiITAMURA, Acta Phyto-
tax. Geobot. 20 (1962) 135. — A. tagala (non CHA-
Misso) Hatus. Fl. Ryukyu (1971) 243; WALKER, FI.
Okin. S. Ryukyu Is. (1976) 424. — A. tagala var.
kankauensis (SASAKI) YAMAZAKI, J. Jap. Bot. 50
(1975) 341, as ‘hankaoensis’.

Undershrub or a twiner up to 5 m high. Branches
terete, c. 3 mm Q, sulcate, glabrous. Leaves charta-
ceous, variable in shape and size even on one speci-
men, ovate or ovate-oblong, sometimes deltoid, reni-
form, or suborbicular, 4—15 by 2.5—12 cm; apex
usually acuminate, rarely acute or obtuse; base cor-
date or shallowly cordate, sometimes sagittate, auri-
cles separate from each other, rounded at the end, si-
nus 1—3 cm deep, glabrous above, minutely puberu-
lous beneath; midrib and basal nerves palmately
5-nerved, the midrib with 1—3 pairs of lateral nerves;
basal nerves 2 pairs, the inner pair ascending to 2/3,
sometimes almost to the apex of the blade, the outer
pair much shorter, spreading toward the margin at
the basal part of the blade, each of them with a few
side branchlets; nerves slightly elevated beneath, dis-
tinct or sometimes rather faint; veins closely reticu-
late, some transverse, distinct or sometimes rather

faint beneath; petiole 2—5S (—7) cm, glabrous. Jnflo-
rescences in the axils of foliage leaves, the very short
rachis (up to c. 16 mm long) with condensed inter-
nodes and bracts, glabrous; bracts ovate or triangu-
lar, c. 1 mm long, glabrous. Pedicel and ovary c. 18
mm, glabrous. Perianth green and dark purple, veins
longitudinal and reticulate, glabrous outside; utricle
subglobose, c. 7 mm g, with a distinct stipe of 3—5
mm, sparsely hairy inside, with 2 glandular, orbicu-
lar bodies (c. 1 mm @); tube straight or sometimes
bent at anthesis almost at a right angle with the utri-
cle and ovary, c. 13 by 2.5 mm, sparsely hairy inside;
limb 1-lipped, obovate-oblong, 23—30 by 10 mm,
longitudinally reflexed at anthesis, apex retuse, up-
per surface glabrous except the slightly hairy margin.
Stamens 6; anthers oblong, c. 1 mm long. Style col-
umn c. 4mm long, 6-lobed; lobes lanceolate, c. 1.5
mm long, with a distinct annular ring at the base.
Capsules short-cylindric, 2—3 by 1.5—2 cm, slightly
6-ridged, glabrous. Seeds triangular, winged, 4—5 by
4 mm (incl. the c. 0.6 mm wide wing), densely verru-
cose on both surfaces, upper surfaces sometimes
covered with a membranous funicle appendage.

Distr. S. Ryukyu Is., Taiwan; in Malesia: Philip-
pines (Luzon, Mindoro, Cebu, Bohol, Salupiri I.),
NE. Sumatra (Pematang Siantar) and SE. Java (Be-
suki Res.: Puger).

Ecol. In forest or along forest edges, sometimes
in rocky situations and on limestone hills, up to c.
300 m. Fi. April, Aug., Oct., Nov., fr. Feb.—June.

Vern. Philippines: wbi-ubihan, Tag.

Notes. The leaves of this species are very variable
in shape, size, texture, etc. The leaf base can be shal-
lowly cordate (with two divergent lobes), or sagittate
(with a deep sinus and two + parallel lobes). The
leaves on the specimens collected from Sumatra and
Java are ovate-oblong and sagittate at the base, but
those from the southern Ryukyus and Taiwan are
often deltoid, ovate, or suborbiculate, and shallowly
cordate at the base. However, there are intermediate
forms among the specimens from the Philippines;
sometimes various leaf forms are even found in one
specimen.

The polymorphism of the leaves in the present spe-
cies can be compared with that found in the well
known Japanese A. kaempferi WILLD.

Some of the leaf forms are similar to those of A.
foveolata, in shape and indumentum underneath,
but that species is quite different in flowers and seed.

A. zollingeriana closely resembles A. tagala, but
can easily be distinguished: in the first the undersur-
face of the leaves is minutely hairy and with distinct
areolation, in the latter it is sparsely short-hairy,
rarely glabrous and with obscure areolation. In flow-
er, fruit and seed the two are different.

The leaves of A. zollingeriana resemble those of
the Japanese A. kaempferi WiLLD.; the undersurface

1984]

of the leaves in the first is rather densely minute-
hairy, in the latter there are loosely appressed hairs.
Fertile specimens of A. kKaempferi differ by absence
of a stipe-like part of the perianth base, perianth
curved in the middle, more or less V-shaped, limb
suborbicular in outline, obscurely 2-lobed, and seed
concave-convex, not winged.

6. Aristolochia minutiflora RipLEY ex GAMBLE, Kew
Bull. (1910) 79, incl. var. dolabrata GAMBLE; KING&
GamsBLE, J. As. Soc. Beng. 75, ii (1912) 31; RIDLEY,
Fl. Mal. Pen. 3 (1924) 18.

Scandent shrub or climber, up to 10 m high.
Branches 3—10 mm a, sulcate, twisted, glabrous.
Leaves thin-chartaceous, lanceolate sometimes ovate
in outline, (5.5—) 12—14 by (2.5—) 5.5—7 cm; apex
acuminate or cuspidate; base cordate, sinus 7—20
mm deep, 12—20 mm wide, auricles rounded at the
base; glabrous above, loosely puberulous beneath;
basal nerves 2 (—3) pairs, palmate, slightly elevated
below, distinct above, inner pair of nerves ascending
obliquely and slightly curved inward, up to c. 2/3 of
the blade; veins usually loosely reticulate, some
transverse, rather fine, distinct beneath, faint above;
petiole 2—6.5 cm, glabrous. /nflorescences in the ax-
ils of foliage leaves, spiciform, up to c. 3.5 cm long,
internodes very short or obscure, sparsely puberu-
lous or almost glabrous; bracts lanceolate, 2-4 mm
long, shortly hairy on both surfaces. Pedicel and
ovary 9—12 mm, sparsely minutely hairy. Perianth
green, red and light grey, straight, sparsely shortly
hairy outside, glabrescent or almost glabrous; utricle
broad-ovoid or subglobose, 3—6 by 2.5—6 mm, not
stiped, sparsely hairy inside, with 2 ellipsoid, glandu-
lar bodies; tube 2.5—5 by | mm, short-fimbriate in-
side; limb 1-lipped, narrow-lanceolate to linear,
11—12 by 2—3 mm, veins loosely reticulate, rather
faint, glandular hairs usually at the lower half of the
upper surface. Sty/e column c. | mm long, obscurely
6-lobed, with a distinct, annular ring. Stamens 6; an-
thers oblong, c. 0.3 mm long. Capsules oblong-obo-
void, 1.7—2.5 by c. 1.2 cm, 6-ridged, distinctly trans-
versely rugose outside (marked by the seeds). Seeds
ovate, c. 5 by 4mm, not winged, granulate on both
surfaces.

Distr. Malesia: Malay Peninsula (Dindings, Pe-
rak, Johore), Borneo (Sarawak: Kelabit Highlands;
Sabah: Mt Kinabalu, Sandakan; NE. Kalimantan:
Mt Buduk Rakik).

Ecol. In primary forest, sometimes found in
swampy forest, or old secondary forest, from low-
land up to 1300 m. F/. March—Aug., /r. Aug., Sept.

7. Aristolochia glaucifolia RipLey, Kew Bull. (1925)

88; Dinc Hou, Blumea 29 (1983) 228, f. 2d.
Twiner up to c. 8 m high. Branches terete, 1.5—3

mm @. Leaves thin-chartaceous, ovate (in outline),

ARISTOLOCHIACEAE (Ding Hou) 93

5—13 by 4—6 cm; apex acuminate; base cordate, auri-
cles almost oblong, 1.5—3 by 1—2 cm, rounded, the
sinus 1.5—3.5 cm deep, glabrous above, lower sur-
face sparsely shortly hairy, sometimes glaucous;
basal nerves | pair, ascending upward to the apex,
each with 1—3 branches along the inner margin of the
auricles, slightly elevated beneath, faint above; veins
loosely transverse or reticulate, distinct beneath, ob-
scure or invisible above; petiole subterete, 3—7 cm,
glabrous. /nflorescences in the axils of foliage leaves,
rarely cauligerous, spiciform, up to 2 cm long, slight-
ly short-hairy; bracts ovate, 2-10 (—20) mm long,
sparsely puberulous on both surfaces. Pedicels and
ovary 6—13 mm long, sparsely puberulous, glabres-
cent. Perianth pale yellowish green and somewhat
purplish at the base, brownish, or dark brown,
straight, with loosely reticulate veins, glabrous out-
side; utricle broad-ellipsoid, 10-12 by c. 5 mm,
sparsely, shortly hairy inside, with 2 glandular, ellip-
tic bodies (1.5—2 mm long); tube cylindric, S—11 by
1—2.5 mm, sparsely hairy inside; limb 1-lipped, nar-
row lanceolate, 15—25 by 3—6 mm, shortly hairy on
the upper surface. Stamens 6; anthers oblong, c. 0.6
mm long. Sty/e column c. 1.5 mm long; lobes 6, tri-
angular, c. 0.5 mm long, with a continuous, annular
ring at their base. Capsules ellipsoid, 2.5—3 by 1.5
cm, slightly 6-ridged. Seeds triangular, 4 by 3.5 mm,
not winged, granular on both surfaces, central part
of the upper surface with a longitudinal ridge.

Distr. Malesia; Sumatra (northern part: Atjeh,
Mt Sinabung, Kabandjahe, Petani Valley, Sibolan-
git, Singgalang; central western part: Batu I., Mt Ke-
rintji, Pajakumbuh; southern part: Bencoolen).

Ecol. In primary forest, sometimes in old sec-
ondary forest, 500-1550 m. Fl. Feb.—Dec., /r.
March, May, Sept.

Note. The plants of this species are easy to recog-
nize by the rather thin-chartaceous, cordate-sagittate
leaves glaucous underneath. According to BRooKs
they are the food plant of Papilio helenae Cum.

8. Aristolochia klossii RipLey, Kew Bull. (1926) 78;
Dinc Hou, Blumea 29 (1983) 231.

Scandent shrub. Branches c. 6 mm @, twisted, sul-
cate, glabrous. Leaves chartaceous, broad-ovate or
deltoid in outline, 13.5—15 by 11.5—15 cm; apex ob-
tuse or acute; base cordate or shallowly cordate,
lobes rounded, the sinus 2—2.5 cm deep, 7-9 cm at
the widest part; glabrous on both surfaces; basal
nerves | pair, curved, ascending to the apex, each
basal nerve branched quite near the base, thus almost
forming another basal pair, elevated and prominent
below, distinct sometimes slightly depressed above;
veins loosely reticulate or transverse, slightly elevat-
ed below, faint above; petiole 4.5—6.5 cm, glabrous.
Inflorescences in the axils of foliage leaves, racemi-
form, up to 6 cm long, sparsely puberulous or almost

94 FLORA MALESIANA

[ser. I, vol. 10!

glabrous; bracts ovate, c. 4mm long, densely shortly
hairy on the margin. Pedicel and ovary 8—20 mm,
glabrous. Perianth deep crimson, white at base, limb
pinkish, edge crimson, straight, with distinct, longi-
tudinal and reticulate veins, glabrous outside, utricle
ellipsoid, 1O—15 by 5—8 mm, with a stipe of 3 mm,
sparsely hairy inside, glandular bodies 2, ellipsoid, c.
2 mm long; tube 16—19 mm long, with scattered,
glandular hairs inside; limb 1-lipped, obovate-
oblong 20—25 by 10—14 mm, apex slightly retuse or
mucronate, with scattered, glandular hairs on the up-
per surface. Stamens 6; anthers ellipsoid or oblong,
c. 0.7 mm long. Style column c. 4.5 mm long,
6-lobed; lobes with basal parts united, projecting and
forming an annular ring. Capsules unknown.

Distr. Malesia: Sumatra (Mentawei Is.: Sipora;
Bandarbaru). Twice collected.

Ecoles Oct.

Notes. The leaves resemble those of A. gaudi-
chaudii from the Moluccas and New Guinea.

More material and especially fruits are desirable.

9. Aristolochia tagala CHAmisso, Linnaea 7 (1832)
207, t. 5, f. 3; Kiotzscu, Monatsb. Akad. Berl.
(1859) 597; DucHaARTRE in DC. Prod. 15, 1 (1864)
480; F.-VitLt. Nov. App. (1880) 174; Vipat, Phan.
Cuming. (1885) 138; Rev. Pl. Vasc. Filip. (1886) 218;
SOLEREDER, Bot. Jahrb. 10 (1889) 464; Fors. &
HemsL. J. Linn. Soc. Bot. 26 (1891) 363; Koorp.
Minah. (1898) 567; MerrR. Publ. Govt. Lab. Philip.
27 (1905) 72; KING & GAMBLE, J. As. Soc. Beng. 75,
ii (1912) 30; Merr. Fl. Manila (1912) 186; Int.
Rumph. (1917) 209; Sp. Blanc. (1918) 135; Back.
Trop. Natuur 8 (1919) 151, f. 7-12, 166; Brown,
Bull. Bur. For. Philip. (1921) 183; C.T. Wuite,
Proc. R. Soc. Queensl. 34 (1922) 30; MErRR. En. Phi-
lip. 2 (1923) 120; RmpLey, Fl. Mal. Pen. 3 (1924) 18,
excl. f. 136 (=A. indica); GAMBLE, Fl. Pres. Madras
pt 7 (1925) 1202; Moore, J. Bot. 63 (1925) Suppl. 83;
HEYNE, Nutt. Pl. (1927) 597; Korpzumi, Fl. Symb.
Orient.-Asiat. (1930) 16; Merr. Gard. Bull. S. S. 8
(1935) 131; ScHmipTin E. & P. Nat. Pfl. Fam. ed. 2,
16b (1935) 241; Burk. Dict. 1 (1935) 239; Merr.
Comm. Lour. (1935) 142; KaANnILALc.s. Fl. Assam 4
(1940) 28; HoEHNE, FI. Brasil. Fasc. 6 (vol. 15, 2)
(1942) 136, t. 120; MERR. & PERRY, J. Arn. Arb. 23
(1942) 384; ibid. 29 (1948) 153; Quis. Medic. PI. Phi-
lip. (1951) 255; HEND. Mal. Wild Fl. (1951) 421, f.
381A & B; L.S. Soir, Proc. R. Soc. Queensl. 68
(1957) 45; Lianc, Acta Phytotax. Sin. 13 (1975) 17,
f. 1,5; BAcK. & BAKH./f. Fl. Java 1 (1963) 163; Kao,
Fl. Hainan 1 (1964) 327, f. 163; H. KENG, Mal. Seed
Pl. (1969) f. 58; ibid. ed. 2 (1978) f. 59; LIANG, Acta
Phytotax. Sin. 13 (1975), 17, f. 1, 5; Lro & Lay, FI.
Taiwan 2 (1976) 576; ANoNyMous, Icon. Cormo-
phyt. Sin. 1 (1972) 548, f. 1096; IGARAsuHI, Food PI.
Papilionidae (1979) t. 12; PERRY, Medic. Pl. E. &

SE. Asia (1980) 47; Dinc Hou, Blumea 29 (1983)
232. — Peponaster major RumpH. Herb. Amb. 5
(1747) 474. — A. acuminata LamMK. Encycl. 1 (1783)
254; WILLD. Sp. Pl. 4 (1805) 157; SPRENG. Syst. Veg.
3 (1826) 751; BL. En. Pl. Jav. 1 (1827) 81; Roxs. FI.
Ind. ed. Carey 3 (1832) 489; WiGut, Icon. Pl. Ind.
Or. (1844) t. 771; Mia. Fl. Ind. Bat. 1, 1 (1858) 1066;
DALZELL & GIBSON, Bombay FI. (1861) 224. — A.
longifolia Roxs. [Hort. Beng. (1814) ‘102’]; Fl. Ind.
ed. Carey 3 (1832) 490. — A. timorensis DECNE,
Ann. Mus. Hist. Nat. Paris III, 3 (1834) 368; Herb.
Timor. Descr. (1835) 40; Mig. Fl. Ind. Bat. 1, 1
(1858) 1066; KLotzscu, Monatsb. Akad. Berl. (1859)
597; DUCHARTRE in DC. Prod. 15, 1 (1864) 481;
BRITTEN in Forbes, Wand. (1885) 515; ENGL. Bot.
Jahrb. 7 (1886) 453; Laut. Bot. Jahrb. 52 (1914) 105.
— A. indica (non L.) Bianco, FI. Filip. (1837) 282;
ed. 2 (1845) 197; ed. 3, 1 (1877) 349; F.-ViLL. Nov.
App. (1880) 174. — A. roxburghiana KLotzscu, Mo-
natsb. Akad. Berl. (1859) 596, excl. WALLICH/N. 2704
(=A. indica); F.-ViLt. Nov. App. (1880) 174; Du-
CHARTRE in DC. Prod. 15, 1 (1864) 480, incl. B
angustifolia DUCHARTRE; Hook. f. Fl. Br. India 5
(1886) 75; SOLEREDER, Bot. Jahrb. 10 (1889) 460;
Laut. Bot. Jahrb. 52 (1914) 106; BoLtpinGu, Zakfl.
Landb. Java (1916) 37; Koorp. Exk. Fl. Java 4 (At-
las) (1926) 590, f. 872; ScHmipT in E. & P. Nat. Pf.
Fam. ed. 2, 16b (1935) 241. — A. moluccana Du-
CHARTRE in DC. Prod. 15, 1 (1864) 438, nom. su-
perfl., new name for A. longifolia Roxs. (1832). —
A. japonica Mig. Ann. Mus. Bot. Lugd.-Bat. 2
(1866) 136; Prol. Fl. Jap. (1866) 68. — A. megalo-
pDhylla K. Scu. in K. Sch. & Hollr. Fl. Kais. Wilh.
Land (1889) 104; Wars. Bot Jahrb. 13 (1891) 300; K.
Scu. Notizbl. Berl.-Dahl. 2 (1898) 113; K. Scu. &
Laut. Fl. Deut. Schutzgeb. Stidsee (1900) 302; RE-
CHINGER, K. Ak. Wiss. M.-N. Kl. Wien 89 (1913)
549; Laut. Bot. Jahrb. 52 (1914) 106; ScHmipTin E.
& P. Nat. Pfl. Fam. ed. 2, 16b (1935) 241; PEEKEL,
Illustr. Fl. Bismarck Arch. (MS) 4 (1947) sub f. 528.
— A. mindanaensis Wars. in Perkins, Fragm. FI.
Philip. (1905) 169; Merr. En. Philip. 2 (1923) 119.
— Fig. If.

Twiner, up to 20 m high. Branches terete, slightly
furrowed, 3—5 mm g, glabrous. Leaves variable in
shape and size, ovate, ovate-oblong, rarely suborbi-
cular, 6—20 (—27) by 4—10 (—16) cm; apex acute or
acuminate; base cordate, auricles rounded, often
connivent, the sinus up to 3.5 cm deep; glabrous or
nearly so above, sparsely shortly hairy, or subgla-
brous beneath; basal nerves 2 pairs, the inner one as-
cending upward to more than 2/3 of the blade, simi-
lar in appearance to the lateral nerves, the outer one
much shorter and weaker, often branched; lateral
nerves 3—S pairs, elevated below, slightly elevated or
faint above; veins loosely reticulate or crossbar-like,
distinct below, obscure above; petiole 2—6 cm,

1984]

ARISTOLOCHIACEAE (Ding Hou) 95

slightly hairy. Jnflorescences in the axils of foliage
leaves, racemiform or paniculate, 2—6 cm long,
slightly hairy, glabrescent, or glabrous; bracts ovate
to lanceolate, up to 10 mm long, puberulous on both
surfaces. Pedicel and ovary 10—18 mm long, sparsely
hairy, glabrescent. Perianth pale yellowish with pur-
ple throat, pale or sordidly green with purple, purp-
lish, or dark reddish brown, straight or slightly
curved, venation faint, sparsely hairy, glabrescent
outside; utricle broad-ovoid or subglobose, 3—9 by
3—7 mm, with a stipe of 1—3 mm, hairy inside, with
2 glandular, ovoid bodies (c. 0.7 mm long); tube
$—10(—15) by c. 2.5 mm, hairy inside; limb 1-lipped,
lanceolate to narrow-lanceolate, 20—30 (—40) by 6—8
mm, apex acute, hairy on the upper surface. Style
column c. 3 mm long, 6-lobed; lobes conical, c. 1.5
mm long, at base forming an annular ring. Stamens
6; anthers oblong, c. 1 mm long. Capsules subglo-
bose, slightly pyriform, or oblong, 3—4 by 2—3 cm,
often 6-ridged, glabrous. Seeds triangular, winged,
8.5—10 mm long and wide (incl. the c. 2 mm wide
wing), granular beneath, much less so above; funicle
with membranous extension covering the upper sur-
face (dry state).

Distr. Widely distributed in India, Sikkim, Sri
Lanka, Bangladesh, Burma, Thailand, Cambodia,
Vietnam, China; Malesia (throughout), New Ire-
land, New Britain, Solomon Is., Australia (Queens-
land). Cultivated in Hort. Bog. sub n. XV-D-40 and
XV-K-AXI-9.

Ecol. In forests and thickets, often at low and
medium altitudes (0O—800 m), sometimes up to c.
1350 m (e.g. New Guinea). Fi. fr. all the year round.

Uses. The powdered roots are said to be tonic,
carminative, and emmenagogic, and a very efficient
remedy for infantile tympanites if they are pulverized
and applied to the abdomen (QuISUMBING, /.c.).

In Ambon, the leaves, ground with curcuma and
warmed, are smeared on the abdomen and the limbs
when they are swollen; or they are made into a paste
for use against such a skin disease as formication
(HeyNE, /.c.; BURKILL, /.c.).

Vern. Sumatra: béngkuh-béngkuh, képing-ké-
ping, olor féengkuh-képing, Simalur. Malay Peninsu-
la: akar ara bukit, akar kétola huta, akar pétola hu-
tan, M. Java: kalajar or kalaijar, S; kapassan, prod-
jon, pujan, J. Timor: wunbewa. Bunguran Is. (=
Groot Natoena): mili utan, M. Philippines (MER-
rit, 1923, /.c.): altdn, malaubi, parolparulan, tala-
taldrum, timbdngan, timbang-timbdngan, Tag.,
goan-goan, Bis., kamkamaulau, \|g., nagerus, taoin-
tdoin, \\k. Celebes: kunit, Manado. Moluccas: ja-
wepplewe, Weda, sasa baru, Ternate. New Guinea:
kobi, Garaina, kolura, Bangwe, mangkapdupdak,
Biak, sisidi, Merauke.

10. Aristolochia coadunata Back. Trop. Natuur 8

(1919) 154, f. 13, 167; Bull. Jard. Bot. Btzg III, 2
(1920) 320; Back. & Baku. f. Fl. Java 1 (1963) 164;
STEEN. Mountain Fl. Java (1972) sub t. 4, 1; DING
Hou, Blumea 29 (1983) 227. — Fig. 12.

a. var. coadunata.

Scandent, high liana, 10-50 m long. Branches
subterete or slightly flattened, 0.5—1.2 cm 9, young
parts densely pubescent, glabrescent. Leaves subco-
riaceous, ovate-oblong to lanceolate, rarely ovate,
7.5—33 by 4—12 cm; apex acuminate, short-acumi-
nate, or acute; base slightly cordate, basal lobes
rounded (and the sinus 0.5—1 cm deep, sometimes
obscure); upper surface pubescent especially on mid-
rib and nerves, glabrescent; undersurface villous or
densely tomentose, glabrescent; basal nerves one
pair, reaching upward to 1/3—1/2 of the blade, later-
al nerves 4—7 pairs, pinnate, veins rather closely reti-
culate; both nerves and veins elevated beneath, dis-
tinct or faint, sometimes depressed or bullate above;
petiole 3—9 cm, pubescent. /nflorescences in axils of
foliage leaves, rarely cauligerous, solitary or fascicu-
late, racemiform, up to 2 cm long, pubescent; bracts
ovate, c. 1.5 mm long, densely pubescent or tomen-
tose. Pedicel and ovary 4—8 mm, pilose. Perianth
dark purple with yellow throat, geniculate, sigmoid,
pubescent outside, venation obscure; utricle ovoid-
tubular, 2.5—3 by 0.7 cm, the apical part bent back-
ward, hairy at the lower half inside; tube cylindric,
3—4.5 by 0.6 cm, closely laterally in contact with the
utricle, the basal part inside slightly projecting into
the utricle cavity, almost glabrous inside; limb rim-
like, 1.5—3 cm @, the rim 0.5—1 cm wide, very ob-
liquely positioned on the tube, obscurely 3-lobed.
Stamens 6; filaments short; anthers oblong, 2—2.5
mm long. Style column 5—7 mm long; lobes 3, trian-
gular, c. 1 mm long. Fruit unknown.

Distr. Malesia: Sumatra (Berastagi; Mt Kerint-
ji), West to East Java (Priangan: Mts Malabar & Pa-
pandajan; Mt Lawu, above Pudjon, probably Mt
Kawi, SE. Mt Smeru, G. Pendil on Mt Idjen).

Ecol. In primary, occasionally in secondary,
mountain forest, 1000-2100 m. Fi. April, June,
Oct., Nov.

Taxon. Closely allied to the Himalayan A. sacca-
ta Wa... from which it differs by the smaller,
non-saccate flowers which have the perianth tube in
close contact with the utricle.

b. var. bosschai Back. Trop. Natuur 8 (1919) 154,
168; Bull. Jard. Bot. Btzg III, 2 (1920) 321; Back. &
BakH. f. Fl. Java 1 (1963) 164.

Similar to var. coadunata, except the perianth
limb entirely sulphureous.

Distr. Malesia; West Java (Preanger: Talun).
Only known from the type.

Ecol. In primary forest, 1600 m

96 FLORA MALESIANA [ser. I, vol. 10!

Fig. 12. Aristolochia coadunata Back. After a coloured drawing made for vAN STEENIS, Mountain Flora of
Java, xX 2/3. West Java, Mt Papandajan, Tegal Pandjang, July 1940 (vAN STEENIS 12625).

1984]

11. Aristolochia momandul K. Scu. in K. Sch. &
Hollr. Fl. Kais. Wilh. Land (1889) 105; K. Scu. &
Laut. Fl. Deut. Schutzgeb. Stidsee (1900) 302;
Laut. Bot. Jahrb. 52 (1914) 106; Dinc Hou, Blumea
29 (1983) 239, f. Sa, 6b, 7c. — A. pithecurus RIDLEY,
J. Bot. 52 (1914) 296; Moore, J. Bot. 61 (1923)
Suppl. 40; Scumiptin E. & P. Nat. Pfl. Fam. ed. 2,
16b (1935) 241; IGARAsHI, Food Pl. Papilionidae
(1979) t. 24. — A. gracilifolia ScHMipT, Bot. Jahrb.
58 (1923) 490; MerrR. & PERRY, J. Arn. Arb. 23
(1942) 383. — A. dictyophlebia MERR. & PERRY, J.
Arn. Arb. 29 (1948) 152. — Fig. 8b.

Twiner up to 6 m, sometimes scrambling on (tall)
tree top up to 30 m. Old stem terete, 2.2—4 cm 9;
branches terete, c. 4 mm 9, short-hairy, glabrescent.
Leaves chartaceous to coriaceous, lanceolate, ellip-
tic, or ovate, (7—) 10—25 (—43) by (2—) 3—11 (—21)
cm; apex acuminate, short-acuminate, rarely cuspi-
date; base subcordate, deeply cordate (especially
plants from rather lower altitude), or slightly con-
cave, rarely rounded or subtruncate (especially when
young, or in plants from rather higher altitude);
sinus 0O—1 (—5) cm; subglabrous or glabrous above,
shortly hairy especially on the midrib, nerves and
veins; nerves elevated and prominent beneath, less so
above; basal nerves | pair, ascending upwards to c.
1/2—2/3 of the blade, branched at base; lateral
nerves 3—5 pairs; veins reticulate or crossbar-like,
prominent beneath, distinct or sometimes faint
above; petiole (1—) 2—3.5 (—7) cm, densely hairy. Jn-
florescences in axils of leaves, or cauligerous, pani-
culiform, rachis 4—19 (or more) cm long, with spa-
cious internodes, densely hairy; bracts minute,
densely hairy. Pedicel and ovary c. 20 mm long,
densely hairy. Perianth straight or with slightly
curved tube, white or pale green with purple vena-
tion, dull wine purple with yellow or orange yellow,
longitudinal veins distinct, reticulations rather faint,
densely hairy outside, with scattered hairs inside;
with a contraction between perianth and ovary, no
stipe; utricle obovoid, c. 10 by 7 mm; tube gradually
enlarged toward apical part, c. 12 by 10 mm; limb an
obovoid body with a long filiform tail composed of
the ends of the lobes; limb 3-lobed (very rarely with
an additional filiform fourth lobe); lobes suborbicu-
lar, c. 10 mm 9, with an apical tail-like appendage
(easily broken off) c. 20 mm. Stamens 6; anthers ob-
long, 1.2—1.7 mm long. Gynostemium c. 3.5 mm
long; style 6-lobed; lobes triangular, 1.5—2 mm long,
each lobe with the basal part slightly extended out-
ward or downward. Capsules golden yellow or bril-
liant orange when ripe, oblong or ellipsoid, 5.5—9 by
3.5—4 cm, strongly 6-ridged, smooth (fine-granular
under a handlens); mostly found indehiscent, but ob-
viously finally dehiscing. Seeds triangular or deltoid,
9-10 mm long and wide, smooth on both surfaces,
not winged.

ARISTOLOCHIACEAE (Ding Hou) 97

Distr. Malesia: Moluccas (Halmahera), West
New Guinea (Sorong, Oransbari, Andai, Jappen,
Biak); Papua New Guinea (Sepik, Madang, Morobe,
Central Distr.), New Britain.

Ecol. In primary forest, beach forest, sometimes
occurring in limestone areas on rocky peak; at low
and medium altitudes, sometimes found up to 1650
m. Fl. Jan.—July, fr. Jan.—June, Sept., Oct.

Uses. Sap from the vine said to be used in ‘pape-
da’ (sago-porridge) (VINK BW 17561, L).

Vern. West New Guinea: ba, Oransbari lang.;
Papua New Guinea: momandul, Madang.

Notes. The leaves of the present species are very
variable, in shape, size, texture, and base, but flow-
ers and fruit are uniform, the tailed buds and lobes
being very characteristic.

As already pointed out by SCHUMANN it is related
to A. deltantha F. v. M. from Queensland, from
which it can be distinguished by the tail-like apex of
the perianth lobes; in A. del/tantha the perianth is
hardly lobed. The leaf variability is in both the same.

A. momandul was described on fruiting material.
In 1914 LAUTERBACH associated flowering material
with it (FoRBEs 621, the type of A. pithecurus Rip-
LEY). Because the species cannot be distinguished
from A. schlechteri on vegetative characters, it will
be essential to have fruit of the latter, to check
whether LAUTERBACH’s conclusion was correct. LAU-
TERBACH erroneously described his material to have
6 perianth lobes.

12. Aristolochia rumphii KosTELetzky, Allg. Med.-
Pharm. Fl. 2 (1883) 465; Merr. Int. Rumph. (1917)
209; HeyNE, Nutt. Pl. (1927) 596; pE Wit, Rumph.
Mem. Vol. (1959) 348; Perry, Medic. Pl. E. & SE.
Asia (1980) 47; Dinc Hou, Blumea 29 (1983) 232, f.
2b. — Radix puluronica RumpPH. Herb. Amb. 5 (1747)
476, t. 177. —A. indica (non L. 1753) L. in Stickman,
Herb. Amb. (1754) 25, quoad Radix puluronica
RuMpPH.; Fitet, Bot. Tuin Weltevr. (1855) 50; Bis-
SCHOP GREVELINK, PI. Ned. Ind. (1883) 268. — Fig.
15e.

Climber. Branches c. 2 mm @, smooth, glabrous.
Leaves thin-chartaceous, oblong, elliptic-oblong,
ovate-oblong, narrow-lanceolate, 7—12.5 by (l—)
3—5.5 cm; apex acuminate; base obtuse, sometimes
slightly cuneate, rarely truncate; glabrous, minutely
shortly hairy beneath, glabrescent; | pair of basal
nerves, ascending upward to about halfway the blade,
lateral nerves 4 or 5 pairs distinct sometimes faint be-
neath, rather faint above; veins loosely transverse or
reticulate, often faint beneath, obscure above; petiole
1.5—2.5 cm, glabrous or sometimes sparsely short-
hairy. /nflorescences in axils of foliage leaves, usually
very short, sometimes up toc. 5 cm long, racemiform
and with distinct internodes; bracts ovate, 1.5—2 mm
long, hairy on the margin, glabrescent, Pedicel and

98 FLORA MALESIANA [ser. I, vol. 10!

a x

=}

Fig. 13. Aristolochia papillifolia Dinc Hou. a. Habit, note lenticels, b. old stem with thick furrowed bark,
both x %, c. CS of ditto, note medullary rays and vascular bundles, x 2, d—e. opening of fruit, x %, f.
immarginate seed, dorsal and ventral (a—c SAN 34658, d—f SAN 17334). Courtesy of Blumea.

1984]

ARISTOLOCHIACEAE (Ding Hou) 99

ovary 10—30 mm, minutely hairy, glabescent. Pert-
anth green with brown limb, straight or with the limb
slightly bent, minutely hairy outside; utricle subglo-
bose, c. 6 mm @, with a distinct stipe of 5 mm, hairy
inside, 2 glandular, subglobose bodies (c. 0.3 mm 9);
tube 17 by 1.5 mm, sparsely hairy inside; limb ob-
long, 27 by 7 mn, hairy above, glabrescent, margin
reflexed. Sty/e column c. 2 mm long, 6-lobed; lobes
lanceolate, c. 1 mm long, basal parts projecting and
forming an annular ring. Stamens 6; anthers oblong,
c. 0.7 mm long. Capsules short cylindric, c. 2.2 by
1.5 cm, 6-ridged. Seeds triangular, 5 by 5—5.5 mm,
verrucose on both surfaces, not winged.

Distr. Malesia: SW. Celebes (Pangkadjene), Ka-
juadi Is. (halfway Flores), Lesser Sunda Islands
(Sumba, Flores, Timor), Moluccas (Ambon, not
seen; Kai and Tenimber Is.).

Ecol. In light forest, grass field and thickets, up
to c. 100 m. Fi. April—June, fr. March—July.

Uses. A decoction of a piece of the root or of
small twigs (less powerful than root) is used to treat
stomach-ache, spasm and constipation, and also in-
termittent fever. On trips travellers often take a piece
of root or lowest part of a stem about the length of
a finger and drink the decoction from it as tea (RUM-
PHIUS, /.c.; HEYNE, /.c.; PERRY, /.c.).

Vern. Akar pulurun, Ambon, tuhe tutunu, Ban-
da, warosbot, Tenimber I.

Notes. One fertile specimen, collected on a hill
above Endeh (Flores) by Father J.J. LOeTERs (n.
2092, L) has one flower and one fruit which match
those of the present species, though its leaves are very
narrow, c. 1 cm wide (fig. 15e).

A. rumphii is closely allied to A. indica from
Southeast Asia (mainly Ceylon and India), similar in
leaf characters and a 1-lipped perianth with a distinct
stipe-like base. It can be distinguished from A. indica
by the few-flowered inflorescences with distinct in-
ternodes, a longer pedicel and ovary, up to 3 cm (in
A. indica up to 1.5 cm), the longer stipe of the utricle,
c. 5 mm (2.5 mm in A. indica), the rather long peri-
anth tube up to 17 mm (against c. 8 mm), smaller
capsules, c. 2—3 cm (against 4—5 cm), and immargi-
nate seeds (distinctly winged in A. indica).

13. Aristolochia papillifolia Dinc Hou, Blumea 28
(1983) 346, f. 3, SA—C. — Fig. 13.

Stout climber, up to 15 m high. Old stem terete,
1.5—2.5 cm o. Branches subterete or slightly flat,
5—7 mm @, glabrous. Leaves subcoriaceous, broad-
ovate or ovate, 13-19 by 9—15.5 cm; apex acute or
short-acuminate; base almost truncate, slightly con-
cave, or rounded; glabrous above, glabrous but pa-
pillate in the areolae beneath; nerves palmate, basal
nerves 2 pairs, each nerve with a few lateral, oblique
branches, the inner pair of nerves reaching the apex,
the outer pair much shorter; nerves elevated and

prominent beneath, distinct above; veins loosely
transverse or reticulate, slightly elevated beneath,
faint above; petiole stout, terete, glabrous, 6—12 cm.
Bracts linear, 3.5—5 mm long, glabrous. Pedicel and
ovary 3—4 cm, glabrous. Flowers cauligerous, fasci-
culate. Perianth straight, veins loosely reticulate, gla-
brous outside; utricle ellipsoid or subglobose, 6—7.5
by 5—6 mm, flanged at the base, hairy inside, with 6
ellipsoid or orbicular, glandular bodies (c. 1.5 mm
long); tube 15—20 by 2 mm, hairy inside; limb
1-lipped, linear, 45—50 by 7—12 mm, papillate on the
upper surface. Stamens 6; anthers oblong, c. 0.7 mm
long. Style column 2 mm long, 6-lobed; lobes del-
toid, c. 1 mm long, basal parts forming a distinct an-
nular ring. Capsules cylindric, 6.5 by 1.2 cm, slightly
6-furrowed, obtuse on both ends. Seeds broad-ovate,
c. 4 by 3 mm, granular on both surfaces, not winged.

Distr. Malesia: Borneo (Sabah: Sandakan, Ton-
god, Tawau Distr.; Sarawak: Gunung Buri).

Ecol. Ridge top in primary forest, hill slope in
disturbed mixed dipterocarp forest, and also in sec-
ondary forest, up to c. 600 m. Fi. May, fr. June, Ju-
ly.

Notes. On the undersurface of the leaves the
areolae are papillose, and cavities containing one
stoma are surrounded by papillate cells (magnitude
x 40); this is a unique character among the Malesian
species.

The 1-lipped perianth has 6 glands inside the utri-
cle, a character only known from A. foveolata anda
West African species.

14. Aristolochia transtillifera DinGc Hou, Blumea 28
(1983) 348, f. 4, 5. — Fig. 2a—b.

Twiner. Branches terete, 3.5—6 mm 9, slightly stri-
ate, glabrous. Leaves subcoriaceous, ovate-oblong,
oblong-elliptic, rarely ovate, 16—20 by 6.5—10 cm;
apex short-acuminate; base obtuse, sometimes
slightly concave, or subtruncate, glabrous; basal
nerves 2 pairs, the inner one ascending to the apex,
elevated and prominent beneath, slightly depressed
above, the outer pair shorter and much weaker, as-
cending close to or along the margin, slightly elevat-
ed beneath, obscure or invisible above; veins distinct-
ly transverse, joined by closely reticulate veinlets,
slightly elevated beneath, obscure or invisible above;
petiole terete, 3—8 cm. Flowers unknown. /nfructes-
cences cauligerous, very short, condensed, bracteate,
knobby; bracts lanceolate, 2—6 mm long, slightly pu-
berulous on both surfaces, glabrescent. Capsules cy-
lindric, 3—3.5 by 1.2 cm, obtuse at both ends, gla-
brous, smooth (but finely verrucose under a hand
lens); pedicel c. 3.5 cm. Seeds triangular, 5 by 4—4.5
mm, subcordate at the base, not winged, finely
verrucose on both surfaces, funicle with a thin ex-
panded appendage covering the upper surface of the
seed,

100

FLORA MALESIANA

[ser. I, vol. 10!

Distr. Malesia: Borneo (Sabah: Beaufort Distr.).
Once collected.

Ecol. Hill side, primary forest, c. 30 m.

Notes. Though the species is as yet only known
in fruit, it stands out by the glabrous, subcoriaceous
leaves with obtuse to sometimes slightly concave base
and clearly trabeculate venation with close reticula-
tions.

It may be related to A. foveolata and A. papillifo-
lia, but differs by these leaf characters.

15. Aristolochia naviculilimba DING Hou, Blumea 28
(1983) 344, f. 2.

Liana, up to 15 m high. Branches subterete, 3—6
mm @, glabrous. Leaves chartaceous, elliptic, 10—15
by 5.5—8 cm; apex shortly acuminate; base rounded
or obtuse, rarely slightly concave; glabrous above,
minutely hairy beneath especially on the venation;
basal nerves 2 pairs, the inner pair and the midrib ele-
vated, prominent beneath, distinct above, ascending
obliquely and slightly curved nearly reaching the
apex of the blade; outer pair of basal nerves rather
weak, running along the margin; the longitudinal
nerves joined by 10—14 loosely transverse or slightly
curved nervules or stronger veins, slightly elevated
beneath, distinct or faint above; veins and veinlets
loosely reticulate, distinct or faint on both surfaces;
petiole c. 3 cm, glabrous. Flowers (detached) cauli-
gerous, on brachyblasts, internodes condensed, ob-
scure; bracts lanceolate or lineate, 3—4.5 mm long,
shortly hairy on both surfaces. Pedicel and ovary c.
3 cm long, glabrous. Perianth dull yellow with dark
purple stripes and markings, curved, distinctly longi-
tudinally 6-nerved or -veined, glabrous outside; utri-
cle subglobose, 10—12 by 9—10 mm, abruptly con-
tracted and collar-like at the base, densely hairy in-
side, with 2 ellipsoid, glandular bodies (3—4 mm
long); tube often at right angles with the utricle, cy-
lindric, c. 15 by 3mm, glabrous inside; limb 1-lipped,
ovate-oblong or lanceolate, naviculiform in side view
when young, 60—90 by 30—35 mm, longitudinally in-
curved, inner surface almost glabrous, except sparse-
ly short-hairy at the basal part. Stamens 6; anthers
oblong, c. 1 mm long. Sty/e column c. 3.5 mm long,

Fig. 14. Aristolochia naviculilimba Dinc Hov. Lat-
eral view of flower, x 2/3 (CLEMENS 20292).

6-lobed, with an annular ring at the base; lobes trian-
gular, c. 1.5 mm long. Capsules unknown.

Distr. Malesia: Borneo (Sarawak: Lundu Distr.,
Mt Poi; W. Kalimantan: Mt Kasian & Mt Liang Ga-
gang); 3 collections.

Ecol. In forest, c. 600 m. Fi/. June, Oct.

Notes. The epithet alludes to the boat-shaped
limb.

Superficially the leaves of this species resemble
those of A. transtillifera in shape, but they are char-
taceous, puberulous beneath, with 10—14 rather
loose transverse cross-veins and reticulate veins.

16. Aristolochia decandra DinGc Hou, Blumea 28
(1983) 343, f. 1. — Fig. 15a—d, 16.

Liana. Old stem slightly flattened, 0.7—1.2 cm @,
rather smooth, glabrous. Branches slightly flattened,
5—7 mm @, smooth, glabrous, often with a series of
3 or 4 buds in the leaf axil. Leaves firmly charta-
ceous, subcrbicular or broad-ovate in outline,
13.5—22.5 by 11.5—18.5 cm; glabrous above, sparse-
ly minutely hairy beneath; apex acuminate; base
deeply cordate, sinus 3—4 cm deep, auricles with al-
most rounded ends, separate from each other, some-
times overlapping; glabrous above, sparsely minute-
ly hairy underneath; basal nerves 2 pairs, the inner
pair ascending to the apical part of the blade, the
outer pair much shorter and giving off at its base a
strong branch resembling a third basal nerve pair;
midrib with 2—3 pairs of lateral nerves; nerves elevat-
ed, prominent beneath, distinct or sometimes rather
faint above; veins loosely transverse and reticulate,
slightly elevated beneath, faint or invisible above;
petiole stout, subterete except the apical part, 10—13
cm long. J/nflorescences in axils of fallen leaves or
cauligerous, paniculiform, up to 15 cm long, puberu-
lous; bracts ovate, c. 2.5 mm long, puberulous on
both surfaces. Pedicel and ovary 4—5 cm, puberu-
lous. Perianth bright yellow or yellowish green,
straight except the apical part of the tube together
with limb curved or bent, with longitudinal and
loosely reticulate veins, puberulous outside; utricle
broad-ellipsoid, 5—6.5 by 3.5—4 cm, flanged at the
base, without a contracted stipe, with 2 glandular, el-
lipsoid bodies (c. 1.5 mm long); tube c. 5 by 1.5—2
cm, hairy inside, especially dense at the apical part,
basal part projecting a band-like syrinx (c. 1.5 mm
wide) into the cavity of utricle; limb 3-lobed, lobes
linear, up to c. 9 by 1 cm, glandular hairy on the in-
ner surface. Stamens 10; anthers oblong, c. 4 mm
long. Style column 10 mm long, 10-lobed; lobes lan-
ceolate, 2-3 mm long, usually hairy in the apical
part. Capsules unknown.

Distr. Malesia: Kalimantan (Lower Serawai R.:
Manga Landu & Lebang Hara, c. 112°30' E, 0°30’
S). Two collections. Cultivated in Hort. Bog. sub n.
XI-A-61, XI-D-32A & 40A, originated from the Bor-

1984] ARISTOLOCHIACEAE (Ding Hou) 101

Fig. 15. Aristolochia decandra Dino Hov. a. Leafy twig; b. young bud and open flower, both x 2; ¢. LS

of lower part of perianth showing the gynostemium inside the utricle and the base of the tube slightly elongat-

ing and projecting into the utricular cavity, nat. size, d. gynostemium, x 5. — A. rumphii KOSTELETZKY. e.

Leafy twig, x 2 (a HANS WINKLER 1256, b after a drawing of a living specimen in Hort. Bog., Dec. 1945,
in L, c—d HANS WINKLER 373, e LoeTers 2092). Courtesy of Blumea.

102

FLORA MALESIANA

[ser. I, vol. 10!

Fig. 16. Aristolochia decandra DinG Hou. CS of flat-

tened stem, medullary rays and wood sections with

vascular bundles elongating in two directions, x 52
(HANS WINKLER 1256).

nean collection of P. DAKKus.

Ecol. Primary forest, on river bank, 80—180 m.
Fl. Nov., Jan.

Vern. Toro bakdi, Dajak.

Note. The only Malesian species with 10 sta-
mens, the others always having 6. In flower it ap-
proaches the West African species of Pararistolo-
chia. Unfortunately, it seems that the species is no
longer extant in the Botanic Gardens at Bogor.

17. Aristolochia samarensis MerR. Philip. J. Sc. 11
(1916) Bot. 178; En. Philip. 2 (1923) 120; ScHmipT in
E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 241.

Apparently erect plant c. 1 m high. Branches te-
rete, striate, glabrous. Leaves thin-chartaceous, el-
liptic, 1S—25 by 6—8.5 cm; apex acuminate, base cu-
neate; glabrous; basal nerves one pair, rather weak
and short, ascending close to the margin up to
1/3—1/2 of the blade; lateral nerves 6 pairs, slightly
elevated beneath, rather faint above; veins loosely re-
ticulate, some crossbar-like, distinct beneath, ob-
scure or invisible above; petiole 6-10 mm, glabrous.
Inflorescences in the axils of leaves, racemiform, ra-
chis up to 1 cm long, with condensed internodes, pu-
berulous; bracts ovate to lanceolate, 4—6 mm long,
puberulous on both surfaces. Flowers not seen (char-
acters based on MERRILL, /.c.). Pedicel and ovary 7
mm, glabrous. Utricle ellipsoid, c. 3.5 cm long; tube
reflexed, 1—1.5 cm long; limb 1-lipped, narrowly ob-
long, 4.5 by 1 cm, apiculate-acuminate. Stamens 6;
anthers ellipsoid, c. 1.5 mm long. S#y/e column c. 6
mm long, 6-lobed; lobes narrowly oblong, c. 3 mm
long. Capsule ovoid, c. 1.5 cm long, prominently
ridged. Seed triangular, 4.5 by 3.5 mm, not winged,
verrucose on both surfaces.

Distr. Malesia: Philippines (Samar: Catubig R.
at Pinipisakan). Once collected.

Ecol. In damp forests near river at low altitude.

18. Aristolochia philippinensis Wars. in Perkins,
Fragm. Fl. Philip. (1905) 170; Merr. En. Philip. 2
(1923) 119; ScHmipTin E. & P. Nat. Pfl. Fam. ed. 2,
16b (1935) 241; Quis. Medic. Pl. Philip. (1951) 254;
IGARASHI, Food PI. Papilionidae (1979) t. 16; PERRY,
Medic. Pl. E. & SE. Asia (1980) 47. — Fig. 2c—d.

Erect shrubby plant up toc. 1 m high. Old stem te-
rete, 4cm 9, slightly irregularly ridged. Branches te-
rete, 5-10 mm @, slightly striate, sparsely puberu-
lous, glabrescent, or glabrous. Leaves membranous
or chartaceous, lanceolate, elliptic, or oblanceolate,
8.5—24 by 3.5—8.5 cm; apex acuminate or cuspidate;
base obtuse, sometimes slightly cuneate; margin en-
tire, rarely remotely minutely toothed; glabrous on
both surfaces, sometimes sparsely short-hairy on the
midrib, nerves and reticulations beneath; basal
nerves | pair, faint, ascending close to the margin at
the basal part of the blade; lateral nerves 5—8 pairs,
slightly elevated on both surfaces; veins loosely reti-
culate, some crossbar-like, distinct sometimes faint
on both surfaces; petiole 4—8 (—20) mm, glabrous.
Inflorescences in axils of foliage leaves, spiciform or
racemiform, rachis up to 2 (—6) cm long, with dis-
tinct internodes, sparsely shortly hairy or glabrous;
bracts lanceolate, 2-4 mm long, shortly hairy on
both surfaces, glabrescent. Pedicel and ovary 6—8
mm, sparsely hairy. Perianth straight, longitudinal
and reticulate veins distinct, glabrous outside; utricle
ovoid or ellipsoid, 6—7 by 5 mm, with a distinct stipe
(3—4 mm) flanged at base; hairy inside, glandular
bodies 2, obscurely orbicular, c. 0.7 mm g; tube c. 15

1984]

by 2 mm, with scattered, glandular trichomes inside;
limb 1-lipped, oblanceolate or narrow-oblanceolate,
18—25 by 3—6 mm, with scattered glandular tri-
chomes, glabrescent. Stamens 6; anthers oblong, c. 1
mm long. Sty/e column c. 4 mm long, 6-lobed; lobes
triangular, 1—1.5 mm long, with an annular ring at
the base. Capsules subglobose, shortly cylindric, or
oblong-ellipsoid, 1.5—2.5 by 1.5 cm, glabrous. Seeds
triangular or deltoid, 4—4.5 by 4 mm, not winged,
verrucose beneath and marginal part above, funicle
broadened and covering the upper surface.

Distr. Malesia: Philippines (Luzon, Mindoro,
Bancalan I., Mindanao).

Ecol. In thickets and forest at low and medium
altitudes, up to 900 m. F/. March, June—Sept., /r.
Jan., March—May, Aug., Sept., Nov., Dec.

Uses. Decoction of the roots is used in the Philip-
pines as a stomachic and emmenagogue (QUISUM-
BING, /.c.).

Vern. Barubo, Neg., ruso-pusoan, Rizal, tam-
bal-balanding, Zambales.

19. Aristolochia humilis Merr. Philip. J. Sc. 13
(1918) Bot. 9; En. Philip. 2 (1923) 119.

Low erect undershrub, up to c. 40 cm high. Stem
terete, 5-10 mm 4, with only 4—7 leaves, young parts
sparsely puberulous, glabrescent. Leaves membra-
nous to chartaceous, elliptic, rarely ovate, 8—25 by
5—11 cm; apex acute, slightly obtuse, or short-
acuminate; base rounded or subacute; glabrous
above, sparsely short hairy on the nerves beneath;
and reticulations; 1 pair of basal nerves ascending
close to the margin upward to c. 1/3 of the blade,
much weaker than other nerves; lateral nerves 5—7
pairs, slightly elevated beneath, distinct sometimes
faint above; veins loosely reticulate, some transverse,
distinct beneath, rather faint above; petiole (2—) 3—4
cm, sparsely shortly hairy. /nflorescences in the axils
of bracts and/or foliage leaves, rachis 2—3 cm long,
with condensed internodes and bracts; bracts ovate,
oblong, or lanceolate, 4-6 mm long, puberulous
outside, glabrous inside. Pedicel and ovary 6—7 mm
long, glabrous. Perianth straight, venation faint, gla-
brous outside, papillate inside; utricle oblong, ellip-
soid or subglobose, c. 6 by 3—5 mm, with a distinct
stipe (c. 2mm) slightly flanged at base; tube S—7 mm
long; limb 1-lipped, oblanceolate, 18 by 4—6 mm.
Stamens 6; anthers oblong, c. 0.7 mm long. Style col-
umn 1.5 mm long, 6-lobed; lobes triangular, c. 0.5
mm long, with an annular ring at the base. Capsules
oblong-ellipsoid, 2—2.5 by 1.5 cm. Seeds triangular,
convex-concave, c. 6 by 4.5 mm, verrucose on margi-
nal part beneath, glabrous, covered above by the
membranous appendage of the funicle, not winged.

Distr. Malesia: Philippines (Luzon: Tayabas).
Twice collected.

Ecol. In damp forests along streams at low alti-

ARISTOLOCHIACEAE (Ding Hou)

103

tude. Fi. fr. May.
Vern. Tangotong-gubat, Tag.

20. Aristolochia macgregorii MERR. Philip. J. Sc. 5
(1910) Bot. 174; En. Philip. 2 (1923) 119. — Fig. 1d.

Erect shrubby plant up to 1 m high. Stems subte-
rete, 3—5 mm Q, striate, pubescent. Leaves charta-
ceous, lanceolate or slightly elliptic, 11—17 by 3—6.5
cm; apex acuminate, rarely acute; base cordate, sinus
0.5—1 cm deep, auricles often overlapping or sur-
rounding the stem; shortly hairy on both surfaces, es-
pecially on midrib, nerves and veins; basal nerves of-
ten 2 pairs, inner pair ascending to halfway the blade,
outer pair very short and weak; lateral nerves 2 or 3
(—6) pairs; nerves slightly raised or nearly flat on
both surfaces; veins often reticulate, some crossbar-
like, flat, distinct, rarely faint on both surfaces; pet-
iole c. 3mm, pubescent. /nflorescences in axils of fo-
liage leaves, spiciform; rachis with condensed inter-
nodes up to 1.5 cm long, pubescent; bracts lanceo-
late, 3—6 mm long, pubescent on both surfaces. Ped-
icel and ovary c. 5 mm, pubescent. Perianth yellow-
ish, erect, with rather faint reticulation, pubescent
outside; utricle subglobose, c. 4 mm 9, basal part
contracted and stipe-like (0.7 mm long); tube cylin-
dric, 10—16 by 2—2.5 mm; limb lanceolate, c. 20 by
3 mm, acuminate, pubescent on the inner surface.
Stamens 6; anthers 1 mm long. Style column short,
very obscurely lobed. Capsules oblong-ellipsoid, c.
1.5 by 1 cm, sparsely short-hairy. Seeds deltoid, c. 3
mm long and wide, not winged, verrucose on both
surfaces.

Distr. Malesia: Philippines (Babuyan Is.: Dalu-
piri 1. & Camiguin I.).

Ecol. In littoral thicket, up to 350 m. Fi. Aug.,
Nov., fr. Aug., Oct., Nov.

21. Aristolochia sericea BLANco, FI. Filip. (1837)
283; ed. 2 (1845) 198; ed. 3, 1 (1877) 350; MerrR.
Publ. Govt. Lab. Philip. 27 (1905) 72; Sp. Blanc.
(1918) 134; Brown, Bull. Bur. For. Philip. 22, 3
(1921) 183; Merr. En. Philip. 2 (1923) 120; ScumipT
inE. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 241; Quis.
Medic. Pl. Philip. (1951) 255; Perry, Medic. Pl. E.
& SE. Asia (1980) 47. — A. imbricata Mast. J. Linn.
Soc. Bot. 14 (1875) 494; Scumiprin E. & P. Nat. Pfl.
Fam. ed. 2, 16b (1935) 241. — Bragantia corymbosa
(non GriFF.) F.-Vitt. Nov. App. (1880) 174. — A.
membranacea Merk. Philip. J. Sc. 14 (1919) 381;
En. Philip. 2 (1923) 119; ScuMipt in E, & P. Nat.
Pfl. Fam. ed. 2, 16b (1935) 241.

Erect shrubby plant up to c. 0.5 m high. Stem te-
rete, c. 3 mm @, densely pubescent, glabrescent.
Leaves thin-chartaceous or membranous, lanceolate
or oblong-lanceolate, 7-15 by 2—5 cm; apex acumi-
nate; base slightly or shallowly cordate, sometimes
the sinus up to 5 (—10) mm deep, rarely obtuse;

104

FLORA MALESIANA

[ser. I, vol. 10!

sparsely shortly hairy above, hairy especially dense
on the venation beneath; basal nerves 2 pairs, inner
pair reaching c. 1/3 of the blade, similar to the lateral
nerves, outer pair weaker and very short, close to the
margin; lateral nerves c. 5 pairs, rather faint beneath,
faint or obscure above; petiole 2—5 mm, densely
hairy. Flowers few (one developing at a time), on
very short, bracteate rachides (less than 10 mm long)
in axils of foliage leaves; bracts lanceolate or elliptic,
3—4 mm, rarely leafy up to 20 mm long, shortly hairy
on both surfaces. Pedicel and ovary 4.5—5 mm,
densely hairy. Perianth straight, sometimes slightly
curved, reticulation rather distinct, hairy outside;
utricle ovoid or ellipsoid, 2.5—5 by 2—2.5 mm, with
a distinct stipe (1.5—2 mm) dilated at the base as a
cap on top of the ovary, hairy inside, with 2 glandu-
lar, ellipsoid bodies (c. 0.6 mm long); tube 5—10 by
1 mm, with short, glandular hairs inside; limb
1-lipped, oblong, 11—15 by 3—4.5 mm, acute or acu-
minate. Stamens 6; anthers ellipsoid, c. 0.6 mm long.
Style column c. 0.7 mm long, 6-lobed; lobes
lanceolate, c. 1 mm, with an annular ring at the base.
Capsules subglobose, c. 1 cm @, densely hairy, usual-
ly glabrescent. Seeds triangular, c. 2.5 by 1.5 mm,
not winged, verrucose on both surfaces; funicle with
membranous extension covering the upper surface.

Distr. Malesia: Philippines (Luzon: Cagayan,
Ilocos Norte, Union, Batangas).

Ecol. Indry thickets at low and medium altitudes
up to 350 m. Fi. Aug., Dec., fr. April, May, July,
Aug., Dec.

Uses. The entire fresh plant is used as a carmina-
tive, emmenagogue and febrifuge. The root, macer-
ated in local spirituous liquors, is administered post-
partum as a uterine tonic; this drug is said to be a vio-
lent abortive (QUISUMBING, /.c.).

22. Aristolochia leytensis Merr. Philip. J. Sc. 10
(1915) Bot. 4; En. Philip. 2 (1923) 119; ScHMipT in
E. & P. Nat. Pfl. Fam. ed. 2, 16b (1935) 241.
Slender vine. Branches terete, sulcate, 2—6 mm 9,
glabrous. Leaves chartaceous or membranous, ovate
or broad-ovate, 12.5—21.5 by 8—12.5 cm; apex acu-
minate; base cordate, sinus (1—) 2—3 cm deep, auri-
cles separate from each other, sometimes overlap-
ping at the base; glabrous on both surfaces, some-
times slightly puberulous on midrib, nerves and veins
beneath when young, glabrescent; basal nerves 1
pair, ascending upward to 2/3—3/4 of the blade,
forked at base; lateral nerves c. 3 pairs; nerves elevat-
ed and prominent beneath, distinct or flat above;
veins loosely reticulate, some crossbar-like, distinct
beneath, rather faint above; petiole (2—) 4—7 cm,
glabrous. /nflorescences in the axils of foliage leaves,
racemiform, rachis up to 3 cm long, with visible in-
ternodes, sparsely puberulous or glabrous; bracts
ovate to lanceolate, 2—3 mm long, puberulous on

both surfaces especially on the margin. Pedicel and
ovary 17—22 mm, puberulous. Perianth straight or
slightly curved, with a distinct stipe (4—5.5 mm), ve-
nation visible, puberulous outside, hairy inside; utri-
cle subglobose, 5—7 mm @, with 2 glandular, subor-
bicular bodies (c. 0.5 mm @); tube c. 20 by 4 mm;
limb 1-lipped, narrow-elliptic, SO—60 mm. Stamens
6; anthers oblong 0.6—1 mm long. Sty/e column c. 4
mm long, 6-lobed; lobes triangular, c. 1 mm long,
with an annular ring around the base. Capsules
(young) subglobose or short-cylindric, c. 1.5 cm @.

Distr. Malesia: Philippines (Leyte, Samar).

Ecol. In thickets and forests near or along
streams at low altitudes. F/. Feb.—March, Aug., fr.
Aug., Sept.

Vern. Maraburakan, S.L. Bis.

23. Aristolochia crassinervia SCHMIDT, Bot. Jahrb.
58 (1923) 491; in E. & P. Nat. Pfl. Fam. ed. 2, 16b
(1935) 241; Merr. & Perry, J. Arn. Arb. 29 (1948)
153; IGARASHI, Food Pl. Papilionidae (1979) t. 23;
Dinc Hou, Blumea 29 (1983) 234, f. 4, 5b. — Fig. 8c,
17.

Fig. 17. Aristolochia crassinervia SCHMIDT, showing

the common basket shape of the dehisced, hanging

capsule in the genus, with seven loose seeds, nat. size
(HUTTON s.7.).

1984]

ARISTOLOCHIACEAE (Ding Hou)

105

Liana, old stem up to c. 1.5 cm @. Branches rather
terete, 4-5 mm 9g, with some prominent lenticels, gla-
brous. Leaves subcoriaceous, (S—) 8—19 by (4—)
5—12 cm; apex acuminate or short-acuminate; base
cordate, with a narrow sinus up to 2.5—3 cm deep;
auricles usually overlapping, rounded at apex; gla-
brous above, minutely hairy beneath; basal nerves 2
(—3) pairs, the inner pair similar to the midrib, as-
cending to near the apex of the blade, the outer one
much shorter, with branches extending upward or to
the auricles, elevated and prominent beneath, dis-
tinct above; lateral nerves hardly distinguishable
from the veins; veins closely reticulate, elevated and
prominent beneath, faint or invisible above; petiole
(2—) 6—8 cm, sparsely shortly hairy or almost gla-
brous. /nflorescences cauligerous, rarely also in axils
of leaves, racemiform, rachis up to 3 cm long, inter-
nodes very short or condensed, sometimes the brac-
teate flowering rachis very short and flowers appear-
ing fascicled; bracts lanceolate, 5-10 mm long,
sparsely minutely hairy on both surfaces, glabres-
cent. Pedicel and ovary c. 20 mm, glabrous. Perianth
white or reddish, no stipe at the base, with a contrac-
tion between perianth and ovary, glabrous outside,
venation rather faint; utricle subglobose, c. 7 mm @,
hairy inside, with 2 glandular, round bodies (c. 0.5
mm @); tube c. 7 by 2 mm, glandular-hairy inside;
limb 1-lipped, linear, c. 20 by 4 mm, with glandular
hairs on the inner surface. Stamens 6; anthers ob-
long, c. 0.7 mm long. Gynostemium c. 1 mm long;
style 6-lobed; lobes triangular, c. 0.5 mm long, the
basal part of each lobe slightly extended outward (c.
0.3 mm long and wide). Capsules yellowish when ma-
ture (NGF 45343), hard, tardily dehiscent, cylindric
or oblong, not ridged or angular, 2.5—4.5 by 2—3
cm, minutely granular. Seeds triangular or deltoid, 7
by 6—7 mm, not winged, rather smooth or obscurely
muriculate on both surfaces; funicle broadened and
covering the upper surface.

Distr. Solomon Is. (Ysabel, Malaita); in Male-
sia: New Guinea (Sepik, Morobe and Milne Bay
Distr.).

Ecol. In primary, sometimes in secondary, for-
ests, on rough, rocky, wet ground, on edge of creek,
or along stream, from lowland up to 1500 m. Fi.
June, fr. June, Aug., Sept., Oct.

Vern. Solomon Is.: kwalokame, Kwara’ae name.

Notes. Among the New Guinea species the leaves
are characteristic: subcoriaceous to coriaceous,
prominently palmately 5 (—7)-nerved, with a promi-
nent closely reticulate venation, and a deeply cordate
base with the auricles often overlapping.

It seems allied to A. foveolata.

24. Aristolochia schlechteri Laut. in K. Sch. & Laut.
Nachtr. Fl. Schutzgeb. (1905) 260; Bot. Jahrb, 52
(1914) 107; Ding Hou, Blumea 29 (1983) 241, f. 6c.

Twiner or scrambling vine. Old stem terete, 1.5 cm
@. Branches terete, 2—5 mm @, pubescent when
young, glabrescent. Leaves subcoriaceous or charta-
ceous, elliptic-lanceolate, oblanceolate, (10—) 17—26
by (2.5—) 5—9 cm; apex acuminate; base cordate with
distinct auricles, or shallowly cordate with obscure
auricles, sinus (0.5—) 1—2 cm deep; glabrous or
sparsely shortly hairy above, short-hairy beneath;
nerves prominent beneath, distinct above; basal
nerves 1 pair, ascending upward to halfway the
blade, lateral nerves 4 or 5 pairs; veins loosely reticu-
late or crossbar-like, slightly elevated beneath, rather
faint above; petiole 2—3.5 cm, shortly hairy. /nflo-
rescences in axils of leaves, or cauligerous, usually |
or 3, simple or spiciform, borne on a knob-like, very
short branch; rachides up to 16 cm long, with spa-
cious internodes, densely shortly hairy; bracts very
small, triangular or lanceolate, c. 1 mm long, densely
hairy. Pedicel and ovary c. 22 mm long, densely
hairy. Perianth slightly curved, whitish or creamy
white with green, brown or purple distinct venation;
with a contraction between perianth and ovary; pu-
bescent outside, with rather scattered glandular hairs
inside; utricle ellipsoid or obovoid, c. 15 by 8-11
mm; tube 17—23 mm long, variable in width (5—8
mm @) in one flower; limb prominently 3 + 3-lobed:
3 triangular lobes (20—25 by 15 mm) and 3 alternate
lobes (each lobe triangular, 1S—20 by 10-15 mm,
apical part, c. 50 mm, laciniate and tail-like when
dry). Stamens 6; anthers oblong, 1.5—2 mm long.
Style column c. 7 mm long, 6-lobed; lobes lanceolate,
2.5 mm long. Capsules not seen.

Distr. Malesia; Papua New Guinea (Sepik, Ma-
dang and Morobe Distr.).

Ecol. In primary forest, beside a river or on
rocky ridge. Fi. Jan., July, Aug., Dec., fr. June.

Vern. New Guinea: pengeramboi, Sepik Distr.,
Waskuk, yogwa, Wagu.

Note: Among Malesian species this is unique in
possessing a 6-lobed perianth. In leaves it cannot be
distinguished from A. momandul. Whether the cap-
sules or seeds are different could as yet not be ascer-
tained.

25. Aristolochia dielsiana Scumipt, Bot. Jahrb. 58
(1923) 490; in E. & P. Nat. Pfl. Fam. ed. 2, 16b
(1935) 241; Dinc Hou, Blumea 29 (1983) 235, f. 6a.

Liana, up to 15—40 m high, c. 2 cm @. Branches c.
5 mm @, shallowly longitudinally sulcate, shortly
hairy. Leaves often coriaceous, broad-ovate, ovate,
lanceolate, rarely narrow-lanceolate, 19-37 by
4.5—23.5 cm (sometimes up to 100 by 70 cm); apex
acuminate, rarely apiculate; base slightly sinuate or
shallowly cordate; + truncate, rarely cordate, sinus
0—1 cm deep; shortly hairy on both surfaces, espe-
cially on midrib, nerves and veins; nerves elevated
and prominent beneath, slightly elevated above;

106

FLORA MALESIANA

[ser. I, vol. 10!

basal nerves | pair, ascending upward to 1/2—3/4 of
the blade, each nerve with a weaker, short branch at
the base; lateral nerves 3—S pairs; veins crossbar-like
or reticulate, slightly elevated and prominent be-
neath, main veins distinct above; petiole 2.5—6 cm,
often densely shortly hairy. /nflorescences cauliger-
ous, | or 2 borne on a brachyblast of the (old) stem,
rachides up to 11 (—29) cm long, with spaced inter-
nodes; bracts very small, densely hairy. Pedicels and
ovary 15—30 mm, densely hairy. Perianth (bud)
straight or slightly curved, greenish white with purple
venation, pale green outside with 3 lobes orange-
yellow and purple inside, or light yellowish purple
with purple veins and becoming dark purple when
open; longitudinal veins distinct, loose reticulations
faint; shortly hairy outside especially on the veins,
glandular hairs scattered inside; utricle obovoid or
ellipsoid, 10—15 by S—10 mm, no glandular bodies
inside visible; tube 30—35 mm long, enlarged toward
the apical part (c. 15 mm 9); limb 3-lobed, lobes tri-
angular to narrow-triangular, 35—SO mm long,
12—14 mm wide at the base, gradually narrowed to-
wards the apex, apical part (25—30 mm) laciniate and
often tail-like when dry. Stamens 6; anthers oblong,
1.7—2 mm long. Gynostemium 5.5 mm long, lower
part (c. 1 mm) stipe-like; style 6-lobed, lobes linear,
2.5—3 mm long, no annular ring at the base. Cap-
sules green or light yellowish green, oblong, up to 20
(—30) by c. 4 cm, indehiscent (?), 6-ridged, smooth,
sparsely shortly hairy, glabrescent. Seeds triangular
or deltoid, 9 by 8.5—9 mm, slightly longitudinally
concave above, convex beneath, smooth on both sur-
faces, not winged.

Distr. Malesia: West New Guinea (Idenburg R.);
Papua New Guinea (Sepik, Central & Northern
Distr.).

Ecol. In primary forest, rarely in secondary for-
est, from the lowland up to 800 (—1200) m. F/. May,
June, July, Oct., Dec., fr. June, July, Dec.

Notes. In flower bud the present species re-
sembles A. schlechteri, but the latter has in anthesis
6 perianth lobes.

It is the host of one of the largest butterfly species
of Rhopalocera, Ornithoptera alexandra, in New
Guinea. Mr. R. STRAATMAN reported upon it, under
the erroneous name ‘A. schlechteri’ (J. Lepidopt.
Soc. 25, 1971, 58—64) that ‘the flower is shaped like
a starfish with three long arms and is dark purple-
brown with a yellow heart. The green fruit is shaped
like a cucumber, 20—30 cm long, strongly ribbed
lengthwise and has a rough skin. It matures slowly
and when fully rotten the seeds fall to the ground and
are carried away by rainwater over generally short
distances, resulting in a number of plants growing in
a restricted area.’

Flower colour is not always reported the same:
HOOGLAND & CRAVEN noted the perianth to be pale

green outside, the lobes orange-yellow and purple in-
side, BRAss said flowers greenish white, tinged and
veined with purple.

26. Aristolochia engleriana ScHMipT in Fedde, Rep.
23 (1927) 288; DinGc Hou, Blumea 29 (1983) 238, f.
5b, 7b. — A. ledermannii ScumiptT, Bot. Jahrb. 58
(1923) 489, non ENGLER 1911.

Small to moderately high twiner. Old stem subte-
rete or slightly flattened, 8-10 mm @, shallowly fur-
rowed. Branches terete, 1.5—2.5 mm @, rather
smooth, shortly hairy when young, glabrescent.
Leaves subcoriaceous or chartaceous, lanceolate,
ovate-elliptic, obovate to oblanceolate, (S—) 8—15
(—19) by (2—) 4.5—6 (—7) cm; apex acuminate, cuspi-
date; base obtuse, rounded, cuneate, or broadly
truncate; glabrous above, sparsely minutely hairy on
midrib, nerves and veins below, glabrescent; nerves
elevated beneath, distinct above; basal nerves 1 pair,
ascending to 1/3—3/4 of the blade; lateral nerves
2—5 pairs, sometimes hardly distinct from veins;
loosely reticulate or crossbar-like veins slightly ele-
vated beneath, often faint above; petiole terete, 5—25
(—35) mm, shortly hairy. /nflorescences in the axils
of leaves, or cauligerous, spiciform, rachides up toc.
4 cm long, internodes distinct, shortly hairy; bracts
minute, densely hairy. Pedicel and ovary 15S—20 mm.
Perianth almost straight except the slightly curved
tube, whitish with violet-brown venation, or orange
yellow (but tube light purplish red with scattered
white spots), longitudinal veins visible or distinct, re-
ticulations loose, obscure, shortly hairy outside,
glandular-hairy inside, with a contraction between
perianth and ovary; utricle obovoid or ellipsoid, c. 15
by 7 mm, no glandular bodies seen; tube 15—20 mm
long, narrowed at basal part (c. 2.5 mm g), the rest c.
6 mm 9g; limb 3-lobed, lobes triangular, 5—6 by 8—10
mm, obtuse or retuse (no tail-like appendage seen).
Stamens 6; anthers oblong, c. 1.5mm. Gynostemium
c. 3.5 mm long, with a distinct stipe-like part (c. 1
mm); style 6-lobed; lobes lanceolate, c. 2mm long, no
annular ring at the base. Capsule bright orange, el-
lipsoid, or pyriform (type description), 6—9.5 by
2—4.5 cm, 6-ridged, smooth. Seeds triangular, c. 11
by 9 mm, smooth on both surfaces, not winged.

Distr. Malesia: West New Guinea (Malingdam);
Papua New Guinea (Sepik, Western, Eastern, and
Southern Highlands Distr.). Fig. 18.

Ecol. Low montane or montane, sometimes mos-
sy forests, (1260—) 1830—2250 m, once at 600—700 m
(Mt Bosavi). F/. March, Sept., fr. Jan., Sept., Oct.

Vern. Papua New Guinea: ya waenuw kunguwp,
Wola language.

Notes. From other New Guinea species easily
recognizable by the 3-nerved leaves with a non-
cordate base (base rounded, obtuse, broadly trunc-
ate, sometimes cuneate) and a pyriform fruit.

Fig. 18. Aristolochia engleriana SCHMIDT, localities
in New Guinea.

IGARASHI(Food PI. Papilionidae, 1979, t. 22) gave
a photograph, under the name A. schlechteri, which
is probably the present species.

27. Aristolochia gaudichaudii DUCHARTRE, Ann. Sc.
Nat. Bot. 2 (1854) 72, t. 6, f. 3-5; Mia. FI. Ind. Bat.
1, 1 (1858) 1067; KLtorzscu, Monatsb. Akad. Berl.
(1859) 597; DucHaRTRE in DC. Prod. 15, 1 (1864)
481; Laut. Bot. Jahrb. 52 (1914) 105; RoEpPKE, Trop.
Natuur 24 (1935) 80, f. 5; Scumipt in E. & P. Nat.
Pfl. Fam. ed. 2, 16b (1935) 241; IGARAsHI, Food PI.
Papilionidae (1979) t. 26 (fig. on the left & top right);
Dinc Hou, Blumea 29 (1983) 238. — A. roxburghia-
na(non KLotzscH) WarB. Bot. Jahrb. 13 (1891) 300.
— Fig. le.

Twiner, sometimes creeper, up to 20 m high. Old
stem terete, rather smooth, c. 1.2 cm 9. Branches te-
rete, c. 4 mm @, striate, glabrous. Leaves charta-
ceous, ovate, broad-ovate, triangular or deltoid in
outline, 11—22 by 10—18 cm; apex acute or short-
acuminate; base subtruncate (especially when
young), subcordate or cordate (usually adult leaves;
with the sinus up to 2.5 cm deep; sometimes with 2
divergent, rounded auricles); glabrous; midrib and
nerves palmate; basal nerves 2 pairs, similar to the
midrib, inner pair ascending to the apex, outer pair
much shorter, elevated beneath, distinct above; later-
al nerves hardly distinguishable from the veins; veins
slightly elevated beneath, distinct or faint above, ma-
jor veins connecting the inner pair of nerves and mid-
rib, + transverse at the lower 1/3 or 1/2 of the blade,
reticulations rather loose; petiole 2.5—8.5 cm, gla-
brous. /nflorescences in the axils of leaves, with ra-
chis up to 6cm long, or cauligerous (often with sever-
al mostly simple branches, arising from a knob or
spot of the stem, up to 14cm long), branches usually
spiciform, with spacious internodes; bracts ovate or
triangular, 0.7—1.5 (—5) mm long, glabrous but ci-
liate on the margin. Pedicel and ovary 10-14 mm,
glabrous. Perianth straight or slightly curved, with

ARISTOLOCHIACEAE (Ding Hou)

107

variable colours recorded as white, yellow, pale
green with pinkish tinge, green to brownish, brown
red or dark brown red, with a distinct stipe (3—4 mm)
slightly dilated at base, longitudinal and reticulate
veins distinct, glabrous outside; utricle broad-ellip-
soid, 8—15 by 5—9 mm, sparsely hairy inside, with 2
glandular, ellipsoid bodies (c. 1.5 mm long); tube
8—12 by 1.5—2 mm, with scattered glandular tri-
chomes inside; limb 1-lipped, obovate or oblanceo-
late, 17—20 by 8—12 mm, with scattered glandular
trichomes on the inner surface. Stamens 6; anthers
oblong, c. | mm long. Sty/e column 3.5—4 mm long,
6-lobed; lobes lanceolate, c. 1.5 mm long, with an an-
nular ring at the base. Capsules oblong, 4—6 by c. 3
cm, glabrous. Seeds (incl. wing) transverse-oblong,
6—11.5 by 12—16 mm; seed proper deltoid 6—8 by
6—8 mm, smooth on both surfaces, with a central,
longitudinal ridge above; wing 2.5—4 mm broad.

Distr. Malesia: Moluccas (Batjan, Ceram) and
New Guinea (Sorong, Manokwari, Fak Fak, Sidei,
Toronta, Sepik) and neighbouring islands: Job I.,
Rawak (Waigeo), Schouten & Biak Is., and New Ire-
land.

Ecol. In primary forest, sometimes in beach, sec-
ondary and swampy forests, clearing ground and
thickets at low altitudes. F/. April, June, July, Sept.,
Oct., Dec., fr. May, Aug., Sept., Oct., Dec.

Vern. New Guinea: daprijo, Irian, surwerro, Pa-
pua.

28. Aristolochia linnemannii Wars. Bot. Jahrb. 13
(1891) 301; K. Scu. & Laut. Fl. Deut. Schutzgeb.
Stidsee (1900) 302; Laur. Bot. Jahrb. 52 (1914) 105;
DinG Hou, Blumea 29 (1983) 239, f. 2f.

Scrambler or climber, c. 6m long. Branches terete,
c. 2.5 mm Q, sulcate, glabrous. Leaves chartaceous,
triangular or deltoid in outline, S—8 by 4.5—6 cm;
apex short-acuminate or acute; base shallowly cor-
date, sinus broad, up to c. | cm deep; glabrous
above, minutely hairy beneath; basal nerves 2 pairs,
slightly elevated beneath, distinct above; inner pair
ascending to the apex, outer pair much shorter, up to
c. 1/3 of the blade; veins reticulate beneath, rather
faint above; petiole 1.5—2.5 cm, glabrous. /n/flores-
cences in the axils of leaves, fasciculate, rachides
condensed, very short, usually 2—3 mm long, some-
times one of the branches up to 10 (—15) mm, gla-
brous; bracts minute, triangular or deltoid, c. | mm
long, glabrous. Pedicel and ovary 15-30 mm, gla-
brous. Perianth oblique or slightly curved, venation
distinct, glabrous outisde, with a distinct stipe (¢. 2.5
mm) slightly expanded at base; utricle subglobose, c.
6 mm @, sparsely hairy inside, with 2 glandular, ellip-
tic bodies (c. 2 mm long); tube c. 17 by 2 mm, with
scattered, glandular hairs inside; limb 1-lipped, ob-
long, c. 20 by 10 mm, obtuse or slightly mucronulate,
margin reflexed, with scattered, glandular hairs on

108

the inner surface, glabrescent. Stamens 6; anthers
oblong, c. 0.7 mm long. Gynostemium c. 2.5 mm
long; style 6-lobed; lobes triangular, c. 0.5 mm long,
basal parts united and extended outward as an annu-
lar ring. Capsules glabrous, broad-ellipsoid or sub-
globose, 2.5 by 1.7—2.5 cm, 6-ribbed or 6-angular,
obtuse. Seeds winged, triangular, c. 6 by 5 mm (incl.
c. | mm wide wing), seed proper with wart-like gran-
ules densely covered beneath, sparsely above, funicle
obscure.

Distr. Malesia: West New Guinea (Div. Hollan-
dia), Papua New Guinea (Morobe Distr.: former
Finschhafen). Two collections.

Ecol. In thickets or along the road at low alti-
tude. Fi. fr. July.

Doubtful species

The following two species of Aristolochia were de-
scribed by O.C. ScHmiptT (see below). The types were
collected at Lordberg, Sepik region, NE. New Gui-
nea, c. 1000 m alt., by LEDERMANN in Nov.-Dec.
1912. These types were lost in B during World War
II. Duplicates of them may be extant but as yet have
not been found. From the New Guinean (flowering)
specimens of Aristolochia examined, I cannot find
specimens to match the descriptions. I have extracted
the essential characters from the original descriptions
as follows:

FLORA MALESIANA

[ser. I, vol. 10!

Aristolochia lauterbachiana ScumiptT, Bot. Jahrb.
58 (1923) 488. — Type: LEDERMANN 9883 (B, lost),
Sepik region, NE. New Guinea.

Leaves unknown. Inflorescences curled, few-flow-
ered, axis densely pilose; bracts minute, subtriangu-
lar, c. 1 mm long and wide, densely pilose. Pedicel c.
13 mm long. Flowers white, red-striate and -spotted,
unilabiate, c. 6.5 cm long, densely pilose; utricle obo-
void, c. 14 mm long; tube enlarged at the apical part,
c. 22 mm long; limb lingulate, c. 20 mm long, apex
narrowed; ovary c. 10 by 1.5 mm, densely pilose.
Fruit unknown.

Aristolochia novoguineénsis SCHMIDT, Bot.
Jahrb. 58 (1923) 489. — Type: LEDERMANN 10362
(B, lost), Sepik region, NE. New Guinea.

Leaves subcoriaceous, lanceolate, 16—18 by 3—5
cm, smooth, apex caudate, up to 2.5 cm long, base
rounded or truncate, nerves 3 at the base; petiole
3—3.5 cm long. Inflorescences curled, multi-
flowered; bracts minute, densely pilose. Pedicel c. 12
mm long. Flowers white, purple-nerved, c. 8.5 cm
long, 1-lipped; utricle ovoid, c. 12 mm long; tube
curved, c. 18 mm long; limb narrow-lingulate, c. 38
mm long; ovary c. 12 by 2 mm, densely pilose. Fruit
unknown.

TRIURIDACEAE (J.P.M. van de Meerendonk, Leyden)’

The Triuridaceae are a small family (c. 6 genera, and c. 45 spp.) of very delicate, saprophytic,
terrestrial, mostly dark-red coloured herbs growing in the deep shade of everwet tropical forest,
entering the subtropics only in Japan and the Bonin Is. They are in Africa confined to restricted
areas in the West and are also in continental Southeast Asia remarkably rare, as yet only known
from two localities in Assam and N. Thailand respectively. Fig. 1. The nearest localities to Indo-
china and China are in Hainan and Botel Tobago Is. (southeast off Taiwan). In Australia they
are only found in the Bellenden Ker Range in NE. Queensland, showing their aversion to dry and
seasonal climates.

By their small stature (10—40 cm), dark colour, and very small flowers they are evasive to collec-
tors; the only one reaching some size (45—140 cm) is Sciaphila purpurea which is found in Peru,
according to GIESEN mainly in termite nests in hollow trunks. During exploration, trip stops,
either for felling or climbing trees, or for culinary or sanitary purposes, offer the best opportunity
to observe them.

Flowering specimens can probably be found throughout the year, as it appeared that of com-
mon species such as Sciaphila arfakiana, specimens have been collected in all months of the year.

Formerly Triuridaceae were usually placed in the affinity with Liliaceae by BENTHAM& HOOKER
and by ENGLER & PRANTL. HUTCHINSON (1934) raised the family to the order Triuridales, along-
side Alismatales to which he also reckoned the saprophytic genus Petrosavia, which usually was
accommodated in Liliaceae, but deviates from Liliaceae in having an apocarpous gynoecium. He
recognized Petrosavia as representing a distinct family Petrosaviaceae.

Recently this controversial matter was further elaborated by CRONQuIsT (1981), who also recog-
nized Triuridaceae in the rank of an order, Triuridales, but more closely associated Petrosaviaceae
with Triuridaceae and finds ‘the resemblance so complete that I would have no hesitation in
placing Petrosavia in the family Triuridaceae on the basis of anatomical evidence.’ This view is
shared by DAHLGREN & CLIFFORD (1982). The removal of Petrosavia from Liliaceae to the affinity
of Triuridaceae is here also supported by MULLER (vide infra) who found that the pollen of Triuri-
daceae shows some similarity to that of Petrosavia and Vallisneria and does not suit that of Lilia-
ceae.

The family was meticulously revised by H. GresEn (1938) who had the rich material (much on
liquid) of Herbarium Bogoriense. In addition, the great value of his work is the fact that he report-
ed in detail on many type specimens, of which some in Berlin are now lost and also on those of
Florence, which were not sent to me on loan. This enabled me to reach a satisfactory interpreta-
tion.

From Malesia GiesEN had some 130 collections at his disposal. The present revision is based
on 300 collections. This increase led to a better insight in the variability of characters and made
it possible to select those that are reliable, which in turn led to a rather heavy reduction in the
number of species and more critical generic and specific delimitations.

As to the genera, Sciaphila (incl. Andruris) is by far the largest (c. 35 spp.), and covers the entire
range. Three small genera (1 or 2 spp. each) are neotropical, a fifth is confined to the Malagasian
area, and the sixth is endemic in the Deccan and Ceylon; both are monotypic.

Literature: A. CRonguist, An integrated system of classification of flowering plants (1981)
1074; R.M.T. DAHLGREN & H.T. CLiFForD, The Monocotyledons (1982) 289, 323, 324; H. Gig-
SEN, Triuridaceae. Pfl. R. Heft 104 (1938); J. HUTCHINSON, The families of flowering plants. 2.
Monocotyledones (1934) 37.

(1) Revised under the supervision of the late Dr. M. Jacobs; made ready for the press with a general intro-
duction by the General Editor.

(109)

110 FLORA MALESIANA [ser. I, vol. 10!

Fig. 1. Approximate range of the family 7riuridaceae.

1. SCIAPHILA

BiuME, Bijdr. 10 (1826) 514; Beccari, Malesia 3 (1890) 329; SCHLTR, Bot.
Jahrb. 49 (1912) 70—84, 3 fig.; GrESEN, Pfl. R. Heft 104 (1938) 30; MEEREN-
DONK, Ident. Lists Males. Specim. n. 63 (1983). — Aphylleia CHAMPION, Calc.
J. Nat. Hist. 7 (1847) 468. — Andruris SCHLTR, Bot. Jahrb. 49 (1912) 71;
GIESEN, Pfl. R. Heft 104 (1938) 15. — Fig. 1-4.

Small and delicate, echlorophyllose, mostly erect, simple or branched herbs
up to c. 45 cm high (in Mal.). Rhizome often with a few scale-like leaves, some-
times branched, mostly with hairs. Stem usually glabrous, sometimes with air-
roots, descending from both sides of the leaves (up to 5 cm of the stem). Leaves
scale-like, scattered, spaced, sessile, mostly appressed, sometimes amplexicaul.
Inflorescences terminal, racemose, all around or most flowers to one side, with
male and female flowers, or male and bisexual flowers, or only bisexual flowers.
Monoecious; female flowers usually in the upper part of the raceme. Flowers
actinomorphic, perianth with 4—10, usually 6, valvate, patent or reflexed, equal
or alternatingly unequal (larger and smaller) segments, connate at base, at the
top glabrous or bearded by uniseriate hairs, or with knob-like appendages. —
o Flowers: with 2—3 or 6 sessile or + sessile, epitepalous (in 7riuris alternitepal-
ous) stamens, in case there are 2 or 3 stamens they are in front of the larger seg-
ments; anthers 1—4-celled, 2—4-lobed, first opening transversally, later also
longitudinally; filaments sometimes far exceeding the dorsifixed anthers, rarely
connate at base. — 9 Flowers: with c. 10—80 obovoid, free ovaries, each with
1 orthotropic, later anatropic ovule; style inserted laterally and adaxially, usually
exceeding the ovary, club- or awl-shaped, in the former case with many hairs
and papillae, in the latter with glabrous apex. — Bisexual flowers: with (2—)
3-6 persistent stamens with clearly visible filaments, anthers 1-celled,
2—3-lobed; ovaries c. 10—50, like in the female flowers, but the style always

1984] TRIURIDACEAE (van de Meerendonk) 111

club-shaped. Fruits obovoid, 3—8 times as large as the ovaries, with persistent,
partly shrivelled style, dehiscent lengthwise from the apex, first abaxially, later
also adaxially. Seed 1, endospermous, elliptic to ovate, the surface netted and
mostly lined, sometimes with a dent. Endosperm absent; embryo anatropous
(first orthotropous).

Distr. About 33 spp.; pantropical and subtropical in Southern Japan and Bonin Is., in Africa only in the
West (Ivory Coast, Nigeria, Cameroun), in continental Southeast Asia only in Assam and N. Thailand, in
Hainan and Botel Tobago Is., throughout Malesia (14 spp.), not in the Central Pacific, but rather well repre-
sented in Micronesia and Melanesia, and Western Polynesia; in Australia only locally in Queensland (Bellen-
den Ker Ra., c. 16° S). Fig. 2.

Ecol. Saprophytes in humous soil between litter of rain-forest, often associated in local saprophyte ‘colo-
nies’ with species of Burmanniaceae, tiny orchids, and Epirixanthes (Polygal.). Mostly at low altitude, ascend-
ing to c. 1200 m, very rarely higher, up to 2200 m altitude.

The root system carries endotrophic mycorrhiza.

Morph. & Anat. Several large papers have been devoted to the morphology and anatomy, notably by
JoHow (1889), PouLsEN (mainly on the embryology) (1906), Wirz (1910), and ToMLINSON (1982).

The anatomy is reduced and stomata are absent. Whether the plants are annual or perennial is not clear;
probably they are annual.

The pollen is that of a Monocot, the grains being inaperturate, as in Scheuchzeria, monosulcate in Sciaphi-
la, and trinucleate as in most Alismatiflorae.

The ovule is orthotropous first, anatropous later. The single integument is two cell layers thick. The seed,
c. 1 mm in size, lacks endosperm and the cells of the testa are filled with air, which might favour wind disper-
sal, anyway at very short distances only, with respect to its ‘concealed’ habit and small size of the plants.

Literature: Jonow, Pringsh. Jahrb. Bot. 20 (1889) 475—525, t. 19—22; PouLsen, Medd. Naturl. Foren.
Kbhyn 49 (1906) 1—16, t. 6; P.B. ToMLINson in Metcalfe (ed.), Anatomy of the Monocotyledons, VII. Helo-
bieae (1982) 466—473, t. 15; Wirz, Flora 101 (1910) 395—446, f. 1—22, t. 4.

150 60 170

60 90 100 no °

Fig. 2. Distribution of Sciaphila BL. east of Africa; outline of range, with the only known localities in India,
Thailand, and Australia represented by a dot. For the Malesian subareas number of endemics above the hy-
phen and number of non-endemics below the hyphen.

112 FLORA MALESIANA [ser. I, vol. 10!

Palyn. Pollen of Sciaphila is trinucleate, boatshaped-ellipsoidal, 25—40 um long with an indistinctly out-
lined colpate aperture. The exine is very thin, probably intectate and covered with microverrucae. In S. arfa-
kiana and S. corniculata the microverrucae are coarser on the apertural side of the grain and finer on the op-
posite side and they are densely spaced in a hexagonal pattern. In S. densiflora, S. tenella and S. winkleri the
microverrucae are of uniform size and more or less densely spaced. The pollen of S. multiflora is transitional
between these two types.

Sciaphila pollen is similar in the microverrucate sculpture to that of Triuris, but the latter is spherical and
inaperturate. The pollen of Hyalisma which is also spherical and inaperturate differs in the rather coarse, di-
morphic gemmate-echinate sculpture.

The pollen of Triuridaceae shows some similarity to that of Petrosavia (Liliaceae) and Vallisneria (Hydro-
charitaceae).

Literature: G. ERDTMAN, Pollen morphology and plant taxonomy, Angiosperms (1952) 439. — J. MULLER.

Chromosomes. Chromosome numbers are few and only known from Sciaphila. OHBA & Stnoto found
for the non-Malesian S. japonica 2n = 48; LARSEN for S. thaidanica from N. Thailand 2n= 28; GREEN & SoL-
BRIG for the New Caledonian S. dolichostyla 2n=44, while So.sric noted for S. densiflora (in sched. P.S.
GREEN 1329) the number as 2n= 22.

Literature: P.S. GREEN & O.T. Sotsric, J. Arn. Arb. 47 (1966) 266—269, f. 1—3; K. LARSEN, Dansk Bot.
Ark. 30 (1963) 249; OuBA & Stnoto, Bot. Mag. Tokyo 38 (1924) 203.

Taxon. Following SCHLECHTER (1912), GIESEN (1938) distinguished between Sciaphila BL. and Andruris
SCHLTR, on account of the absence cg. presence of what he called an awl-shaped prolonged connective. Ac-
tually the anther is dorsally attached near the base of the usually very long filament. The apical part of the
latter is easily shed, as GreESEN himself already noted, and hence the structure is often difficult to recognize.

Using this feature as a character on generic level would in my opinion lead to artificial distinctions. This
becomes clearly evident in comparing the type specimens and/or descriptions for example of Sciaphila arfa-
kiana and Andruris anisophylla, and of S. tuberculata and A. clemensiae, which are all otherwise identical;
in fact they represent only a single species, Sciaphila arfakiana.

GigsEN had subdivided the genus into a number of sections and subsections which can be retained. The spe-
cies accepted for Malesia are accommodated in the key almost all according to his subdivision.

Notes. As essential characters for specific distinction are mainly found in the structure of the androecium,
collectors should gather ample material and ensure that male flowers are represented, and check whether
plants with bisexual flowers are extant. Hitherto these are only found in S. maculata and S. tenella. The struc-
ture of the stamens can be best observed in mature buds or very young flowers.

The great influx of material since GiEsEN’s monograph has led to a rather heavy reduction in the number
of accepted species in Malesia and adjacent countries. GrESEN recognized for Malesia 49 spp. and 2 doubtful
ones; one, S. buruensis being added later. In the present revision I recognize 14 spp., plus a doubtful one.

In the ‘Identification Lists of Malesian Specimens’ n. 63, published simultaneously with this revision, all
names in Sciaphila of Malesia and adjacent regions in the West Pacific are listed, with indication of their types
and disposition.

KEY £0 THE SPECIES

1. Plants with bisexual flowers. Mostly also male flowers present towards the apex. Sect. Hermaphroditantha
subsect. Polyandra.
Pav iale ang: DISEXUAl hOWELS With 3 SLAMECNS =. +. eee Sees es cote Pee emer Sees che ee 1. S. maculata
2. Male flowers with 6 stamens. Bisexual flowers with 3—6, but generally 6, stamens ...... 2. S. tenella
1. Flowers unisexual (the female towards the base, the male towards the apex).
3. Perianth of the male flower consisting of 4—8 equal segments. Sect. Oliganthera subsect. Quadrilobatae
(incl. also 14. S. micranthera).
4. Stamens 2. Male perianth segments at the apex with a knob................. 3. S. quadribullifera
4. Stamens 3. Male perianth segments at the apex bearded or glabrous.
5. Male flowers always with 6 perianth segments. Style awl-shaped, in general 2—3 times as long as the car-
(2 aed ae Bie ese icicaeParike.. oii 6 > CIRC RORS CASE MENCI SRN NCE) ROR RASS. CAME Pan IOS YA te 4. S. corniculata
5. Male flowers with 4—8 perianth segments. Style club-shaped, generally as long as the carpel
5. S. secundiflora
3. Perianth of the male flower consisting of 3 larger segments alternating with 3 smaller ones.
6. At least 3, mostly 6, of the male perianth segments at the apex with a stipitate globose to ellipsoid knob.
7. Female perianth segments with a minute knob at the apex (look at buds)........... 6. S. wariana

1984]

TRIURIDACEAE (van de Meerendonk)

bi3

7. Female perianth segments at the apex without appendages.
8. Only the 3 smaller male perianth segments at the apex with a stipe and a small ellipsoid knob. The 3 lar-

ger perianth segments without appendages ..

7. S. nana

8. All male perianth segments with a stipe and knob.........0...0.0. 0000 ceceecee 8. S. arfakiana
6. Male perianth segments at the apex without a stipe or knob, but long-bearded.

9. Stamens 6. Sect. Hexanthera.

10. Young anthers 3-lobed. Flowers all around the stem, rarely most flowers to one side (secund)

10. Young anthers 2-lobed. Flowers generally all to one side (secund) .........

9. Stamens 3.
11. Young anthers 2- or 3-lobed.

12. Anthers 2-lobed. Sect. Oliganthera subsect. Bilobatae..... 0.0.0... eee eee

9. S. densiflora
10. S. corallophyton

11. S. winkleri

12. Anthers 3-lobed. Sect. Oliganthera subsect. Trilobatae.

Pom edicels ie, 5—8 mm long ....:. .76e es «
fae Pedicels '‘c) 2—4mmlongs inc): heed...

12. S. consimilis
13. S. multiflora

11. Young anthers 4-lobed. (Belongs to sect. Oliganthera subsect. Quadrilobatae, see lead 3)

1. Sciaphila maculata Miers, Proc. Linn. Soc. 2
(1850) 72 (n.v.), repr. Ann. Mag. Nat. Hist. II, 7
(1851) 324; Trans. Linn. Soc. 21 (1852) 48; Bru. in
Hook. J. Bot. Kew Misc. 7 (1855) 10; Mia. Fl. Ind.
Bat. 3 (1856) 232; F. v. M. in Walp. Ann. 5 (1860)
917; ScHNIzL. Iconographia | (Suppl.) (1860-67) pl.
57: f. 27, 28; VipAL, Rev. Pl. Vasc. Filip. (1886) 282;
ENGL. in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 238, f.
179: A—F; Becc. Malesia 3 (1890) 331; Merr. En.
Philip. 1 (1923) 28; Gresen, Pfl. R. Heft 104 (1938)
39, f. 7: 1-3. — S. affinis Becc. Malesia 3 (1890)
331, pl. 39: 14-18; Ript. J. Str. Br. R. As. Soc. 7.
33 (1900) 197; Mat. Fl. Mal. Pen. (Monoc.) 2 (1907)
126; J. Fed. Mal. St. Mus. 6 (1915) 188; Merr. En.
Born. (1921) 38; Ript. Fl. Mal. Pen. 4 (1924) 364;
Gresen, Pfl. R. Heft 104 (1938) 37; HENb. Mal.
Wild Fl. (Monoc.) (1954) 203, f. 121. — S. herma-
phrodita Scuutr, Bot. Jahrb. 49 (1912) 76, f. 3:
K-—O; J.J. SmitH, Nova Guinea 14 (1927) 325; Gte-
SEN, Pfl. R. Heft 104 (1938) 38, f. 7: 9-10. — S. mi-
nuta SCHLTR, Bot. Jahrb. 49 (1912) 84, f. 2: O-S;
GieEseEN, Pfl. R. Heft 104 (1938) 68. — S. decipiens
Back. Handb. Fl. Java | (1925) 66; STEEN. Trop.
Natuur 23 (1934) 51; Gresen, Pfl. R. Heft 104 (1938)
37; Back. & Baku. f/f. Fl. Java 3 (1968) 8.

Erect herb, c. 3—16 cm high, mostly simple, some-
times branched at the base. Roots filiform, c.
0.2—0.3 mm in CS, seldom branched, glabrous. Stem
c. 0.4—0.9 (—1.1) mm in CS, mostly glabrous, inter-
nodes c. 3-29 mm. Leaves oblong, acuminate to
acute, c. 1—2.5 by 0.5—0.9 mm, sometimes semi-am-
plexicaul, appressed, but top often patent. Raceme c.
2-8 cm; flowers c. 6—40, all around, sometimes sec-
und. Bracts lanceolate, acute, c. 0.8—2 by c. 0.2—0.5
mm, appressed to the pedicel. Pedicels c. 2~9 mm, c.
0.15—0.25 mm in CS, patent at c. 45-60 (—70)°,
straight for almost the whole length. — o& Flowers
(sometimes absent): perianth segments 6, 3 larger
ones alternating with 3 smaller ones, all completely

14. S. micranthera

reflexed and bearded at the top; larger segments
long-triangular, acuminate, c. 0.75 by c. 0.15 mm;
smaller segments triangular, acute, c. 0.65 by c.
0.20—0.25 mm. Stamens 3, c. 0.2 mm; filaments
short; anthers 3-lobed. — Bisexual flowers: perianth
similar to that of the male flower but smaller differ-
ences between large and small perianth segments.
Stamens 3, very rarely 2, c. 0.2—0.3 mm; filaments
short; anthers 2—3-lobed. Carpels c. 10-30, c.
0.3—0.6 mm long; style inserted laterally at the base
or just above the base, when young just exceeding the
carpel, club-shaped, the apex beset with hairs and pa-
pillae.

Distr. Malesia: Malaya (all parts), Borneo (Sara-
wak), New Guinea (NE. part).

Ecol. Rain-forest (sometimes on ridges), on hu-
mus or between dead leaves, 100—1200 m. F/. April,
July, Nov., Dec.

Notes. When fresh plant wine-red, crimson or
dark-purplish red; flowers red or purplish red.

It is possible to distinguish between S. maculata
(incl. S. minuta) and S. affinis (incl. S. hermaphrodi-
ta and S. decipiens) because male flowers are absent
in the latter. In the Identification List such specimens
have been marked (f). In my opinion, all are conspe-
cific.

I did not see the types of S. affinis and S. deci-
piens, but GiEsEN, who did, thought them to be con-
specific.

2. Sciaphila tenella Bi. Bijdr. 10 (1826) 515; Mus.
Bot. 1 (1851) 321, f. 48; Miers, Trans. Linn, Soc, 21
(1852) 48; Bru. in Hook. J. Bot. Kew Mise. 7 (1855)
10; Mig. Fl. Ind. Bat. 3 (1856) 232; F. v. M.in Walp.
Ann. 5 (1860) 917; ScHNizi. lconographia | (Suppl.)
(1860-67) pl. 57: t. 13-16, 19-25; Brcc. Malesia 3
(1890) 331; Janse, Ann. Jard. Bot. Btzg 14 (1896) 85;
Hemst. Ann. Bot. 21 (1907) 75, pl. 10: 11-17;
Went, Nova Guinea 8 (1909) 165; Koorp. Exk. Fl.

114

FLORA MALESIANA

[ser. I, vol. 10!

Java | (1911) 96; Merr. En. Born. (1921) 38; BAcK.
Handb. Fl. Java 1 (1925) 66; J.J. Smirn, Nova Gui-
nea 14 (1927) 326; STEEN. Trop. Natuur 23 (1934) 51;
GIESEN, Pfl. R. Heft 104 (1938) 40, f. 7: 11—12, incl.
var. robusta GIESEN et var. voigtii GIESEN, /.c. 41;
Merr. & CHUN, Sunyatsenia 5 (1940) 15; Back. &
BAKH./. Fl. Java 3 (1968) 8; ANoNyMous, FI. Haina-
nica 4 (1977) 63, f. 985. — Aphylleia erubescens
CHAMP. Calc. J. Nat. Hist. 7 (1847) 468. — S. eru-
bescens (CHAMP.) Migrs, Trans. Linn. Soc. 21 (1852)
48: Bru. in Hook. J. Bot. Kew Misc. 7 (1855) 10;
Mia. Fl. Ind. Bat. 3 (1856) 232; F. v. M. in Walp.
Ann. 5 (1860) 917; ScHNizL. Iconographia | (Suppl.)
(1860-67) pl. 57: f. 27; THw.En. Pl. Zeyl. (1861) 294;
ENGLER in E. & P. Nat. Pfl. Fam. 2, 1 (1889) 238, f.
179: G—H; Hook. f. Fl. Br. India 6 (1893) 558; in
Trimen, Fl. Ceyl. 4 (1898) 368; Aston, Fl. Ceyl.
Suppl. (1931) 298; Gresen, Pfl. R. Heft 104 (1938)
41, f. 8: 1—3. — S. subhermaphrodita J.J. Smit,
Nova Guinea 14 (1927) 326, pl. 36: 4. — S. torricel-
lensis K. Sco. & SCHLTR in K. Sch. & Laut. FI.
Schutzgeb. Nachtr. (1905) 54, pl. 2; GresENn, Pfl. R.
Heft 104 (1938) 42, f. 8: 4, 5, 9. — S. pumila GIESEN,
Pfl. R. Heft 104 (1938) 39, f. 7: 4-6. — Fig. 3 B15.

Erect herb, at the base a little flexuous, rarely
branched, (1.5—) 5—24 cm high. Roots filiform, c.
0.1—0.3 mm in CS, sometimes branched, glabrous or
with a few hairs. Stem c. (0.3—) 0.5—1.5 mm in CS,
glabrous, internodes c. 3—25 mm long. Leaves ovate
to oblong, acute to acuminate, 1—3 by 0.7—2.0
(—2.5) mm, amplexicaul or semi-amplexicaul, ap-
pressed. Raceme c. 1—16 cm long, with c. 5—50 flow-
ers, flowers more or less all around. Bracts oblong-
ovate to oblong, acute, c. 1—2 (—3) by c. 0.3—0.9
mm, mostly appressed to the pedicel, often the top a
bit patent, rarely patent to the pedicel at 20—30°.
Pedicels c. 2—7 (—8—15) mm long and c. 0.10—0.35
mm in CS, patent at (30—) 50—90°, straight or re-
curved for a smaller part to halfway. — o Flowers:
perianth segments 6, all bearded at the top and com-
pletely reflexed, 3 larger segments alternating with 3
smaller, the larger ones long-triangular, acute, c.
0.6—1.5 (—1.8) by 0.2—0.4 (—0.6) mm; the smaller
ones triangular, acute, 0.5—1.1 (—1.5) by c. 0.2—0.6
mm. Stamens 6, c. 0.2—0.3 mm, filaments c. 0.1 mm
long, connate at the base; anthers 3-lobed. — Bisex-
ual flowers c. 1—3 mm in size; perianth like in the
male flower, but larger segments c. 1.1—2 by c.
0.3—0.7 mm; smaller segments 0.8—1.6 by 0.3—0.6
mm. Stamens 3—6, probably always 6, but easily
broken off; filaments long; anthers (2—) 3-lobed, c.
0.2—0.3 mm. Carpels c. 15—50, obovoid, c. 0.2—0.5
mm long when young, the upper half with tubercles;
style inserted at the base, more or less as long as the
carpel, the apex with hairs and papillae.

Distr. Ceylon and Malesia: Sumatra (Eastcoast,
Bencoolen), Malaya (Pahang, Johore, Singapore),

W. Java, Borneo (Sarawak, Sabah), Philippines
(Mindanao), Celebes (Central part and SE. Penin-
sula), Moluccas (Obi I.), New Guinea (all parts ex-
cept the SW., also in New Britain and Bougainville
I.) and the Solomon Is. (Guadalcanal and San Cris-
tobal Is.).

Ecol. (Solitary) plant, in shade of dense (some-
times somewhat disturbed) rain-forest, often on hill-
sides, on clay, chalk or porous nickel-rich soil, some-
times ultra-basic soil, at various altitudes between 15
and 2250 m. Fi. May—Feb.

Notes. Fresh plant red, purple or pinkish to
coral-pink, flowers red or bright pink, fruits red or
pinkish.

GIESEN distinguished between S. tenella and S.
erubescens by the presence or absence of hairs at the
apex of the perianth segments. But he thought it pos-
sible that their absence on the type specimen of S.
erubescens was due to the fact that it possessed only
very old flowers. In other specimens I found young
flowers with, and old flowers without hairs, on the
same plant. Therefore I decided that only one species
is concerned.

In the absence of male flowers several specimens
are suggestive of S. picta from South America. For
the time being, we regard these as S. fene/la with fe-
male flowers only; in the Identification List such spe-
cimens are marked with (f).

3. Sciaphila quadribullifera J.J. SMitH, Nova Guinea
14 (1927) 324, pl. 35: 1; GresEN, Pfl. R. Heft 104
(1938) SGmiiste

Branched, erect at the base, somewhat flexuous
herb, 6—15 mm high, glabrous all over. Roots fili-
form, c. 0.2—0.3 (—0.4 in liquid) mm in CS, with
many long (c. 0.4 mm) hairs, sometimes branched.
Stem c. 0.4—0.8 mm in CS, internodes c. 3—23 mm
long. Leaves oblong, acute to acuminate, c. 1—3 by
c. 0.5—0.9 mm, not amplexicaul, appressed. Raceme
c. 0.5—1.5 cm long, flowers c. 5—28, all around.
Bracts lanceolate, acute, c. 1—2 by c. 0.3—0.4 mm,
appressed to the pedicel. Pedicels straight, c. 1.5—3.5
mm long and c. 0.2—0.3 mm in CS, patent at c.
30—45°. — o Flowers: perianth segments 4, equal,
oblong-ovate, acuminate, at apex with a stipitate glo-
bose knob, c. 0.6—0.8 by c. 0.4—0.5 mm. Stamens 2,
inserted in front of two opposite perianth segments;
filaments short; anthers 4-celled, 4-lobed. — 9
Flowers: perianth as in the male flower, but segments
acute, 0.6—0.85 by c. 0.35—0.6 mm, apex without a
stipe and knob. Carpels c. 30—50, c. 0.4—0.6 mm,
upper half with tubercles; style inserted laterally
about halfway, when young exceeding the carpel, al-
most club-shaped, apex with papillae.

Distr. Malesia: New Guinea (NW. and NE.
part).

Ecol. In forest, 300—1000 m.

1984] TRIURIDACEAE (van de Meerendonk) 115

A3

Fig. 3. Sciaphila secundiflora THw. ex Bru. Al. Female flower at anthesis, x 10, A2. carpel with style, x
20, A3. fruits, x 5, A4. seed, x 20, AS. young male flower, with stamens closed, x 10, A6. male flower,
with stamens dehisced, x 10. — S. tenella Bi. B/. Bisexual flower, the stamens recurved and seen at their
apex, B2. young stamen, seen from inside, B3. the same, from outside, B4. the same, from above, BS. stamen,
dehisced, from inside, all x 10. — S. arfakiana Brecc. C/. Female flower, x 10, C2. carpel with style, x
20, C3. male flower with young stamens, x 10, C4. young stamen, from outside, C5. the same, from inside,
C6. older stamen, dehisced, from inside, all x 20(A/, 2, 5, 6 VAN ROYEN & SLEUMER 6459, A3, 4 VERSTEEG
1231, B/ BSIP 1104, B2—5 NGF 29310, C/—6 ScuMutTz 3655).

A2

4. Sciaphila corniculata Becc. Malesia 3 (1890) 336,
pl. 39: 5—13; Giesen, Pfl. R. Heft 104 (1938) 56. —
S. neo-caledonica Scuutr, Bot. Jahrb. 39 (1906) 19;
Gutttaum. Bull. Soc. Bot. Fr. 84 (1937) 256; Gik-
SEN, Pfl. R. Heft 104 (1938) 54; Guittaum. Fl. Nouv.
Caléd. (1948) 22. — S. oligochaete Scuitr, Bot.
Jahrb. 49 (1912) 82, f. 3: E~J; Gresen, Pfl. R. Heft
104 (1938) 54, f. 12: 8. — S. gatiensis Scuvtr, Bot.
Jahrb. 49 (1912) 84, f. 2: X-A’; J.J. Smiru, Nova

Guinea 14 (1927) 325; Giesen, Pfl. R. Heft 104
(1938) 56. — S. conferta J.J. SMirn, Nova Guinea 14
(1927) 324, pl. 35: 2; Giesen, Pfl. R. Heft 104 (1938)
54.

Erect, branched herb, 2.5—13 cm high. Roots fili-
form, 0.15—2 mm in CS, with a few hairs. Stem gla-
brous, c. 0.3—0.6 mm in CS, internodes c, 3—20 mm
long. Leaves oblong, acute, c. 0.8—1.5 by c, 0.4—0.7
mm, not amplexicaul, appressed. Raceme c. 0.5~—3.5

116

FLORA MALESIANA

[ser. I, vol. 10!

cm long. Flowers c. 3—9 (—18), all around. Bracts
oblong to oblong-lanceolate, acute, c. 1—1.5 by c.
0.3—0.7 mm, appressed to the pedicel. Pedicels
straight, c. 1—2 (—3) mm long and c. 0.15—0.25 mm
in CS, patent at (10O—) 30—45 (—90)°, flowers not
hanging. — oO Flowers: perianth segments 6, all
equal, oblong, acute, c. 0.5—0.8 by c. 0.25—0.4 mm,
at the top bearded, patent to reflexed. Stamens 3, c.
0.25—0.3 mm; filaments short, at the base connate;
anthers 4-celled, 4-lobed. — 9 Flowers: perianth as
in the male flower but segments 0.5—1.2 mm long
and the top glabrous. Carpels c. 25—60, c. 0.2—0.3
mm long (when young); style inserted laterally, awl-
shaped, in general 2—3 times as long as the carpel,
with tubercles over its whole length.

Distr. Solomons (Kolombangara I.) and Ma-
lesia: New Guinea (NW. and NE. parts, Waigeo I.)
and Moluccas (Obi and Aru Is.).

Ecol. Mixed primary forest and Sapotaceae-
dominated forest, sometimes on limestone rocks,
growing on humus or among dead leaves, jn shade,
75—600 m. Fi. Nov.—Feb., June.

Notes. Fresh plant violet or scarlet. Leaves dark
wine-red or black, flowers carmine, fruits whitish,
with wine-red spots.

WENT Sr came to the erroneous conclusion that S.
corniculata was conspecific with S. nana, and this
was copied by KoorpeErs and Mrs. KOORDERS-SCHU-

_ MACHER; they accepted the name S. corniculata al-
though this is younger than S. nana, and claimed the
species for Java, where it does not occur. See under
S. nana.

5. Sciaphila secundiflora THw. ex Bru. in Hook. J.
Bot. Kew Misc. 7 (1855) 10; Mia. Fl. Ind. Bat. 3
(1856) 232; THw. En. Pl. Zeyl. (1861) 294; Hook. /f.
Fl. Br. India 6 (1893) 558; in Trimen, Fl. Ceyl. 4
(1898) 368; Makino, Bot. Mag. Tokyo 14 (1905) 141;
ALsTON, Fl. Ceyl. 6 (Suppl.) (1931) 298; GrEsEN, Pfl.
R. Heft 104 (1938) 60, f. 14: 1-3. — S. major BEcc.
Malesia 3 (1890) 332, pl. 40: 1-11; RENDLE, J. Bot.
39 (1901) 178; RipL. Mat. Fl. Mal. Pen. (Monoc.) 2
(1907) 126; HEemsL.in Hook. Ic. PI. 29 (1907) t. 2850:
f. 1-6; Rip. J. Fed. Mal. St. Mus. 6 (1915) 188;
Merr. En. Born. (1921) 38; Rip. Fl. Mal. Pen. 4
(1924) 364; GresENn, Pfl. R. Heft 104 (1938) 59. — S.
sumatrana BeEcc. Malesia 3 (1890) 333, t. 40: 12—20;
Ript. J. Fed. Mal. St. Mus. 8 (1917) 119; GIESEN,
Pfl. R. Heft 104 (1938) 63. — S. papuana BeEcc. Ma-
lesia 3 (1890) 335, t. 41: 1—5; J.J. SmitH, Nova Gui-
nea 14 (1927) 325; GrEsEN, Pfl. R. Heft 104 (1938)
60, f. 14: 4-6. — S. macra K. ScH. & SCHLTR in K.
Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 55, pl. 2,
non SCHLTR (1912) which is S. multiflora; GIESEN,
Pfl. R. Heft 104 (1938) 61, f. 14: 8—11. — S. monti-
cola K. Scu. & SCHLTRin K. Sch. & Laut. Fl. Schutz-
geb. Nachtr. (1905) 55; J.J. Smirw, Nova Guinea 14

(1927) 325; GiEsEN, Pfl. R. Heft 104 (1938) 61. — S.
versteegiana WENT, Nova Guinea 8 (1909) 165, pl.
47; J.J. SmitH, Nova Guinea 14 (1927) 325; GIESEN,
Pfl. R. Heft 104 (1938) 63, f. 14: 12-15. — S.
pilulifera SCHLTR, Bot. Jahrb. 49 (1912) 77, f. 1:
Z—C'; J.J. Smit, Nova Guinea 14 (1927) 325. — S.
maboroensis SCHLTR, Bot. Jahrb. 49 (1912) 78, f. 1:
V—-Y; J.J. SmitH, Nova Guinea 14 (1927) 325. — S.
brachystyla SCHLTR, Bot. Jahrb. 49 (1912) 80, f. 2:
J—N; GresEN, Pfl. R. Heft 104 (1938) 61, f. 14: 7. —
S. werneri SCHLTR, Bot. Jahrb. 49 (1912) 80, f. 2:
E—H; Gresen, Pfl. R. Heft 104 (1938) 61. — S. aste-
rias Rip. J. Fed. Mal. St. Mus. 6 (1915) 188; FI.
Mal. Pen. 4 (1924) 365; HENb. J. Mal. Br. R. As.
Soc. 17 (1939) 82.:— S. inornata PETcH, J. Ind. Bot.
Soc. 3 (1923) 226; Aston, Fl. Ceyl. 6 (Suppl.) (1931)
299; GIESEN, Pfl. R. Heft 104 (1938) 67. — Fig. 3
Al-6.

Erect herb, 6—33 cm high, sometimes somewhat
ascending, mostly branched, but often one of the
twigs at a ramification broken off. Roots filiform, c.
0.2—0.3 (—1) mm in CS, with hairs. Stem glabrous,
c. 0.3—1.5 mm in CS, internodes c. 5—50 mm long.
Leaves oblong to ovate, acute, c. 1.5—3 (—4) by c.
0.5—1.5 (—2) mm, sometimes semi-amplexicaul, ap-
pressed or patent to 25°. Raceme c. 0.5—19 cm long.
Flowers c. 3—35, more or less all around. Bracts ob-
long-lanceolate to lanceolate, acute, scale-like, c.
1—3 (—4) by c. 0.3—0.7 (—1) mm, appressed to the
pedicel, or sometimes patent to 10°. Pedicels c. 1—5
(—6) by c. 0.1—0.3 (—0.5) mm in CS, patent at c.
(30—) 45—90°, mostly straight, sometimes very
slightly recurved. — o Flowers: perianth segments
4, or 6 or 7, rarely 5 or 8, equal, patent, long-triangu-
lar, acute, c. 1.5—5 by c. 0.2—0.6 (—2.2) mm, some-
times at about halfway contracted into a long narrow
point, apex glabrous. Stamens 2—3, c. 0.5—0.6 mm;
filaments very short (stamens almost sessile); anthers
4-celled, 4-lobed. — 9 Flowers: perianth segments
(4—) 5-10, patent, equal, c. 2.8 by c. 0.3—0.7 (—1)
mm, oblong to lanceolate, often at about halfway
contracted into an awl-shaped point, apex glabrous.
Carpels c. 20—80, obovoid, c. 0.35—0.6 (—0.8) mm
long, the upper half with many tubercles; style club-
shaped, inserted laterally at the base or about half-
way, the apex with many hairs and papillae.

Distr. Ceylon, Hongkong; in Malesia: N. Suma-
tra (Atjeh), Malaya (all parts), Borneo (Sarawak, Sa-
bah, W. & E. Kalimantan), New Guinea (N. & SE.
parts, Japen and Mios Num Is.), New Britain, New
Ireland, Solomons (San Cristobal).

Ecol. Plant of (damp rocky) rain-forest, often on
rocky terrain, sometimes dominated by Pandanus or
Agathis, on limestone hills but also known from ke-
rangas forest, 15—1250 m. Fi. July—Feb.

Notes. Fresh plant white (when young), red, pale
mauve or purplish; flowers white, mauve red or pur-

1984]

TRIURIDACEAE (van de Meerendonk)

117

ple (perianth segments sometimes with dark bor-
ders), anthers white, fruits red.

PETCH (1923, see above) regarded S. inornata as
‘most closely allied to S. secundiflora’ and ‘resemb-
ling S. sumatrana’ but, knowing about the characters
he used distinguishing between his species and the
two others and from his description of S. inornata
(though I did not see the type), I think it safe to com-
bine them.

With regard to several species here combined,
which were still kept apart by GrEsEN, his own key
even does not work.

I did not see the type of S. asterias Riwi., but I did
see RrpLey 16312 which, according to GIESEN, is al-
most cotypical with the type.

6. Sciaphila wariana (SCHLTR) MEERENDONK, comb.
noy. — Andruris wariana SCHLTR, Bot. Jahrb. 49
(1912) 71, f. 1: A—E; Gresen, Pfl. R. Heft 104 (1938)
22, f. 3: 15—17; Tuyama, Bot. Mag. Tokyo 52 (1938)
61.

Erect, branched herb, 8—15 cm high. Roots fili-
form, c. 0.2—0.4 mm in CS, with long (to 1 mm)
hairs. Stem glabrous, 0.4—0.7 mm in CS, internodes
4—15 mm long. Leaves oblong-lanceolate, acute,
1—1.5 mm by c. 0.2—0.35 mm, not amplexicaul, ap-
pressed, but the top often patent. Raceme c. 1—2 cm
long. Flowers c. 10—15, all around. Bracts lanceo-
late, acute, c. 1—1.5 by c. 0.20—0.25 mm, appressed
or patent to 10°. Pedicels straight, c. 6—8 mm long
and c. 0.1—0.15 mm in CS, patent at 45—60°. — o
Flowers: perianth segments 6, patent, 3 larger ones
alternating with 3 smaller ones, all oblong and at the
apex with a stipitate (stipe c. 0.15 mm long) subglo-
bose knob; the larger ones c. 0.8—0.9 by c. 0.2—0.25
mm; smaller segments c. 0.7 by c. 0.2 mm. Stamens
3, c. 0.15—0.20 mm; anthers almost sessile, 4-celled,
4-lobed, filament mostly clearly exceeding the an-
ther. — 9 Flowers: perianth segments 6, 0.6—0.8 by
c. 0.2 mm, completely reflexed, all equal in shape
and size, oblong-lanceolate, the apex with a stipitate
(stipe c. 0.03—0.05 mm) very minute knob (mostly
only well visible in buds or young flowers). Carpels
c. 30—40, c. 0.2 mm long (without style); style insert-
ed laterally, c. 0.4—0.9 mm long, the apex acute.

Distr. Malesia: New Guinea (NE. part: Goromia
at Waria R.; Lordberg at S. Hunstein Mts), 3 collec-
tions.

Ecol. In forests, 350-1000 m.

7. Sciaphila nana Bi. Mus. Bot. | (1851) 322, f. 48;
Bru. in Hook. J. Bot. Kew Misc. 7 (1855) 10; Mia.
Fl. Ind. Bat. 3 (1856) 232; F. v. M.in Walp. Ann. 5
(1860) 917; Becc. Malesia 3 (1890) 338; PouLsen,
Medd. Naturh. Foren. Kbhvn (1906) 1; Went, Versl.
Verg. Kon. Ak. Wet., Wis- & Nat. Afd. (1909) 698;
Back. Handb. FI. Java 1 (1925) 65; Sreen. Trop.

Natuur 23 (1934) 50; GreseENn, Pfl. R. Heft 104 (1938)
18, f. 2: 4-10; Back. & Baku. f. Fl. Java 3 (1968) 7.
— S. corniculata (non BrEcc.) WENT, Versl. Verg.
Kon. Ak. Wet., Wis- & Nat. Afd. (1909) 698;
Koorp. Exk. Fl. Java 1 (1911) 96; Koorp.-SCHUM.
Syst. Verz. I, §1 (1912) 6. — Andruris gracillima GtE-
SEN, Pfl. R. Heft 104 (1938) 18, f. 2: 1-3. — Andru-
ris nana (B.) GIESEN, /.c. — Andruris loheri GIESEN,
l.c. 19, f. 3: 1-4.

Erect, mostly branched herb, c. S—15 cm high.
Roots filiform, c. 0.2 mm in CS, with hairs. Stem c.
0.2—0.8 mm in CS, glabrous, internodes c. 3-18 mm
long. Leaves not amplexicaul, appressed, oblong to
lanceolate, acute, c. 1—2 by c. 0.3—0.7 mm. Raceme
c. 0.5—5 cm long. Flowers c. 7—35, all around.
Bracts oblong-lanceolate to lanceolate, acute, c.
0.6—1.3 by c. 0.2—0.3 mm, appressed to the pedicel
or patent to 25° toit, in the latter case mostly perpen-
dicular to the stem. Pedicels c. 2—7 mm long and c.
0.1—0.2 mm in CS, patent at c. 30—60 (—70)°,
straight, sometimes slightly recurved at the top. — &
Flowers: perianth segments 6, 3 larger alternating
with 3 smaller, all oblong and patent to reflexed; the
larger segments acute, without appendages, c. 0.7—1
by c. 0.3—0.35 mm; the smaller segments at the apex
with a stipitate, small, ellipsoid knob, c. 0.5—0.8 by
c. 0.2—0.3 mm. Stamens 3, c. 0.3 mm; filaments
often exceeding the 4-celled, 4-lobed anthers. — Q
Flowers: perianth segments patent, (4—) 5—6, equal,
oblong, acute, c. 0.5—0.6 by c. 0.2—0.35 mm, apex
without appendages. Carpels c. 20—40 (—70), c.
0.3—0.35 mm; style inserted laterally near the top,
awl-shaped, c. 0.4—0.8 mm long, apex acute.

Distr. Malesia: Sumatra (Banka), Malaya (Pe-
rak, Pahang), W. Java, Philippines (Luzon).

Ecol. Dense forest, sometimes under bamboo,
250—500 m, once at c. 1150 m. Fi. Aug.

Vern. Tjengtleng, S.

Notes. Stem and flowers of fresh plant purple.

WENT Sr (see above) came to the conclusion that
West Javanese specimens belonged to S. corniculata
and hinted at the conspecificity of that species with
S. nana.

8. Sciaphila arfakiana Becc. Malesia 3 (1890) 337, t.
41: 6—14; Giesen, Pfl. R. Heft 104 (1938) 57, f. 13:
4. — S. crinita Becc. Malesia 3 (1890) 338, pl. 42:
1—9; Scuttr, Bot. Jahrb. 49 (1912) 71. — S. anda-
jensis Becc. Malesia 3 (1890) 339, pl. 42: 10-14;
Went, Nova Guinea 8 (1909) 166; J.J. Smrrn, ibid.
14 (1927) 323. — S. clemensae Hemsv. Hook. Ic. PI.
29 (1907) pl. 2850: f. 7—14; Merr. En. Born. (1921)
38; En. Philip. 1 (1923) 28; Back. Handb, Fl. Java
1 (1925) 65; Steen. Trop. Natuur 23 (1934) 51, f. 9;
Back. & BAKu./. Fl. Java 3 (1968) 7. — S. australa-
sica Hemst. Kew Bull. (1912) 44; Domin, Bibl. Bot.
85 (1926) 256. — Andruris crinita (Becc.) SCHLTR,

118

FLORA MALESIANA

[ser. I, vol. 10!

Bot. Jahrb. 49 (1912) 71; Tuyama, Bot. Mag. Tokyo
52 (1938) 22. — Andruris andajensis (BECC.)
SCHLTR, Bot. Jahrb. 49 (1912) 71; Tuyama, Bot.
Mag. Tokyo 52 (1938) 61; GrEsEN, Pfl. R. Heft 104
(1938) 28. — Andruris celebica SCHLTR, Bot. Jahrb.
49 (1912) 72, f. 1: F—L; Tuyama, Bot. Mag. Tokyo
52 (1938) 61. — Andruris tenella SCHLTR, Bot.
Jahrb. 49 (1912) 74, f. 1: M—Q; Tuyama, Bot. Mag.
Tokyo 52 (1938) 61. — S. inaequalis SCHLTR, Bot.
Jahrb. 49 (1912) 77, f. 1: R-—U; J.J. Smitry, Nova
Guinea 14 (1927) 324; Gresen, Pfl. R. Heft 104
(1938) 58, f. 13: S—7. — S. atroviolacea SCHLTR, Bot.
Jahrb. 49 (1912) 79, f. 2: A—D; GrgseEn, Pfl. R. Heft
104 (1938) 57, f. 13: 2-3. — S. vitiensis A.C. SMITH,
Bish. Mus. Bull. 141 (1936) 15, f. 5; GresEn, Pfl. R.
Heft 104 (1938) 28. — Andruris anisophylla GIESEN,
Pfl. R. Heft 104 (1938) 23, f. 4: 2-6. — Andruris cle-
mensae (HEMSL.) GIESEN, Pfl. R. Heft 104 (1938) 23,
f. 4: 7—9, incl. var. borneensis GIESEN, /.c. 25. — An-
druris australasica (HEMSL.) GIESEN, Pfl. R. Heft 104
(1938) 25, f. 4: 10—13. — Andruris elegans GIESEN,
l.c. 25, f. 5: 1-4; Hosokawa, J. Jap. Bot. 16 (1940)
540. — Andruris javanica GIESEN, Pfl. R. Heft 104
(1938) 27, t. 5: 5-9. — Andruris vitiensis (A.C.
SMITH) GIESEN, /.c. 28; A.C. SmiTH, Sargentia 1
(1942) 5; Bull. Torrey Bot. Club 70 (1943) 534; Par-
HAM, Plants Fiji (1964) 257. — S. tuberculata GiE-
SEN, Pfl. R. Heft 104 (1938) 57, f. 12: 9-12. — S. va-
lida GiESEN, I.c. 59, f. 13: 8-11. — Andruris pala-
wensis TUYAMA, Bot. Mag. Tokyo 52 (1938) 63. —
Andruris buruensis J.J. SMitH, Bull. Jard. Bot. Btzg
III, 16 (1939) 111. — Fig. 3 C1—-6.

Erect, simple or branched, sometimes a bit flex-
uous herb, c. (2—) 4—28 cm high. Roots filiform, c.
0.1—0.4 mm in CS, with hairs. Stem c. 0.3—1 (—1.5)
mm in CS, glabrous, internodes c. 4—33 mm long.
Leaves oblong to lanceolate, appressed, not amplexi-
caul, c. 1—3 by 0.2—1 mm, acute to acuminate. Ra-
ceme c. 0.5—14 cm long; flowers c. 5—65, all around,
very rarely all flowers to one side. Bracts oblong-lan-
ceolate, acute, c. 0.5—2.5 by c. 0.1—0.5 mm, ap-
pressed to the pedicel (and the top mostly patent) or
patent from the pedicel to 40° (and in that case most-
ly perpendicular to the stem). Pedicels c. 2—21 mm
long and c. 0.1—0.3 (—0.4) mm in CS, patent at
20—90° (mostly 30—45°), straight or the apical part
recurved. — Oo Flowers: perianth segments 6, 3 lar-
ger ones alternating with 3 smaller ones, all oblong-
ovate, the apex with a stipitate globose to ellipsoid
knob; larger segments c. 0.7—1.7 by c. 0.3—0.5 mm,
smaller segments c. 0.6—1.4 by c. 0.35—0.5 mm. Sta-
mens 3, c. 0.3—0.5 mm; filaments clearly exceeding
the 4-celled, 4-lobed anthers (but the acute apex of
the filament often broken off). — 9 Flowers: peri-
anth segments 6, more or less equal, oblong to trian-
gular, acute, c. 0.5—1 by c. 0.25—0.50 (—0.65) mm,
apex without appendages, but often thickened. Car-

pels c. (1O—) 20—40 (—70), c. 0.2—0.5 mm long; style
awl-shaped, acute, inserted laterally, c. 0.6—1.8 mm
long, amply exceeding the carpel.

Distr. Micronesia (Palau), W. Polynesia (Fiji
Is.: Viti Levu, Vanua Levu, Vanua Mblalavu), Solo-
mons (Bougainville); in Malesia: New Guinea (all
parts except the SW.; also in New Britain and Manus
Is.), Philippines (Mindanao), Moluccas (Ceram,
Ambon), Celebes (N. Peninsula and Central part),
Lesser Sunda Is. (Flores), West Java, Borneo (Sa-
bah, W., S. & E. Kalimantan), Malaya (Kelantan,
Pahang), and Central W. Sumatra.

Ecol. Rain-forest (low montane or montane),
often on a hill, cliff, ridge, crest or in a river valley,
on sandstone and other bedrock, in shade of trees or
fern clumps; on humus, one time on a termite hill.
Sometimes associated with Corsia, Burmannia or
Epirixanthes; 100—2130 m. Fil. Jan.—Dec.

Notes. Fresh plant pink, pale mauve, red or pur-
ple; flowers light brown, red or purple with white or
yellowish anthers, fruit reddish blue, red-purple or
pale pink (later orange-brown). The plant flowers
and then dies; scarce.

In BUWALDA 6161 (from Ambon) and vAN ROYEN
& SLEUMER 6268 (New Guinea: Mt Cycloop) the style
does not clearly exceed the carpel.

9. Sciaphila densiflora SCHLTR, Bot. Jahrb. 49 (1912)
87, f. 3: U—X; GiEsEN, Pfl. R. Heft 104 (1938) 46, f.
9: 1-2. — S. reflexa SCHLTR, Bot. Jahrb. 49 (1912)
87; GIESEN, Pfl. R. Heft 104 (1938) 48, f. 9: 9-10. —
S. longipes ScHutr, Bot. Jahrb. 49 (1912) 88; J.J.
SmiTH, Nova Guinea 14 (1927) 326, pl. 36: 3; GIESEN,
Pfl. R. Heft 104 (1938) 46, f. 9: 3. — S. trichopoda
SCHLTR, Bot. Jahrb. 49 (1912) 89, f. 3: P—T; GiEsEN,
Pfl. R. Heft 104 (1938) 48, f. 9: 7-8. — S. flexuosa
GIESEN, Pfl. R. Heft 104 (1938) 45, f. 8: 15—17. —
S. nutans GIESEN, /.c. 46, f. 9: 4-6. — Fig. 4.
Erect herb, seldom branched and if so, then at the
base, c. 6—43 cm high. Roots filiform, c. 0.15—0.30
mm in CS, sometimes branched, with hairs. Stem c.
0.5—1.2 mm in CS, glabrous, internodes c. 3—55 mm
long. Leaves oblong-ovate to ovate, acute, c. 1.5—5
by c. 0.6—2.5 mm, mostly semi-amplexicaul, ap-
pressed. Racemes 1—21.5 cm long, with c. 7—120
flowers, from very dense (100 flowers on 12 cm) to
rather lax (35 flowers on 14 cm). Flowers all around,
sometimes with a tendency towards one side. Bracts
lanceolate, acute, c. 1—2 (—3.5) by c. 0.4—1 (—1.5)
mm, mostly appressed to the pedicel, the top often a
little (10—20°) patent. Pedicels 3—25 mm long and
0.1—0.2 mm in CS, (45—) 60—90° patent, recurved,
straight for a smaller or larger part of the length,
sometimes a bit sinuous, the flowers always hanging
down. — oO’ Flowers: perianth segments 6, 3 larger
ones alternating with 3 smaller ones, all completely
reflexed and at the top long-bearded; larger segments

1984]

TRIURIDACEAE (van de Meerendonk)

119

Bl

Fig. 4. Sciaphila densiflora Scurtr. D/. Habit, « 1,

D2. male flower, x 10, D3. mature stamen, from

outside, x 10, D4, the same, from inside, x 10(D/
Brass 28255, D2—4 BW 8553).

long-triangular, (0.6—) 1.3—2 by (0.2—) 0.3—0.5 mm,
the top abruptly contracting into a long narrow point
(c. 1/3 of the length); smaller segments long-triangu-
lar, acute, (0.5—) 0.8—1.5 by c. 0.2—0.4 mm wide.
Stamens 6, c. 0.2 mm; filaments short; anthers
3-lobed. — Q Flowers: perianth like in the male
flower, but the larger segments c. 1—2.5 by c.
0.3—0.7 mm and the smaller segments 0.7—1.5 by c.
0.4—0.6 mm. Carpels c. 15—40, 0.3—0.55 mm; style
club-shaped, inserted laterally at the base or about
halfway, exceeding the carpel when young; apex with
many hairs and papillae.

Distr. Ceylon; in Malesia: Lesser Sunda Is. (Ti-
mor, Flores), Borneo (Sarawak, W. & E. Kaliman-
tan; Natuna Is.), Philippines (Luzon), Moluccas
(Halmaheira), New Guinea (all parts except the SW.
part; also in Normanby, Rossel and Sudest Is.) and
New Caledonia.

Ecol. Local and uncommon plant in rain-forest,
often on ridge crests, dry ridges, limestone rocks,
clay or sandy or poor stony soils, rooting in raw hu-
mus or deep leaf litter, mostly in shade, 100—1200 m,
once at c. 1950 m. Fi. Feb.—Nov.

Vern. New Guinea: ware, Uruaru lang., Purari
River.

Notes. Fresh plant coral-pink, purplish or red;
flowers red with yellow stamens, fruits red.

In NGF 19532 (K, liquid) the pedicels are almost
parallel to the stem.

10. Sciaphila corallophyton K. Scu. & SCHLTR in K.
Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 54, t. 2:
A: a—d; ScuHitr, Bot. Jahrb. 49 (1912) 76, 89; Gir-
SEN, Pfl. R. Heft 104 (1938) 45, f. 8: 10—12, incl. var.
gracilis GIESEN, /.c.; HANSEN, Dansk Bot. Ark. 25
(1969) 88. — S. dolichostyla Scuitr, Bot. Jahrb. 39
(1906) 19; ScHinz in Sarasin & Roux, Nova Caledo-
nica, Bot. 1 (1920) 59; GumLLaum. Bull. Soc. Bot. Fr.
84 (1937) 256; GreEsEN, Pfl. R. Heft 104 (1938) 45, f.
8: 13—14; Gutttaum. Fl. Nouv. Caléd. (1948) 22;
Mém. Mus. Hist. Nat. Paris n.s. Bot. 8 (1959) 189.

Erect, mostly simple herb, c. 5S—23 cm high. Roots
filiform, c. 0.15—0.25 mm in CS, with a few hairs.
Stem c. 0.5—1.5 (—2) cm in CS, glabrous, internodes
c. 4—20 mm long. Leaves oblong-lanceolate, acute,
c. (l—) 1.5—2.5 by c. 0.5—1.5 mm, not amplexicaul,
appressed to the pedicels. Raceme c. 0.5—12 cm long,
with c, 4—32 flowers, generally most of the flowers
to one side. Bracts lanceolate, acute, c. 1—2 by c.
0.4—0.8 mm, appressed to the pedicel, but the top
patent. Pedicels c. 2~5 mm long, c. 0.2—0.4 mm in
CS, patent at c. 35—45°, recurved but sometimes
straight for half the length or less. — o Flowers: per-
ianth segments 6, 3 larger ones alternating with 3
smaller ones, all reflexed and at the top bearded; the
larger segments oblong-lanceolate, acute, c, 1.1—1.7
by c. 0.3-0.5 mm; smaller segments oblong, acute,

120

c. 1—1.4mm by c. 0.3—0.6 mm. Stamens 6; filaments
short, anthers 2-lobed. — 9 Flowers: perianth like
in the male flower, but the top of the segments gla-
brous and the larger segments 1.3—1.8 by 0.4—0.6
mm; smaller segments c. 1.1—1.4 by c. 0.4—0.6 mm.
Carpels c. 30—50, c. 0.3—0.4 mm; style club-shaped,
inserted laterally near the base, exceeding the carpel
when young, at apex with many hairs and papillae.

Distr. Micronesia: Carolines (Ponape), Melane-
sia (New Caledonia); in Malesia: New Guinea (NE.
part).

Ecol. In forest, on rockwalls, along clay streams,
400—2100 m. Fi. Jan.—Dec.

11. Sciaphila winkleri SCHLTR, Bot. Jahrb. 48 (1912)
88; MerRrR. En. Born. (1921) 38; GrEsEn, Pfl. R. Heft
104 (1938) 52, f. 11: S—7. — S. hydrophila ScHLTrR,
Bot. Jahrb. 49 (1912) 85, f. 2: T—W; GigsEn, Pfl. R.
Heft 104 (1938) 51, f. 11: 1—4.

Erect, branched herb, c. 3—14 cm high. Roots c.
0.2—0.3 mm in CS, with a few hairs. Stem c. 0.3—0.5
mm in CS, glabrous, internodes c. 3—21 mm long.
Leaves oblong, acute, c. 1—2 (—2.5) by c. 0.5—0.9
mm, semi-amplexicaul, appressed or patent at c. 10°.
Raceme c. 0.5—9.5 cm long, with c. 2—40 flowers,
mostly all flowers to one side. Bracts oblong-lanceo-
late, acute, sessile, c. 1—1.5 (—2) by c. 0.3—0.5 mm,
appressed to the pedicel. Pedicels c. 1—2 (—3) mm
long and c. 0.1 (—0.25, liquid material) mm in CS, c.
60—90° patent, recurved. — o Flowers: perianth
segments 6, 3 larger alternating with 3 smaller ones,
all reflexed and at the top bearded; larger segments
oblong, acute, c. 0.5—0.8 by c. 0.3 mm; smaller seg-
ments oblong, acute, c. 0.4—0.6 by c. 0.25 mm. Sta-
mens 3, c. 0.2—0.3 mm, filaments short and at the
base connate, anthers 2-lobed. — 9 Flowers: peri-
anth like in the male flower but larger segments c.
1—1.2 by c. 0.5—0.6 mm, smaller segments c. 0.7—1
mm by c. 0.4 mm. Carpels c. 40—80, obovoid, c.
0.3—0.7 mm; style club-shaped, inserted laterally at
the base or about halfway, when young mostly just
exceeding the carpel; apex with many hairs and papil-
lae.

Distr. Malesia: Borneo (Sarawak, W., E. & S.
Kalimantan), New Guinea (NW. and NE. parts).

Ecol. Rain-forest, rooting between decaying
leaves or in humus, in deep shade, 80—180 m.

Note. Fresh plant red.

12. Sciaphila consimilis BL. Mus. Bot. 1 (1851) 322;
Bru. in Hook. J. Bot. Kew Misc. 7 (1855) 10; Mia.
Fl. Ind. Bat. 3 (1856) 232; F. v. M.in Walp. Ann. 5
(1860) 917; GresEn, Pfl. R. Heft 104 (1938) 51, f. 10:
7-9.

Simple, erect but especially at the base a bit flexu-
ous herb, c. 5.5—22 cm high. Roots c. 0.15—0.5 mm
in CS, with hairs. Stem c. 0.2—0.5 mm in CS, gla-

FLORA MALESIANA

[ser. I, vol. 10!

brous, internodes c. 5—18 mm long. Leaves oblong-
lanceolate, acute, c. 1—3 by c. 0.4—1.3 mm, semi-
amplexicaul or not, appressed, the top mostly a bit
patent. Raceme c. 2—14 cm long, flowers c. 15—70,
all around. Bracts lanceolate, acute, c. 1—2 by c.
0.2—0.4 mm, sessile, mostly appressed to the pedicel,
sometimes 10—30° patent. Pedicels c. 5S—8 mm long
and c. 0.1 mm in CS, 45—90° patent, recurved or
often curled. — o Flowers: perianth segments 6, 3
larger ones alternating with 3 smaller ones, all re-
flexed and at the top long-bearded; larger segments
oblong-lanceolate, acute, c. 0.75-0.9 by c.
0.25—0.45 mm, smaller segments (oblong-)lanceo-
late, acute, c. 0.5—0.7 by c. 0.2—0.3 mm. Stamens 3,
c. 0.25—0.3 mm; filaments short and at the base con-
nate; anthers 3-lobed. — 9 Flowers: perianth like in
the male flower, but the top glabrous or with very
few, short hairs and larger segments 0.7—1 by
0.3—0.45 mm, smaller segments 0.5—0.8 by 0.25—0.3
mm. Carpels c. 15—30, 0.2—0.25 mm; style club-
shaped, inserted laterally at the base or about half-
way, exceeding the carpel when young; apex with
many hairs and papillae.

Distr. West Polynesia (Fiji Is.: Vanua Levu); in
Malesia: Philippines (Luzon, Mindanao).

Ecol. In forest, amongst thick carpet of leaves.

Note. Stem curved whilst. growing from under
leaves to get to the light. Stems and fruits of fresh
plant red.

13. Sciaphila multiflora GrEsEN, Pfl. R. Heft 104
(1938) 49, f. 10: 1-2. — S. macra SCHLTR, Bot.
Jahrb. 49 (1912) 86, f. 3: A—D, non K. Scu. &
SCHLTR in K. Sch. & Laut. (1905), which is S. secun-
diflora. — S. mindanaensis GIESEN, Pfl. R. Heft 104
(1938) 51, f. 10: 3-6. — S. stemmermannii Foss. &
SACHET, Pac. Sci. 34 (1980) 15, f. 1-2.

Erect, sometimes a bit flexuous herb, 6—40 cm
high, branched (mostly at the base). Roots c. 0.2—0.3
mm in CS, glabrous, seldom branched, sometimes
with a few hairs. Stem c. 0.4—1:3 mm in CS, gla-
brous, or with a few hairs, internodes c. (5S—) 12—50
mm long. Leaves oblong-ovate to oblong-lanceolate,
acute, c. 1.5—3 by 0.8—1.2 mm, not amplexicaul, ap-
pressed. Raceme c. (2—) 7—31 cm long; flowers c.
8—40, all around or more or less to one side. Bracts
oblong-lanceolate, acute, c. 1—2 by 0.4—0.8 mm, ses-
sile, appressed to the pedicel but the top mostly a bit
patent. Pedicels c. 2—4 mm long and c. 0.1—0.3 mm
in CS, 45—90° patent, recurved. — o Flowers: peri-
anth segments 6, 3 larger ones alternating with 3
smaller ones, all reflexed and at the top bearded; lar-
ger segments oblong, acuminate, c. 0.8—1.3 by c.
0.3—0.5 mm, smaller ones oblong, acuminate, c.
0.6—1.1 by c. 0.3—0.4 mm. Stamens 3, c. 0.4 mm, fi-
lament short and at the base connate, anthers
3-lobed. — 9 Flowers: perianth like in the male

1984]

flower but the apices glabrous or bearded, larger seg-
ments c. 0.9—1.5 by c. 0.3—0.8 mm, smaller seg-
ments c. 1.1—1.2 by c. 0.4—0.6 mm. Carpels c.
10—40, c. 0.2—0.35 mm; style club-shaped, inserted
laterally at the base or about halfway, exceeding the
carpel when young; apex with many hairs and papil-
lae.

Distr. Micronesia (Carolines: Palau); in Malesia:
New Guinea (E. part: Waria area; Milne Bay Distr.)
and Philippines (Mindanao).

Ecol. Plant of primary forest, sometimes on
steep slopes; 30—800 m. F/. May, July.

Note. Fresh plant reddish purple, with tinged
pink or purple flowers, anthers yellow; fruits dark-
purple.

14. Sciaphila micranthera GIEsEN, Pfl. R. Heft 104
(1938) 54, f. 12: 1-4.

Erect herb, c. 7—13 cm high, simple or branched at
the base. Roots c. 0.15—0.2 mm in CS, with a few
hairs. Stem c. 0.3—0.7 mm in CS, glabrous, inter-
nodes c. S—10 mm long. Leaves oblong to lanceolate,
acute, c. 1—3 by c. 0.4 mm, not amplexicaul, ap-
pressed or 20° patent. Raceme c. 2.5—11 cm long;
flowers c. 15—90, more or less all around. Bracts lan-
ceolate, acute to acuminate, c. 1—2 by c. 0.15—0.3
mm, more or less appressed to the pedicel. Pedicels
c. (S—) 7-15 mm long and c. 0.15—0.2 mm in CS,

TRIURIDACEAE (van de Meerendonk)

121

more or less perpendicular to the rachis, straight to
slightly recurved, sometimes somewhat sinuous. —
o Flowers: perianth segments 6, 3 larger ones alter-
nating with 3 smaller ones, all reflexed and at the
apex bearded; the larger ones oblong-lanceolate,
somewhat obtuse, c. 1 by 0.25 mm; the smaller ones
oblong-lanceolate, obtuse, c. 0.8 by c. 0.2 mm. Sta-
mens 3, c. 0.25—0.3 mm, almost sessile; filaments
very short, anthers 4-celled, 4-lobed. — 9 Flowers:
perianth like in the male flowers; carpels c. 15—25, c.
0.3—0.35 mm; style inserted laterally about halfway,
exceeding the carpel, club-shaped, the apex with
hairs and papillae.

Distr. Malesia: Borneo (Sarawak and W. Kali-
mantan: Bt. Kenepai).

Ecol. Primary forest, growing on thick humus,
300 m.

Note. Fresh plant dark-red all over.

Doubtful

Sciaphila papillosa Becc. Malesia 3 (1890) 334, pl.
39: 1—4; GreEsEN, Pfl. R. Heft 104 (1938) 67.

Based on a specimen of BEccari from NW. New
Guinea, Vogelkop Peninsula, Hatam, Mt Arfak in
vil-1875 (FI?, n.v.). BECCARI has not seen any male
flowers, so it is not possible to identify this species.

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FLORA MALESIANA

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and the Rijksherbarium, Leyden, Holland,
executed by Foundation Flora Malesiana

Scientific Communications
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Series I— Spermatophyta (Flowering Plants)

. Cyclopaedia of collectors & collections. 1950. pp. clii+ 639 (microfiche edition)

. Malesian vegetation

. Malesian plant geography

. General chapters and revisions. 1948—1954. pp. ccix + 631 (microfiche edition)

. Bibliography, specific delimitation & revisions. 1955—1958. pp. cccxlii+ 596

. Systematic revisions. 1960-1972. pp. 20+ 1023 (microfiche edition)

. Systematic revisions. 1971—1976. pp. 18+876

. Cyclopaedia of Collectors, Suppl. 2. Systematic revisions. 1974—1978. pp. 19+577
. Systematic revisions. 1979—1983. pp. 48 + 600

. Systematic revisions. part 1. 1984. pp. 121

Series II — Pteridophyta (Ferns & Fern Allies)

. Taxonomical Revision. 1959-1982. pp. (20) + xxiv + 600

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