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MALESIANA

SERIES I - SPERMATOPHYTA
Flowering Plants

Vol. 10, part 2

Revisions

INDEX TO REVISED FAMILIES

PAL COTACEAC i cles 6 «bree e's 4: 3,592. Dipsacacede Wert vo ds 4: 290
Actinidiaceae s. str. ..... 4: 37 Dipterocarpaceae........ 9: 237
FATZOACCUGS: fh Me wind nsw ore x dhe 4: 267 . Droseracede 22+\.. SST 5 Sv 597
Alismataceae .... 5: 317; 6: 915 Elaeagnaceae .......... 10: 151
Alseuosmiaceae ........ 10335. Blatniacenet tines. aascrta 4: 203
Amaranthaceae Epacridacede ast iva. 6: 422
As 69,°593° Ge 915y ericacede nis ect 6: 469, 943
Anacardiaceae .......... 8: 395). Erythroxylaceae:..'. 24... 5: 543
Ancistrocladaceae ....... Be Give, LRARACEAGL area nate oie & 7: 265
Aponogetonaceae . 4: 11; 7: 213. Flacourtiaceae
A valtaceae-Ts gis is's hs Saccleta Ol DP 15. S052) 62-943- 5 75° 827
Aristolochiaceae ....... 102753) Flavellaniacede:.i.426. 4: 245
Balanophoraceae........ TS. 18S) WIGCTANIACEAE cisievie atte 6: 445
BHSEMACEAG! oh nile. .(ctersict cass Ae. S008, | GmetacedSunae sis 4: 336; 6: 944 -
BataCede tes fern sie fins 2 93414) Gonystylacede) iia vies a2 4: 349
Betulaceae ...... 5: 207; 6: 917 Goodeniaceae
BIONOMIACEHC co aie whe ahi. 8: 114 Bt 539585075, 8.9495 7s B27
Bixdcede’s) Str. 2. dace uss 4: 239° ‘Haemodoraceaé..... 5)... Soul
Burmanniaceae...... Asm Sie s92. a) EAIOLARACCAC s sinen casein. 13 239
Burseraceae Hamamelidaceae........ eos
pe 209). 5075.6: 9175" 721820. “Hippocrateaceae:.7.5...°. ..: 6: 389
BWEOIISCEAG cee Sees 5:0 < 5: 118 Hydrocharitaceae
‘ Byblidaceae ............ 7: 135 5: 381; 6: 952
Gallitrichacede «ssa... 4: 251 MHydrophyllaceae........ 4: 207
Campanulaceae .... 6: 107, 928 Hypericaceae ........... $y 41
PaniaDACEdC rosa... crete Ath 258 WUGAGHIACEAG wie Sulcus ears anal
Capparacede., 2c chi.3 3)2,2 Of s in Os (0 (ce Te aa ane ae S25, 77
Caprifoliaceae Wustan@aceae: 2. cha. n>. 6: 143
BAS. SOSsOTHOZ BY) | WUNCACEAG)s« 5 os ba ticbe ahs 4: 210
Cardiopteridaceae ....... Usa, 9307, UNCAPINACEAE: <2 aei..63 52 Ae Si!
(elastracese. 6: 227,389, 930. . Labiatae’. . <i. ae. 8: 301
Centrolepidaceae........ oman PICCACCAGS fate cide aiideinee Temas
Ceratophyllaceae........ cS BY va (er (Se 7: 219
Chenopodiaceae Lentibulariaceae ........ 83275
ae 99-8 594-6: 932) Liliaceae. ! 33 ks 2 c.503 9: 189
Chloranthaceae ........ ROS 1256 Woraniaceae ) 4.05.61: 6: 293, 953
IPIFIRTACGAG . tots: 23s sis Os 7: 139 Lophopyxidaceae........ 7: 89
Cochlospermaceae....... 4: 61 Malpighiaceae .......... Seales
Combretaceae Martyniaceae ........... 4: 216
4: 533; 5: 564; 6: 932 Menispermaceae ....... 10: 157
Connaraceae .... 5: 495; 6: 933 Monimiaceae.......... 10: 255
Convolvulaceae.... 4: 388, 599; Moringaceae............ 4: 45
eh 56sn0s, AG, TeiSooF Mi yOpordceae )) 2s sa. one 4: 265
SOMmMGede tin -w tu bad Bee OS e IVINTICACEAS sates tus wssevacte LPH
Corynocarpaceae 4: 262; 5: 557 Najadaceae............. 6: 157
Crassulaceseistis 23.30.00 4: 197 = Nyctaginjaceae. | 202.30 00 6: 450
Crypteroniaceae......... S218) IN VSSAGEAR ch ers eee 4: 29
Cyperaceae...... Tae aao5: Sr TOT 4s OCNMACEACr ies hint so sae de OF
WAtISGaCedGe tins sisjesie. ss 4:°382" SQ lacdcede <s oe. eee AOe tuk
Dichapetalaceae . 5: 305; 6: 941 Omagraceae............. 8: 98
Dilleniaceae ..... 45-141 139824". Opiliaceae. 3, he 10: 31
Dioscoreaceae .......... 4: 293: — (Omalidatéae >i asc ue RAs

Papaveraceae .......6.6e 5: 114
Passifloraceae ........45 7: 405
Pedaliatede . :... a7 4: 216
Pentaphragmataceae..... 4: 517
Pentaphylaceae ......... 5$) 12)
Philydraceae. . (ci aan A'S
Phytolaccacede.. .2/2aa0e 4: 229
Pittosporaceae ; 3... Sa. 5: 345
Plumbaginaceae......... 4: 107
Podostemaceae ... 4: 65; 6: 963
Polemoniaceae .- 322.0 4: 195
Pontederiacede: : sa.aeen 4: 255
Portulacaceae. . > os. eeee Je. 121
Primulaceae .. .... ssseeees 6: 173
Proteaceae ...\.: 54 eee 5: 147
Punicaceae ... +... .s:aee 4: 226
Restionaceae...... . ...ueeae 5: 416
Rhizophoraceae
5: 429; 6: 965; 8: 550
Salicaceae ........ 7) see 5: 107
Salvadoraceae ..cc9aeee 4: 225
Sarcospermaceae ........ 4: 32
Saururacede .... /. saa 4: 47
Scyphostegiaceae
5: 297; 62°967.
Simaroubaceae ..... 6: 193, 968
Sonneratiaceae
4: 280, 513; 6: 973
Spargariiaceae |. sae 45.233.
Sphenocleaceae ......... Asta 2).
Sphenostemaceae....... 10: 145
Stackhousiaceae......... 4:55)
Staphyleaceae........2.: : 49
Stylidiaceae: 42. 5) Seaman 4: 529
Styracaceéae.: «. 2. eee 4: 49
Symplocaceae ».. 7 aoe 2-205
Taccaceae |.) 2 ae 7: 806
Thymelaeaceae
4: 349; 6: 1, 976; 7: 830
Trapaceae :.). 5... 4; 43
Trigoniaceae . ..:; aaa 4: 59
Trimeniaceae -2..: saeeee 10: 327
Triuridaceae .0. 52 aaa 10: 109
Turneraceae... .\. sauna 4: 235
Typhaceae........ ae oe eee
Ulmaceae ...::.. ae 8: 31

Umbelliferae

Valerianaceae........... 4: 253

Violaceae.......... 7: 179, 831

Xyridaceae .......+ 4: 366 | S98)
Zygophyllaceae ...... a a =

THE LuESTHER T. MERTZ LIBRARY

THE NEW YORK BOTANICAL GARDEN

4: 113, 595;
5: SSSo0ue 983; 7: 830 if

CHLORANTHACEAE (B. Verdcourt, Kew)

Erect or straggling herbs, shrubs or trees, sometimes monoecious or dioecious,
the herbs sometimes rhizomatous; branches sometimes jointed at the nodes,
sometimes without vessels (Sarcandra). Leaves simple, decussate or sometimes
whorled in fours, serrate, crenate or dentate, the teeth often thickened at the
apex, penninerved, usually petiolate; petioles more or less connected at the base
at least by a transverse line or connate into a distinct sheath; in Ascarina often
alternating with leafless internodes which have the petiolar sheath; stipules mi-
nute to fairly conspicuous, subulate, borne on the petiole bases or sheath, occa-
sionally pectinate. Flowers much reduced, without perianth, fully unisexual or
essentially bisexual with the reduced anther-bearing organ adnate to the side of
the ovary; arranged in spicate, paniculate, or capitate axillary or terminal inflo-
rescences. — Male flowers bracteate or not, apparently consisting of 1—5 sta-
mens, or in Hedyosmum consisting of numerous anthers in a cone-like struc-
ture; if 3 then the whole forming a fused 3-lobed organ sometimes enveloping
the female flower by its edges, the central anther with 2 or aborted loculi and
the laterals with single loculi, simply lobed or with connectives slightly to con-
siderably produced so that the whole organ is 3-fingered; if with only 2 anther
locelli then these on either side of a thickened filament plus connective. — Fe-
male flowers naked or enclosed by a cupular bract, the perianth adnate to the
ovary, often minutely or shortly dentate at the apex and the ovary thus inferior;
ovary 1-locular; stigma sessile or style short; truncate, 2-lipped, depressed or
subcapitate (or horseshoe-shaped in one species), rarely linear or clavate. Ovule
solitary, orthotropous, pendulous, bitegmic and crassinucellate. Drupes fleshy,
small, ovoid or globose, sometimes more or less 3-sided in Hedyosmum, free or
united into a mass by the bracts; endocarp hardened and crustaceous. Seeds sub-
globose, exarillate, with copious fleshy or oily endosperm and minute embryo,
the cotyledons divaricate or scarcely formed.

Distribution. Four genera with about 80 species. Since VESTER’s (1940) small-scale map the
family (Ascarina) has been found in Madagascar. It is mainly tropical but Ascarina extends south
to North Island of New Zealand (fig. 6) and Ch/oranthus and Sarcandra extend north to Japan,
China, Korea and the eastern U.S.S.R. (Ussuri).

Ascarina occurs in the Pacific and reaches New Guinea and the Philippines with a distinct sec-
tion Madascarina in NE. Madagascar; Chloranthus and Sarcandra are widely distributed in Male-
sia, India, Indochina and China. Hedyosmum occurs in the New World from Mexico to Brazil
and Peru and in the West Indies with one species occurring in the Old World in S. China, W. Su-
matra, Borneo and Celebes (fig. 8).

The family is now absent from Africa, W. Asia, Australia, and much of America.

HumpBert & Capuron (1955) when describing Ascarinopsis (= Ascarina sect. Madascarina)
speak of it as part of the most ancient floristic element in Madagascar, a survival from the Creta-
ceous flora and AuBREVILLE (1976) considers it as an Australo-Papuan element similar to Hibber-
tia, Dillenia, Evodia, Protium (& America), Macadamia, Elaeocarpus, Weinmannia (& America),
Bubbia, etc.

The complete absence of Chloranthaceae from tropical Africa at the present time is paralleled
in many other groups with trans-Pacific distributions. The discovery of the fossil pollen type Cla-
vatipollenites there could indicate that something very like Ascarina once occurred in Africa. The

(123)

124 FLORA MALESIANA [ser. I, vol. 102

family might appear to owe its present distribution to a Gondwanaland origin in the Early Creta-
ceous or even earlier, the absence from Australia and Africa being attributed to climatic vicissi-
tudes. Clavatipollenites (see later) is known from the Early Cretaceous of the U.S.A. (Maryland),
England, Israel, Patagonia, South & Central Africa, Brazil, Australia, efc.; and probable Oligo-
cene-Early Miocene deposits of South Africa (Cape Province) (COETZEE, 1981); and if all these
refer to Ascarina or some closely allied genus then a different course of distribution is indicated.
It seems the family may have been well distributed and common in the past but it is equally ap-
parent that migrations involving any kind of stringent climatic deterioration are not feasible.
Ascarina lucida for example was formerly (10,000—5,000 BP) abundant in New Zealand but is
now much reduced due to increase of frost and drought (MCGLONE & Moar, 1977).

References: AUBREVILLE, Adansonia II, 15 (1976) 302; Cortzer, S. Afr. J. Sc. 77 (1981) 341;
COETZEE & MUuLLER, Ann. Mo. Bot. Gard. 71 (1984) 1092, f. 11; HumMBERT & CAPuRON, C. R.
Ac. Sc. Paris 240 (1955) 28, fig.; MCGLONE & Moar, New Zeal. J. Bot. 15 (1977) 485—489;
VesTER, Bot. Arch. 41 (1940) 349, f. 144.

Fossils. As far as I am aware no undoubted fossils of parts of the plants other than pollen
are known, but leaves with chloranthoid characters have been found in the Lower Cretaceous Po-
tomac Group (UpcHURCH, 1984). Several Lower Cretaceous pollen types have been referred to
the family, but can hardly refer to the recent genera. Pollen of recent genera is also known from
various strata. These are dealt with elsewhere (see p. 126 and 143).

Reference: UpcHurcH, Amer. J. Bot. 71 (1984) 192—202.

Ecology. The species are all moist evergreen forest species, many ascending into submontane
forests. They occur from 0 to 3300 m in Malesia.

Dispersal. The white-fruited Chloranthus erectus (= officinalis) is dispersed by birds according
to RipLey (Disp. 1930: 410) and the red-fruited Sarcandra glabra must also be.

Pollination. VAN DER HAMMEN & GONZALEZ (1960) have shown that Hedyosmum is wind-
pollinated and has a high pollen production, but it has usually been assumed that the forest-
dwelling Chloranthus and Sarcandra are insect-pollinated but I have traced no recorded observa-
tions. Some collectors mention scent.

Reference: VAN DER HAMMEN & GONZALEZ, Leidse Geol. Meded. 25 (1960) 261—315.

Floral morphology. Payeras long ago as 1857 investigated the floral morphology of Ch/o-
ranthus spicatus and found that during the early stages of development the median lobe of the
anther-bearing organ appears first, soon followed by the two laterals which are distinct in origin
but immediately join up to form the 3-lobed organ and later still the ovary arises as a half-moon-
shaped outgrowth with the curved side towards the bract. ARMouR (1906) investigated the mor-
phology of the flower. The minute scale at the base of the anther-bearing organ on the ovary in
some species has been looked on as a perianth but it is not vascularised and probably simply an
outgrowth. The anther-bearing organ in ‘C. chinensis’ (probably C. erectus = officinalis) has
been described as bearing four anther-lobes each composed of two pollen-sacs, usually regarded
as corresponding to three stamens, of which the median one has two anther-lobes and the lateral
ones reduced to one and the traces are consistent with this, but development gives no evidence
of reduction of the lateral stamens, nor whether the flowers are to be regarded as reduced or the
reverse. In Sarcandra glabra the anther-bearing organ is usually described as a single stamen, the
position of the two anther-lobes resembling that of a normal Angiosperm stamen but the presence
of two traces suggests a derivation from an organ similar to that of Chloranthus by total reduction
of the median anther-lobes and reduction in width.

These curious structures have been looked on as separate male and female flowers joined and
simulating a bisexual flower, e.g. by HOOKER /., but most authors (e.g. ARMOUR) have considered
the flowers to be bisexual and Swamy & BAILEy’s (1950, 1953) studies of the vascularisation of
the structure support this. They could, however, be considered as very reduced inflorescences de-
rived from mixed cymules of male and female flowers such as occur in Ascarina, but there is no
evidence from vascularisation to support this.

1986] CHLORANTHACEAE (Verdcourt) 125

In Sarcandra the bract has a single trace; the carpel a double or single median strand and ventral
strands close or separated; staminal trace double and joining median traces or single and free to
below the bract or one free and one joined.

EnpreEss (1971) investigated the female flowers of Hedyosmum mexicanum and the following
is taken from his own summary. The flowers are free and not partly fused with each other nor
with the inflorescence axis. The perianth region extends not only to the free 3-lobed perianth-tube
and three double wings on the flower ridges, but also to the periphery of the whole flower below
the style. The 3-lobed perianth is initiated as the first floral organ, contrary to the occasional small
protrusions below the stamen attachment in Chloranthus. The fruit is a kind of drupe, the wings
of the flowers forming the outer subfleshy part and the periphery of the flower body the inner
hard part. The ovary wall tissue degenerates around the growing and ripening seed, the fruit wall
thus consisting mainly or completely of perianth tissue. The gynoecium lacks distinct signs of
pseudomonomery and seems to be truly monocarpellate. It is distinctly ascidiform at least up to
the style-base with an oblique or transverse ventral slit, the stylar canal and with ventral median
placentation appearing markedly laminar at anthesis. Except for the 3 bundles of the atropous
bitegmic and crassinucellate ovule there are no independent ovary bundles.

Leroy (1981, 1983a, 1983b) considers the male structures, formerly universally interpreted as
inflorescences of bractless flowers much reduced to single stamens, to be strobiloid male flowers,
each bearing several hundred spirally arranged stamens and closely resembling a gymnospermous
cone-like male flower. This reinterpretation considered in conjunction with monosulcate pollen
similar to pollen known from the Lower Cretaceous and adaptation to wind pollination suggests
that the male Hedyosmum flower is one of the most primitive Angiosperm flowers still existing.
Male Ascarina flowers with either 3—5 stamens or 1—3 stamens and Hedyosmum male flowers
with very numerous stamens are homologous and it is suggested easily derived from a common
ancestor.

References: ARMouR, New Phytol. 5 (1906) 49—55, pl. 3—4; ENpress, Bot. Jahrb. 91 (1971)
39—60; Leroy, XIII Int. Bot. Congr. Abstr. (1981) 136; Taxon 32 (1983) 169—175; C. R. Ac. Se.
Paris 296 (1983) 747—752; PAYER, Traité d’organogénie comparée de la fleur, Paris 1 (1857) 422;
Swamy, J. Arn. Arb. 34 (1953) 385—399; Swamy & Bamey, J. Arn. Arb. 31 (1950) 121-125.

Anatomy. This has been very thoroughly investigated by Swamy (1953) and SwaMy & BAILEY
(1950). An outstanding feature is the lack of vessels in the xylem in Sarcandra. The other genera
have vessels but they are relatively unspecialised, Ch/oranthus being the least advanced. In Sar-
candra tracheary elements in the secondary xylem are arranged in + undisturbed radial seriations
as seen in transverse section; tracheids in the region of the first year’s growth measure nearly 1.9
mm and have very extensive overlapping ends similar to other vesselless dicotyledons indicating
a cambium of very primitive type, and unusually long fusiform initials. The wood in Hedyosmum
is of a very unspecialised type; parenchyma paratracheal as incomplete sheaths around the vessels;
rays sometimes up to | mm wide, multiseriate rays composed of almost entirely upright cells;
fibres with simple pits and occasionally septate wall rather thin; in Ascarina the parenchyma is
apotracheal and the multiseriate rays of square to procumbent cells. Ascarina and Hedyosmum
have nodes typically of the unilacunar type with 2 vascular strands in the leaf of Ascarina and
5 in Hedyosmum, the lateral pairs larger; Ch/oranthus and Sarcandra have modified unilacunar
nodes with 5 vascular strands in the petiole, 2 much larger and extending most of the length of
the midrib, the intermediate small trace disappearing about half-way but formed by the fusion
of 2 minor branches of the larger traces at nodal level; the two small lateral strands come from
a different gap. In Hedyosmum the stipular sheath formed from the connate petiole bases consists
of collenchymatous tissue and supports the stem during intercalary growth. Lateral branches are
initiated in the leaf-axils but are attached to the parent axis above the node at maturity; cork
formed on the inner surface of the sheath brings it into intimate contact with the stem. There is
a pulvinus on the stem at the upper margin of the sheath. In 1. arborescens and related species
the nodal sheath develops by pushing beyond the apical growing point and surrounding it, tightly

126 FLORA MALESIANA [ser. I, vol. 102

closed above the bud and affording it protection. In Sarcandra and Chloranthus the stomata have
| —2 subsidiary cells oriented parallel to the guard cells whilst in Ascarina and Hedyosmum there
is a rosette of 4—6 ordinary epidermal cells.

BARANOVA (1983) reported that the laterocytic type of stomatal apparatus occurs in Chloran-
thus, Sarcandra and sometimes Hedyosmum along with other types. This type is known from a
very heterogeneous mixture of families.

MELVILLE (1962) stated the 2 leaf traces in Ascarina can unite at various levels in the petiole or
lamina and form a single vein, but in Chloranthus and Sarcandra each of the initial pair of traces
forks, resulting in bundles, the middle pair reuniting to give a final triple trace. In Hedyosmum
a trace of 5 bundles results from the bifurcation of the two outer bundles of such a triple trace.
He points out these types are also to be found in both the Pteridosperms and Cordaitales.

Mucilage canals are present in the petiole, larger veins and also in the margins of the pith in
Hedyosmum, in some species also containing sphaerocrystalline masses. Small clustered crystals
are recorded in the inner part of the cortex. Stone cells are scattered in the cortex of younger stems
of Sarcandra with larger groups in the pith; in older pith these cells form conspicuous transverse
diaphragms alternating with plates of parenchymatic cells, but these diaphragms are absent from
Chloranthus. Ethereal oil cells occur in the mesophyll of the leaf.

References: BARANOVA, Brittonia 35 (1983) 93-102; MELviLLE, Kew Bull. 16 (1962) 39—40;
Swamy, J. Arn. Arb. 34 (1953) 385—399; Swamy & Barley, J. Arn. Arb. 31 (1950) 121—125.

Palynology. WArKER (1976) summarised the palynology of this very eurypalynous family as
follows: “pollen grains anasulcate, inaperturate, with ‘colpoid complexes’ or colpoid streaks,
polycolpoidate or polycolpate, heteropolar, apolar or isopolar; boat-shaped-elliptic, globose-
oblate or globose, tectate or semi-tectate; more or less psilate, fossulate, scabrate, rugulate or re-
ticulate; monads; medium-sized to small”. He noted that polycolpate grains derived from mono-
sulcate ones occur only in this family and in Aristolochiaceae, stating that the polycolpate/poly-
porate pollen found in the two families must be considered unique in its clear monosulcate deriva-
tion. KUPRIANOVA (1981) has given much data on the pollen of several species. Grains of Chloran-
thaceae are remarkably similar to those of the oldest known fossil Angiosperm pollen: Clavatipol-
lenites COUPER at first known only from the Lower Cretaceous of England, Israel (N. Negev),
Maryland (U.S.A.), and Patagonia. KUPRIANOVA (1967) was the first to claim that Clavatipolle-
nites and Ascarina are congeneric; COUPER (1960) had earlier noted the strong similarity, and it
is clear that there is at least close relationship. A very strong case has been made for the conspeci-
ficity of Clavatipollenites evittii (California, Maestrichtian) and Ascarina lucida. MULLER (1981)
summarised the fossil pollen records of the Clavatipollenites-Ascarina complex. In New Zealand
there are records from the Maestrichtian to the present day, bridging the gap between Cretaceous
and recent. A virtually continuous record from the Albian to the Eocene has recently been dis-
covered in Australia where elimination from the continent may have been due to increasing aridi-
ty. A record has also been published from the 90° ridge in the Indian Ocean from Oligocene de-
posits. In Europe there are records from the Aptian, Albian and Cenomanian with probably ex-
tension to the Barremian. LAING (in Ferguson & Muller, 1970) gave a Barremian record from
England. Apart from the eastern U.S.A. record there have been finds from California and Chile
(Maestrichtian), Bahamas (Cenomanian), Falkland Plateau (Palaeocene), Central Africa (Al-
bian, Aptian and possibly Barremian), South Africa (probably Oligocene to early Miocene), and
Brazil (and once again possibly eliminated from the latter three by climatic deteriorations). DoYLE
(1977) commented on the various types of Clavatipollenites pollen and compared the finely
clavate-retipilate forms from the Lower Cretaceous with Ascarina, the coarser clavate irregularly
aperturate ones with Hedyosmum and the reticulate nearly inaperturate type with Sarcandra.
Considered in conjunction with the distribution of the recent species, a purely Gondwanaland dis-
tribution does not cover all the main facts of fossil distribution and either some of the latter are
misidentified or a Pangaea type theory now largely discarded by most workers must be resorted
to. The family must have been widespread and common in the Lower Cretaceous, only a few spe-
cies lingering on to the present day.

1986] CHLORANTHACEAE (Verdcourt) 7

References: CoETZEE, S. Afr. J. Sc. 77 (1981) 341; Coetzee & MULLER, Ann. Mo. Bot. Gard.
71 (1984) 1092, f. 11; Couper, New Zeal. Geol. Surv. Palaeont. Bull. 32 (1960) 1—87; Doy_e,
Bull. Cent. Rech. Explor. Prod. Elf-Aquitaine (1977) 451—473; KuprRiaANova, Pollen et Spores
9 (1967) 95-100; Bot. Zhurn. USSR 26 (1981) 3—15; LatNc in Ferguson & Muller, Linn. Soc.
Symp. Ser. 1 (1976) 15—25; MULLER, Bot. Rev. 47 (1981) 9-12; WALKER in Beck, Origin & Early
Evolution of Angiosperms (1976) 269.

Chromosomes. EHRENDORFER et a/. (1968) compiled lists of numbers for primitive families
and gave Hedyosmum arborescens n = 8; Sarcandra glabra 2n = 30; Ascarina rubricaulis 2n =
28; Chloranthus serratus n = 14 &2n = 30; C. spicatus 2n = 30; C. japonicus 2n = 30; C. fortu-
nei 2n = 60. He stated that from these data it can be seen that there are still true diploids in the
family: x = 8, further secondary polyploid base numbers x2 = 14 and 15 and continuing infragen-
eric polyploidy (4 x,). EHRENDORFER stated (in Beck, 1976) that “Chloranthaceae demonstrate
progressive elimination of diploids (n = 8), major representation on the 4 x level (n = 14 and
15, the latter from 7 + 8?) and occasional origin of 8 x (n = 30)”, corresponding in part with
the Piperales.

References: EHRENDORFER et a/. Taxon 17 (1968) 337—468; in Beck, Origin & Early Evolution
of Angiosperms (1976) 228—229.

Phytochemistry. Evento the collector the aromatic smell suggests relationship with the Pi-
peraceae. In 1964 HEGNAUER complained that the chemistry of this family had scarcely been
touched. BATE-SMITH (1962) found in the hydrolysed leaf-extract of Ch/loranthus erectus (= offi-
cinalis) none of the otherwise very widespread phenols save for B-coumaric acid and in this respect
it resembles many Piperaceae. The ethereal oils which undoubtedly exist have so far not been
chemically investigated.

Kupsitzk1& REZNIK (1966) in their investigation of flavonoids as systematic characters investi-
gated 4 species of Chloranthus, 7 of Hedyosmum and 5 of Ascarina and found them to contain
traces to massive amounts of the two derivatives of flavanole, quercetin and kampferol, the for-
mer dominating. SHio & Hicucui (1981) have demonstrated the presence of a gymnosperm type
of lignin, guaiacyl lignin in Sarcandra glabra and Chloranthus spicatus.

References: BATE-SmitH, J. Linn. Soc. Bot. 58 (1962) 95—173; HEGNAUER, Chemotaxonomie
der Pflanzen 3 (1964) 428; Kupirzk1& REZNIK, Beitr. Biol. Pfl. 42 (1966) 445—470; SHio & HiGu-
CHI, Wood Res. Kyoto Univ. 67 (1981) 43—46 [from For. Prod. Abstr. 5 (1982) no 2121].

Taxonomy. Early placings of Ch/loranthus were various and mostly wide off the mark. LIND-
LEY, who first recognised the family (1821) placed them in Piperales and his clear view has been
supported by many authors (BENTHAM & Hooker, 1880, 1883; ENGLER, 1894; ARMouR, 1906;
Swamy, 1953; Howarp, 1970; BEHNKE, 1975; BURGER, 1977) with arguments from various dis-
ciplines (morphology, anatomy, phytochemistry, pollen). After digesting a wholesale literature
on the subject (1985), I believe this comes at least close to the correct affinity. The most probable
conclusion from the floral structure appears to be that Ch/oranthaceae is a group of Piperales pre-
senting in some points, especially in the structure of the ovary, primitive characters in common
with the majority of the Archichlamydeae, while in other aspects modifications of the flowers are
shown. A.C. Smiru (1972) found them more primitive than other elements of the Laurales and
repeated (1981) that Chloranthaceae is best assigned to its own order, Chloranthales, validated
by Leroy (1983). HuTCHINSON (1973) and Cronguist (1981) retain the family in Piperales, but
this then remains a matter of choice.

It is certainly clear that the small, insignificant family Chloranthaceae is one of great impor-
tance in the study of primitive flowering plants whose ancestry points to high age, probably even
to the Early Cretaceous, judging from the affinity of the pollen to Clavatipollenites. Unusual ge-
neric ranges also point towards early time, viz. one species of otherwise neotropical Hedyosmum
in Indo-Malesia and one species of Ascarina in Madagascar even representing a separate section.
_ References: Armour, New Phytol. 5 (1906) 49-55, pl. 3 & 4; Bennke, Ann. Mo. Bot. Gard.
62 (1975) 647—663; BenTHAM & Hooker /., Gen. PI. 3 (1880) 133—135, (1883) 1220-1221; Bur-

128 FLORA MALESIANA [ser. I, vol. 102

GER, Bot. Rev. 43 (1977) 345—393; CRONQUIST, Integr. Syst. Class. (1981) 83—85; ENGLER, Nat.
Pfl. Fam. 3, 1 (1894) 12—14; Howarp, Bot. J. Linn. Soc. 63, Suppl. 1 (1970) 195—214; Hutcuin-
SON, Fam. FI. Pl. ed. 3 (1973) 513—519; LeRoy, Taxon 32 (1983) 169—175; LinDLEy, J. Collect.
Bot. (1821) sub t. 17; A.C. Smiru, J. Ind. Bot. Soc. Jubilee Vol. SOA (1972) 215—226; Fl. Vit.
Nova 2 (1981) 97; Swamy, J. Arn. Arb. 34 (1953) 375—408; VERDCOURT, Kew Bull. 40 (1985)
218—224.

Generic delimitation. This is very decisive. The genera with fully unisexual flowers or
dioecious inflorescences with marked protrandry, Ascarina (including Ascarinopsis) and Hedyos-
mum are clearly distinct from Chloranthus and Sarcandra in which the male floral parts are adna-
te to the ovary. In Ascarina both male and female flowers can be bracteate, but the male flower
has only 1—5 anthers, whereas in Hedyosmum the male flowers have numerous anthers arranged
in a cone with basal involucre and the female flowers are enclosed in highly developed cupular
bracts. Morphologically Chloranthus and Sarcandra are more closely inter-related than the other
two. Sarcandra is well characterised by its single stamen with anther cells on a club-shaped organ
and particularly by its vessel-less xylem; Ch/oranthus can occasionally have a single stamen but
then it is of a different structure, e.g. as in C. multistachys PEI. HUTCHINSON and others main-
tained Tricercandra A. GRAy distinct from Chloranthus on account of the fused male floral parts
being produced into long narrow lobes (‘connectives’) but there are intermediates between this
and the simple 3-lobed state found in typical Chloranthus which has been even more emphasised
by new discoveries in China; it is not even possible to maintain it as a section. Ascarinopsis can
on the other hand be kept as a section of Ascarina there being some distinctive characters such
as 2—5 stamens and the curious disposition of two leafless nodes between the leafy ones.

Uses. The Chinese grow Chloranthus spicatus in pots for its fragrant aromatic leaves and it
was formerly grown ona large scale in Java in one montane locality and used for imparting a scent
to native tea and is apparently still used on a small scale for this purpose. BURKILL (1935) reported
that leaves and flowers are left in contact with the tea whilst it is drying; they are later discarded
or left to add bulk to the tea. Its use has been claimed to be deleterious. Among medicinal uses
are as a poultice on carbuncles and boils, as a diaphoretic and excitant, as a cure for malaria but
said to be poisonous in overdose, for back-ache and as a tea-like drink for treating coughs. Ch/o-
ranthus erectus (= officinalis) is also used medicinally and as a tea, both the leaves and the roots
being used. HEYNE (1927) gives extensive medicinal uses for the latter. It has a sudorific effect
and is used in cases of fever and as a restorative during some phases of venereal disease; also as
a stimulant and mixed with Cinnamomum bark as an antispasmodic during parturition.

DosEDLA reported (GILLI, 1979) that formerly the leaves of Chloranthus erectus (= officinalis)
were eaten together with pork as an offering in House Tambaran in the Hagen area of New
Guinea. Areas where the plant grew were avoided; today the whole plant which stains the hands
green on touching is taboo. VELDKAmpP (in sched.) reported that in S. Borneo the branches are
boiled and the slightly peppery tasting concoction drunk by women to prevent conception.

Sarcandra glabra has also been used to scent tea or add bulk and used in Indonesia for the same
purposes as Chloranthus erectus. In the Philippines an infusion is said to be useful in treating
headaches, asthma and ‘internal pains’. It is also used in Chinese medicine for bone fractures,
contusions and in the form of a leaf-decoction as an astringent treatment for vomiting. FRAKE
reported (PNH 37989 & 38204) that on Mindanao the leaves are pounded and applied to ulcers
and Cuat(S 35411) that the warm leaves are used for sprains. HEYNE also mentions that when
tea-planting started in Java the cultivation of Chloranthus was forbidden by the Dutch.

References: BURKILL, Dict. Econ. Prod. Mal. Pen. 1 (1935) 535; Gitui, Ann. Naturh. Mus.
Wien 84 (1979) 426; Heyneg, Nutt. Pl. Ned. Ind. (1927) 533—534; Perry & METzGER, Medic. Pl.
E. & SE. Asia (1980) 78—79.

Note. The drawings on the four plates were made by Mrs. MAUREEN CHURCH, at Kew.

1986] CHLORANTHACEAE (Verdcourt) 129

KEY TO THE GENERA

1. Flowers essentially bisexual, the male part adnate to the ovary.

2. Anther-loculi 4 (in Mal.) on a 3-lobed organ interpreted as 3 fused stamens with central 2-locular
anther and a I-locular anther on each side. Leaves of widespread species usually finely serrate. Fruit white,
greenish or yellowish, rarely tinged violet or pinkish, the tip sometimes dark violet-purple

1. Chloranthus

2. Anther-loculi 2 on a narrowly clavate organ, i.e. apparently one stamen with a 2-locular anther. Leaves

Pmuimeveny COAUSelyiserrate>.Pruit:red., rarely. black sc sins -tci3-. ee - o e 2. Sarcandra
1. Flowers truly unisexual or apparently so (in Mal.).
3. Male flowers bracteate with 1—2 (—5) stamens; female flowers naked. Shrubs or trees 2.5—24 m tall

3. Ascarina
3. Male flowers with involucre and numerous stamens in a cone-like structure; female flowers in rounded
ovate bracts with long caudate apices. Herbs or subshrubs.....................02- 4. Hedyosmum

1. CHLORANTHUS

SWARTZ, Phil. Trans. R. Soc. 77 (1787) 359; Sotms-Laus. in DC. Prod. 16, 1
(1869) 473; BENTH. in B. & H. Gen. PI. 3 (1880) 134; ARMour, New Phytol. 5
(1906) 49, pl. 3 & 4, p.p.; Swamy, J. Arn. Arb. 34 (1953) 375; MAEKAWA, J.
Jap. Bot. 45 (1970) 289, f. 1—5; FANG Wu-Kuo, Acta Phytotax. Sin. 18 (1980)
220. — Nigrina THUNB. Nov. Gen. (1783) 58; Fl. Jap. (1784) 65; Nova Acta Soc.
Sc. Upsal. 7 (1815) 142, t. 5. — Stropha Noronna, Verh. Bat. Gen. K. W. ed.
15 (1791) Art. IV.4 & ed. 2 (1827) 66, nomen; cf. STEEN. in Essays in Biohistory
efc., Regn. Veget. 71 (1970) 376. — Cryphaea Bucu.-Ham. Edinb. J. Sc. 2
(1825) 11, non Monr in Weber (1814), nec Bripet (1822). — Fig. 1-3.

Shrubs or perennial erect or straggling herbs, glabrous and aromatic; stems
jointed at the nodes. Leaves decussate, or sometimes subverticillate in whorls
of 4, serrate; petiole bases connected by a transverse line or shortly connate;
stipules mostly small. Spikes terminal, slender, sometimes branched, sometimes
leafy at the base, the flowers each in the axil of persistent mostly subopposite
bracts along the spicate opposite side branches. Flowers essentially bisexual, the
male part usually a 3-lobed organ adnate to the ovary and sometimes enveloping
it forming a fleshy mass, the lobes (‘connectives’) short and broad or longer or
distinctly narrow and elongate, in some species almost or quite free, or in one
species not noticeably lobed and reduced to + nothing but 1—3 anthers; anthers
| —3, the introrse locelli variously arranged, either 3 ‘anthers’, one median with
2 approximate, or + separated locelli and one on each lateral edge with single
locelli, or 2 lateral 1-locellate anthers and usually no median one (but this can
vary in a population), or 1—3 anthers joined and the lobed organ practically or
quite obsolete with one 2-locellate anther and usually two lateral 1-locellate an-
thers. Ovary naked; stigma subsessile, truncate. Drupes usually white, fleshy.
Seeds subglobose, minutely apiculate, narrowed below, invested by the thin
fibrous endocarp, the seed-coat with a lignified endotestal palisade.

Distr. About 20 spp. extending from Japan, China and extreme East U.S.S.R. to India, Ceylon and Male-
sia as far as New Guinea.

130 FLORA MALESIANA [ser. I, vol. 102

Fig. 1. Chloranthus erectus (BUCH.-HAM.) VERDCOURT. a. Flowering branchlet, x 2/3, b. portion of young

nflorescence, x6, c. young flower, x 16, d. portion of older inflorescence, x6, e. mature flower, x 16.7:

anthers, x 20, g. ovary, X30, A. LS ovary, x 30, i. fruiting branchlet, x 2/3, j. fruits, x 4 (a—c ANDERSON
S 20818, d—h vAN ROYEN 3626, i—j CHAI S 35523).

1986] CHLORANTHACEAE (Verdcourt) 131

Taxon. Various authors have advanced subdivisions but there is no unanimity of opinion and this will
have to wait for a monograph of the entire genus.

Palyn. Grains subspherical averaging about 22.5 by 30 um, polycolpate with 3—8 colpi but often much
modified reaching the poles in a few species but often short and appearing as slender unsculptured wavy meri-
dional streaks and in some species only distinguishable as very faint tenuous short zigzag lines; exine with fine-
ly reticulate-pitted sculpture.

Floral morph. (see also general section). The staminal organ of Chloranthus has commonly been inter-
preted as 3 connate stamens and the available evidence suggests that it actually developed in this way from
the concrescence of 3 stamens rather than the expansion of a single stamen and acquisition of supernumerary
locelli (Swamy, 1953). Where the central lobe of the male flower usually bears no anther cells, e.g. in C. japo-
nicus, there is actually a good deal of variation showing an almost continuous range from rudiments to almost
perfect anther cells in the same spike (MAEKAWA, 1970).

KEY TO THE SPECIES

1. Perennial unbranched herbs or subshrubby herbs with + subverticillate leaves in fours congested at top of
stem. Anther-bearing organ with 3 distinct narrow lobes ............... 0.0000 ee cece: 3. C. henryi

1. Branched subshrubs or shrubs with decussate leaves. Anther-bearing organ shortly 3-lobed or 3-crenate.
2. Leaves usually long acuminate, 8—29 by 3—13 cm, usually finely shallowly glandular-serrate. Fruits and
SME MNPMMNRN I SPS 20 iy 1s CN Cra) pa HS ee GOT es Fo inh = RE ENG, ORS EOE CELT a ERE 1. C. erectus

2. Leaves usually + obtuse, 5—12.5 by 2—4 cm, crenate-serrate. Fruits and stamens green or yellowish (culti-

MIE B.S) 3, 2 gOS go alo ate a fae eal Sh al Shs BORE

1. Chloranthus erectus (BUCH.-HAM.) VERDCOURT,
Kew Bull. 40 (1985) 217. — C. erectus SwEET, Hort.
Suburb. Lond. (1818) 28, nomen; WALL. Cat. (1832)
230, n. 6881, nomen. — Cryphaea erecta BucH.-
Ham. Edinb. J. Sc. 2 (1825) 11, t. 2. — C. officinalis
Bi. En. Pl. Jav. 1 (1827) 79; Fl. Jav., Chloranthaceae
(1829) 10, t. 1; CorpEMoy, Adansonia 3 (1863) 298;
Soims-Lavs. in DC. Prod. 16 (1869) 474; Hook. /.
Fl. Br. India 5 (1886) 100; Forspes & HEMSLEY, J.
Linn. Soc. Bot. 26 (1891) 369; K. Scu. & Laut. FI.
Deut. Schutzgeb. Siidsee (1900) 263; Nachtr. (1905)
241; Lecomte, Fl. Gén. I.-C. 5 (1910) 94; PULLE, No-
va Guinea 8 (1911) 623; GamB_E, J. As. Soc. Beng.
75 (1912) 33; Koorp. Exk. Fl. Java 2 (1912) 40;
Grspss, J. Linn. Soc. Bot. 42 (1914) 129; Merr. En.
Philip. 2 (1923) 21; Rip. J. Mal. Br. R. As. Soc. 87
(1923) 89; Burk.& Hortr. Gard. Bull. S. S. 3 (1923)
69; Riwt. Fl. Mal. Pen. 3 (1924) 52; Burk. & HEND.
Gard. Bull. S. S. 3 (1925) 411; RENpDLE, J. Bot.,
Suppl. 63 (1925) 85; BartLett, Pap. Mich. Ac. Sc.
Arts Lett. 6 (1926) 48; RipL. Kew Bull. (1926) 78;
Henp. Gard. Bull. S. S. 4 (1928) 308; Mam in Fed-
de, Rep. 34 (1934) 270; Burk. Dict. (1935) 535; Pe,
Sinensia 6 (1935) 667, f. 1; METCALF, Fl. Fukien 1
(1942) 39; HeNnpb. Mal. Nat. J. 6 (1951) 443, f. 400;
Hara, Fl. East. Himal. 1 (1966) 44; Mever, Bot.
Bull. Herb. For. Dept. Sandakan 10 (1968) 32. — ?
C. elatior R. Br. ex Sims, Bot. Mag. (1820) 48 sub t.
2190, nomen; Link, En. PI. Berol. 1 (1821) 140; Cor-
pemoy, Adansonia 3 (1863) 299; Back. & Baku. /.
Fl. Java 1 (1963) 175; Kena, Orders & Fam. Mal.
Seed Pl. (1969) 122, f. 73; ed. 2 (1978) f. 74; Hara,
Fl. East. Himal. 2 (1971) 14; C.Y. Wu, Fl. Yunna-

2. C. spicatus

nica | (1977) 18, f. 5/1—3; Qin & SHain Fang, FI. Si-
chuanica | (1981) 138, f. 58; Wu Kuo-FAna, FI. Rei-
publ. Pop. Sin. 20 (1982) 84, t. 26/6-10. — C.
inconspicuus (non SWARTZ) BLANCO, FI. Filip. ed. 2
(1845) 54. — C. salicifolius PREs1, Epim. Bot. (1851)
231. — Fig. 1-3.

Shrubby herb, subshrub or small shrub, 0.3—3 m,
rarely epiphytic, + aromatic when crushed; nodes
distinctly swollen, sometimes purplish. Leaves bright
green, glossy above in life, oblong-lanceolate to ellip-
tic- or ovate-oblong, 8—29 by 3—13 cm, usually long-
acuminate at the apex, cuneate at the base, rather
thin, the margin rather finely shallowly (rarely more
coarsely) glandular-serrate, with c. 9 pairs of nerves,
sometimes purple beneath; petiole 0.75—1.7 cm; stip-
ules minute, subulate. /nflorescences scented, with
5—13 spikes 2.5—5 cm long; bracts sheathing; ovate,
acute. Anther-bearing organ yellow, between green
and white or violet-white, 3-lobed, 1.2—1.6 mm long;
median anther 2-locellate; lateral anthers 1-locellate.
Ovary enclosed by the male part. Fruit white, cream
or rarely tinged violet or pinkish, the tip sometimes
dark violet-purple, succulent, glossy, 5-7 mm o.
Seed yellowish white.

Distr. Continental SE. Asia from Nepal to Yun-
nan and Andaman Is., and throughout Malesia to
New Guinea (incl. New Britain and New Ireland).

Ecol. Primary and secondary forest, including
Pandanus and palm, Araucaria, and Nothofagus-
Castanopsis in montane forest, often on limestone,
sometimes riverain or in boggy areas, mainly low-
land, (20—)50—1450(—2535) m. Fl. Jan.—Dec. Rip-
LeY (Disp., 1930) stated that it is bird dispersed.

FLORA MALESIANA

[ser. I, vol. 102

Fig. 2. Chloranthus erectus (BUCH.-HaM.) VERDCOURT in fruit near Njarumkop, NW. Kalimantan (A. ELsE-
NER 30, 5-8-1964).

Taxon. The epithet e/atior has come into wide
usage but there is no BROWN specimen at the BM or
any LINK specimen now extant at B, so it is not pos-
sible to be certain what was actually intended, since
the description given by Link is fragmentary, his
type being sterile. | agree with VAN STEENIS (in Jitt.)
that the name is better discarded in favour of one
which is unambiguous and of which the type is avail-
able.

BUCHANAN-HAMILTON’s faulty drawing and de-
scription have misled authors as to the identity of
Cryphaea erecta, but the examination of the type (E)
left no doubt.

Vern. Sumatra: garaman gadjah, harostulang,
lelada-rima, Alas, sébah, si-eném-berugi, Ss.-e.-
béngi, s.-e.-bérnih, Karo, si-pukhor-lisak; Java: ka-
ras tulang, kerastulang, kras tulang, manik, muni,
pulu dengen, sukattam, ujah, ujahan; Borneo:
bétuk, Sarawak, Kelabit, dikut-dikut, langut-langut,
luai, Kinabatangan, /alamak, Murut, /Jambayau, La-
buk, /ud, Bangyi, mongkitir, Mub; Philippines: bab-
bainm di dup dup hug, pangutug, If., banwa-
limatok, Palawan, barau-barau, Luzon-Laguna
Prov., bélekbut, sunulampong, vimug, Sub., buro-

bonlao, Negros, tul-an hinbad, Samar, tunggdo
Tagb.; New Guinea: kaha, Bulolo-Garaina, kuggé-
bdlts, Hagen, kundiewdia, Madang, Saidor, moluh-
muluh, silibe, Dani, namakebuya, Kainantu-Okapa,
rumukembalo, Kagua, sohkumo, Orokaiva-
Mumuni, tjalangan, Enga, Poio; New Britain: an-
gore.

Note. Judging by dried material specimens from
New Ireland have larger fruits but the constancy of
this needs checking in the field. Sterile material of
Chloranthus erectus (= officinalis) with more
toothed leaves than usual can be difficult to distin-
guish, but Sarcandra glabra usually dries with more
reddish brown coloured foliage.

2. Chloranthus spicatus (THUNB.) MAKINO, Bot.
Mag. Tokyo 16 (1902) 180; Merr. Trans. Am. Phil.
Soc. n.s. 24 (1935) 127; Pet, Sinensia 6 (1935) 668, f.
2; Bot. Bull. Acad. Sinica 1 (1947) 114; STEwarp,
Man. Vasc. Pl. Lower Yangtze Valley (1958) 70, f.
43; MANSFIELD, Die Kulturpfl. Beih. 2 (1959) 13;
Back. & Baku. f. Fl. Java 1 (1963) 176; Ouwt, FI.
Japan (1965) 361; C.Y. Wu, Fl. Yunnanica 1 (1977)
18, f. 5/4—6; Qin & Sua in Fang, FI. Sichuanica 1

CHLORANTHACEAE (Verdcourt) 133

Fig. 3. Chloranthus erectus (BUCH.-HAM.) VERD-
court. Njarumkop, NW. Kalimantan (A. ELSENER
30, 15-8-1964).

(1981) 139; Wu Kuo-Fana, FI. Reipubl. Pop. Sin. 20
(1982) 83, t. 26/1—5. — C. inconspicuus SWARTZ,
Phil. Trans. R. Soc. 77 (1787) 359, t. 15; L’HERIT.
Sert. Angl. (1788) 35, t. 2; BENTH. Fl. Hongk. (1861)
334; Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 129;
Soims-LauB. in DC. Prod. 16 1869) 474; ForBEs &
Hems ey, J. Linn. Soc. Bot. 26 (1891) 368; ENGLER
in E. & P. Nat. Pfl. Fam. 3, 1 (1894) 13, f. 12; LeE-
comtE, Fl. Gén. I.-C. 5 (1910) 95; Koorp. Exk. FI.
Java 2 (1912) 42, f. 3; Britren, J. Bot. 55 (1917) 344;
Burk. Dict. (1935) 535. — Nigrina spicata THUNB.
Nov. Gen. (1783) 58; Fl. Jap. (1784) 65. — Nigrina
spicifera LAMK, Ill. Gen. 1 (1792) 295 & (1791) t. 71.
— C. indicus WiGut, Ic. PI. Ind. Or. 6 (1853) t. 1945.
— C. obtusifolius Mig. Fl. Ind. Bat. 1, 1 (1856) 802.

Small glabrous shrub, 0.5—1.5 m, with ascending
or + spreading branches. Leaves oblong-ovate or
ovate-elliptic to elliptic, 4—-13.5 by 2—8.5 cm, rather
obtuse or at least not or scarcely acuminate at the
apex, cuneate at the base, rather coarsely crenate-
serrate, nerves 4—6 pairs; petiole 0.4—1.2 cm; stip-
ules membranous, linear, 2—3 mm, mucronate. /n-
florescences terminal with 10-20 ascending spikes

2—5 cm long; peduncles 3—8 cm; bracts 1.5 mm;

>

bracteoles | mm. Anthers 3, the central with 2 locelli
and the laterals 1-locellate, the cells + 0.5 mm long.
Fruit green or yellowish, 4 by 2 mm, narrowed at the
base.

Distr. China but widely cultivated elsewhere in
eastern Asia; in Malesia cultivated in Java (e.g. Para-
kansalak) and Sumatra (West Coast); 700—920 m.
Fl. Jan.—Dec.

3. Chloranthus henryi HeEmsLey, J. Linn. Soc. Bot.
26 (1891) 367; HAND.-Mazz. Symb. Sin. 7 (1929)
156; Pel, Sinensia 6 (1935) 679, f. 6; Bot. Bull. Acad.
Sinica 1 (1947) 115; Wu Kuo-Fana, FI. Reipubl.
Pop. Sin. 20 (1982) 92, t. 28/9—13. — C. philippi-
nensis MerRR. Philip. J. Sc. 7 (1912) Bot. 259; En.
Philip. 2 (1923) 22. — C. verticillatus MERR. Philip.
J. Sc. 10 (1915) Bot. 3; En. Philip. 2 (1923) 22. —C.
oldhamii (non Soims-Laus.) MERR. & Quis. Philip.
J. Sc. 82 (1953) 324.

Herb 20—40 cm; stem simple, glabrous or pubes-
cent towards the apex, with 3
ing paired scales 6—9 mm long, which are caducous
or perhaps not developed in some specimens; root-
stock said to be aromatic. Leaves in whorls of (?3—)4
at apex of stem, elliptic to broadly elliptic or obo-
vate-elliptic, 8—15.5 by 3.5—9.5 cm, narrowly long-
acuminate at apex, cuneate at base, margin sharply
serrate, serrations terminated by prominent thick
glands, glabrous, or shortly pubescent on the 5—9
pairs of nerves beneath; petiole 2—7 mm, glabrous or
pubescent. /nflorescence terminal; peduncle slender,
7.5—13 cm long with 2—3(—4) spikes 2—6 cm long;
secondary peduncles 0.5—1.2 cm, sparsely pubes-
cent; bracts ovate, + 1.5 mm long. Male part at-
tached about the middle of the ovary, white; central
lobe 2.2—3 by 1 mm (4 by 1.2 in original description
of C. verticillatus), lateral lobes 1.5—1.8 by 0.6—0.8
mm, all rounded at apex; anther cells about half as
long as lobes, 2 on central lobe, one on each lateral
lobe, 0.8—1 mm long. Ovary + ovoid, narrowed
from near middle to a + triangular apex, 1.5(—2?)
by | mm. Fruits white?

Distr. China; in Malesia: Philippines (Luzon:
Ifugao, Mt Polis; Bayninan, Banaue, Cagayan, Abu-
lug R.).

Ecol. Forest; banks of
250—1200 m. Fi. Jan.—Feb.

Vern. hayad, If.

Note. When describing C. verticillatus MERRILI
suggested it might be nearest to C. henryi, but later
MERRILL and QuISUMBING synonymised it with C.
oldhamii Soims-Laus. MERRILL also compared C,
philippinensis with C. henryi but stated the former
had much smaller flowers. Chloranthus henryi is var-
iable particularly in regard to the ratio of the length
of the anther cells to that of the lobes of the male part
which vary from about a 1/2 to more usually 1/4 to

5 leafless nodes bear-

irrigation canals;

134

FLORA MALESIANA

[ser. I, vol. 102

1/3, but some Chinese material is very similar to that
from the Philippines and for the present I am content
to consider them conspecific. The above description

better range of material from China and the Philip-
pines may show constant differences and that sub-
species can be recognised.

does, however, refer only to Philippine material. A

2. SARCANDRA

GARDNER, Calc. J. Nat. Hist. 6 (1846) 348; Swamy & BamLey, J. Arn. Arb. 31
(1950) 127; VERDCOURT, Kew Bull. 39 (1984) 66. — Fig. 4.

Shrubs or shrublets with nodose jointed branchlets. Primary and secondary
xylem without vessels. Leaves decussate or sometimes appearing subverticillate
where nodes are congested, pinnately nerved, usually coarsely serrate, serra-
tions thickened at apex; petiolar sheath short, with short setaceous stipules on
its margin. Flowers essentially bisexual, in lax terminal spikes with c. 3 main
branches, each with up to c. 12 flowers; bracts boat-shaped. Male part reduced
to a club-shaped or + discoid organ usually interpreted as a filament and con-
nective of a single stamen bearing 2 locelli, adnate about 2/3 of the way up the
abaxial side of the ovary and with a cushion-like fold below the point of attach-
ment; locelli latrorse or introrse, opening lengthwise, usually separated but
sometimes touching at their tips. Ovary ovoid, 1-locular; ovule pendulous, bi-
tegminal, crassinucellate; stigma sessile depressed and subcapitate. Drupe near-
ly invariably red, 1-seeded, obovoid, bearing the scar of the fallen male part on
its anterior face; pericarp succulent, the epidermis without stomata. Seed pen-
dulous with membranous testa; innermost cell layer of outer integument devel-
oping into a lignified palisade of radially elongate cells with 1 —several crystals,
the cell cavity becoming filled with an internal reticulum of lignin.

Distr. Two species in Ceylon, India, Burma, Indochina to China and Japan, one of which in Malesia:
Malay Peninsula, Sumatra, Java, Borneo, Lesser Sunda Is., Philippines, New Guinea. A map (now out of
date) was published by SUEssENGUTH & GALL, Mitt. Bot. Staatssamml. Munchen | (1953) 351.

Taxon. There is no doubt that Swamyis correct in maintaining this genus, well-characterised by the vessel-
less xylem, characteristically shaped male part of the flower with 2 locelli and with 1 or 2 traces rather than
3, presence of stone cells, dorsal carpel traces, characteristic pollen, nearly invariably red fruits etc. Confusion
has occurred in the nomenclature of the species occurring in S. India and Ceylon (VERDCOURT, 1984).

Palyn. Grains spherical about 28 um diameter, acolpate, the exine coarsely reticulate.

Morph. It is logical to treat the flowers as bisexual flowers since the two staminal traces usually join with
the median carpel traces although flower is really a vague term for combinations of structures of different
Origins; variants exist, however, which have a single staminal trace free at least to below the bract.

1. Sarcandra glabra (THUNB.) NAKAI, FI. Sylv. Ko-
reana 18 (1930) 17, t. 2; Swamy & BatLey, J. Arn.

(1794) 331. — Chloranthus monander R.Br. ex SIMs,
Bot. Mag. (1821) t. 2190. — Ardisia glabra (THUNB.)

Arb. 31 (1950) 117; Onw1, Fl. Japan (1965) 361;
Mever, Bot. Bull. Herb. For. Dept. Sandakan 10
(1968) 33; Hortkawa, Atlas Fl. Japan 1 (1972) map
28; Lru TANG-Suut, Fl. Taiwan 2 (1976) 566, t. 410;
C.Y. Wu, Fl. Yunnanica 1 (1977) 21, f. 6/5—7; Qin
& SHAw in Fang, FI. Sichuanica 1 (1981) 135, f. 57;
Wu Kuo-Fana, FI. Reipubl. Pop. Sin. 20 (1982) 79,
f. 25/1—5; VERDcouRT, Kew Bull. 40 (1985) 216. —
Bladhia glabra THuns. Trans. Linn. Soc. Lond. 2

DC. Trans. Linn. Soc. Lond. 17 (1834) 123. — Chlo-
ranthus brachystachys sensu BENTH. Fl. Hongk.
(1861) 334, p.p.; Sotms-Laus. in DC. Prod. 16
(1869) 475; ForBpes& HEMSLEY, J. Linn. Soc. Bot. 26
(1891) 367, p.p.; REHDER, J. Arn. Arb. 10 (1929)
111, p.p., non Bu. s.str. — Chloranthus denticulatus
Corpemoy, Adansonia 3 (1863) 296. — Chloranthus
glaber (THuUNB.) Makino, Bot. Mag. Tokyo 26
(1912) 386. — Fig. 4.

1986] CHLORANTHACEAE (Verdcourt) 135

Fig. 4. Sarcandra glabra (T Hun.) NAKAI. a. Flowering branchlet, x 2/3, b. part of inflorescence, x 6, c. flo-
wer, back view, x 10, d. flower, front view, x 10, e. stigma, = 16, f. LS ovary, x 16, g. fruiting branchlet,
* 2/3, h. fruits, x2 (a—f PurseGctove 4097, g KALKMAN 5319, A Nooresoom & Cua 1825, spirit material).

136

FLORA MALESIANA

[ser. I, vol. 102

For further references and synonymy see under
ssp. brachystachys.

Glabrous shrub or half-woody herb, 0.6—3 m tall;
stems up to 1.5 cm @ with + swollen nodes, longitu-
dinally ridged when dry; bark + smooth. Leaves
elliptic-oblanceolate, lanceolate or narrowly
elongate-elliptic, 2-20 by 1—8 cm, long-acuminate,
cuneate at the base, + subcoriaceous, coarsely or
shallowly very sharply serrate or dentate-serrate, the
teeth with thickened points; lateral nerves in 5—10
pairs; stipules small, linear-subulate, c. 1.5 mm long;
petiole 0.4—1.7 cm. Jnflorescences greenish or +
white, 3—8 cm long, the spikes rather dense, 1—3.5
(—5) cm long; paired bracts 3 mm long; bracteoles
oblong, 1 mm long, sometimes + trifid. Male part
1.3—2 mm long, 1—1.3 mm wide, the anther cells ex-
tending from half to the whole length or almost so,
c. 1.3 mm long. Female part flask-shaped or subglo-
bose, 1—1.5 mm long; stigma up to 1 mm wide. Fruit
at first yellow, becoming red or bright orange, very
rarely black, 4-7 mm a (dry), shining. Seed pale, yel-
lowish or cream.

Distr. Continental SE. & E. Asia, throughout
Malesia.

Note. The typical ssp. glabra occurs in N. & Cen-
tral China, Korea, Japan and the Ryukyu Islands.
Some specimens from NW. India and S. China are
somewhat intermediate.

KEY TO THE SUBSPECIES

1. Male structure with anther cells about as long as
the structure itself, the non-antheriferous part
pga yh oosecoocat ssp. brachystachys

1. Male structure with anther cells much shorter, the
non-antheriferous part (/.e. apparent ‘filament’)
well-developed. E. Asia........... ssp. glabra

ssp. brachystachys (BL.) VERDCOURT, Kew Bull. 40
(1985) 216. — Ascarina serrata Bu. En. Pl. Jav. 1
(1827) 80. — Chloranthus brachystachys Bu. FI.
Jav., Chloranthaceae (1829) 13, t. 2; BENTH. FI.
Hongk. (1861) 334; Sotms-Laus. in DC. Prod. 16
(1869) 479; Forses & HEMSLEY, J. Linn. Soc. Bot. 26
(1891) 367; Srapr, Trans. Linn. Soc. Lond. II, 4
(1894) 217; C.B. Ros. Philip. J. Sc. 4 (1909) Bot. 70;
Lecomte, Fl. Gén. I.-C. 5 (1910) 95; Koorp. Exk.
Fl. Java 2 (1912) 41, f. 4; Gress, J. Linn. Soc. Bot.
42 (1914) 129; Ript. J. Mal. Br. R. As. Soc. 8 (1923)
89; Merr. En. Philip. 2 (1923) 21; Burk. & HoLtr.
Gard. Bull. S. S. 3 (1923) 70; Rmpi. Fl. Mal. Pen. 3
(1924) 53, f. 140; RENDLE, J. Bot. Suppl. 63 (1925)
85; BARTLETT, Pap. Mich. Ac. Sc. Arts Lett. 6 (1926)
48; REHDER, J. Arn. Arb. 10 (1929) 111, p.p.; MALM
in Fedde, Rep. 34 (1934) 270; Burk. Dict. (1935) 535.

— Chloranthus hainanensis Pei, Sinensia 6 (1935)
674. — S. hainanensis (PEI) Swamy & BAILey, J.
Arn. Arb. 31 (1956) 128, f. 4/15—17; C.Y. Wu, FI.
Yunnanica | (1977) 21, f. 6/1—4; Wu Kuo-FAna, FI.
Reipubl. Pop. Sin. 20 (1982) 80, f. 25/6-11. —
Chloranthus glaber sensu Back. & BAKH./f. Fl. Java
1 (1963) 175, non (THUNB.) MAKINO S.str. — S. gla-
bra auctt. mult.; MEUVER, Bot. Bull. Herb. For. Dept.
Sandakan 10 (1968) 33; KENG, Orders & Fam. Mal.
Seed Pl. (1969) 123, f. 74 & ed. 2 (1978) f. 75.

Male structure with anther cells almost equalling
it, .e. the non-anthiferous part much reduced.

var. brachystachys

Proits red. ~

Distr. NE. India (Assam, Manipur, Naga Hills),
Bangladesh, Burma, Thailand, N. Vietnam, S. Chi-
na (incl. Hainan), and throughout Malesia.

Ecol. Evergreen, both lowland and lower mon-
tane forest, secondary forest, heath forest, moist
ground ridge forest, sometimes by water, also eroded
limestone slopes with thin ‘mos’ soil, ridge podso-
lized soil, shingle banks and stream beds, shaly
slopes; 135—2550 m. F/. Feb., April—June; fr. Jan.—
May, Aug.—Nov.

Vern. Sumatra: kaju aek, k. duri-duri, k. si détu
aek, k. si marsangkat, si détu rapping, si marduri-
duri, si rapping bulung, Batak; Java: atukan, péngan
dukiki; Lesser Sunda Is.: /awi-nata, nio-kodé (=
monkey’s coconut), Flores; Philippines: baga-baga,
puti-selimbangun, Sub., bagen-hoho, holog di na-
muyug, If., apot, damoko, emem, gapas, umuum,
Ig., baga-baga, puti-selimbangun, Sub., gumok,
Buk., kari-kari, Bag., total, Yak., em-em, N. Luzon;
New Guinea: brok, brok-mugup, Sepik, Telefomin,
kaguna, tsui ‘alagebo, S. Highlands, Kutubu.

Notes. Field notes and spirit material show that
the fruit is much larger in life, e.g. 9-15 by 7—10
mm.

Sterile material of Chloranthus erectus with much
more coarsely toothed leaves than usual can be diffi-
cult to distinguish, but Sarcandra glabra usually
dries with more reddish brown coloured foliage rath-
er than a grey-green colour.

var. melanocarpa (RIDL.) VERDCOURT, Kew Bull. 40
(1985) 217. — Chloranthus brachystachys BL. var.
melanocarpus Riw1. J. Mal. Br. R. As. Soc. 87 (1923)
89.

Differs in having black fruits.

Distr. Malesia: N. Sumatra (Berastagi, West
Hill; Lake Toba, Gunung Batu, Lopang, 10 km ESE
of Prapat).

Ecol. Montane rain-forest, 1400—1500 m.

Note. I had dismissed RmDLEy’s variety as a casu-
al variant, but its recollection in the same general

1986] CHLORANTHACEAE (Verdcourt) 137

area suggests a population of black-fruited speci- red throughout its range the occurrence of such a va-
mens exists; since the fruit is uniformly described as riety seems worth emphasising.

3. ASCARINA

J.R. & G. Forst. Char. Gen. (1775) 59; ed. 2 (1776) 117; CorpEMoy, Adansonia
3 (1863) 302; Sotms-LauB. in DC. Prod. 16 (1869) 477; BENTH. in B. & H. Gen.
Pl. 3 (1880) 134; Swamy, Proc. Nat. Inst. Sc. India 19 (1953) 371; v. BALGooy,
Blumea, Suppl. 5: Pacif. Pl. Areas 2 (1966) map 64; A.C. Smirn, J. Arn. Arb.
57 (1976) 405; Moore, J. New Zeal. Bot. 15 (1977) 491; JEREmiE, Adansonia II,
20 (1980) 273; A.C. SmitH, FI. Vit. Nova 2 (1981) 98; JEREmiE, Fl. Nouv. Caléd.
et Dep. 11 (1982) 171; RAw.inGs, J. New Zeal. Bot. 12 (1974) 564. — Ascari-
nopsis HUMBERT & CAPURON, C. R. Ac. Sc. Paris 240 (1955) 28, fig.; AUBREV.
Adansonia II, 15 (1976) 302. — Fig. 5.

Shrubs or small or sometimes quite large trees, usually apparently dioecious
or monoecious, glabrous, aromatic, the branches jointed at the nodes and the
bases of the internodes sometimes swollen. Leaves decussate, often coriaceous,
obtusely serrate, the serrations often gland-tipped; petiolar sheath very short;
stipules very small, subulate; in sect. Madascarina JEREMIE and some species of
sect. Ascarina there are two intermediary aphyllous nodes with eventually deci-
duous sheaths 2—6 mm long between successive pairs of leaves the stipular collar
made up of the petiole-bases bearing 2 pairs of small teeth under 5 mm long.
Flowers basically in much condensed biparous sessile cymes borne in the axil of
a bract with an abaxial male flower and | or 2 adaxial female flowers with bracts
and bracteoles (if 2-flowered), essentially representing a bisexual flower; some-
times 1—2 stamens are associated with 1—2 rudimentary or functional adaxial
carpels which develop much later; but mostly inflorescences reduced to single
male flowers or 1 —2 female flowers and appearing monoecious or dioecious. —
Male flowers bracteate, reduced to 1—2 subsessile anthers with parallel linear
locelli opening lengthwise or in sect. Madascarina with 2—5 (usually 3) sessile
stamens with bilocular anthers. — Female flowers consisting of a naked sessile
ovoid-globose ovary without outer bracts or with 2 often caducous inner bracts
and if flowers paired then with bracteoles also; stigma sessile, truncate or
2-lipped or in sect. Madascarina horseshoe-shaped. Fruits purplish grey turning
black, obovoid, with thin succulent exocarp and stony smooth or verrucose un-
specialised endocarp; epidermis without stomata. Seeds ovoid, flattened, with
smooth testa, the seed coat with unlignified endotestal palisade but with ligni-
fied fibrous exotegmen.

Distr. 12 species in Madagascar, Pacific Islands (widespread from Solomons to the Marquesas), New
Caledonia and New Zealand; in Malesia 4 spp.: Borneo, Celebes, Philippines, New Guinea. Fig. 6.

Ecol. Mostly in montane or submontane rain-forest, 450-3300 m; in both lowland and montane rain-
forest in New Zealand and at lower and medium altitudes elsewhere in the Pacific.

Palyn. Grains spherical, about 30 4m diameter, monocolpate, with fine faint pitted-reticulate sculpture.

138 FLORA MALESIANA [ser. I, vol. 102

Fig. 5. Ascarina philippinensis C.B. Ros. a. Male flowering branchlet, x 2/3, b. stipules, x 4, c. portion of

male inflorescence, x 8, d. base of anthers showing bract, x 16, e. apiculate apex of anther, x 14, f. TS an-

ther, x 16, g. female flowering branchlet, x 2/3, h. female inflorescence, x4, i. portion of older female in-

florescence, x6, j. female fruiting branchlet, x 2/3, k. LS ovary, x16, /. fruits, x4, m. seed, x6, n. leaf,

x 2/3 (a—f VAN ROYEN & SLEUMER 6075, g—h Ramos & EDANO BS 30647, i VAN ROYEN & SLEUMER 8127, j—m
VAN ROYEN & SLEUMER 5898, 1 VAN BaALGooy?).

1986] CHLORANTHACEAE (Verdcourt) 139

Taxon. The species of sect. Ascarina fall into two groups, a predominantly western Pacific group with
a single bract subtending the flower and male flowers_with two stamens and the other mostly eastern Pacific
group with 3 bracts and | stamen. -

Floral biology. Ascarina species have usually been assumed to be dioecious but A.C. SmiTH (1976)
pointed out that he had seen specimens which were monoecious with male and female inflorescences on the
same branchlet. RAWLINGs (1974) published an observation made in 1972 by J. Down that a solitary tree of
A. lucida produced copious pollen and later was loaded with drupes thus disproving that the genus is dioe-
cious. It was noted that flowers were produced in the early spring, developing very slowly during August and
September and that the drupes did not mature until the following spring and did not fall until after new flowers
had formed. Even earlier in 1971 J. GopLey studied a plant in flower and suspected it might be a hermaphro-
dite with very strong protandry. Moore (1977) showed that A. /ucida exhibits a peculiar type of monoecism
where a male flower is represented by a single stamen and commonly accompanied by two female flowers
which mature distinctly later or in other compound inflorescences a stamen and single accompanying ovary
mature almost simultaneously. In essence there is a reduced biparous sessile cyme which resembles a bisexual
flower and has in fact been accepted as a single flower. CoRDEMoy (1863) long ago suggested that the ‘single
flower’ of CAloranthus is in reality a little glomerulus, a biparous sessile cyme. JEREMIE (1980) also comments
on this subject and states that generally male and female flowers are on different plants but exceptionally cer-
tain specimens present flowers of 1 or 2 stamens and one or two rudimentary but sometimes functional carpels
in an adaxial position /.e. falsely hermaphrodite with + marked protandry but true monoecism also occurs.
SmiTH (1981) has accepted that this condensed biparous cyme with male and female flowers is the basic inflor-
escence in the genus. A survey of the herbarium material of A. philippinensis has not revealed anything but
dioecious plants.

KEY TO THE SPECIES

1. Leaves rounded or only very shortly acuminate at the apex. Flowers with a single subtending bract; male
flowers with 2 stamens. Endocarp smooth.

2. Leaves subsessile, the petiole short and thick, about 2 mm long; blades narrowed to rounded or subcordate
Rapes Rat S ret Z, aris oS Se Tae bodes WOR Ges 2 ie 5 d,s Saw oe See ee ae ee 3. A. subsessilis

2. Leaves distinctly petiolate, the petioles slender, 0.4—1.5 cm long; blades cuneate at base.

3. Fruits 1.2—2 mm long, with thinner more finely wrinkled exocarp (dry material), in more lax terminal
and axillary inflorescences, the distances between the fruits greater and inflorescence-branches more
MMAR Te ess 2 ets WOR ATS... DROS PEs Se oe 2. A. maheshwarii

3. Fruits 2—3 mm long, with thicker more coarsely wrinkled exocarp, in lax to usually very densely congest-
ed terminal inflorescences, the distances between the fruits shorter and inflorescence-branches usually
SCRROIES SOS 31521515 Fs Dok Sah He ae POL 1. A. philippinensis

1. Leaves distinctly long-acuminate at the apex. Flowers with 3 subtending bracts; male flowers with | stamen.
paint watty-papillate:.’. 06.06.8020 ee... TUR eR Rae 4. A. diffusa

1. Ascarina philippinensis C.B. Ros. Philip. J. Sc. 4
(1909) Bot. 70; Merr. En. Philip. 2 (1923) 22; Swa-
my, Proc. Nat. Inst. Sc. India 19 (1953) 375, f. 2;
Meuer, Bot. Bull. Herb. For. Dept. Sandakan 10
(1968) 32, fig.; A.C. Smirn, J. Arn. Arb. 57 (1976)
409; v. Roven, Alpine Fl. New Guinea 3 (1979) 1256.
— A. reticulata Merr. Philip. J. Sc. 12 (1917) Bot.
263; En. Philip. 2 (1923) 22; Herne in Fedde, Rep. 54
(1951) 226.

Small to medium-sized tree, 7.5—24 m, @ to c. 35
(—60) cm, sometimes a shrub or treelet 2—4 m at high
altitudes; twigs very brittle; bark very variable,
white, grey or dark reddish brown, very rough and
flaky to slightly to distinctly fissured, the fissures
widely spaced, deep, bordered with yellowish scar
tissue; inner bark straw to orange or pale brown, fi-
brous; blaze off-white to pale brown; cambium yel-

low; wood pinkish straw, turning + orange, the rays
well marked, with no exudate or sapwood described
as yellow or straw-coloured and heartwood purplish
or dull brownish. Leaves very variable, elliptic, ob-
long, ovate or obovate, 3.2—14(—18) by 1.5—8.5
(—10) cm, rounded or obtuse to very shortly acumi-
nate at apex, cuneate at base, rather fleshy or leathe-
ry, glossy, + paler beneath, crenate with thickened
tips to crenations; nerves 8—11 pairs; petiole 0.4—1.5
cm; stipules linear to ovate-boat-shaped, 1—2.5 mm
long, often caducous; intermediate leafless nodes of-
ten present, with sheaths up to 6.5 mm long. /n/flores-
cences terminal compound spikes with several
branches usually spreading from a central axis,
1.5—3 cm long, the final branches spiciform with
4—8 flowers; bracts ovate, up to 1.5—3 by 1—2 mm.
— Male flowers whitish or pale greenish yellow with

140

FLORA MALESIANA

[ser. I, vol. 102

Fig. 6. The disjunct distribution of the genus Ascarina Forst. The single species in Madagascar (not indicated
here) is the sole representative of the sect. Madascarina LEROY & JEREMIE (Ascarinopsis HUMB. & CAPURON).
Localities of sect. Ascarina after Pacific Plant Areas 2, map 64.

2 collateral stamens and rarely a third adaxial one;
anthers c. 3 by 1—1.5 mm, the projecting connective
subacuminate. — Female flowers and fruits congest-
ed, the inflorescence-branches usually 0.5—2 cm
long, the flowers usually not separated by more than
2 mm; ovary green; stigma brownish. Fruits with spi-
cy aroma, green turning purple-black. Endocarps
straw-coloured, sublenticular, 2.5 by 2 by 1.3 mm,
keeled.

Distr. Malesia: N. & Central Borneo, Celebes,
Philippines (Luzon, Panay, Mindanao), New Guinea
(incl. New Britain & Manus I.).

Ecol. In New Guinea usually Nothofagus domi-
nated rain-forest, understorey in montane forest,
450—2850(—3300, on Mt Wilhelm) m. Male fl. June,
July, Sept.; ripe fr. June—Aug.

Vern. New Guinea: gwa, Wagu, jamarapara,
Kapaukoe, kengepa, Hagen, Wankl, kogile gol, Si-
nasina lang., Nimai dialect, kKokun, pohn, tipitan-
gan, Togoba, kondomapi, S. Highlands, Mt Ne,
maskinen, Mendi, moyok, Washuk, obilak, Enga,
penge mange, S. Highlands, SE. Mt Giluwe.

2. Ascarina maheshwarii Swamy, Proc. Nat. Inst.
Sc. India 19 (1953) 377, f. 3; CoRNER, Phil. Trans. R.
Soc. Lond. B 255 (1969) 576; A.C. Situ, J. Arn.
Arb. 57 (1976) 411.

Small to medium-sized, dioecious tree 3—15 m,
with broad trunk 15—55 cm g; rarely with thick but-
tresses to 1 m; bark pale grey to dark brown, smooth
or fissured; blaze dull brown; wood soft, white or
straw-coloured. Leaves elliptic to somewhat obo-
vate, 6—14 by 3—8 cm, shortly acuminate at the apex,
narrowly cuneate at the base, crenate or crenate-
dentate, the tips of the crenations with brown to
black thickenings (hydathodes), dark green and shi-
ny; nerves 12—20 pairs; petiole 0.4—2 cm; stipules tri-
angular-subulate, 0.5—1.5 mm long from a narrow
sheath; leafless nodes with short to cylindrical
sheaths 3—10 mm long bearing leaf vestiges about 1
mm long. Jnflorescences green, terminal but often
also borne in axils of upper leaves (up to 4 nodes be-
low apex); branches 5—8, slender, (1—)2.5—3.5 cm,
each unbranched or again branched, the flowers well
spaced with internodes 2—8 mm; lower bracts of in-
florescence similar to sheaths at leafless nodes, the
upper longer, 0.5—1.5 mm long; bracteoles support-
ing the flowers deltoid, 0.5 mm long or almost obso-
lete. — Male flowers with 2 yellowish anthers 2—3
mm long. — Female flowers with cream ovary and
unequally bilobed sessile stigma. Fruits drying
yellow-brown but said to be black (WHITMORE),
translucent green or greenish cream, ovoid, 1.2—2

1986]

mm long, exocarp thin and finely wrinkled (dry
state); endorcarp + _ discoid, straw-coloured,
smooth.

Distr. Solomon Is.; in Malesia: Papua New Gui-
nea (Milne Bay, Morima Range), incl. Manus & Bou-
gainville Is.

Ecol. Primary rain-forest, also open forest and
hillside forest, 450—750 m. Male fl. July, fr. June,
Oct.

Note. Although SmirH (1976) speaks of this ‘very
distinct species’ characterised particularly by having
additional axillary inflorescences, slender long in-
florescence branches with spaced flowers and smaller
fruits with less wrinkled exocarp, examination of ex-
tensive material of A. philippinensis shows that the
situation is not so simple. Ascarina maheshwarii is
very uniform in the Solomon Is., even on different is-
lands, but all the characters can be found in various
specimens of A. philippinensis which is very variable
over its wide range; small fruits are found for instan-
ce In Rau 158 (Onim) and Vinas & W1AKaBu LAE
59447 but both have typical philippinensis foliage;
long infloresence branches occur in some material
from the Cycloop Mts which have, however, larger
more wrinkled fruits and foliage of philippinensis.

3. Ascarina subsessilis VERDCOURT, Kew Bull. 40
(1985) 213, f. 1.

Medium-sized, dioecious tree 15—24 m, 8—35
(—50) cm 9; bark pale grey-brown to dark brown, fis-
sured; wood straw-coloured or light brown. Leaves
elliptic or oblong-elliptic, rarely narrowly obovate,
5—14 by 3—9 cm, rounded at the apex, narrowed,
rounded or subcordate at the base, shallowly cren-
ate, rather fleshy in life, drying thick and with mi-
nutely rugulose surface, the costa sometimes dividing
the leaf somewhat asymmetrically, subsessile; nerves
10—15 pairs; petiole thick, c. 2 mm; stipules minute,
c. 1 mm long; leafless nodes present with sheaths 2—6
mm long bearing lateral leaf-vestiges 0.8—1.5 mm
long, rugulose in dry state. Inflorescences terminal,
branched and occasionally a simple unbranched one
from a lower node. Male 2 cm long, 3-branched at
the base, the main branch with side branchlets c. 5
mm long; anthers c. 3 mm long. — Female 1.5—4 cm
long with 3—5 main branches each with 5—7 rather
thick branchlets 1.5—2.5(—3) cm; lower bracts simi-
lar to the leaf-vestiges at leafless nodes, the upper
again similar but with the lateral projections longer,
lanceolate, longitudinally folded, 4.5 mm long; brac-
teoles supporting flowers deltoid, scarcely 1 mm
long; female flowers not seen. Fruits greenish white
(presumably becoming dark when ripe?), ovoid, +
congested; exocarp strongly wrinkled in the dry
state; endocarp c. 1 mm long (mature?), smooth.
Distr. Malesia: Papua New Guinea (Morobe,
Southern, Western & Eastern Highlands). Cf. Verp-

CHLORANTHACEAE (Verdcourt)

141

court, l.c., map 1.
Ecol. Montane rain-forest, partly secondary for-
est on limestone ridge, 1830—3200 m. Male fl. Aug.,

fr., Nov., Jan., June—Sept.

Vern. New Guinea: porn, Hagen, tomil, Sinasina
lang., Nimai dialect, wabaliok, wabilak, Enga, wa-
baljok, Enga-Kepilam, masnene Mendi.

4. Ascarina diffusa A.C. Smirn, J. Arn. Arb. 57
(1976) 415, f. 5—10; Fl. Vit. Nova 2 (1981) 99, f. 35a
& b. — A. lanceolata (non Hook.f.) auct.; SWAMY,
Proc. Nat. Inst. Sc. India 19 (1953) 377, p.p.

Shrub to small or medium-sized tree, dioecious or
monoecious, 4—15(—25) m tall with open spreading
crown; trunk 7—10 cm @; young stems reddish; bark
light grey to brown, smooth; blaze red and wood
white, soft. Leaves elliptic-lanceolate to lanceolate,
4—17 by 1.5—4.5 cm, distinctly acuminate at the
apex, cuneate at the base, minutely to coarsely ser-
rate, save near the base, thinly coriaceous; nerves
12—25 pairs; petioles slender, 0.5—2.7 cm; stipules
1—3 mm long on leaf sheaths 1—5 mm long; leafless
nodes occur. /nflorescences green, yellowish or pur-
ple, many-flowered, lax, terminal and in the upper
axils; main axis 3-branched, the central branch again
2—3-branched but some axillary inflorescences sim-
ple; individual branches slender, 3—12 cm long, the
flowers closely placed, each subtended by 3 brac-
teoles scarcely 1 mm long; lower main bracts of inflo-
rescence up toc. 1 cm long consisting of a sheath with
lateral projections; upper bracts ovate-acuminate, 3
mm long./nflorescence glomerules consisting basi-
cally of an abaxial male flower and adaxial female
flower but more usually unisexual. — Male flowers
with | stamen; anthers 2—4 mm long, scarcely | mm
wide (reported purple in Fiji specimen), with a mi-
nute deltoid apical projection. — Female flowers:
ovary green, ovoid, 1.5 mm long; stigma between
translucent and white or purple, 0.8—1.3 mm wide.
Fruit ellipsoid, 2—3 by 1.2—2.5 mm; exocarp thin
and slightly wrinkled in dry state; endocarp rounded
ellipsoid, + 1.5 by 1.2 mm, compressed, the margin
keeled, surfaces warty-papillate.

Distr. Solomon Is., New Hebrides, Fiji, Samoa,
Cook Is.; in Malesia; Papua New Guinea (New Brit-
ain: Mt Talawe; Bougainville).

Ecol. Rain-forest, cloud-forest with Pandanus,
900—1800 m. Male fl. Feb., Aug., fr. May.

Vern. New Britain: pot, Talasea.

Note. There is considerable difference between
the foliage and inflorescence length of the single New
Britain specimen seen and those from Bougainville (3
specimens), but bearing in mind the variation
throughout its range this seems of no significance.
Fropin observed flower buds (young fruits?) red-
violet whitish at tip and inflorescence purple but else-
where the inflorescence is described as yellowish,

142 FLORA MALESIANA [ser. I, vol. 102

Fig. 7. Hedyosmum orientale MERR. & CHUN.a. Female flowering plant, x 2/3, b. female inflorescence, x 4,

c. female flower and bract, x 6, d. female flower, stigma removed, x 8, e. perianth spread out, x 10, f. ovary,

x 10, g. ovary opened to show ovule, x 16, A. TS ovary, x 10, i. stigma, x 10, /. portion of stigma lobe to

show multicellular hairs, x 24, k. part of male plant, nat. size, /. young male flower, x 2, m. portion of ex-

panded male flower, x 8, n. front view of anther, x 12, 0. side view of anther, x 12, p. TS of anther, x 16
(a—j S 16539, kK—p Burtt 12775, spirit material).

1986] CHLORANTHACEAE (Verdcourt) 143

4. HEDYOSMUM

Swartz, Prod. Veg. Ind. Occ. (1788) 84; CorpEMoy, Adansonia 3 (1863) 302;
Soims-Lavus. in DC. Prod. 16 (1869) 479; BENTH. in B. & H. Gen. PI. 3 (1880)
135; ENGLERin E. & P. Nat. Pfl. Fam. 3, 1 (1894) 13; Swamy, J. Arn. Arb. 34
(1953) 402; ENpREss, Bot. Jahrb. 91 (1971) 39; LeRoy, Abstr. XIII Int. Bot.
Congress (1981) 136; Taxon 32 (1983) 169; C. R. Ac. Sc. Paris 296 (1983) 747.
— Fig. 7.

Herbs, shrubs or trees, monoecious or dioecious; branches often jointed at
the nodes, often exuding a gelatinous aromatic exudate when cut. Leaves decus-
sate, mostly serrate, the serrations often tipped with glands (? hydathodes); pet-
iolar sheath mostly with marginal subulate or pectinate stipules. /nflorescences
axillary or terminal, sometimes united with the stem near their base; flowers tru-
ly unisexual; individual inflorescences unisexual but the compound inflorescen-
ces often with female flowers above and male flowers on lower branches. In
some species the flowers are partly fused amongst themselves and with the axis.
— Male flowers cone-like, solitary or paniculate, with involucres and very nu-
merous bilocellate anthers, filaments absent or very short; locelli parallel open-
ing lengthwise, at first 2-celled; connective shortly appendaged or subpeltate
above the loculi. — Female flowers variously capitate or paniculate, distinctly
bracteate; perianth-tube adnate to the ovary, limb very short, 3-toothed; style
very short, stigmatose at apex or stigma sessile, rarely linear or clavate, often
caducous; outer and inner integuments of ovary 3 cells thick. Drupes free, or
united by the bracts into a dense mass, globose or ovoid, often 3-sided or
3-ribbed, sometimes crowned with persistent perianth lobes, the fleshy wall
formed partly by the accrescent perianth; exocarp juicy; endocarp hard; seed
coat unspecialised.

Distr. About 35 spp., almost entirely in the New World from Mexico to Peru and Brazil and West Indies,
a single species in SE. Asia, extending into W. Malesia.

Palyn. ErpTMAN (1952) described the pollen of H. brasiliense as 1-sulcate or 1-sulcoidate (?), sometimes
.provided with a number of thin branched and irregularly placed apertural areas, which in optical section can
convey the impression of a polycolpoidate grain. According to Swamy (1953) the grains tend to be larger than
in other genera of the family up to 35 4m and are generally polycolpate resembling Ch/oranthus but the fur-
rows are often localised at a pole or other locus and the ends joined or reduction in number of the furrows
resulting in acolpate grains occurs in some species; the exine is finely reticulate-pitted. MULLER (1981) reported
that pollen grains of this genus have been found in Upper Miocene deposits in Mexico, Guyana and Colombia
and suspects DoyLe may have been correct in comparing irregularly aperturate forms from the Lower Creta-
ceous with this genus.

References: ExpDTMAn, Pollen morphology and plant taxonomy (1952) 111—112; MULLER, Bot. Rev. 47
(1981) 9—12; Swamy, J. Arn. Arb. 34 (1953) 392—393.

Floral morph. Leroy(1981, 1983) has suggested that what has usually been considered to be an inflores-
cence of unistaminate, naked, ebracteate, male flowers is in fact itself a strobiloid male flower bearing several
hundred spirally arranged stamens, an extremely primitive structure which correlates with the ‘Lower Creta-
ceous’ type monosulcate pollen and adaptation to wind pollination. I have accepted this interpretation as be-
ing the more plausible.

Dr P. Rupatt of the Jodrell Laboratory (Kew) has cut sections of the axis of material collected by B.L.
Burtt in Sarawak. It contains 8 bundles and only single bundles enter the stamens, which is not at variance

144 FLORA MALESIANA [ser. I, vol. 102

with the theory. ENDREss’ comments (1971) on the female flower have already been mentioned in the general
section (see p. 125).

BURGER (1977) considered that both the female flower (the most complex in the family) and the male (the
most reduced) were derived from simple flowers resembling those of Chloranthus and Sarcandra, an idea of
course totally at variance with that of LeRoy. The 3-lobed covering of the female flowers of Hedyosmum truly
suggests reduction and fusion from an ancestor with a 3-parted perianth. In some species this tissue becomes
succulent and white in fruit, an aril-like adaptation for dispersal by birds. It may have arisen by modification
of 3 adnate staminodes and the dorsal and lateral positions of the ribs and alternation of the vascular bundles
of the ribs with those of the ovary supports this idea.

References: BURGER, Bot. Rev. 43 (1977) 361; ENpREss, Bot. Jahrb. 9 (1971) 39—60; LeRoy, XIII Int. Bot.

Congr. Abstracts (1981) 136; Taxon 32 (1983) 169-175; C. R. Ac. Sc. Paris 296 (1983) 747—752.

1. Hedyosmum orientale MeRR. & CHUN, Sunyatse-
nia 5 (1940) 36, t. 5; Wu Kuo-FAna, FI. Reipubl.
Pop. Sin. 20 (1982) 95, f. 30. — H. nutans (non Sw.)
Merr. Lingn. Sc. J. 5 (1927) 29. — Hedyosmum sp.
STEEN. Bull. Jard. Bot. Btzg II, 13 (1933) 182.

Dioecious glabrous herb or subshrub, 1—2.5 m
tall; stems long, often straggling at the base, brittle
and juicy above, up to 2 cm g, smelling like ginger-
root; branches often drooping; upper internodes of-
ten condensed, particularly in young plants so that
petiolar sheaths overlap. Leaves lanceolate to linear-
oblong, 9—22 by 1.5—4 cm, thin, long acuminate at
the apex, narrowly cuneate at the base, the margins
rather closely serrate, the serrations apically thick-
ened, more or less obtuse; lateral nerves 20—30 pairs,
fine, + raised; petioles 0.6—2(—3.5) cm, longer on
one side, each pair joined to form a sheath 6—15 by
6—8 mm. — Male flowers few in panicles, the floral
axes 2—4.3 cm, the stalks c. 1 cm; very young flowers
cone-like, about 1 cm long with a basal oblique ring
6 mm wide of c. 13 acute or bifid bractlets; stamens
about 300, anther locule 1.2 mm (2 mm in vivo) long,
the connective appendage compressed, acute, 0.5—1
mm long, asymmetrically incurved. — Female inflor-
escences green, paniculate, c. 5 by 2 cm, the few
branches 1—1.5 cm, + few-flowered; bracts rounded
ovate, c. S—6 by 5 mm with a long caudate apex 3—6
mm long, c. 1 mm wide; sepals 3, triangular, 1—1.2
mm long and wide, entire, crenate or + dentate.
Ovary somewhat 3-angled, genuinely monocarpel-
late; stigma greenish with red tips, essentially lanceo-
late in outline, 2—2.2 mm long, irregularly lobulate,
covered with several-celled hairs; tip of ovary with
triangular impressed area with mammillate centre.
Fruit ellipsoid, 3—4 mm long, + 3-angled, crowned
by the calyx.

Distr. S. China (Hainan; Kwangsi-Tonkin bor-
der; S. Vietnam: Kontum); in Malesia: Sumatra
Westcoast Residency (Lubuk Sulasi; Laras Talang),

N. Bencoolen (Bt Daun), Borneo (West: Bt Tibang;
Sarawak: Hose Mts, base of Bt Kajang, Bt Sarpan-
dai; Bt Kenawang: Usun Apau), and Central Celebes
(Palu Distr.: near Lake Lindu, G. Njilalaki, and west
slope of Mt Poroka, Timbu). Fig. 8.

Fig. 8. Localities of Hedyosmum orientale MERR. &,
CHUN.

Ecol. Forest on dacite hillside, in ridge forest and
foot of wet cliff, with conifers and Fagaceae, 1000—
2365 m. Fi. March—April, fr. March—July.

Note. In the BM there is a HorsFIELD specimen
of the genus, labelled in sched. H. sumatranum,
which is the earliest collected specimen from the Old
World.

SPHENOSTEMONACEAE (C.G.G.J. van Steenis, Leyden)

The taxonomic position and rank of the only genus Sphenostemon has a chequered history. In
the course of time it has, under various names, been attributed to the Aquifoliaceae (by BAILLON,
as Sphenostemon, 1875), to the Icacinaceae (as a species of Phlebocalymna, by F. VoN MUELLER,
1875), to the Guttiferae (as Nouhuysia, by LAUTERBACH, 1912), and to the Trimeniaceae (by
Grpss, as Idenburgia, 1917).

Battey & Swamy (1953) and BarLey (1956) examined the anatomy and concluded that the genus
could not belong to either Guttiferae or Trimeniaceae cq. Monimiaceae, but they gave no clear
alternative. When I summarised the complete generic synonymy (1955), I found it likely to retain
Sphenostemon in Aquifoliaceae.

An other opinion approached that of F. voN MUELLER, viz. that by INGLE & DADSwWELL (1961)
who suggested, on the strength of the wood anatomy, a likeness with Platea in the Icacinaceae,
and possibly also an affinity to Polyosma (Saxifragaceae).

HUTCHINSON (1959) and Airy SHAW (1972) stuck to the Monimiaceous affinity, and I must ad-
mit that there is a distinct resemblance, in androecium in particular, with Trimenia, but this is
overruled by the anatomical and other differences. They felt possibly also strengthened by the fact
that LOESENER (1942) had expelled Sphenostemon from the Aquifoliaceae, and had suggested af-
finity with Theaceae or Ochnaceae, or as representative of a separate family.

In a good overview BERNARDI(1964) concluded that Sphenostemon should remain in A quifolia-
ceae. In this he is followed by CRonquistT (1981).

A very thorough anatomical research of Aquifoliaceae led Baas (1975) to the conclusion that
Sphenostemon is anatomically allied to both A quifoliaceae and Icacinaceae, probably more to the
latter. In fact, in comparing the macromorphological characters it appears that all of them occur
in Icacinaceae. He proposed that the genus should be accommodated in a family of its own, an
idea already advanced by Arry SHAw (1972), allied to both Aquifoliaceae and Icacinaceae; this
view is also held by THORNE (1983). The removal of Sphenostemon from Aquifoliaceae is well
sustained by the seed structure. The fruit was mostly defined as a drupe containing a pyrene. Balr-
LEY (1956) showed that the sclerified tissue is, however, not derived from the endocarp and that
the fleshy envelope of it is really the whole of the pericarp. He accepted the sclerified tissue as
derived from the testa. I expressed my doubt about this interpretation to Dr. W.A. VAN HEEL
(Leyden), who found that the sclerified tissue surrounding the seed is of chalazal nature and that
the seed belongs to a type characterized by CoRNER (1976) as pachychalazal, a peculiar feature
occurring in a limited number of families, amongst them /cacinaceae, but not Aquifoliaceae. Al-
though not too fond of split families, I feel this new observation gives additional strength to recog-
nize Sphenostemon representing a family of its own.

DicKIsoN & BAAs (1977) noted a remarkable similarity in vegetative anatomy and some gross
morphological features between Sphenostemon and Paracryphiaceae, a monotypic family from
New Caledonia. This is compatible with the gradually accepted transfer of Aquifoliaceae, Icaci-
naceae, and Sphenostemonaceae from the heterogeneous order of the Celastrales to the Theales.

References: P. Baas, Blumea 22 (1975) 311—407, especially 339-340; H.E. Barton, Bull.
Mens. Soc. Linn. Paris no 7 (1875) 53; Adansonia 11 (1875) 307; I.W. Batmey, J. Arn. Arb. 37
(1956) 360—365, 9 fig.; I.W. Bamey & B.G.L. Swamy, J. Arn. Arb. 34 (1953) 77—87, fig. 1-3,
tab. 1—3; L. BERNARDI, Candollea 19 (1964) 199-205; E.J.H. Corner, The seeds of dicotyledons
(1976) 5—6; A. Cronguist, Integrated system, efc. (1981) 720; W.C. Dickison& P. Baas, Blumea
23 (1977) 417—438, especially 429—431; L.S. Grsss, Contrib. Flora & Phytogr. Arfak Mts (1917)
136; J. Hutcutnson, Families of flowering plants, ed. 2, 1 (1959) 139; The genera of flowering
plants 1 (1964) 124; H.D. INGLE& H.E. Dapswett, The anatomy of the secondary xylem of SW.
Pacific tree species and their taxonomy, in mimeogr. 12 pp. pamphlet offered in a symposium
of the 10th Pacif. Sci. Congr., Hawaii (1961) especially 5—7, table 1; C. LAUTERBACH, Nova Gui-
nea 8 (1912) 844; Tu. Logsener, in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 36; F. von MUELLER,
Fragmenta phytographiae Australiae 9 (1875) 151; H.K. Airy SHaw, Kew Bull. 27 (1972)
325-326; in Willis, Dict. ed. 8 (1973) 1087; C.G.G.J. VAN Steents, Svensk Bot. Tidskr. 49 (1955)
19—23; R.F. Tuorne, Nordic J. Bot. 3 (1983) 103.

(145)

146 FLORA MALESIANA [ser. I, vol. 102

1. SPHENOSTEMON

BAILL. Bull. Mens. Soc. Linn. Paris no 7 (1875) 53; Adansonia 11 (1875) 307;
STEEN. Svensk Bot. Tidskr. 49 (1955) 19. — Nouhuysia Laut. Nova Guinea 8
(1912) 844; ENGLER in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 197; STEEN. Acta
Bot. Neerl. 1 (1952) 94, f. 2; Hatrus. Bot. Mag. Tokyo 65 (1952) 109. — Iden-
burgia Grpps, Fl. Phyt. Arfak (1917) 136, f. 10, 11; GmG& Scutitr, Bot. Jahrb.
58 (1923) 246, f. 2; PERKINS, Gatt. Monim. (1925) 23, f. 14. — Fig. 1.

Shrubs or small trees. Leaves simple coriaceous, exstipulate, almost entire to
distinctly glandular-dentate, penninerved, scattered, subopposite to pseudo-
whorled, articulate at base. Racemes terminal or axillary, bracteate at base, at
most as long as the leaves. Floral bracts early caducous. Flowers bisexual, ac-
tinomorphic, the pedicel articulate at the base, all opening about simultaneous-
ly, white. Sepals 4, free, decussate, widely imbricate, mostly hooded, sometimes
less convex, rounded, outer ones mostly + saccate at base, and with prominent
midrib. Petals free, 0 or 4 similar to the sepals but of more fleshy texture, all
caducous at anthesis. Disk 0. Stamens 1-seriate, 4, 6, or 8—13, + sessile, after
being exposed appearing as a globular body, sometimes with a few (1 —3) flimsy
appressed persistent appendages added (? staminodes), at anthesis free and ex-
panded; connective firm, brown outside; anther-cells introrse and lengthwise
dehiscent. Ovary superior, sessile on a thickened receptacle, often with grooves
from the pressure of the stamens in bud, + fusiform or cylindric, capped by a
fleshy, cap-shaped, sessile, slightly bisected stigma; cells 2, each with one pen-
dent, apically attached, apotrophic ovule. Pseudo-drupe broad-ellipsoid to sub-
globular, not rarely asymmetric, crowned by the stigma. Pericarp fleshy, finally
black. Seed(s) 1 and having the shape of the fruit, or 2 and then plano-convex.
Embryo small, surrounded by a thick, chalazal envelope, of which the outer lay-
er is hard and bony and whether or not ruminate functioning as and superficially
resembling the structure of a pyrene; this bony layer sometimes ridged outside
and star-shaped in CS.

Distr. New Caledonia, N. Queensland and East Malesia: New Guinea (incl. also New Britain, New Ireland
and adjacent isles), Moluccas (Ceram) and Central Celebes. Four species in New Caledonia, | in Queensland
and 3 in New Guinea.

Ecol. Montane rain-forest.

Morph. The New Caledonian species differ from the others in having 4 petals; all of them have few sta-
mens (4 or 6) and often subentire leaves. According to Baas (1975) they are also distinct from the species of
ser. Apetalae in their vegetative anatomy. As exposed in the introduction, the structure of the seed is different
from that hitherto assumed as a pyrene with the bony tissue derived from the endocarp; or, as BAILEY (1956)
assumed, from the testa. As a matter of fact the fleshy ‘pericarp’ is on its inside lined by an epidermis against
the bony tissue. The bony tissue, continued inside by soft tissue, is again demarcated against the seed proper;
the envelope of the seed is derived from the chalaza which, during development of the fruit, completely sur-
rounds the seed. In ser. Apetalae the bony outer part of the pachychalaza extends ruminations in the soft part
of it.

However, in the New Caledonian S. pachycladus of ser. Sphenostemon, the bony outer part of the pachy-
chalaza is circular, without ruminations. Unfortunately I have not succeeded to obtain ripe fruit of the other
three New Caledonian species to check the structure of their seed, although I found a slight indication of it
in S. oppositifolius Hur.

1986]

SPHENOSTEMONACEAE (van Steenis)

147

Taxon. The differences found between the New Caledonian species and those from East Malesia and
Queensland induce me to raise the series to the rank of section, as follows.

Section Sphenostemon

Ser. Sphenostemon; STEEN. Svensk Bot. Tidskr. 49 (1955) 21.
Petals 4. Stamens 4 or 6. Sclerified outer tissue of the pachychalaza in fruit circular, not rumi-
nated. Leaves with a hypodermis and closed vascular system in midrib.

Distr. Four species in New Caledonia.

Section Apetalae (STEEN.) STEEN., sfat. nov.

Ser. Apetalae STEEN. Svensk Bot. Tidskr. 49 (1955) 22.

Petals absent. Stamens 6 or 9—13. Sclerified outer tissue of the pachychalaza in mature fruit
ruminated in its softer inner tissue. Leaves without a hypodermis and with a simple open vascular
system. Anatomy different from species of sect. Sphenostemon.

Distr. North Queensland and East Malesia.

KEY TO THE SPECIES

1. Stamens (5—)6. Inflorescences short, 1—2 cm. Pedicels even in fruit only c. 1/2 cm. Leaves usually not con-

spicuously toothed to almost entire.

2. Racemes puberulous, few-flowered. Stigma grooved, cupular, appressed to fruit apex, 2-3 mm @. Habit
rather delicate, twigs thin. Leaves ovate- to obovate-lanceolate, acute-acuminate to caudate, 5—11 by

1 3/4—3 1/4 cm; petiole 4—7 mm .............

Pe nraen ieee nr eS htaba ce ay Bie a8 2. S. arfakensis

2. Racemes mostly glabrous. Stigma punctate in flower, flat and + elevated in fruit, hardly 3/4 to 1 mm
wide, not cap-shaped and grooved. Habit not delicate, twigs rather sturdy. Leaves mostly oblanceolate,
rather short- and mostly blunt-acuminate, 5—13 by 2 1/2—4 cm; petiole 10-20 mm. 3. S. lobosporus

1. Stamens 8—13. Inflorescences 1 1/2—11 cm in fruit. Pedicels in flower 1/2—1 1/2 cm, in fruit up to 2 cm.
Stigma sessile, grooved and cap-shaped, appressed to fruit apex, 2-3 mm @. Leaves almost always con-

1 a 7 Mle Spas i ela aa allele teat ie

1. Sphenostemon papuanus (LAuT.) STEEN. & Erprt-
MAN, Svensk Bot. Tidskr. 49 (1955) 22; v. RoYEn, Al-
pine Fl. New Guinea 3 (1982) 1252, f. 399. — Nou-
huysia papuana Laut. Nova Guinea 8 (1912) 844;
Steen. Acta Bot. Neerl. 1 (1952) 97, f. 2; Hatus.
Bot. Mag. Tokyo 65 (1952) 110. — Jdenburgia novo-
guineensis Gipps, Fl. Phyt. Arfak (1917) 137, f. 10.
— Idenburgia pachyphylla Guc & ScHLTrR, Bot.
Jahrb. 58 (1923) 246, f.2 A~M. — Idenburgia elaeo-
carpoides Guc & Scuutr, /.c. 247, f. 2 N—-X. —
Idenburgia pauciflora A.C. Smiru, J. Arn. Arb. 22
(1941) 234. — Nouhuysia pauciflora (A.C. Smirx)
Steen. Acta Bot. Neerl. 1 (1952) 97; Harus. Bot.
Mag. Tokyo 65 (1952) 109. — Nouhuysia novogui-
neensis (Gipes) Hatus. Bot. Mag. Tokyo 65 (1952)
109. — Nouhuysia pachyphylla (Gi.G & ScuitrR) Ha-
tus. Le. — Nouhuysia elaeocarpoides (Guc &
Scuitr) Hatus. 1c. — S. pauciflorum (A.C. Siti)

sey ey ery ye: ot: eee Gen

STEEN. & ERDTMAN, Svensk Bot. Tidskr. 49 (1955)
22. — Fig. 1.

Treelet or tree, (2—)5—25 m tall, 4—5S0 cm @, glab-
rous or innovations and inflorescences short-hairy or
with small brown scales; branchlets rather sturdy;
blaze and wood mostly orange brown. Leaves
pseudo-opposite to pseudo-verticillate (3—7 leaves),
coriaceous, dentate, elliptic to lanceolate or oblan-
ceolate to obovate, rounded to acuminate at apex,
distinctly nerved and veined, venation above some-
times shallowly sulcate, underneath pale or whitish
green in vivo, 2 1/2—19 by 1 1/2—7 cm; petiole
1/2—3 cm. Racemes terminal and axillary or on short
shoots, | 1/2—11 cm at base mostly bracteate; bracts
4-10 by 3—5 mm, ciliate. Flowers white. Pedicels
3—15 mm, in fruit up to 20 mm. Sepals convex to
hooded, the outer + saccate at base, midrib elevat-
ed. Petals 0. Stamens 8~—13, close together, thick, +

148

FLORA MALESIANA

[ser. I, vol. 102

ae my "es i

SY w®,

Ds
eS 2

Fig. 1. Sphenostemon papuanus (LAuT.) STEEN. & ERDTMAN. a. Habit, x 1/2, b. inflorescence, nat. size, c.
ripe bud, d. flower during anthesis, e. pistillum, all x3, f. anther, dorsal side, x 5 (RUTTEN 2240).

triangular in CS, the connective dark brown, occa-
sionally with 1—3 flimsy appendages adhering to the
stamen-globe, later the stamens spreading. Ovary fu-
siform, thick, 2 mm; stigma sessile, slightly bilobed,
cap-shaped, appressed, both with impressions from
the stamens. Fruit broad-ellipsoid or obovoid, rarely
oblique, very rarely with a stipe-like base, 1 1/2—2
1/2 by 3/4—1 3/4 cm, via red finally black. Seeds
1—2, broadly ellipsoid, (if 2) plano-convex, smooth
or ribbed (lobed in CS); stigmatic cap 3—4 mm o.

Distr. Malesia: Central Celebes, Moluccas (Ce-
ram), and New Guinea (incl. Goodenough, Fergus-
son, New Britain and New Ireland Is.).

Ecol. A subsidiary small tree in primary mixed
montane forest, or mossy forest, often associated
with Nothofagus, Quintinia, Elaeocarpus, Myrta-
ceae and Libocedrus, also in old secondary forest,
sometimes fire-induced; (SO0O—)800—3000(—3300) m,
rather common. F/. June—Jan., fr. Jan.—Dec., not
seldom //. and fr. together; c. 130 collections.

Vern. Though no uses are mentioned the plant is

well known, carrying a large number of names in
New Guinea: bukhane, Mt Ne, W. Highl., kemena-
bubodereh, kimnababoderreh, Chimbu: Masul, ke-
ramura, wehnagenaja, Mairi: Watabung, kibamo,
Kutubu, W. Highl., kKonge, Wahgi: Minj, kup, Ha-
gan: Togoba, mandam’, Sila lang., medaboh, Mt
Ambua, mem, Mendi, meme, Mendi, Tomba,
mem/(i), Enga, menseh?, Tanah Merah, mime, Meri-
manta, mogoro, mokoro, morogl, mororo, Kapau-
ku, omipa, Asaro: Kefamo, ounatrok, Hindenburg
Ra., Sepik, patiba, Hagen, Minj, soka, Naho lang.
New Britain: napun, newala, Mt Talawe.

Notes. By the large increase of specimens avail-
able (in 1952 c. 24, now c. 130) the variability has
shown up and S. pauciflorum cannot be upheld. The
number of tangible differential characters used in the
key in 1952 has also become more vague.

This variability extends to the indument; several
specimens (8) carry small brown scales on the ra-
cemes and innovations, in 7 the racemes and innova-
tions are hairy to puberulous, the hairs sometimes

1986]

mixed with scales, 5 specimens have more or less bul-
late leaves and are hairy, some 8 specimens have
ridged seeds, 3 have both ridged seeds and some
hairs. In 8 specimens I found 1, 2 or 3 flimsy appen-
dages which adhere to the anther-globe. Formerly I
assumed that they could represent reduced petals (4
true petals are found in the New Caledonian species)
but as these appendages are not decussate and persist
with the stamen whorl, I am now inclined to regard
them as reduced stamens. These independent varia-
tions do not allow to distinguish further taxa.

2. Sphenostemon arfakensis (Gipps) STEEN. & ERDT-
MAN, Svensk Bot. Tidskr. 49 (1955) 22. — Idenburgia
arfakensis Grpps, Fl. Phyt. Arfak (1917) 139, f. 11.
— Nouhuysia arfakensis (GtpBs) STEEN. Acta Bot.
Neerl. 1 (1952) 97; Hatus. Bot. Mag. Tokyo 65
(1952) 110.

Shrub or small tree, 5 m. Twigs rather delicate.
Leaves lanceolate, to oblanceolate, distinctly acute-
acuminate, rather obscurely toothed, to almost en-
tire, tending to be opposite, not in pseudo-whorls,
5—11 by 1 3/4— 3 1/4 cm; petiole thin, 4-7 mm. Ra-
cemes short, puberulous, few-flowered, c. 2—3 cm.
Buds 5 mm. Sepals shallow-convex, not hooded. Sta-
mens 6. Stigma grooved, cupular, appressed to the
apex of the ovary, 2—3 mm @. Fruit 10 by 7 mm.

Distr. Malesia: New Guinea (Arfak Mts), 2 col-
lections.

Ecol. On open summit of Mt Kubré, c. 2700 m.
Fl. Dec.

Notes. Though some dozen collections of Sphe-
nostemon were made on the Arfak Mts, the only spe-
cimen known, besides the type (Grpss 6003), is Kos-
TERMANS 2217, found near Angi Gita Lake at 1800 m.

By the stigma it resembles S. papuanus and differs
from S. lobosporus, but the 6 stamens point the re-

SPHENOSTEMONACEAE (van Steenis)

149

verse way. By its delicate habit it differs from both.

3. Sphenostemon lobosporus (F.v.M.) L.S. Situ,
Proc. R. Soc. Queensl. 68 (1957) 43. — Phleboca-
lymna lobospora F.v.M. Fragm. 9 (1875) 151.

Tree, 5—9 m high, c. 20 cm o. Leaves elliptic, obo-
vate, or oblanceolate, mostly almost entire, opposite
or 3 or 4 in pseudo-whorls, marginal teeth usually
few and faint, apex acuminate, 5—10 by 2—4 cm; pet-
iole 1—1 1/2 cm. Racemes short, few-flowered, 1—2
cm, mostly glabrous. Pedicels 3—5 mm in fruit.
Bracts 4 1/2—5 mm. Stamens (5—)6. Ovary slender,
hardly ridged; stigma small knob-shaped. Fruit as in
the other species, but crowned by a punctate stigma
which is also in fruit non-appressed and hardly 1 mm
Qo.

Distr. N. Queensland (Cook & S. Kennedy Dis-
tr.); in Malesia: SE. New Guinea (Milne Bay Distr.:
Simpson Ra.; Mt Dayman; W. Highl. Prov.: Jimi
valley); 3 collections.

Ecol. Subsidiary tree in mixed, montane rain-
forest, 1200—1500 m (in Queensland 500—1150 m).
Fl. Aug., fr. June, Aug.

Notes. Of the three specimens from Papua, one
(SCHODDE 5550) has flowers and fruit which enabled
to examine the stamens: one flower had 6 and ano-
ther 5 + a staminode. This specimen agrees also in
the inconspicuously dentate leaf margin. The other
sheet (Brass 23154) has only fruits, but the stigma is
typical for S. Jobosporus; however, it has distinctly
dentate leaves and racemes up to 5 cm in fruit, which
is atypical for S. /obosporus. The third specimen
(For. Coll. Herb. Bulolo 8458) has hairy infructes-
cences, the two others being glabrous, reducing dif-
ferences with S. arfakensis to the stigma only!

In Queensland the species attains a height of 6—24
m and a stem diameter of 50 cm.

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a

ELAEAGNACEAE (J.F. Veldkamp, Leyden)'

This small family has a typical northern hemisphere range; it is absent from South America, ex-
tends in Eurasia only to the Mediterranean and in Southeast Asia to Malesia and NE. Queensland.
There are 3 genera of which E/aeagnus occurs throughout the range (20—50 spp.), Shepherdia oc-
curs only in North America (c. 3 spp.), and Hippophaé occurs throughout Eurasia (c. 3 spp.).
The habitat is chiefly in steppes and along coasts, but in SE. & E. Asia, Malesia and N. Queens-
land Elaeagnus is found as a substage liana in the everwet rain-forest, showing no special prefer-
ence for seasonal climates.

A characteristic feature is the universal occurrence of an often dense indument of scales and
frequent occurrence of short-shoot thorns.

About the affinity three opinions prevail. BENTHAM & Hooker /. (1880) placed the family near
the Thymelaeaceae and this position in the Thymelaeales was still upheld by MELCHIOR (1964) and
supported by Rao (1974). Most authors, e.g. RENDLE (1952) follow VON WETTSTEIN (1911), who
accommodated the family in the Myrtiflorae. CRONQUuIST (1981) included it in his Rosidae-order
Proteales, but wondered whether this was not an artificial place. He also pointed out affinities
to the Thymelaeaceae which he included in the Rosidae-order Myrtales, but remarked (/.c. 603)
that “As a putative member of the Myrtales, the Elaeagnaceae would stand out like a sore thumb
on anatomical as well as floral morphological grounds, but an evolutionary relationship via the
Thymelaeaceae cannot be ruled out on the basis of present evidence. For the present it will do
no harm to retain the Proteaceae and Elaeagnaceae in the same order. When more evidence is
available it may become necessary to restore the order E/aeagnales and insert it in a position fol-
lowing the Myrtales.”

The latter view was held by TAKHTAJAN (1969), who in 1980, however, placed the Elaeagnales
with the Elaeagnaceae as its only family next to the RAamnales in the Celastranae, which super-
order he placed next to the Proteanae, which include the Proteales. The Thymelaeaceae he put
in the Malvanae and the Myrtales in the Myrtanae. Such a relationship with the Rhamnales was
also proposed by HUTCHINSON (1926, 1959, 1973) and THORNE (1983).

DAHLGREN (1975) recognizes Elaeagnales but does not make a decision about affinity other than
those mentioned.

Pollen structure does not lead to an unequivocal opinion about affinity (LEIns, 1967).

Fossil pollen considered to represent Elaeagnus dates from the Oligocene (MULLER, 1981).

References: BENTHAM & Hooker, Genera Plantarum 3 (1880) 203; Cronguist, An integrated
system efc. (1981) 603, 606; DAHLGREN, Bot. Notis. 128 (1975) 119—147, especially p. 134; Hut-
CHINSON, Fam. FI. Plants | (1926) 245; ed. 2 (1959) 342; ed. 3 (1973) 424; Lerns, Grana Palynol.
7 (1967) 390—399; Metcuior, Syllabus der Pflanzenfamilien ed. 12, 2 (1964) 320; MULLER, Bot.
Review 47 (1981) 87; V.S. Rao, J. Ind. Bot. Soc. 53 (1974) 156—161; RENDLE, Classification FI.
Pl. 2 (1952) 372; TAKHTAJAN, Flowering Plants (1969) 229; Bot. Review 46 (1980) 225—359; THorR-
NE, Nordic J. Bot. 3 (1983) 105; von WetTsTEIN, Handb. Syst. Bot. (1911) 669.

ELAEAGNUS

Tourn. ex LINNE, Sp. PI. 1 (1753) 121; Gen. Pl. ed. 5 (1754) 57; SCHLECHTEND.
in DC. Prod. 14, 2 (1857) 606; Maxim. Mél. Bot. 7 (1870) 559; Serv. Bull. Herb.
Boiss. II, 8 (1908) 381; Beih. Bot. Centralbl. 25, 2 (1909) 1 (‘Mon. Eléagn.); ’T
Hart & VeELDK. Blumea 26 (1980) 393. — Fig. 1, 2.

Shrublets, shrubs, rarely trees, or (in Mal.) woody climbers, branching mono-
podially, with stellate scales; older parts usually armed with thorns derived from
short-shoots. Leaves spiral, simple, entire. F/owers usually bisexual, in axillary

(1) Introduction by the General Editor.

(151)

152 FLORA MALESIANA [ser. I, vol. 102

inflorescences, or pseudo-terminal, cymose, actinomorphic, 4(—5—8)-merous,
often scented. Bracteoles absent. Perianth simple; tube + cylindric, qua-
drangular, constricted above the ovary, then inflated (‘limb’); perianth-
segments (in Mal.) 4, valvate. Disk usually inconspicuous, intra-staminal. Stfa-
mens (in Mal.) 4, alternitepalous, inserted in the throat; anthers dorso-versatile,
introrse with 2 longitudinal slits. Ovary superior, 1-locular; style 1; stigma uni-
lateral, elongated. Ovule 1, basal, anatropous. Fruit a drupe, enclosed in the en-
larged fleshy perianth-tube, usually with 8 longitudinal ribs; exocarp fleshy,
mesocarp bony or leathery, endocarp woolly pubescent inside (in Mal.). Seed
1; endosperm absent (in Mal.), rarely scanty; embryo straight.

Distr. About 20—?45 spp. over the northern hemisphere through Bo Southeast Asia and Malesia to
N. Queensland; in Malesia 2 spp.

Ecol. Primary and secondary rain-forest, without a preference for dry regions or a seasonal climate, from
the lowland up to c. 2100 m.

Taxon. SERVETTAZ (1908, 1909) made an excessively detailed classification of the genus, splitting up the
classical species into a number of others, subspecies, and varieties. This work is often more an impediment
than a help in the study of the taxonomy of Elaeagnus. The delimitating characters used by SERVETTAZ have
usually been drawn from too few specimens, and his use of the shape, size, colour, and consistency of the
leaves and the colour of the flowers has proven to be of little value, as these generally are individual features
of the specimens, but not of taxa. He had a very confusing way of citing his material. In Malesia he distin-
guished 6 spp. and a number of infraspecific taxa. Later authors have recognized only one, joining all into
E. latifolia, or two, E. conferta and E. triflora. | agree with the latter opinion, although it may be remarked
that the differences between these two species and E. /atifolia are only slight. The differences seem to be clear-
cut when flowers are available, but vegetative material is impossible to identify. None of SERVETTAZ’ infraspe-
cific taxa could be maintained.

The Malesian species of Elaeagnus belong to sect. Elaeagnus (sect. Sempervirentes, nom. inval., SERVET-
TAZ, 1909).

Uses. The Malesian species are usually misidentified as E. /atifolia and the possibly different uses of the
species can therefore not be disentangled. HEYNE (Nutt. Pl. 1927, 1152) mentioned the presence of more or
less edible, sourish fruits (areuj susumunding or areuj dudurenan) and of extremely acid ones (areuj dudure-
nan). This may be due either to the presence of various races of one or both species, or it may be of specific
significance.

Field notes. Flowers fragrant, white, cream, pale yellow, whitish and brown dotted, pale inside. Fruit
pinkish to glossy red or pale brown, when ripe juicy and sweet to very acid. Measurements of the fruit are
taken from dry specimens in the herbarium; they are much smaller than in the living state; collectors hardly
ever measure them in the field.

KEY TO THE SPECIES

The term ‘limb’ means the inflated, quadrangular part of the perianth between the constriction of the tube
and the perianth segments.

1. Flowers tubulose-campanulate; limb 4.5—7 mm long. Style straight, sparsely scaly at base only. Perianth
segments (2—)3—4 by 2—3.25 mm. Continental Southeast Asia ...................00000- E. latifolia

1. Flowers trumpet-shaped; limb 1—4 mm long. Style at apex hook-shaped.
2. Limb 3.5—4 mm long, sometimes thickened at base. Style densely stellate-scaly, especially near the base.
Renan sepmients! (i — leo —2 sb yp les — 2) mime erie ree eae rt ae ee ee 1. E. conferta
2. Limb 1—3(—4) mm long. Style glabrous, rarely with some stellate scales at base. Perianth segments
7A —5)pby 2—Suminiad..iyf) nas eens) Aon erat: Sel Serco beeen ee 2. E. triflora

1986]

ELAEAGNACEAE (Veldkamp)

153

1. Elaeagnus conferta Roxs. [Hort. Beng. (1814) 11,
nomen] Fl. Ind. 1 (1820) 460; ed. Carey 1 (1832) 440;
A. Ricw. Mém. Soc. Linn. Paris 1 (1823) 385, 405;
SCHLECHTEND. in DC. Prod. 14, 2 (1857) 612; Lin-
naea 30 (1859—60) 367; ibid. 32 (1863) 301; Kurz,
For. Fl. Burma 2 (1877) 331; Serv. Bull. Herb.
Boiss. II, 8 (1908) 389, incl. ssp. javanica (BL.) SERV.
et ssp. dendroidea (SCHLECHTEND.) SERV.; Beih. Bot.
Centralbl. 25, 2 (1909) 89, 91, f. 5.1, incl. ssp. eucon-
ferta SERV., var. calcuttensis SERV., ? var. malaccen-
sis SERV., var. septentrionalis SERV., et var. silheten-
sis SERV.; Lecomte, Fl. Gén. I.-C. 5 (1915) 181;
Back. & Baku. f. Fl. Java 2 (1965) 86; ’r Hart &
VELDK. Blumea 26 (1980) 396, with full synonymy
and discussion. — EF. javanica BL. Bijdr. (1826) 638;
SCHLECHTEND. in DC. Prod. 14, 2 (1857) 614; Lin-
naea 30 (1859-60) 377; Back. in Heyne, Nutt. PI.
(1927) 1152. — E. arborea Roxs. var. dendroidea
SCHLECHTEND. in DC. Prod. 14, 2 (1857) 612. — E.
gaudichaudiana SCHLECHTEND. /.c. 612; Linnaea 30
(1859—60) 370; Serv. Bull. Herb. Boiss. II, 8 (1908)
390; Beih. Bot. Centralbl. 25, 2 (1909) 103; Merr.
Trans. Am. Phil. Soc. II, 24 (1935) 279. — E. den-
droidea SCHLECHTEND. Linnaea 30 (1859-60) 362.
— Fig. la—d.

Evergreen woody climber, occasionally erect, up
to 12 m, innovations silvery scaly. Axillary buds 2,
collateral. Leaf elliptic to obovate, 6.5—11(—12.5) by
3.5—5(—5.5) cm, base obtuse, sometimes rounded,
apex acute to acuminate; nerves 5—8 pairs, loop-
shaped; blade above with a slightly immersed midrib,
below brown to silvery scaly. Petiole 8—11 mm. Jn-
florescences 1—5-(or 6-)flowered fascicles; bracts up
to 2 mm long. Pedicels 1—2(—3) mm. Flowers
trumpet-shaped, 6—8 mm long; tube 1.5—2(—2.5) by
c. 1 mm g; limb 3.5—4 by 1.5—2.5(—3) mm @, about
twice as long as the perianth-segments; these trian-
gular to broadly ovate, (1—)1.5—2 by 1.5—2 mm, in-
side with stellate scales. Filaments filiform, broadly
winged towards the base, 0.5—1.5 mm long; anthers
(0.5—)1(—1.5) mm long. Ovary 1—2 by c. 0.5 mm 9,
glabrous. Style terminally hook-shaped, exserted for
(0.5—)1—2 mm above the throat, densely stel-
late-scaly; stigma 1—2 mm long. Fruit ellipsoid, up to
35 by 15 mm ao (spirit); seed and peanut-shaped em-
bryo to 20 by 7 mm 9; radicle up to 3 mm long.

Distr. Continental SE. Asia: Nepal, India (As-
sam), through Bangladesh, Burma (Pegu), Indochi-
na, Andamans & Nicobars, to Malesia: Malaya (P.
Penang, Perak), Sumatra, throughout Java (Bogor,
Priangan, Semarang, Madiun, Surabaya).

Ecol. Rather rarely recorded, in bamboo and
mixed deciduous, and in evergreen forests; 400—2000
m.

Vern. Java: areuj dudurenan, a. susumunding,
kakaduan, meligi, S.

Notes. ’t Hart& VetpkKamp(1 980) at length dis-

cussed the typification and location of the types of
the complicated synonymy. Elaeagnus gaudichaudi-
ana has been included, the type of which came from
Indochina. MERRILL (1935) suggested in his evalua-
tion of LourREtRo’s species (Flora Cochinchinensis,
1790) its conspecificity with Octarillum fruticosum
Lour. (E£. fruticosa (LourR.) CHEVAL.), which he
claimed to be the only representative of Elaeagnus in
Indochina. As the LouRErRo specimen in the BM is
sterile, and SERVETTAZ distinguished 3 species in
Indochina, this conclusion cannot be accepted. It
may be further noted that Lourerro described his
species with solitary flowers, while E. conferta usual-
ly has several-flowered fascicles which are only occa-
sionally reduced to one flower. Elaeagnus conferta is
thus the oldest name available of certain application.

Field notes. Usually climbing, rarely a tree,
with reddish brown bole. Flowers yellowish, pendu-
lous. Fruit orange-red when ripe, juicy, delicious but
acid, up to 4 by 2 cm a.

2. Elaeagnus triflora Roxs. [Hort. Beng. (1814) 11,
nomen] FI. Ind. 1 (1820) 459; ed. Carey 1 (1832) 439;
SERV. Bull. Herb. Boiss. II, 8 (1908) 390, incl. ssp. ri-
gida SERV., ssp. obsoleta SERV., ssp. polymorpha
SERV. ef ssp. tetragonia SERV.; Beih. Bot. Centralbl.
25, 2 (1909) 104, incl. var. brevipes SERV. et var. lon-
gipes SERV.; BAcK. & BAKH./f. Fl. Java 2 (1965) 86;
*T Hart & VELDK. Blumea 26 (1980) 398. — E. /ati-
folia (non L.) A. Ricw. Mém. Soc. Linn. Paris 1
(1823) 386, 404; Mor. Syst. Verz. (1846) 70; Mia. Pl.
Jungh. 2 (1852) 173, incl. forms; ZoLi Syst. Verz. 2
(1854) 117; SCHLECHTEND. in DC. Prod. 14, 2 (1857)
610, p.p., incl. var. triflora SCHLECHTEND.; Linnaea
30 (1859-60) 347; Kurz, Nat. Tijd. N. I. 27 (1864)
172; BENTH. FI. Austr. 6 (1873) 39; Hook.f. Fl. Br.
India 5 (1886) 202, p.p.; Fors. & HeEmst. J. Linn.
Soc. Bot. Lond. 26 (1894) 403, p.p.; Gicin E. & P.
Nat. Pfl. Fam. III, 6a (1894) 251, p.p.; GAMBLE, J.
As. Soc. Beng. 75, ii (1912) 267; Koorp. Exk. Fl. Ja-
va 2 (1912) 658; Ripi. J. Fed. Mal. St. Mus. 8 (1917)
81; Merr. Sp. Blanc. (1918) 279; RENDLE, J. Bot. 63
(1925) Suppl. 90, p.p.?; C.T. Wuite, Contr. Arn.
Arb. 4 (1933) 75; von MALM in Fedde, Rep. 34 (1934)
282. — E. ferruginea A. Ricu. Mém. Soc. Linn. Pa-
ris 1 (1823) 387, 404; Mig. Pl. Jungh. 2 (1852) 173;
SCHLECHTEND. in DC. Prod. 14, 2 (1857) 610; Lin-
naea 30 (1859-60) 350; ibid. 32 (1863) 303; Serv.
Bull. Herb. Boiss. I, 8 (1908) 390, incl. ssp. suma-
trana SERV.; Beih. Bot. Centralbl. 25, 2 (1909) 110,
incl. var. richardia Serv. et var. atrovirens SERV. —
E. philippinensis Perrotret, Mém. Soc. Linn. Paris
3 (1824) 114; Merr. Sp. Blanc. (1918) 279; En. Phi-
lip. 3 (1923) 134; Quis. Medic. PI. Philip. (1951) 638;
Liu & Lat, Quart. J. Taiwan Mus. 33 (1980) 247. —
E. rigida Bu. Bijdr. (1826) 639; Mig. Pl. Jungh. 2
(1852) 173; SCHLECHTEND. in DC. Prod. 14, 2 (1857)

154

FLORA MALESIANA

V.O384

[ser. I, vol. 102

ORROGE aieae lose 1
if * ae v ihc 3y
ree Se
ms

Nien Ls SN Oa

ae

pe

ae eae

Fig. 1. Elaeagnus conferta Roxs.a. Habit, x 1/2, b, c. flowers, x3, d. scale, x 25. — E. triflora Roxs. var.
brevilimbata ’Tt Hart. e. Flower, x3. — E. triflora Roxs. var. triflora. f. Flower, x3 (a—d ARENS 34, e
HooGLanp 4537, f LORZING 8523). ;

614; Linnaea 30 (1859-60) 376. — E. angustifolia
(nonL.) BLANCO, FI. Filip. 1 (1837) 74; ed. 2 (1845) 53;
ed. 3, 1 (1877) 100. — E. perrottetii SCHLECHTEND. in
DC. Prod. 14, 2 (1857) 613, nom. superfl.; Mia. FI.
Ind. Bat. 1, 1 (1858) 981. — E. cumingii SCHLECH-
TEND. in DC. Prod. 14, 2 (1857) 613; SERV. Bull.
Herb. Boiss. II, 8 (1908) 391, incl. ssp. perrottetii
SERV.et ssp. philippinensis SERV. —E. rostrata SERV.
Bull. Herb. Boiss. II, 8 (1908) 392; Beih. Bot. Central-
bl. 25, 2 (1909) 113, f. 5: 40—42. — E. zollingeri Serv.
Bull. Herb. Boiss. II, 8 (1908) 392; Beih. Bot. Central-
bl. 25, 2 (1909) 112, f. 5: 36—39. — Fig. le, f.

KEY TO THE VARIETIES

1. Limb 2—3(—4) mm long, subequal to 1.5 times as
long as the segments. Style glabrous or rarely with
some stellate scales at base ..... a. var. triflora

1. Limb 1—1.5 mm long, about half as long as the
perianth segments. Style glabrous

b. var. brevilimbata

a. var. triflora — Fig. 1f.
Evergreen woody climber, occasionally shrubby,
up to 10 m high, innovations silvery scaly. Axillary

1986]

ELAEAGNACEAE (Veldkamp)

buds 2, collateral. Leaf elliptic to oblong,
1.5—10(—12) by (0.3—)1—4(—6) cm, base obtuse,
sometimes rounded, apex acuminate, sometimes
acute; nerves 5—8 pairs, loop-shaped; blade above
with a slightly immersed midrib, below brown to sil-
very scaly. Petiole (4—)5—6(—7) mm. Jnflorescences
1—3(—8)-flowered fascicles; bracts up to 2 mm long.
Pedicels 2—5(—7) mm, elongated in fruit to 8 mm.
Flowers trumpet-shaped, 6—9 mm _ long; tube
1.5—2.5 by c. 1 mm g; limb 2—3(—4) by (1.5—)

Pig. 2. Stem thorns on the stem of an Elaeagnus liane
(West Java, Tjibodas; van Sreenis 11162), x 1/2.

155

2—2.5(—3) mm g, slightly shorter than to 1.5 times as
long as the perianth segments; these broadly ovate,
(2—)2.5—4(—5) by 2—3 mn, inside sometimes with
stellate scales. Filaments filiform to cone-shaped,
0.3—1 mm long; anthers (0.5—)1(—1.5) mm long.
Ovary 1—2 by 0.5(—1) mm g, glabrous. Style fili-
form, terminally hook-shaped, exserted for
(0.5—)1—2 mm above the throat, glabrous, rarely
with a few scattered stellate scales at base; stigma
1—2 mm long. Fruit ellipsoid, up to 4 by 2 cm (in spir-
it), seed and peanut-like embryo up to 17 by 6 mm;
radicula up to 3 mm long.

Distr. N. Queensland, throughout Malesia, incl.
also New Britain. Also in Botel Tobago, SE. of Tai-
wan. Not in New South Wales.

It cannot be ruled out that this species occurs in
continental Asia as well under some other name. AlI-
though not all names have been checked for that
area, so far none were discovered which should have
been included here.

Ecol. Primary rain-forest, swamp forest, second-
ary scrub with Gleichenia, sometimes gregarious,
from the lowland up to 1800 m.

Uses. The ripe fruit is given to children suffering
from amoebic dysentry (Suuit, 1934, cited by Qut-
SUMBING, 1951, sub E. philippinensis). CHOPRA
(1933, ex QUISUMBING, /.c.) stated that the flowers
are astringent and cardiac.

Vern. Sumatra: hail-hail, Toba-Batak, kail-kail,
Karo-Batak. Java: (areuj or daun) dudurénan, em-
pos, kakduan, kitjepot, leutik, S, berbikuda, godong
wadang, ketadak, Md. Bali: kelintju. Philippines:
bantap, Sul., banekan, Bon., kopapei, padias, lg.,
lagot, Buk., malaimus, P. Bis.

Note. Field note. Shrub or woody climber. Flow-
ers strongly scented.

b. var. brevilimbata ’r HART, Blumea 26 (1980) 400
(‘brevilimbatus’). — E. triflora Roxs. ssp. tetrago-
nia (non Serv.): MERR. & PERRY, J. Arn. Arb. 22
(1941) 267. — Fig. le.

Leaf with (5—)6—11 pairs of nerves; petiole (2—)
3—4(—5S) mm long. I/nflorescences 1—4 (or 5)-
flowered fascicles. Flowers 4—7.5 mm long; tube
1.5—2 by c. 1 mm g; limb 1—1.5 by 1.5—2 mma, +
half as long as the perianth segments; these broadly
ovate, 2—3 by (1.5—)2(—2.5) mm, stellate-scaly in-
side. Fruit 15—17 mm long. Seed up to 12 by 4.5 mm;
embryo up to 9 mm long.

Distr. N. Queensland and in Malesia: through-
out New Guinea.

Ecol. Primary rain-forest, Castanopsis-forest,
swampy secondary forest; (0O—)450—1600(—2125) m
alt.

Note. Field notes. Climber or scrambling shrub,
to 30 m high. Flowers grey green, white, yellow, very
fragrant. Fruit red, fleshy.

156 FLORA MALESIANA [ser. I, vol. 10

Uncertain

Elaeagnus conferta Roxs. var. pallescens SERV.
Beih. Bot. Centralbl. 25, 2 (1909) 96. — This is based
on KorTHALss.n. from Sumatra (in L), but its iden-
tity remains uncertain as the specimens are sterile.

MENISPERMACEAE (L.L. Forman, Kew)' ”

Dioecious woody or sometimes herbaceous climbers, rarely erect shrubs or trees
(Cocculus sp. in Mal.); tubers sometimes present (Sfephania spp.); sometimes
producing exudate or rarely latex (Fibraurea; Tinomiscium). Wood often with
concentric rings or arcs of vascular bundles separated radially by interfascicular
rays, or vascular bundles in one ring; wood sometimes yellow. Young shoots of-
ten tendrilliform. Young stems usually drying longitudinally striate. Stipules ab-
sent. Leaves spiral, simple (rarely trifoliolate extra-Mal.), often palmatinerved
at base and sometimes peltate, or penninerved, margin usually entire, some-
times broadly crenate (rarely dentate extra-Mal.), sometimes deeply 3—S5S-lobed;
petiole often swollen at base, sometimes also at apex, sometimes leaving a raised
discoid scar on the stem. /nflorescences axillary or on defoliate branches or cau-
liflorous; solitary or fasciculate, various in form, often cymes, thyrses or
pseudoracemes, branching of cymes rarely umbelliform (Stephania spp.), flow-
ers rarely in a disciform capitulum (Sfephania spp.); female usually fewer-
flowered than male, female rarely with accrescent bracts (Cissampelos spp.).
Flowers small, usually green, yellow or white, actinomorphic or female some-
times zygomorphic. Sepals usually in 1—2(—4) whorls of 3, or 1 whorl of 4, the
outer whorl(s) smallest, imbricate but the innermost whorl sometimes valvate
and sometimes + connate, sepals rarely spirally arranged (Hypserpa); in female
sometimes reduced to 1 or 2. Petals mostly 3—6 in 1 or 2 whorls or 0, free or
sometimes + connate, usually smaller than the sepals, rarely larger (Sarcopeta-
lum), the lateral edges or lobes often inflexed and sometimes clasping the oppo-
site stamen, often glandular within; in female sometimes reduced to 1 or 2. Sta-
mens mostly 3 or 6, sometimes 9 or up to c. 40, often free and opposite a petal,
or variously connate, sometimes forming a peltate synandrium, connective
sometimes adaxially or abaxially thickened, rarely terminally prolonged (Ma-
crococculus); anthers introrse to extrorse with dehiscence longitudinal to trans-
verse. Staminodes sometimes present in female, usually subulate. Carpels free,
usually 3 or 6, sometimes | or to 12 (to c. 30 in extra-Mal. Tiliacora), sometimes
borne on a short gynophore; style terminal when present; stigma often sessile,
reflexed and lobed or divided. Pistillodes 0 in male. Ovules 2 reducing to | in
development, attached ventrally. Fruits of 1—6 (—10) drupes sometimes borne
on an enlarged + globose, discoid or columnar carpophore which is rarely
shortly branched (Anamirta, Tiliacora). Drupes sometimes narrowed at base in-
to a Stipe, style-scar terminal, ventral or close to base; exocarp membranous to
coriaceous, mesocarp fleshy; endocarp usually bony, rarely papyraceous to
crustaceous (Pycnarrhena spp.), rugose, tuberculate, spiny, ridged or variously
ornamented on at least the dorsal surface, sometimes smooth or surface fibrous,

(1) With some assistance by the General Editor.

(2) A number of figures are reproduced from precursors in the Kew Bulletin by permission of the Controller
of Her Majesty’s Stationary Office. They have been marked Courtesy Kew Bull.
Ten plates have been drawn by Miss Eleanor Catherine at Kew, at the expense of Foundation Flora Male-
siana.

(157)

158 FLORA MALESIANA [ser. I, vol. 102

usually with a condyle, i.e. a ventral sometimes hollow intrusion into the seed-
cavity around which the seed is curved, or a ventral groove, cavity or chamber;
the condyle when hollow often 2-chambered and with 2 lateral or ventral aper-
tures, or condyle septiform or lamelliform, then sometimes centrally perforate.
Seed often horseshoe-shaped or subannular, sometimes straight and + broadly
ellipsoidal or deeply cup-shaped; endosperm present or absent, sometimes rumi-
nate. Embryo usually either elongate and with semiterete or flattened con-
tiguous cotyledons or flat and very thin with divaricate foliaceous cotyledons,
sometimes broadly ellipsoidal with thick contiguous cotyledons, rarely cotyle-
dons much folded (Arcangelisia); radicle very small.

Distribution. The family is almost entirely tropical, the exceptions being Menispermum, a
northern temperate genus with 2 disjunct species in North America and Northern Asia, and a few
species of Cocculus which extend into North America and temperate Asia.

There are 73 genera in the family and of these 30 occur in Asia, 30 in Africa, 22 in America
and 10 in Australia to the Pacific. Of the 25 Malesian genera 20 occur in continental Asia, and
6 occur in Africa of which 2 (Cissampelos and Cocculus) are also in America. Of the Malesian
genera 9 are shared with Australia and of these 6 extend into Asia; Legnephora is limited to Cen-
tral and East Malesia, Carronia and Sarcopetalum occur in New Guinea. Only 2 of the Malesian
genera are endemic, Chlaenandra and Macrococculus, both in New Guinea.

Fossils. As can be expected from a widely distributed, pantropical family with no special
means of dispersal and belonging to assumed primitive ancestry, early fossils are extant. The most
certainly well-identified taxa are from the fruit which is characteristic and date from the Eocene.
I assume they will be later located in the Upper Cretaceous. Leaf fossils have been described from
the Lower Cretaceous to the Oligocene, but these records must be regarded with caution since the
leaf morphology of Menispermaceae occurs in many other families.

Fossilized endocarps assigned to or allied to modern Southeast Asian genera of Menisperma-
ceae have been found in the Early Eocene (Y presian) London Clay Flora and other Lower Tertiary
floras of Southern England. Species of Tinospora were described by REmD & CHANDLER (1933)
and CHANDLER (1961, 1962). A species of Diploclisia appears in CHANDLER (1961: 161—162, t.
16/14—17), a Parabaena in CHANDLER (1964) and a Tinomiscium in CHANDLER (1961: 149—150,
f. 17, t. 15/18—21). Davisicarpum which resembles Limacia in its larger lateral condylar cavities
is described in CHANDLER (1978: 20, t. 3/6—8) together with Atriaecarpum on p. 21, t. 4/4—10,
which resembles Tinosporeae; another fossil genus of the latter affinity is Microtinomiscium in
REID & CHANDLER (1933: 164, t. 4/5—6).

R.A. Scott (1956) has discussed the fossil endocarps of Tinosporeae found in the Eocene Clar-
no formation in Oregon, U.S.A. These differ from the living representatives in having very thick
walls containing lacunae. Scott suggested that the endocarps of modern Parabaena, especially
the complex P. megalocarpa, can be derived from the fossil Chandlera by reduction of the outer
endocarp wall, and similarly modern Tinospora and Odontocarya (tropical America) endocarps
can be derived from the fossil Odontocaryioides by reduction.

Leaf fossils identified with modern Malesian genera, especially Cocculus and Cissampelos, are
known from the Lower Cretaceous to the Oligocene. Many of the records are listed in THANIKAI-
MONI (1984) and among these are Anamirta in Alaska (Mid Eocene) and ?Arcangelisia in Wash-
ington, U.S.A. (Mid Eocene).

Fossil pollen seems not to have been recorded, according to MULLER (1981).

References: CHANDLER, Flora Pipe Clay series Dorset (1962) 61—62, t. 8/4—10; Lower Tertiary
Flora Southern England 1 (1961) 154, t. 32/40; ibid. 4 (1964) 112, t. 2/17—19; ibid. 5 (1978) 20,
t. 3/6—8; MULLER, Fossil pollen records of extant Angiosperms; Bot. Rev. 47 (1981) 141; Reimp

1986] MENISPERMACEAE (Forman) 159

& CHANDLER, London Clay Flora (1933) 165—167, t. 4/7—12; Scott, Evolution 10 (1956) 74-81;
THANIKAIMONIL Trav. Sect. Sc. Techn. Inst. Francais Pondichéry 18 (1984) 129—132.

Ecology. Mostly climbing in lowland to montane forest, often where the canopy is interrupt-
ed, e.g. along river-banks, margins of forest or in clearings. Cocculus orbiculatus mostly occurs
near the sea-shore in Malesia. It has been demonstrated in Thailand by H. BANZIGER (1982) that
various Menispermaceae, but especially Tinospora, form an important link in the biological chain
which leads to extensive damage to certain fruit crops, especially longan (Dimocarpus longan) and
citrus, including mandarin. The damage is caused principally by the noctuid moth Othreis ful-
lonia, whose larvae feed mainly on leaves of Tinospora, or on other Menispermaceae when Ti-
nospora is not available. The species of Tinospora involved are typical components of secondary
vegetation where forest has been destroyed or disturbed. Destruction of forest in Thailand thus
encourages the spread of Tinospora which in turn promotes the increase of the moth which dam-
ages the fruit crops.

Pollination. Since the plants are dioecious and often grow very remote from one another, the
problem arises as to how pollination is effected. According to BANZIGER’s observations (pers.
comm.) in Thailand, the pollinators of Menispermaceae are small Diptera and Hymenoptera and
possibly also small Coleoptera and Lepidoptera. These insects are undoubtedly attracted by the
scent of the flowers. Nectar is produced apparently by the petals which are often very small,
nectary-like and glandular. In the case of Anamirta cocculus, which bears flowers in great pro-
fusion, the scent can be detected by man at a distance of 50 m. The leaves of at least some species,
e.g. Cyclea barbata and Stephania japonica, produce a fragrance which becomes evident when
they are kept in a closed container. It is therefore possible that the leaves may also play a rdéle
in attracting insects.

Altitude. In Malesia Menispermaceae is a typical lowland family, most species ascending to the
hills below 1000 m altitude. Still one third of the species ascends above that altitude, 27 in all.
Among those 4 are found as high as 1200 m, 1 up to 1300 m, 1 to 1400 m, 3 to 1500 m, 4 to 1600
m, 3 to 1700 m, 5 to 1800 m, 4 to 2000 m, 1 to 2100 m, and one is even found at 2800 m altitude.
Except for 5 of the 27, these ranges extend continuously from the lowland or lower hills up to
submontane altitude, e.g. Stephania capitata which ranges from the lowland to 2000 m. The 5
species which do not occur in the lowlands are: A/bertisia megacarpa, 1500—1600 m, Cyclea kina-
baluensis, 1700—2800 m (Mt Kinabalu), Legnephora acuta, 1750 m, Pycnarrhena tumefacta,
1200—2000 m (Mt Kinabalu), and Stephania montana, 1300—2700 m. The number of montane
to lower-subalpine species is thus very small.

Seasonal climate. The great majority of species are bound to the everwet vegetation types or
are indifferent to climatic conditions. Only three are not and occur exclusively in regions subject
to a seasonal climate. They are: Anamirta cocculus, which shows a distinct preference for a sea-
sonal climate and is therefore absent from the everwet Sunda shelf belt (Malaya, Borneo, West
Java, Philippines, and only once collected in N. Sumatra). The second species is Sarcopetalum
harveyanum, an Australian species also recorded from SW. New Guinea. The third is Tinospora
subcordata, collected in Timor, the Tenimbar Is. and S. New Guinea (Merauke eastwards to
Tarara).

Soils. Under the ecology of the species the soils are mentioned on which specimens were collect-
ed, according to the field notes. There seems often no distinct preference, e.g. for Pycnarrhena
tumefacta (in E. Borneo) is mentioned: loam and limestone; sandy; lime and sandstone.

Only a few species show preference, e.g. for limestone and calcareous rocks: Cocculus trilobus,
Anamirta cocculus (in E. Java), Tinospora trilobata, T. merrilliana, T. dissitiflora (coral shores),
T. glabra (also on the sandy beach, probably calcareous), 7. baenzigeri (Christmas I.), Tinomis-
cium petiolare (Java).

Dispersal. Positive records on dispersal are almost absent, obviously due to lack of observations
and botanists interested in this matter. Beccarinoted that fruits of Macrococculus pomiferus and
Chlaenandra ovata are eaten by cassowaries. Ripiey (1930) mentioned that the orange-yellow

160 FLORA MALESIANA [ser. I, vol. 102

drupes of Fibraurea tinctoria (= F. chloroleuca) are transported by civet-cats and birds and that
the drupes of Limacia oblonga (= L. velutina) are dispersed by civet-cats. According to Dr. M.
LEIGHTON (personal comm.) fruits of Arcangelisia flava and Coscinium fenestratum are eaten and
dispersed by orang-utans, gibbons and macaques.

Also man may have in recent time contributed to dispersal on intention, knowing the medicinal
properties of species. Thus it is here suggested that drupes of Tinospora crispa and T. baenzigeri
may have been introduced by Singapore migrant workers in Christmas I. (Indian Ocean) working
for the phosphate mines.

References: BANZIGER, Mitt. Schweiz. Entom. Ges. 55 (1982) 213—240; RipLey, Dispersal of
plants (1930).

Morphology. Tuberous roots. Many menisperms produce thickened roots or tubers but un-
fortunately these are scarcely known. In field work they should be given attention.

Habit. An exception to the general climbing habit of the family is Cocculus laurifolius, which
is an erect shrub or small tree.

Stems. The young stems when dry are often longitudinally ridged, the ridges corresponding to
the vascular rays. The bark on old stems is sometimes characteristic for certain genera or species:
in Tinospora it is usually parchment-like, becoming detached; in Fibraurea it becomes pale buff-
coloured, contrasting with the blackish petioles. The stems often bear prominent cup-like thicken-
ings from which the petioles arise.

White latex exudes from the cut stems of Fibraurea and Tinomiscium and it has been reported
by Rip.ey in Cocculus orbiculatus.

The wood is yellow, owing to berberine, in Arcangelisia, Coscinium, and Fibraurea.

Leaves. The petiole is often swollen and geniculate at the base and sometimes swollen at the
apex. The swollen regions have the function of turning the lamina to face the maximum light, an
important need when the plant is climbing its way through dense levels in the forest canopy. The
anatomical nature of these structures was investigated by CzZAPEK (1909), RUDOLPH (1909), SPER-
LICH (1910).

The leaves are always clearly peltate in Stephania; they are peltate or not in Cissampelos, Cosci-
nium, Cyclea, Diploclisia, and Sarcopetalum; in Haematocarpus they can be slightly peltate or
not peltate; in the other genera they are not peltate (in Mal. spp.).

The nervation is usually tripli- or more nerved to palmatinerved at the base, but in A/bertisia,
Carronia, Macrococculus and Pycnarrhena the leaves are mostly penninerved but sometimes the
lowermost nerves are crowded at the base.

Domatia occur in TJinospora spp. (pockets of glandular patches) and in Arcangelisia (hollow
with margin of the aperture hairy) and Anamirta (hairy patches).

Inflorescences. The basic unit in the family is a cyme, as in Cissampelos and Pericampylus. The
cymes are reduced and fascicled in A/bertisia, Macrococculus and Pycnarrhena; they are often
racemosely arranged in a pseudopanicle, i.e. a thyrse, and when the cymes are reduced to a single
flower a pseudoraceme results as in Pachygone, Sarcopetalum, Tiliacora, Tinomiscium and Tino-
spora spp. In Coscinium the cymes are condensed to dense heads of flowers racemosely arranged.
The cymes are umbelliform in Sfephania spp. and these are sometimes racemosely arranged; in
Stephania capitata and S. dictyoneura the cyme is condensed to a disciform capitulum with the
flowers sessile on a fleshy discoid base.

The inflorescences are often axillary, but sometimes in the axils of fallen leaves; in Diploclisia
(Mal. spp.) and Macrococculus they arise only from old, leafless stems. In some species, e.g. Ste-
phania spp., the position can vary from axillary to ramiflorous.

Flowers. The flowers although small show considerable variety. They are generally composed
of trimerous whorls with one or more whorls of each organ. Evolutionary trends of a similar na-
ture involving fusion and reduction are seen in the sepals, petals and stamens. The sepals of the
innermost whorl are valvate in Carronia, Limacia and Tiliacora but connate in a thick, fleshy tube
in Albertisia. The single whorl of sepals in Cyc/ea can be free or connate. The usually minute pet-

1986] MENISPERMACEAE (Forman) 161

als are often glandular and appear to function as nectaries. With their lateral edges often incurved,
sometimes clasping the opposite stamens, they form small pockets which apparently hold nectar.
The petals are connate in Cissampelos, free or connate in Cyclea, but absent in Anamirta, Arcan-
gelisia, Coscinium, and Fibraurea, rarely absent in Pycnarrhena. Sarcopetalum is unusual in the
family in having petals larger than the sepals. Asymmetrical female flowers with the sepals and
petals reduced to | or 2 occur in Cissampelos and some Stephania spp. The carpels which are al-
ways free, are reduced to | in Cissampelos, Cyclea and Stephania.

The androecium displays considerable diversity in the family, both in the form of the stamens
and the degree of fusion. The stamens are free in about half the genera, while the filaments are
connate to varying degrees in the rest. In Coscinium, only the inner 3 stamens are connate while
in Macrococculus it is the outer ring of 6 which are slightly joined at the base. The stamens are
completely fused into a peltate synandrium in Cissampelos, Cyclea, Parabaena and Stephania.
Dehiscence of the anthers varies from vertical to oblique to horizontal, but at the same time the
apical part of the stamen may be curved over. In Tinomiscium the small anthers are sometimes
immersed in the thick connective. In Fibraurea the prominent collar below the anther may repre-
sent a petal fused to the filament.

Drupes. The fruits provide some of the most useful characters for generic classification in the
family. The fruit consists of 1 or more drupes depending on the number of carpels. Curvature
of the drupes is frequently evident; it is caused by the greater enlargement during development
of the dorsal side compared with the ventral side. This process can result in the apparent bending
over on the ventral side of the apex towards the base, bringing the style close to the base. This
process produces a horseshoe-shaped endocarp and seed in the Menispermeae. The curvature can
also be lateral with the lateral edges curving round towards the ventral side. Curvature in both
directions results in a hollow boat-shaped or cup-shaped endocarp (e.g. most Coscinieae and
Tinosporeae). Either type of curvature results in a ventral intrusion into the endocarp: this ventral
part of the endocarp which intrudes into the seed-cavity or forms a ventral cavity is known as the
condyle, which in its various forms provides useful characters to distinguish genera.

Curvature of the pericarp and endocarp (with seed) can occur independently, for example in
Albertisia where the style-scar is close to the base of the drupe, indicating curvature, the endocarp
and seed remain straight. On the other hand, in Chlaenandra where the style-scar is terminal, the
endocarp and seed are deeply cup-shaped and curved around a deeply intrusive, clavate condyle.

The curvature in growth of the drupes of many genera is probably a device allowing a number
of free carpels to enlargen considerably in development while remaining attached to the very small
receptacle. Another means to the same end is provided by the development in many genera of a
carpophore which separates the drupes of one flower. At anthesis the tiny carpels may sit on an
insignificant gynophore which as the carpels develop enlargens into the carpophore. In many ge-
nera this may be no more than a subglobose lump from which the drupes diverge. In Macrococcu-
lus the carpophore is discoid with up to 10 drupes radiating from the margin, while in Tiliacora
and Anamirta it develops a short branch below each drupe. Sometimes the base of the drupe itself
is narrowed into a stipe as in Haematocarpus, Macrococculus and Tinomiscium. In Stephania
capitata some dozens of minute sessile female flowers are closely packed on a discoid inflores-
cence, and as each drupe develops it becomes elevated on a long stalk, which is probably a carpo-
phore.

Endocarp. The form of the endocarp, its degree of curvature, if any, and the nature of the con-
dyle provide important generic characters. The endocarp is frequently ornamented or sculpted,
especially on the convex dorsal surface, with a great variety of projections or patterns which are
often useful in delimiting species, if not genera.

In the horseshoe-shaped to subannular endocarps of the Menispermeae, lateral dome-shaped
extensions over both sides of the central septum produce a 2-chambered condyle in Hypserpa and
Limacia. This process is partially developed in some Cocculus spp., while loss of the central sep-
tum leads to a 1-chambered condyle in Cyclea. The 2 lateral chambers just mentioned correspond

162 FLORA MALESIANA [ser. I, vol. 102

to the 2 chambers or channels in the ventrally deeply intrusive condyles of Anamirta, Coscinium
and Chlaenandra, where lateral curvature of the seed-cavity has been carried much further. A sim-
ilar structure but in a much less developed form is seen in some Tinosporeae where the ventral
concavity is sometimes divided longitudinally into 2 by a median ridge.

Seed. The large straight ellipsoidal seed without endosperm and with large thick cotyledons is
presumably the primitive state and is exemplified by A/bertisia, Macrococculus and Pycnarrhena
(all Tiliacoreae). The more derived seeds are imbedded in copious endosperm; they are curved and
narrow with elongate collateral cotyledons as in most Menispermeae, or they have very thin folia-
ceous divaricate cotyledons as in Tinosporeae and Coscinieae (part).

The primitive seed-types seen in Pycnarrhena associated with a straight drupe having the style-
scar terminal and usually a very thin crustaceous endocarp suggests that the drupe of Pycnarrhena
is the earliest type extant in the family. Pycnarrhena also has a primitive leaf-type in the family,
the nervation being pinnate and not 3- or more-nerved at the base.

References: CzAPEK, Ber. Deut. Bot. Ges. 27 (1909) 404—407; Rupbo pH, /.c. 411—421; SPER-
LicH, ibid. 28 (1910) 57—59; Untersuchungen an Blattgelenken..I. Reihe (1910).

Pollen morphology. The pollen is small (rarely exceeding 20 um on polar axis). Spheroidal
to prolate in shape; single, isopolar; tricolporate, tricolpate, occasionally triporate and, more
rarely, apparently inaperturate. The ornamentation ranges from perforate to coarsely reticulate,
sometimes the muri are granular-papillose or transversely ridged (segmented). There is a well-
developed columellate sexine approximately twice as thick as the nexine. In transmission electron
microscopy differential staining shows there is a distinct foot layer and endexine present.

Pollen morphology does not delimit tribal boundaries but is useful in distinguishing genera,
groups of genera and, sometimes, species. The triporate pollen of Stephania has a coarsely reticu-
late tectum with sparsely distributed coarse granules in the lumina; there is a distinct endopore.
The pollen of Coscinium is also triporate but the endoapertures are indistinct and the reticulum
has small, dense granules in the lumina. Cissampelos and Cyclea have angulaperturate, oblate-
spheroidal, tricolporate pollen. The apertures are covered by a smooth operculum.

The apertures of Tiliacora are very obscure or absent. Tiliacora triandra can be separated easily
from the other species by its granular-papillose muri. Macrococculus has distinctive rugulate or-
namentation.

Some species of Tinospora have distinct aperture margins which fuse on the poles. This pollen
type has a coarsely reticulate tectum and lolongate (i.e. elongated in the plane of the polar axis)
endoapertures. Tinomiscium and Fibraurea are the only other Malesian genera for which lolon-
gate endoapertures have been recorded.

The genera Albertisia, Carronia, Aspidocarya, Chlaenandra and Parabaena are tricolpate.
More or less spheroidal or broadly lalongate (i.e. elongated in the plane of the equatorial axis)
endoapertures of varying size are present in the other genera of the region. These genera have per-
forate to reticulate ornamentation and are not always easy to distinguish.

References: FERGUSON, Pollen morphology of the tribe Triclisieae of the Menispermaceae in
relation to its taxonomy: Kew Bull. 30 (1975) 49—75; Pollen morphology of the tribe Coscinieae
of the Menispermaceae in relation to its taxonomy: Kew Bull. 32 (1978) 339—346; Har ey, Pollen
morphology and taxonomy of the tribe Fibraureae (Menispermaceae): Kew Bull. 40 (1985)
553—565; HARLEY & FERGUSON, Pollen morphology and taxonomy of the tribe Menispermeae
(Menispermaceae): Kew Bull. 37 (1982) 353—366; THANIKAIMONI, Morphologie des pollens des
Menispermacees: Trav. Sect. Sc. Techn. Inst. Francais Pondichéry 5 (4) (1968) 1—56. — I.K. FEr-
GUSON (Kew).

Cytology. The cytology of the Menispermaceae is relatively little-known and chromosome
counts have been made in only a few species in seven of the 25 known Malesian genera, viz. Tilia-
cora, Tinospora, Anamirta, Cocculus, Cissampelos, Cyclea and Stephania. The chromosomes of
these genera, and in fact of the entire family, are very small with the diploid numbers 2n = 24
or 26 being the most frequent and 2n = 22 being less common (THANIKAIMONI, 1984). Polyploidy

1986] MENISPERMACEAE (Forman) 163

is relatively rare, with the tetraploid numbers 2n = 48 being recorded in Cyclea peltata by
MarTHEw (1958) and 2n = 50 in Cocculus orbiculatus (= C. trilobus) by NAKAJIMA (1937), the
latter being wrongly quoted as 2n = 52 by THANIKAIMONL

Interestingly, in male plants of Cocculus orbiculatus a single heteromorphic bivalent was ob-
served at meiosis by NAKAJIMA. This report is doubly significant, firstly because sex chromosome
heteromorphy is generally uncommon, even in the dioecious plants among which it should be ex-
pected to occur universally, secondly and more importantly because only a single heteromorphic
chromosome pair was observed by NAKasiMa, indicating that considerable structural changes
must have occurred in the chromosomes of this species since it first arose as an allotetraploid,
probably from an original hybridization between 2n = 24 and 2n = 26 plants.

Chromosome number and morphology can sometimes provide information on the closeness of
relationship of genera or even of families, but because true chromosome homology cannot be con-
firmed between members of different families due to their widespread genetic incompatibility it
is difficult and often unreliable to propose inter-family relationships simply on the basis of simi-
larities of chromosome shape, size and number. Nevertheless, RAVEN (1975) has used this infor-
mation to indicate a chromosomal affinity between the Menispermaceae and the taxonomically
related Lardizabalaceae, in which diploid chromosome numbers of 2n = 28, 30 and 32 are com-
mon. The Berberidaceae, another family in the Ranunculales which is classified near to the Me-
nispermaceae, appears from a gross chromosomal standpoint to be related less closely to the
Menispermaceae than the Lardizabalaceae, having 2n = 12, 16 and 28 as the most common
chromosome numbers (FEDOROV, 1969).

The suggestion by ForMAN (see below) that Sabia (Sabiaceae) is a near ally of the Menisperma-
ceae finds some support from information available on its chromosome numbers, since SUGIURA
(1936) found 2n = 24 small chromosomes in S. japonica.

References: Feporov, Chromosome numbers of flowering plants (1969) (in Russian);
MATHEW, Proc. Indian Acad. Sc. B 47 (1958) 274—286; Nakasma, Cytologia, Fujii Jub. Vol.
(1937) 282—292; Raven, Ann. Missouri Bot. Gard. 62 (1975) 724—764; Sucrura, Cytologia 7
(1936) 544—595; THANIKAIMON], Trav. Sect. Sc. Techn. Inst. Francais Pondichéry 18 (1984) 95.
— P.E. BRANDHAM (Kew).

Affinities. There is general agreement that the Menispermaceae belong in the order Ranun-
culales (= Berberidales) close to the Lardizabalaceae and Berberidaceae. This view is followed
in the recent classifications of families by TAKHTAJAN (1980), CRONQuIST (1981), DAHLGREN
(1983) and THORNE (1983). The Menispermaceae share with both Lardizabalaceae and Berberida-
ceae small, mostly 3-merous flowers with stamens often 6 and opposite the often nectariform pet-
als. The Menispermaceae share with the Lardizabalaceae a mostly twining woody climbing habit,
unisexual flowers, carpels distinct and often 3 or more; shared with Berberidaceae is the presence
of benzyl-isoquinoline alkaloids (including berberine) and aporphine alkaloids.

Another probable relative of Menispermaceae is Sabia Colebr., as originally suggested by
Biume (1851) when he first described Sabiaceae, then monogeneric. Sabia is a genus of climbing
shrubs with curved drupes containing sculpted endocarps. The flowers which are bisexual and
pentamerous (unlike Menispermaceae) are remarkable in having the stamens, petals and sepals
all opposite one another. This arrangement can easily be derived from the doubly trimerous, i.e.
hexamerous flowers, which often occur in the Menispermaceae and in which, owing to the alter-
nating trimerous whorls of sepals, petals and stamens these organs are all opposite one another.
Miers while working on Menispermaceae placed Sabia between that family and Lardizabalaceae
(see LINDLEY, 1853). Hooker /. & THOMSON (1855) considered the genus intermediate between
Schizandraceae and Menispermaceae. The subsequent, and probably mistaken, inclusion of Me-
liosma and Ophiocaryon in Sabiaceae \ed later authors to place Sabiaceae in the Sapindales, but
in his recent system of family classification CRoNQuistT (1981) tentatively placed Sabiaceae in the
Ranunculales near Menispermaceae.

References: Buume, Mus. Bot. Lugd. Bat. 1 (1851) 369; Cronaguist, An integrated system of

164 FLORA MALESIANA [ser. I, vol. 102

classification of flowering plants (1981) 140; DAHLGREN, Nordic J. Bot. 3 (1983) 143; HooKER
f. & THomson, Flora Indica (1855) 208; LiInDLEy, Vegetable Kingdom ed. 3 (1855) 467a—467b;
TAKHTAJAN, Bot. Review 46 (1980) 261; THORNE, Nordic J. Bot. 3 (1983) 85—117.

Taxonomy. In his excellent monograph Diets (1910) divided the family into 8 tribes, which
are not completely separable. In his key to the tribes Dre.s was obliged to key out 2 genera and
part of another separately, and furthermore some of his distinctions between tribes do not hold.
Of the 8 tribes, 5 occur in Malesia, but 2 of these, Fibraureeae and Tinosporeae should probably
be combined, as suggested by BARNEBY (1972). Previously BARNEBY and KRUKOFF accepted DIEL’s
tribes in their work on American genera as did TROUPIN in his monograph of African Menisper-
maceae (1962). Although I do not consider these tribes to be altogether satisfactory, for conve-
nience I retained them in my series of papers on the family (1956 seg.). There is a clear need for
a complete review of the generic classification and delimitation in the family from a multidisci-
plinary approach, but until such reassessment is carried out on a world-wide basis it does not seem
worthwhile making partial and possibly temporary adjustments to the existing tribal framework,
although provisionally I include Fibraureeae under Tinosporeae. For the correct names of the 8
tribes see FORMAN (1982).

Generic delimitation also raises problems, especially when a world-wide view is taken. The gen-
era in one continent can be keyed out fairly readily, but when genera are compared between conti-
nents some of the supposed distinctions break down. Thus Tinospora in the Old World is prob-
ably not distinct from Odontocarya in America and Chasmanthera in Africa. Until the genera
have been reviewed on a world-wide basis I prefer largely to maintain the status quo, apart from
sinking Epinetrum (Africa) into Albertisia and Fawcettia (Australia) into Tinospora. These ad-
justments do not affect other genera whereas further piecemeal changes although desirable would
cause further problems.

The tribes in Asia are characterized by the following combinations of characters.

Coscinieae — Sepals imbricate. Petals 0. Stamens either all or only the inner 3 connate. Carpels
3—6. Drupe with style-scar sublateral towards base or lateral. Endocarp smooth or fibrillo-pilose,
subglobose with condyle obsolete, or subhemispherical with condyle deeply intrusive and
2-chambered. Endosperm present, sometimes ruminate. Seed broadly ellipsoidal or cup-shaped.
Embryo with thin foliaceous divaricate cotyledons which are sometimes much folded.

Menispermeae — Sepals usually free in 1—few whorls or sometimes connate when in 1 whorl,
the innermost whorl sometimes valvate, or sepals spiral. Petals (O—)3—6(—9), sometimes connate.
Female flowers with perianth sometimes reduced to 1—2 parts. Stamens free or partly connate or
united into a peltate synandrium. Carpels 1—6. Drupe strongly curved with style near base. Endo-
carp with + horseshoe-shaped dorsal region usually ornamented with projections or transverse
ridges; condyle deeply intrusive, either lamelliform and + obovate with the seed-cavity curved
around its margin or hollow with 1—2 chambers, sometimes perforate. Endosperm usually pres-
ent, but absent in Pachygone. Seed elongate, strongly curved. Embryo elongate and curved with
narrow contiguous cotyledons.

Tiliacoreae — Sepals imbricate or inner whorl valvate and sometimes connate. Petals rarely ab-
sent (Mal. spp.). Stamens free or connate. Carpels 3—10 (Mal. spp.). Drupe with style-scar near
base or lateral. Endocarp smooth, wrinkled, rugose or coarsely reticulate; straight and condyle
absent or curved with condyle intrusive and septiform. Endosperm usually absent, but present and
ruminate in Tiliacora. Seed ellipsoidal, straight. Embryo with thick accumbent cotyledons or
elongate and strongly curved with elongate contiguous cotyledons.

Tinosporeae (incl. Fibraureeae) — Sepals imbricate, rarely connate at the base. Petals 6 or 0.
Stamens free or united into a peltate synandrium. Carpels 3(—4). Drupe with style-scar terminal.
Endocarp spiny, verrucose, rugose or smooth; condyle a ventral hollow or longitudinal groove
or deeply intrusive and clavate. Endosperm present, sometimes ventrally ruminate. Seed usually
straight and ventrally hollowed or grooved, sometimes cup-shaped. Embryo with foliaceous di-
varicate or imbricate cotyledons.

1986] MENISPERMACEAE (Forman) 165

References: BARNEBY, Mem. N.Y. Bot. Gard. 22 (1972) 144—148; Drets, Pfl. R. Heft 46 (1910);
ForRMAN, The Menispermaceae of Malesia and adjacent areas I— XII: Kew Bull. (1956—1984); The
correct names for the tribes of Menispermaceae. Kew Bull. 37 (1982) 368; TRoupin, Monogr.
Menisp. afric.; Acad. R. Sc. d’Outre-Mer, Cl. Sc. & Médic., Mém., N.S. vol. 13, fasc. 2 (1962).

Comparative phytochemistry. All members of the family seem to produce phenylala-
nine- and tyrosine-derived isoquinoline alkaloids (HEGNAUER, 1969, 1973; THORNBER, 1970; SI-
won, 1982). Aporphines, bisbenzylisoquinolines, and quaternary and intensely coloured proto-
berberines such as berberine and its allies are most typical of the family. In some genera these
more usual types of isoquinoline alkaloids are accompanied by less common or even rare types
of benzylisoquinoline-related alkaloids. Such types of Menispermaceous alkaloids are the hasuba-
nans, the azafluoranthenes and related tropolo-isoquinolines, and the dibenzazonines and related
Erythrina alkaloids. Moreover, in recent time, pavine-type and aristolactam-type alkaloids were
detected in the family. Alkaloid chemistry clearly allocates Menispermaceae to Polycarpicae with
the position of one of its more specialized members. Other groups of constituents which seem to
be rather characteristic of the family are the bitter and more or less toxic principles, which are
sesquiterpenoids like picrotoxin or diterpenoids such as columbin and tinophyllone (HEGNAUER,
1969, 1973). It is perhaps not solely accidental that quaternary protoberberine alkaloids like ber-
berine, columbamine, jatrorrhizine and palmatine and diterpenoid bitter principles such as tino-
phyllone also occur in some genera of Rutaceae. A third group of phyto-constituents, the cycli-
tols, is known to be accumulated by members of several genera of Menispermaceous plants; it
is represented by the diastereoisomeric cyclohexanepentols (+ )-quercitol (quercitol) and
(—)-quercitol (viburnitol); they are presently known to occur in the genera Cissampelos, Coccu-
lus, Cyclea, Legnephora, Menispermum, Pachygone, Stephania, Tiliacora, and Triclisia. The
phenolic constituents were studied only superficially hitherto. Leaves contain flavonols or fla-
vones, or both, but seem to lack representatives with trihydroxylated B-ring and true tannins. All
other classes of phyto-constituents were neglected by phytochemists. Nevertheless, some inci-
dental observations might prove in future to be taxonomically relevant. In this respect 5-octade-
cenoic acid is an important fatty acid of seed triglycerides of Dioscoreophyllum cumminsii (HEG-
NAUER, 1969, 1973), and occurrence of the cyanogenic glucoside taxiphyllin in Stephania japonica
(unpublished observation) should be mentioned. The rare cis-5-octadecenoic acid is one of the
main fatty acids of seed oils in the Ranunculaceous genera Aquilegia and Thalictrum, and taxi-
phyllin and biogenetically related tyrosine-derived glycosides are the usual cyanogenic compounds
of Gymnosperms, Monocots, and Polycarpicae. A cyanide group is also present in the non-cyano-
genic glucoside menisdaurin of Menispermum dauricum; this type of compound, however, seems
to be rather erratically distributed in Angiosperms.

References: HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 73-95, 428—431, 456; ibid.
6 (1973) 783; suppl. in prep.; Stwon, A pharmacognostic study of some Indonesian medicinal
plants of the family Menispermaceae; Thesis Univ. Leiden (1982); THORNBER, Phytochemistry 9
(1970) 157—187. — R. HEGNAUER.

Uses. From the paragraph on phytochemistry it appears that the family abounds in species
with many different alkaloids and HeyNe (1927), BurK«1 (1935) and QuisUMBING (1951) have
mentioned that many species are used medicinally. Japanese chemists have been very interested
in the past; also in the neotropics much research has been performed, e.g. by KRUKOFF c.s. Notes
on these uses, for all kinds of illnesses, external and internal, have been recorded here under the
following species (see there):

Anamirta cocculus (L.) W. & A. Limacia oblonga Hook. f. & TH.
Cissampelos pareira L. Pericampylus glaucus (LAMK) MERR.
Cocculus trilobus DC. Stephania capitata (BL.) SPRENG.
Coscinium fenestratum (GAERTN.) COLEBR. Tinospora crispa (L.) Hook. f. & Tu.
Fibraurea tinctoria Lour. Tinospora glabra (BuRM. f.) MERR.

Minor uses mentioned are: extracting yellow dye from the plants, use as a fish-poison, use of

166 FLORA MALESIANA [ser. I, vol. 102

the stems for basketry, making belts, etc. Tiliacora triandra is used as a flavouring in cooking in
Thailand. Fruit of Anamirta cocculus was in the past century extensively used in the adulteration
of beer. Of a few species the fruits are edible, e.g. of Albertisia papuana, Limacia oblonga.

References: BuRKILL, Dictionary of the economic products of the Malay Peninsula (1935);
Heyne, Nuttige Planten van Nederlandsch-Indié (1927); QuisumBiING, Medicinal plants of the
Philippines (1951).

Possible confusion with other families has occurred occasionally when specimens are in
the sterile state, even with Liliaceae. See under excluded names.

Confusion could occur with Aristolochia (Aristolochiaceae), the leaves of which have similar
venation, but they do not produce an abscission layer and wither away without leaving a leaf-scar
(cf. Dinc Hou, Blumea 29, 1983, 22).

According to SLEUMER (FI. Mal. I, 7, 1971, 80) the leaves of Phytocrene (Icacinaceae) are simi-
lar in shape and venation to those of various genera of Menispermaceae; also the petiole emerges
from a shallow cup-like thickening of the stem which also often occurs in Menispermaceae. In
the latter, however, the petiole is generally swollen in the uppermost and maybe also in its basal
part, which is never the case in Phytocrene.

Confusion could occur with Passifloraceae, which have, however, tendrils; the leaves in Adenia
possess in addition 2 large basal glands on the leaves. Cucurbitaceae have also tendrils. Confusion
could also occur with Dioscorea, but the leaves in this genus have mostly distinctly trabeculate
cross-venation. Sabia (Sabiaceae) has pinnate venation, which is rare in Menispermaceae.

More difficult are Miquelia (Icacinaceae) and Erythropalum (Olacaceae); both have tripli-
nerved-pinnate venation. Cardiopteris (Cardiopteraceae or Icacinaceae) has however very similar
leaves.

Hints to Collectors. Since the plants of this family are always dioecious, it is necessary
to search for both male and female individuals, which may be quite distant from one another.
Female inflorescences are often fewer-flowered than the male, which makes them less conspi-
cuous. Fruits are very important, especially for the endocarp characters.

When the inflorescences are on leafless stems, make certain that the foliage being collected real-
ly comes from the same climber and not from another growing with it. Note any uncertainty in
this respect.

The colour of the wood in the stems of the bigger lianes should be noted as well as the presence
and colour of any latex or sap.

Attention should be given to the underground parts; presence or absence of tubers or stolons
should be recorded.

Spot-characters to aid identification of Malesian Menispermaceae
I. Vegetative characters

Erect tree or shrub: Cocculus laurifolius.

Wood yellow: Arcangelisia, Coscinium, Fibraurea.

Stems strongly tuberculate: Tinospora crispa.

Bark detaching as a parchment-like layer: Tinospora spp.

Bark pale contrasting with blackish petioles: Fibraurea (often).

White latex present: Fibraurea, Tinomiscium (evident in herbarium specimens when any part is
broken).

Leaves very finely and closely striate on upper surface: Tinomiscium.

Leaves whitish tomentellous beneath, large: Coscinium.

Domatia (pocket or aperturate) or glandular patches in basal and/or main nerve axils, leaves lar-
ge: Anamirta, Arcangelisia (upper surface of leaf papillose over insertion of petiole), Tinospo-

ra spp.
Petiole rugulose, leaves narrow: Tiliacora.

1986] MENISPERMACEAE (Forman) 167

Nervation penninerved (base with lowermost nerves sometimes crowded but not clearly 3-nerved):
Albertisia, Carronia, Macrococculus and Pycnarrhena.

Il. Flowering characters

Inflorescence a fascicle: A/bertisia, Macrococculus (on old wood), Pycnarrhena.

Inflorescence pseudoracemose: Pachygone, Sarcopetalum, Tiliacora, Tinomiscium (red-hairy),
Tinospora spp.

Umbelliform cymes and peltate leaves: Stephania spp.

Flowers sessile and crowded on a fleshy disciform head: Stephania capitata and S. dictyoneura.

Inner sepals connate in a thick fleshy tube: A/bertisia.

Inner sepals valvate: Carronia, Limacia, Tiliacora.

Petals larger than sepals: Sarcopetalum.

Stamens completely united into a peltate synandrium: Cissampelos, Cyclea, Parabaena,
Stephania.

Stamens with filaments + connate: A/bertisia, Anamirta, Arcangelisia, Coscinium (inner only),
Hypserpa spp., Macrococculus (outer only), Pycnarrhena, Sarcopetalum.

Female perianth zygomorphic: Cissampelos, Stephania spp.

Ill. Fruit characters

Carpophore bearing 3—9 drupes with a short branch below each drupe: Anamirta, Tiliacora.

Carpophore clavate up to 4 cm long, bearing 3 transversely subovoid drupes over 2 cm long: Arc-
angelisia.

Drupe narrowed at the base into a stipe: Haematocarpus, Macrococculus, Tinomiscium.

Drupes stalked and crowded, many arising from a disciform capitulum: Stephania capitata and
probably S. dictyoneura (fruits unknown).

Drupe with style-scar terminal (Tinosporeae): Chlaenandra, Coscinium, Fibraurea, Tinomis-
cium, Tinospora.

Drupes subglobose, pale-tomentellous, 2.8—3 cm @: Coscinium.

Endocarp globose, c. 4 cm @, densely covered with branched spines: Chlaenandra.

Endosperm absent: Carronia, Haematocarpus, Macrococculus, Pachygone, Tiliacora.

Notes on using the generic keys. In addition to the strictly contrasting characters, some
further characters are included in the male and female keys as an aid to identification.

Fruits are necessary when using the female key. Some details of the endocarp, including the
shape of the seed-cavity, are sometimes evident from the external appearance of the dried drupe
when the pericarp is thin and dries close to the endocarp. In other cases it is necessary to expose
at least part of the endocarp.

It may be necessary to cut the drupe in half, usually transversely in relation to the base, to see
details of the condyle, endosperm and cotyledons.

The condyle is a ventral development of the endocarp which intrudes into the seed-cavity or
forms a ventral cavity.

168 FLORA MALESIANA [ser. I, vol. 102

SCIENTIFIC KEY TO THE GENERA

1. Seeds without endosperm.

2. Condyle obsolete or septiform. TILIACOREAE (except Tiliacora).
3. Condyle obsolete. Seed straight, broadly ellipsoidal. Stamens often more than 6 and + connate.
4. Drupe with style-scar below the apparent apex. Sepals all free.

5. Drupes not narrowed at base into a stipe, 3—6 on a + globose carpophore. Endocarp usually papyra-
ceous to crustaceous and + smooth, sometimes thick, bony and verrucose. Stamens 6—18, usually
connate for most of length; connectivenot prolonged’....:°:..:.....--.-....+. 1. Pycnarrhena

. Drupes narrowed at base into a stipe, 5—10 radiating from the margin of a disciform carpophore.
Endocarp thick and bony, dorsally covered with a coarse open network of ridges, ventrally smooth.
Stamens 9, 3 inner free surrounded by 6 slightly joined at base; connective prolonged into a tongue

2. Macrococculus
4. Drupe with style-scar near base. Inner sepals connate in a fleshy tubular calyx. Stamens c. 18—24, con-

a

Natesin ayCOMIGAINGH ISL Cleary cere ces sre ere oct Pe each ec Aid ah caer a ia, 3. Albertisia

3. Condyle an elongate septum around which the seed is sharply bent. Stamens 6, free.
Gs InneL ScpalsnvalVvate caves: PEMnInNenVedinncmrs tasters cients, is o/4 ceo nie sve ela os eee eee 4. Carronia
Gu Sepalsialimmonicateslsecavesithipimenvedsmrerr etre isciccn sche ee serie eicins 5. Haematocarpus

2. Condyle hollow with a central aperture on both sides of endocarp. (MENISPERMEAE p.p.) 15. Pachygone
1. Seeds with endosperm.
7. Drupe with style-scar terminal. Embryo with foliaceous cotyledons. TrnosporEAE (incl. Fibraureeae).
8. Petals 3 or 6.
9. Stamens free.
10. Endocarp globose, covered with series of branched spines. Condyle clavate, 2-chambered, deeply in-
truding into the seed cavity. Stamens with flattened elliptic filaments broader than the anthers
7. Chlaenandra

10. Endocarp not spiny. Stamens with + terete filaments not broader than the anthers.

11. Endocarp knobbly, papillose or smooth with a ventral aperture or elongate depression; condyle
sometimes forming a ventral cavity. Endosperm usually ventrally ruminate. Plant without a latici-
hEKOustSyStem=alseavesmnOtestniaterabOvers esa ase] oo haere nee one one tile ee 8. Tinospora

11. Endocarp rugose to rugulose, strongly compressed dorsiventrally without a ventral aperture or de-
pression; condyle obsolete. Endosperm not ruminate. A laticiferous system present throughout the

Plant. Weeaves tinelysandicloselyasiniatesaDOVe: cv ecr ec. Wels om cie eer etal 10. Tinomiscium
9. Stamens united in a peltate synandrium. Endocarp spiny, the condyle forming a ventral concavity or an
enclosedmnflatedsch amb eles cuarertaee at ous oi ays oc Paes house shes) Ss aus Fetes cic, Suc col ene etsacness eo, Ree 9. Parabaena

8. Petals 0. Endocarp subellipsoidal, smooth with condyle forming a narrow ventral groove 11. Fibraurea
7. Drupe with style-scar near base or on one side.
12. Embryo with foliaceous, divaricate cotyledons. Seed broadly ellipsoidal or deeply cup-shaped. Petals 0.
COSCINEAE.
13. Condyle obsolete. Seed broadly ellipsoidal. Carpophore clavate. Anthers 9—12 in a sessile cluster
12. Arcangelisia
13. Condyle deeply intruding into the seed-cavity. Seed deeply cup-shaped.
14. Condyle deeply 2-lobed, hollow. Carpophore shortly branched below the drupes. Anthers 30—35 ina
Stalkediclistenslmt Onescencennyampaniclestor 1s araei se ei ekas ta tein) eer eee 13. Anamirta
14. Condyle + peltate, unlobed. Carpophore globose. Stamens 6, the outer 3 free with 1-locular introrse
anthers, the inner 3 with connate filaments and 2-locular latrorse anthers. Inflorescence a raceme of
PeaunclediGallssolenOwekSErerratte rntbrt racic cr mite cre cisions ictus «tasters 14. Coscinium
12. Embryo elongate with narrow, semi-terete or flattened cotyledons lying close together. Seed elongate,
strongly curved, + horseshoe-shaped. Petals present, at least in male flowers.
15. Endosperm ruminate. Condyle septiform (TILIACOREAE P.P.) .......-.0 eee cece eens 6. Tiliacora
15. Endosperm not ruminate. Condyle usually obovate in outline with the seed curved around its margin,
sometimes hollow and 1—2-chambered. MENISPERMEAE (except Pachygone).
16. Sepals spirally arranged, imbricate, variable in number, 7—12. Stamens 9—40, free or connate. Car-
BelSeZ Sa (INANE SS P75) apners i (cs apoxss colats have SOCAL ete Avice Pek a ee 16. Hypserpa
16. Sepals in whorls of 3 or in one united whorl.
17. Carpels 3—6. Stamens free or only filaments connate.
18. Inner sepals valvate. Condyle hollow, 2-chambered .................00.00ce00: 17. Limacia

1986] MENISPERMACEAE (Forman) 169

18. Inner sepals imbricate. Condyle + lamelliform.
19. Petals thick and fleshy, much larger than the sepals. Stamens with connate filaments
18. Sarcopetalum
19. Petals smaller than sepals or absent in female flowers. Stamens free.
20. Petals entire, sometimes absent in female flowers.
21. Condyle obovate to rotund in outline. Inflorescences axillary, cymes or thyrses.

22. Petals absent in female flowers. Anthers dehiscing with transverse + introrse slits, the cells
separated abaxially by the thickened connective. Endocarp with an entire dorsal median wing
or ridge and prominent lateral curved or cucullate crests bordering the condyle, the dorsal re-
pion smooth’ (Mal. spp"): . Saka eae: eee ae 19. Legnephora

22. Petals present in female flowers. Anthers dehiscing with longitudinal latrorse slits, the cells
parallel and close together. Endocarp without a dorsal median wing or ridge and lateral
crests, dorsally covered with short pointed processes ................. 20. Pericampylus

21. Condyle elongate, very narrow with the seed sharply bent around it. Inflorescences from old
woody stems (Mal. spp.), long racemes of cymes (Mal. spp.) ............. 21. Diploclisia

20. Petals bifid or emarginate. Endocarp dorsally verruculose or transversely ridged, with an aper-
ture on both lateral faces leading to a lateral cavity ........................ 22. Cocculus

17. Carpel 1. Stamens completely connate in a peltate synandrium with the anthers horizontal around
the margin.
23. Female inflorescences with accrescent bracts (Mal. sp.). Flowers 4-merous. Petals connate
23. Cissampelos
23. Female inflorescences without accrescent bracts.
24. Inflorescences thyrsoid or pseudoracemose. Condyle hollow, 1-chambered. Flowers 4-merous.

Petals*free*or ‘connaten inane ie eT A. ee ee eee 24. Cyclea
24. Inflorescences composed of umbelliform cymes or disciform capitula. Condyle lamelliform.
Flowers 3—4-merous! Petalsiires: es es Se ee re eee 25. Stephania

KEY TO THE GENERA BASED ON MALE PLANTS

1. Upper surface of leaves very finely and closely striate. Inflorescences racemose on leafless stems, usually
MMM MIMORSINETIFOSE: ao. 5.8 6 80k pa Sows dss ER ER UO ee ee 10. Tinomiscium
1. Upper surface of leaves not striate.
2. Leaves triplinerved, the main basal pair of nerves extending almost to the leaf-apex.
3. Woody climber. Sepals and petals marked with reddish lines or spots. Inflorescence a panicle or raceme
5. Haematocarpus
3. Erect shrub or small tree. Sepals and petals not marked with lines or spots. Inflorescence a cyme or short
RAerePaO PCP INES) | es aca. ARE et, Sets AR See. 22. Cocculus laurifolius
2. Leaves not triplinerved.
4. Inner 3 sepals connate in a fleshy tubular calyx, minute outer sepals free. Stamens c. 18—24, connate
peraneoital clusters . 633s ceh ca niet 5 hati. Gah ld ae Pee 3. Albertisia
4. Sepals free or if connate then calyx of one whorl only.
5. Petals thick and fleshy, much larger than sepals. Inflorescences pseudoracemose. Leaves peltate or sub-
Ta ly ea OE ORRIN RETR Gr ee ee 18. Sarcopetalum
5. Petals smaller than sepals or absent.
6. Sepals 7—12, spirally arranged, not in distinct whorls. Stamens 9—40, free or connate. Inflorescences
eymiose or thyroid |. vs vii we. LSB 1 he eee OE Ae eee ee 16. Hypserpa
6. Sepals in one or more distinct whorls.
7. Connective prolonged into a flattened tongue. Stamens 9, the outer 6 joined at base, the inner 3 free.
mulorescenices fasciculate on old stems .2 ...0 ses ee sa een ee eaten 2. Macrococculus
7. Stamens without a prolonged connective.
8. Stamens united or at least partly connate at the base, or only the inner 3 stamens connate.
9. Stamens completely united in a peltate or clavate synandrium, the anthers in a horizontal ring.
10. Inflorescences composed of umbelliform cymes or disciform capitula. Leaves peltate
25. Stephania
10, Inflorescences cymose, thyrsoid or pseudoracemose. Leaves peltate or not.

170 FLORA MALESIANA [ser. I, vol. 102

11. Inflorescence a peduncled, corymbose cyme. Sepals 4. Petals connate in a cup. Leaves peltate
23. Cissampelos

11. Inflorescences thyrsoid or pseudoracemose, or if cymose then sepals 6 in 2 whorls and petals 6.
12. Sepals 4(—5), free or connate. Petals 4, free or connate. Inflorescences thyrsoid or pseudorace-

mose, Leaves sometimes: peltatesays-e in eee eee eee ee oe 24. Cyclea
12. Sepals 6, free in 2 whorls. Petals 6, free. Inflorescences thyrsoid or cymose. Leaves not
peltate wear UR ee ee. A Oe ee 9, Parabaena

9. Stamens with the filaments + connate or only the inner stamens connate.
13. Inflorescence a raceme of peduncled balls of flowers. Stamens 6, the outer 3 free, the inner 3
joined. Leaves tomentellous (often whitish) below .......................- 14. Coscinium
13. Inflorescence a panicle, cyme or fascicle.
14. Lower surface of leaves with domatia in axils of main nerves. Leaves palmately nerved at base.
Inflorescence a panicle. Anthers in a globose cluster.

15. Flowers pedicellate. Anthers 30—35 in a stalked cluster................... 13. Anamirta
15. Flowers sessile or subsessile. Anthers 9—12 in a sessile cluster........... 12. Arcangelisia
14. Leaves lacking domatia, penninerved or subpalmately nerved at base. Inflorescence a cyme or
fascicle. Stamens 6—18 with the filaments + joined..................... 1. Pycnarrhena

8. Stamens all free.
16. Petals absent. Stamens with thick filaments, a prominent collar evident below the anther. Inflo-
Lescencerallaxapanicles WiOOdl yellows... ae noone oe a. aioe Sol ae se ee 11. Fibraurea
16. Petals present. Stamens without a prominent collar below the anther. Wood not yellow.
17. Inner 3 sepals valvate.
18. Sepals glabrous or subglabrous. Inflorescence a pseudoraceme. Petiole rugulose 6. Tiliacora
18. Sepals tomentellous to densely pilose. Petiole not rugulose.
OMintiorescenceiaicyme onlaxipseudopanicle Speer). a. seee ees eee 17. Limacia
19. Inflorescence narrowly thyrsoid, the branches subspicate................... 4. Carronia
17. Inner 3 sepals imbricate.
20. Main basal nerves and their main outer lateral branches extending to the leaf-margin.
21. Inflorescences on old leafless stems, long racemes of cymes.............. 21. Diploclisia
21. Inflorescences axillary.
22. Anthers dehiscing with longitudinal latrorse slits, the cells parallel and close together. Leaves

ustially i broadlyrand(shallowlytcrenates<.cse.aca0 = seen) eee 20. Pericampylus
22. Anthers dehiscing with transverse + introrse slits, the cells separated abaxially by the thick-
ened connective. Leaves usually entire or subentire ................... 19. Legnephora

20. Main basal nerves and their main outer lateral branches breaking up or anastomosing well with-
in the leaf-margin.
23. Inflorescence a large panicle up to c. 50 cm long with lateral branches up to 15 cm, borne on
old leafless stems. Stamens with flattened elliptic filaments broader than the anthers
7. Chlaenandra
23. Inflorescences much smaller and narrower, pseudoracemose, pseudospicate, narrowly thyr-
soid or narrowly pseudopaniculate, axillary or on leafless stems. Stamens with filaments not
broader than the anthers.
24. Petals bifid. Inflorescence a cyme or a raceme of cymes....... 22. Cocculus orbiculatus
24. Petals entire.
25. Inflorescences yellowish tomentose to pubescent, always axillary and pseudoracemose.

Leaf-base rounded to truncate or very obtuse (rarely subcordate)....... 15. Pachygone
25. Inflorescences glabrous, puberulous or hispidulous, axillary or on leafless stems, various in
form. eaf-baseoftencordatem. .n5 ehh. ee eee eae ie eee 8. Tinospora

KEY TO THE GENERA BASED ON FEMALE PLANTS

1. Upper surface of leaves very finely and closely striate. Inflorescences racemose on leafless stems, usually
LELEUCINEDTIS-1OMIENLOSEM otis Set fee coe il. eS a ee, Ue. See 10. Tinomiscium

1. Upper surface of leaves not striate.

2. Leaves triplinerved, the main basal pair of nerves extending almost to the leaf-apex.

1986] MENISPERMACEAE (Forman) 171

3. Woody climber. Drupes ellipsoidal, 40—45 mm long. Sepals and petals streaked with reddish lines or

BENGE es. Scie ss a mean views ayo ROK wines osicw O06 nee eee eale 5. Haematocarpus
3. Erect shrub or small tree. Drupes rotund in outline, c. 4 mm long. Sepals and petals not marked with
MAESIOL SPOUS Cree eee te ce re there, Ree SO PARED OL Regen Genet arnS 22. Cocculus laurifolius

2. Leaves not triplinerved.

4. Inflorescences with accrescent rotund bracts becoming up to 1.5 cm long and persisting at fruiting stage,
often obscuring the 5 mm long pubescent drupes. Leaves peltate, tomentose to puberulous. Slender
<TOT Ss eo ee Be Si TONES Te TPE Te et ON RT ASI, OCR a a 23. Cissampelos

4. Inflorescences without accrescent bracts.

5. Inflorescences composed of umbelliform cymes or disciform capitula. Drupe with style-scar near base,
4—11 mm long. Endocarp compressed, + obovate in outline with a sometimes perforate lamelliform
condyle surrounded by a horseshoe-shaped seed cavity, dorsally ornamented with projections of various
SemmtraCe(FATISVETSE TIG RES 2). (Ak LR Se SE aR ee 25. Stephania

5. Inflorescences not of umbelliform cymes or disciform capitula.

6. Seed and seed cavity straight or broad and concave or deeply cupular.
7. Seed and seed cavity deeply cup-shaped, surrounding the thickly clavate, sometimes 2-lobed, deeply
intrusive condyle. Drupes subglobose. Leaves palmately nerved at base.
8. Lower surface of leaves with domatia in axils of main nerves. Drupes 9-11 mm @ 13. Anamirta
8. Leaves without domatia. Drupes over 25 mm g.
9. Drupes 40—45 mm @, glabrous. Endocarp covered with branched spines. Leaves glabrous
7. Chlaenandra
9. Drupes 28—30 mm 9, tomentellous. Endocarp not spiny. Leaves tomentellous (usually whitish) on
fewer suriace’. 2°. f.0 2 .'. SBR Re, Se ee Roe Lee 14. Coscinium
7. Seed and seed cavity either straight, + ellipsoidal and endocarp with condyle absent, inconspicuous
or forming a longitudinal groove, or seed dorsiventrally flattened and concave and endocarp with
condyle forming a ventral cavity, concavity or inflated chamber.
10. Seed + ellipsoidal filled by large embryo with thick cotyledons, endosperm absent. Leaves penni-
nerved, sometimes with some subbasal nerves.

11. Drupes narrowed at base into a pronounced stipe, 5— 10 radiating from margin of a disciform car-
pophore. Endocarp 3—6 cm long, bony, covered dorsally with a coarse open network of ridges,
ventrally smooth. Inflorescences fasciculate on protuberances on old wood. Leaves densely re-
PHO. L BINS Ge GAL CRO PE A eS 2. Macrococculus

11. Drupes with base not narrowed into a stipe, 3—6 on a knob-like carpophore. Endocarp either
Papyraceous to crustaceous and + smooth to wrinkled or bony and verrucose.

12. Drupes subcompressed-ellipsoidal, tomentellous with style-scar near base, 2.5—6 cm long. Inner

sepais:connate into‘aifleshy tubes: 4. - 6: 2 AL eae oe ie 3. Albertisia
12. Drupes subglobose or broadly ellipsoidal, glabrous puberulous or tomentellous with style-scar
below the apparent apex, 0.8—3 cm long. Sepals all free.................. 1. Pycnarrhena

10. Seed largely filled with endosperm which surrounds the very thin embryo. Leaves palmately nerved
at base or with basal nerves diverging from midrib slightly above the base.

13. Lower surface of leaves with hollow domatia in axils of main nerves, leaf papillose on upper sur-

face around insertion of petiole. Drupes 2.2—5.5 cm long with style-scar on one side, borne on

a clavate carpophore up to 4 cm long. Endocarp covered with a fibrous layer, condyle incon-

SOON OF BONE ois Perce ne scared eh Kinin ated ele antec y seein ere 12. Arcangelisia

13. Lower surface of leaves without domatia or only present in basal nerve-axils. Drupes with termi-
nal style-scar.

14. Basal nerves of the leaf diverging from the mid-rib several mm above the base. Endocarp rather
smooth, subellipsoidal, 2.0—2.5 cm long, the condyle forming a ventral narrow longitudinal
groove. Inflorescence a lax panicle. Wood yellow ...........0seseeeeueeees 11. Fibraurea

14. Basal nerves arising from the base of the leaf. Endocarp variously ornamented or smooth, often
+ rotund in outline but sometimes subellipsoidal. Wood not yellow.

15. Endocarp spiny, the condyle forming a ventral concavity or an enclosed inflated chamber.
Badosperm not rowinate 750s Pe AY cab lets. p Perel aye bane te Raw sTy 9. Parabaena
15. Endocarp rugose, papillose or smooth with the condyle forming either a ventral cavity with a
central external aperture, or a ventral elongate depression. Endosperm usually ventrally rumi-
DN cccvcccosvcsaPacecti reve Terns a Hep ae cha ee esha a eore Meee eee nes 8. Tinospora

6. Seed and seed cavity elongate and strongly curved, semi-annular to horseshoe-shaped.

172 FLORA MALESIANA [ser. I, vol. 102

16. Drupes 6—7 mm long borne on a shortly branched 3—4 mm long carpophore. Carpels 8—9. Petioles

TUS OSE: «eo scionyis a Gi'v's es avons eon de deh aualole cfveae pteiaeloamaaes ob evens huloneeue aces erence eae 6. Tiliacora
16. Drupes not on a conspicuous carpophore. Carpels 1—6. Petioles not rugulose.
17. Leaves penninerved. Drupes and narrowly thyrsoid inflorescences tomentellous ... 4. Carronia

17. Leaves with base palmately 3—7-nerved. Drupes glabrous to pubescent.

18. Petals fleshy, larger than the sepals. Drupes semicircular in outline, (4—)6 mm long. Endocarp
ornamented dorsally with pointed processes and/or transverse ridges, laterally concave. Seed and
seed cavity semi-annular. Leaves subpeltate or peltate with petiole inserted up to 3 mm from basal
margin, fine reticulation prominent especially on lower surface. Woody stems verruculose

18. Sarcopetalum

18. Petals, if present, much smaller than the sepals. Seed and seed cavity mostly + horseshoe-shaped.

19. Inflorescences from old woody stems, consisting of very long racemes of cymes, up to c. 50 cm
long. Drupes narrowly obovate in outline, 15—30 mm long. Seed and seed cavity deeply horse-
shoe-shapeduwithsthe: 2) anmism==eparallelifa cc myuseye ayaa Qieetsiia stele) ala -2s 8 eee 21. Diploclisia
19. Inflorescences axillary. Drupes obovate to rotund in outline.
20. Endocarp with an entire dorsal median wing or ridge and laterally spreading curved or cucullate
crests bordering the condyle, the dorsal region smooth (Mal. spp.). Petals absent
19. Legnephora
20. Endocarp without a dorsal median wing or prominent ridge and lateral crests, the dorsal region
sometimes ornamented, e.g. verruculose, ridged or spinulate. Petals present.
21. Seed without endosperm, cotyledons large and thick, collateral. Endocarp rather smooth with
ausinalliicentralypertorationionibothilateralitaces# 2 eases a see eee 15. Pachygone
21. Seed with endosperm surrounding the narrow embryo.
22. Drupes 14—25 mm long. Dorsal region of endocarp smooth or only slightly rugose. Condyle
hollowe2-chambenredsdnnersepalsavalvaters. e. sareras o5clsiees foie > eee 17. Limacia
22. Drupes less than 12 mm long. Dorsal region of endocarp often ornamented. Condyle lamelli-
form or hollow. Inner sepals imbricate.

23. Leaves with main basal nerves and their main outer lateral branches leading directly to the
margin, which is usually broadly and shallowly crenate. Endocarp laterally concave, dorsal-
ly covered with short pointed processes, condyle lamelliform, imperforate

20. Pericampylus

23. Leaves with main nerves breaking up well within the entire margin. Endocarp with thick,
hollow condyle, often perforate.

24. Condyle 1-chambered with or without a small lateral perforation on each side. Endocarp
with a ventral perforation, dorsal surface with + papilliform processes and/or transverse
ridges in longitudinal rows. Drupes 5—8 mm long. Carpel 1. Sepals 2 or 3. Petals entire.
Weavestsometimeswpeltateyaan yee 6 of ner. ieee ne ae aoe 24. Cyclea

24. Condyle with 2 distinct lateral chambers each with a large lateral aperture. Endocarp not
ventrally perforate. Carpels (2—)3 or 6. Sepals 6—12. Leaves never peltate.

25. Drupes 6—12 mm long. Dorsal surface of endocarp rugose. Carpels (2—)3. Sepals spiral,
(Om ectalsventiner. ioe, setts aa Seine he toe ek Sa 1 Re ee 16. Hypserpa
25. Drupes 4—5 mm long. Dorsal surface of endocarp closely and irregularly wrinkled-
papillose. Carpels 6. Sepals 6(—9) in 2(—3) whorls. Petals bifid 22. Cocculus orbiculatus

1. PYCNARRHENA

Miers [Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 44, nomen] ex Hook.f. & TH. FI.
Ind. 1 (1855) 206; Mrers, Ann. Mag. Nat. Hist. ser. 3, 20 (1867) 11; Contr. Bot.
3 (1871) 351; Diets, Pfl. R. Heft 46 (1910) 48; Forman, Kew Bull. 26 (1972) 405;
ibid. 30 (1975) 97, 99; ibid. 33 (1979) 568. — Antitaxis Miers [Ann. Mag. Nat.
Hist. ser. 2, 7 (1851) 44, nomen], ibid. ser. 3, 20 (1867) 12; Contr. Bot. 3 (1871)
354, t. 142. — Batania Hatus. Mem. Fac. Agric. Kagoshima Univ. 5 (1966) 29.
— Fig. 1.

1986] MENISPERMACEAE (Forman) 173

Lianes or scandent shrubs. Leaves with petioles swollen at the base and usual-
ly at apex, leaving a prominent cup-like scar on the stem; lamina usually penni-
nerved and + elliptic, sometimes (in Mal., P. insignis) with lower nerves crowd-
ed at base and ovate. /nflorescences axillary or ramiflorous, mostly cymose, the
peduncles sometimes 1-flowered and fascicled. — Male flowers: sepals 6—15 in
whorls of 3, imbricate, the outer ones minute, inner ones largest (in P. ozantha
the innermost sometimes smaller), rotund and concave; petals 3—6, or 0 in P.
ozantha, minute, mostly broadly obovate; stamens variable in number, 6—18,
tightly clustered with the filaments usually connate for most of their length
(shortly connate in P. /ucida). — Female flowers: sepals and petals as in male;
staminodes 0; carpels 3—6 with recurved stigmas. Drupes + subglobose with
style-scar on the ventral side below the apex, arising from the margin of a swol-
len + globose carpophore; endocarp usually thin, papyraceous to crustaceous
and smooth to wrinkled, but in P. ozantha bony and verrucose; endosperm ab-
sent; seed broadly ellipsoidal, cotyledons large and thick, very slightly curved.

Distr. S. China and SE. Asia throughout Malesia to Queensland; 9 spp.

Notes. The genus Pycnarrhena has proved very difficult to revise owing to the inadequacy of the material
available. Of the nine species recognized, the fruits of one are unknown, while the flowers (of both sexes) and
inflorescences are unknown for another. In addition, female flowers are unknown for four species. Since
fruits and inflorescences provide the most useful characters for separating species of this genus, the lack of
some of this essential information is a major disadvantage in framing a taxonomic revision. Moreover, there
is considerable variation between specimens in their foliage, and without other correlated characters, it has
been most puzzling in certain instances to decide exactly where to draw the boundary between species. For
these reasons I have been unable to frame a practical key which will work in general for single specimens,
i.e. male or female, but the information incorporated in it will limit considerably the different possibilities
when naming.

KEY TO THE SPECIES

Sem emictte,. DASE DEOACIY COTUALE «rus, 0. 895 0,554 Si + + 2d « oie bln Eo eer te se eden Sete peas 1. P. insignis
1. Leaves + elliptic, base rounded to acute.
2. Petals present, clearly differentiated from the sepals. Carpels 3(—4). Endocarp crustaceous, smooth to
wrinkled.
3. Male inflorescences fasciculate, the peduncles mostly 1-flowered.
4. Drupes glabrous.
5. Male flowers laxly fascicled, pedicels 3—5(—10) mm; lamina 6—9(—12) by 2.5—4(—S.5) cm, drying +

flat, lateral nerves not markedly impressed on upper surface ............00es0 e000 2. P. lucida
5. Male flowers densely clustered, pedicels up to 2 mm; lamina 10—17 by 3.5—7 cm, drying subbullate,
main lateral nerves impressed on upper surface. Bengal, E. Himalaya, ?7N Burma... P. pleniflora
4. Drupes minutely tomentellous or puberulous. Male flowers on pedicels S—10 mm. Lamina 8—12(—18)
Naas Mme BOD TEN 00» a 5 oi gimse inst tapardseoe ¢ wErieek © it eee Ao aiaedien Brin ae gene es 3. P. longifolia

3. Male inflorescences cymose (female peduncles sometimes 1-flowered),
6. Drupes minutely puberulous. Leaves often markedly bullate ...............++5. 4. P. manillensis

6. Drupes glabrous. Leaves usually not bullate.
7. Drupes 8—10 mmo. Infructescences branched, slender, elongate up to7 cm long 5. P. novoguineensis
7. Drupes 15—30 mm 9, drying smooth (when ripe). Infructescences usually with rather thick peduncles,
usually unbranched and fascicled when axillary, but branched when ramiflorous . 6. P. tumefacta
2. Clearly differentiated petals absent (occasionally 1). Carpels 4-6. Drupes 12-20 mm @, drying shrivelled,
giaprous. Endocarp thick, Very Nard, VELrUcose « «+0406 4:4-0\¢.09-0.9)4 wn. Oe ts pid nee inmstnlens 7. P. ozantha

174

FLORA MALESIANA

[ser. I, vol. 102

1. Pycnarrhena insignis (HATUS.) FORMAN, Kew Bull.
33 (1979) 568. — Batania insignis Hatus. Mem. Fac.
Agric. Kagoshima Univ. 5 (1966) 29, 70, f. 1.

Scandent shrub c. 10 m. Young stems striate, glab-
rous, 2—5 mm @, bearing raised, discoid petiole-
scars; older stems rather smooth or obscurely wrin-
kled. Leaves with glabrous petioles (2—)3—4 cm,
broadened at the very base, geniculate and drying mi-
nutely wrinkled towards the apex; lamina ovate, base
broadly and shallowly cordate, apex shortly and ob-
tusely acuminate, 9—14.5 by 6.5—10.5 cm, lateral
nerves 6—8 pairs, the lowest 2—3 pairs arising close
together near the base, the midrib, lateral nerves and
conspicuous reticulate tertiary venation slightly
raised on both surfaces, glabrous, thinly coriaceous.
Flowers and inflorescences unknown. Infructescence
axillary, subterminal or arising from the axils of fal-
len leaves, cymose, c. 4cm long, at first minutely pu-
berulous, peduncles slender, S—10 mm, terminating
in a depressed-subglobose carpophore 6-7 mm @
bearing around its margin c. 6 drupes or their scars.
Drupes red, + broadly ellipsoidal, drying deeply
wrinkled, c. 2 cm long, glabrous, exocarp thinly co-
riaceous, mesocarp drying very thin, endocarp char-
taceous; embryo reniform, 9-10 mm long, cotyle-
dons very thick, slightly curved.

Distr. Malesia: Philippines (Batan I.: Mahatao).

Ecol. Thickets, 100 m.

Note. CuTLer, who examined the leaf-anatomy
in the genus (Kew Bull. 30, 1975, 41—48, pl.) exam-
ined the leaf-anatomy of the species and found that
it agrees in all essential characters with that of the ge-
nus.

2. Pycnarrhena lucida (TEUSM. & BINN.) Mig. Ann.
Mus. Bot. Lugd.-Bat. 4 (1868) 87; Becc. Malesia 1
(1877) 159; Boeri. Cat. Hort. Bog. 1 (1899) 43;
Diets, Pfl. R. Heft 46 (1910) 49; Back. Schoolfl.
(1911) 39; Koorp. Exk. Fl. Java 2 (1912) 230; YAama-
Moto, J. Soc. Trop. Agric. 16 (1944) 34; Back. &
BAKH. f. Fl. Java 1 (1963) 154; FORMAN, Kew Bull.
26 (1972) 408. — Cocculus lucidus TEsM. & BINN.
Nat. Tijd. N. I. 4 (1853) 397; Ned. Kruidk. Arch. 3
(1855) 394; Mig. Fl. Ind. Bat. 1, 2 (1858) 82. — Anti-
taxis fasciculata Miers [Ann. Mag. Nat. Hist. ser. 3,
20 (1867) 14, nomen], Contr. Bot. 3 (1871) 356;
Hook. f. & Tu. Fl. Br. India 1 (1872) 106; Kino, J.
As. Soc. Beng. 58, ii (1889) 388. — Antitaxis calocar-
pa Kurz, J. Bot. 13 (1875) 324; Fl. Burma 1 (1877)
57. — P. fasciculata (Miers) DrEts, Pfl. R. Heft 46
(1910) 50; Ri. Fl. Mal. Pen. 1 (1922) 108. — P. ca-
locarpa (Kurz) Drets, Pfl. R. Heft 46 (1910) 51.
Leaves with slender petioles, puberulous at first,
later glabrescent, 0.8—1.7 cm; lamina elliptic, apex
acuminate, base acute to obtuse, 6—9(—12) by 2.5—4
(—5.5) cm, main lateral nerves c. 6 pairs, reticulation
conspicuous and prominent on both surfaces, glab-

rous (rarely puberulous along midrib and main
nerves on lower surface). Inflorescences axillary or
ramiflorous, fasciculate; peduncles 1- (or few-) flo-
wered, 3—5(—10) mm, usually puberulous. — Male
flowers: sepals 9-12, outer ones minute, puberu-
lous, inner 3—6 larger, rotund to broadly rotund,
1.25 by 1.25—1.75 mm, glabrous; petals 3, cunei-
form, 0.5—0.75 mm long; stamens 6—12, 0.5 mm,
shortly connate. — Female flowers unknown.
Drupes red, globose, 8—15 mm @, glabrous; endo-
carp crustaceous.

Distr. Hainan, Cambodia, Thailand, Andaman
& Nicobar Is.; in Malesia: Malaya (Perak, Malacca),
Central W. Sumatra, W. Java (incl. P. Panaitan in
Sunda Strait).

Ecol. Forests, up to 500 m, in Casuarina equiseti-
folia stands near the shore on P. Panaitan, on lime-
stone in Peninsular Thailand.

Note. A single collection from W. Java (Koor-
DERS 26732) with flowers twice the normal size for
this species is the type of P. montana Back. In other
respects this collection resembles P. /ucida, but its
taxonomic status remains in some doubt. See at the
end of the genus.

3. Pycnarrhena longifolia (DECNE ex Miq.) BECcc.
Malesia 1 (1877) 160; Diets, Pfl. R. Heft 46 (1910)
52; FORMAN, Kew Bull. 26 (1972) 410. — Cocculus
longifolius DECNE ex Mig. Ann. Mus. Bot. Lugd.-
Bat. 4 (1868) 84. — Antitaxis longifolius (DECNE ex
Mia.) Miers, Contr. Bot. 3 (1871) 357. — Antitaxis
cauliflora Miers [Ann. Mag. Nat. Hist. ser. 3, 20
(1867) 14, nomen], Contr. Bot. 3 (1871) 356. — P.
cauliflora (MteERs) DiEts, Pfl. R. Heft 46 (1910) 52;
Back. Schoolfl. (1911) 40; Hat. f. Meded. Rijks-
herb. 12 (1912) 10; Koorp. Exk. Fl. Java 2 (1912)
230; YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 33;
Back. & Baku. f. Fl. Java 1 (1963) 154.

Leaves with petioles usually puberulous, 1.8—2.5
cm; lamina elliptic to narrowly elliptic to lanceolate-
elliptic, apex abruptly to gradually acuminate, base
obtuse to rounded, 8—12 (—18) by 3—6 (—8) cm, lat-
eral nerves 6—8 pairs, midrib often puberulous be-
low, surfaces otherwise glabrous. Inflorescences
ramiflorous, fasciculate, peduncles 1-flowered. —
Male flowers with minutely puberulous pedicels,
5—10 mm; outer sepals 2—4, minute; inner sepals yel-
low, 3, concave, rotund, 2 mm long, thick and fleshy
except towards margin; petals 3, oblate, 0.75—1 mm
long; stamens 7—12, tightly clustered, c. 0.5 mm
long. — Female flowers: outer sepals 1—2, minute,
puberulous; inner sepals yellow, 4—6, 2.25 mm long;
petals 3, + broadly obovate, 1.5—2 mm, glabrous,
thick; carpels 3—4, 1 mm long, subellipsoidal, style
short, recurved. Drupes borne on unbranched pu-
berulous to subglabrous peduncles 7—20 mm, glo-
bose, 12—15 mm g, minutely tomentellous or puber-

1986]

ulous; endocarp crustaceous.

Distr. Malesia: Central & S. Sumatra (also Sebesi
I. and P. Panaitan in Sunda Strait), Central Java,
Lesser Sunda Is. (Lombok, Sumba, Sumbawa, Flo-
res, Timor).

Ecol. Forests, up to 700 m.

Vern. Sumba: punduk.

4. Pycnarrhena manillensis Vmat, Rev. P|. Vasc. Fi-
lip. (1886) 45; Diets, Pfl. R. Heft 46 (1910) 52; in El-
mer, Leafl. Philip. Bot. 4(1911) 1163; Merr. Fl. Ma-
nila (1912) 203; W.H. Brown, Minor Prod. Philip.
For. 3 (1921) 186; Merr. En. Philip. 2 (1923) 144;
QuiBILAN & SAnTos, Nat. Appl. Sc. Bull. Philip. 3
(1933) 353; Quis. Medic. PI. Philip. (1951) 297; For-
MAN, Kew Bull. 26 (1972) 411; PANCHO, Vasc. Fl. Mt
Makiling 1 (1983) 275, f. 82. — P. elliptica DiE1s, Pfl.
R. Heft 46 (1910) 54; in Elmer, Leafl. Philip. Bot. 4
(1911) 1163; Merr. En. Philip. 2 (1923) 144; Philip.
J. Sc. 29 (1926) 367; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 33.

Leaves with puberulous to subglabrous petioles,
1—3 cm; lamina + elliptic to oblong-elliptic (some-
times narrowly oblong-elliptic in Mindanao), usually
+ bullate, apex acuminate, base obtuse to rounded
(or subcordate), (9—)11—18 by (2.5—)4—8 cm, lateral
nerves, c. 6 pairs, strongly impressed above and con-
spicuously joined near the margin, the fine reticula-
tion raised on both surfaces, upper surface glabrous,
beneath usually puberulous. — Male inflorescence
axillary, cymose, well-branched, many flowered,
2.5—4 cm long, puberulous. — Male flowers with pu-
berulous pedicels up to 3 mm; outer sepals 2—5, min-
ute, puberulous; inner sepals 4—7, rotund to broadly
elliptic, 1.5—2 mm long, glabrous or lightly puberu-
lous outside; petals 2—4, obovate, 0.75—1 mm long;
stamens 10—15, 0.75—1 mm long. — Female flowers
unknown. /nfructescence: peduncles usually bearing
1 to 3 fruits, axillary and very short, 0.5—1 cm, or
ramiflorous and up to 2 cm. Drupes globose or sub-
reniform, 1—1.5 cm long, minutely puberulous;
endocarp crustaceous.

Distr. Malesia: Philippines (Luzon, Panay, Sa-
mar, Leyte, Alabat, Negros, Mindanao, Basilan, Ca-
motes, Sulu Is.).

Ecol. Thickets and forests, up to 1600 m. FI.
May-—Nov., /r. July—March.

Uses. Infusions prepared from the roots and
powdered bark are used medicinally for a variety of
ailments. Various alkaloids have been extracted from
this species: see QuisumBING (1951, /.c.), BRUCH-
HaAuseN et al. (Arch. Pharm. 293, 1960, 454, 785) and
THornber (Phytochem. 9, 1970, 164, 172).

Vern. Philippines: ambal, mamangal, Tag., ha-
lnot, C. Bis., huluot, Bis.

Note. The anatomy of the leaves and stem have

MENISPERMACEAE (Forman)

175

been described by Drpasupm (Proc. 8th Pac. Sc.
Congr. 1957, 348—380).

5. Pycnarrhena novoguineensis Mig. Ann. Mus. Bot.
Lugd.-Bat. 4 (1868) 87; Becc. Malesia 1 (1877) 158;
Drts, Pfl. R. Heft 46 (1910) 54; KANEH. & HATus.
Bot. Mag. Tokyo 56 (1942) 472; FoRMAN, Kew Bull.
26 (1972) 412. — P. australiana F. v. M., Vict. Nat.
3 (1886) 61; F.M. BarLey, Queensl. FI. 1 (1899) 35. —
P. grandis K.Scu.& Laut. Fl. Deut. Schutzgeb. Sud-
see (1900) 313; Diets, Pfl. R. Heft 46 (1910) 54; Nova
Guinea 14 (1923) 80. — P. sayeri Diets, Pfl. R. Heft
46 (1910) 55.

Leaves with glabrous to sparsely puberulous peti-
oles 1.5—3.5 cm; lamina elliptic or oblong-elliptic,
apex acuminate, base obtuse to rounded, 18—28 by
4.5—8(—9.5) cm, lateral nerves 6—9 pairs, often im-
pressed above, venation obscure or prominent on
both surfaces. /nflorescences axillary or ramiflorous,
branched, 1.5—7 cm long, puberulous (the ramiflor-
ous inflorescences are larger and more branched than
the axillary ones). — Male flowers with pedicels up to
3 mm, or sessile; outer sepals 1—2, minute; inner sep-
als 3—6, + rotund, concave, 2—2.25 mm long; petals
3—4 obovate, 1 mm long; stamens 9—16, the cluster
c. 0.5 mm long. — Female flowers with pedicels, sep-
als and petals + as in male flowers; carpels 3, subel-
lipsoidal, 1 mm _ long. Jnfructescence slender,
branched, up to 7 cm, often rather elongate. Drupes
yellow, globose, 0.8—1 cm @, glabrous; endocarp
crustaceous.

Distr. Australia (Queensland); in Malesia: New
Guinea (in West common, in East only | coll.), incl.
Misool I.

Ecol. Forests at low altitude, up to 50 m. Accord-
ing to a field note the male flowers are heavily scent-
ed.

6. Pycnarrhena tumefacta Miers [Ann. Mag. Nat.
Hist. ser. 3, 20 (1867) 12, nomen], Contr. Bot. 3
(1871) 353; Becc. Malesia | (1877) 159; Diets, Pfl. R.
Heft 46 (1910) 53; Merr. En. Born. (1921) 248; Ya-
MAMOTO, J. Soc. Trop. Agric. 16 (1944) 34; FoRMAN,
Kew Bull. 26 (1972) 414. — Cocculus celebicus
Boer-. Cat. Hort. Bog. (1899) 40. — P. borneensis
Diets, Pfl. R. Heft 46 (1910) 53; Merr. En. Born.
(1921) 248; YaMAmoro, J. Soc. Trop. Agric. 16 (1944)
32. — P. celebica (Borrt.) Diets, Pfl. R. Heft 46
(1910) 53; Merr. Philip. J. Sc. 7 (1912) Bot. 266; En.
Philip. 2 (1923) 144; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 33. — P. merrillii Dievs, Pfl. R. Heft 46
(1910) 52; in Elmer, Leafl. Philip. Bot. 4 (1911) 1163;
Merr. En. Philip. 2 (1923) 144. — P. membranifolia
Mere. Philip. J. Sc. 20 (1922) 382; En. Philip. 2
(1923) 144. — P. balabacensis YAMAmoro, Trans.
Nat. Hist. Soc. Taiwan 34 (1944) 224, t. 1. — P. bata-
nensis YAMAMOTO, /.c. 226, t. 3. — P. castanopsidifo-

176 FLORA MALESIANA [ser. I, vol. 102

Fig. 1. Pycnarrhena tumefacta MIERs. a. Habit, x 2/3, b. male inflorescences, xX 2/3, c.. male flower, x 8,

d. inner sepal, e. petal, f. stamens, all x 10, g. female flower, front sepals and petals removed, x 8, A. part

of infructescence, r. TS of drupe, j. part of endocarp, k. seed, all nat. size. — Pp. ozantha Drts. |. Drupe,

m. endocarp, n. TS of drupe, all nat. size (a—b RSNB 4633, c—f DE VoGEL 4449, g RSNB 4436, h—-k SAN
9413, /-n NGF 24162).

1986]

lia YAMAMOTO, /.c. t. 2 (‘castanopsisifolia’). — Fig.
la—k.

Leaves with glabrous to tomentellous petioles,
1—6 cm, sometimes markedly swollen and geniculate
at apex; lamina narrowly to broadly elliptic, oblong-
elliptic or oblanceolate-elliptic, apex acuminate, base
rounded, obtuse or acute, 10—32 by 3.5—14 cm,
main lateral nerves 6—9 pairs, venation often promi-
nent on both surfaces, glabrous apart from midrib
on lower surface which is sometimes puberulous, es-
pecially near the base. — Male inflorescences axillary
or ramiflorous, cymose, few to several clustered to-
gether, 1—3 cm long, puberulous. — Male flowers
sessile or on puberulous pedicels up to 3 mm; sepals
pale green to white or yellow, outer sepals 2—4, mi-
nute, often puberulous; inner sepals 4—6, + rotund,
concave, 1.5—2.25 mm long, glabrous; petals 3—6,
+ broadly obovate 0.75—1.5 mm long; stamens
6—18 in a cluster c. 0.75 mm long. — Female inflo-
rescences similar to male but less branched, or un-
branched (i.e. 1-flowered peduncles). — Female
flowers (from RSNB 4436, Mt Kinabalu) on puberu-
lous pedicels up to 15 mm long; outer sepals 3,
broadly triangular, 1 mm long; inner sepals 4, +
broadly elliptic, rather thick, 2.5—3.5 mm long; pet-
als 3, broadly obovate, thin, 2 mm long; carpels 3—4,
subellipsoidal, 1 mm long, stigma recurved. (One im-
mature female flower of JAHERI 1679 had 6 inner sep-
als, 2 petals and 4 carpels.) Jnfructescence when axil-
lary usually fasciculate and peduncles unbranched,
peduncles rather thick, 1—3 cm; when ramiflorous
then branched and up to 7 cm. Drupes yellow to red,
globose to broadly subellipsoidal, 1.5—3 cm long,
glabrous; endocarp crustaceous, finely wrinkled.

Distr. Melanesia (Solomons); in Malesia: New
Guinea (West, ?East), S. Moluccas (Kei Is., Halma-
hera), N. Moluccas (Obi I.), Celebes (Sangihe Is.),
Philippines (Mindanao, Mindoro, Luzon, Palawan),
Borneo (incl. Banguey I.), and Kangean Is. (Kaju
Waru and Sipandjang Is.) off E. Java.

Ecol. Forests and thickets, up to 400 m, around
Mt Kinabalu in N. Borneo at 1200—2000 m. The sub-
stratum in N. Borneo was defined as loam, sandstone
and limestone. In Sarawak in peat-swamp forest. In
Halmahera in disturbed primary forest on dry hill-
side with loose porous clayey soil. In N. Moluccas
(Obi I.) on transition between coral sand beach and
red porous nickel soil. Fl. Jan.—Sept., Dec., fr.
Jan.—Dec.

Vern. Sarawak: akah badi, aker tulang. Halma-
hera: ogumi besi. Mindanao: dulupat. W. New Gui-
nea: fekan.

Notes. Some rather distinctive forms are appar-
ent, which may later prove to be distinct species, but
with the incomplete material so far available they are
regarded here as all conspecific. Specimens from the
peat-swamp forests of Sarawak have large, coarsely

MENISPERMACEAE (Forman)

| ea

reticulate leaves, and those from Mt Kinabalu have
leaves with long petioles. In the Philippines, apart
from the more typical, larger-leaved form, there is
also a small-leaved plant exemplified by the synonym
P. castanopsidifolia.

The extent of branching of the inflorescences de-
pends on their position on the plant: those on
younger, leafy stems are less branched than those on
stems which are old and leafless. There is therefore
a considerable difference between simple, short, axil-
lary infructescences and those which are branched,
elongate and ramiflorous. Herbarium specimens
normally only show one or the other of these types.

The flowers are very fragrant.

The specimens cited in FORMAN (1972) from New
Hebrides, New Ireland and New Britain belong to P.
ozantha.

7. Pycnarrhena ozantha DicLs, Bot. Jahrb. 52 (1915)
187; FoRMAN, Kew Bull. 26 (1972) 416. — P. papua-
na KANEH. & Hatus. Bot. Mag. Tokyo 56 (1942)
473. — Fig. 1l—n.

Young stems and petioles puberulous, later gla-
brescent. Leaves with petioles (0.8—)1.2—2.3 cm; la-
mina mostly elliptic to oblong-elliptic, base rounded
to cuneate, apex acuminate, 9—22 by 3—8 cm, lateral
nerves 6—8 pairs, linking towards the margin, reticu-
lation prominent below, glabrous apart from midrib
below. Inflorescences axillary or ramiflorous, fasci-
culate (or few-flowered racemes teste Diets). — Male
flowers on puberulous pedicels 4—7 mm: sepals yel-
low, c. 10, glabrous, the larger inner ones rotund,
2—2.5 mm long, sometimes the innermost smaller;
petals absent (occasionally 1); stamens 9-15, 1 mm
tightly clustered. — Female flowers: sepals 12—15;
petals absent; carpels 4—6, 1 mm. Drupe yellow to
red, on slender 7—15 mm peduncle (incl. carpo-
phore), subglobose, 1.2—2 cm @, drying shrivelled
with the surface forming irregular ridges, glabrous;
endocarp bony, slightly curved, subreniform, bear-
ing c. 5 dorsal rows of warty protuberances.

Distr. New Hebrides (Vanuatu); in Malesia: New
Guinea, New Britain, New Ireland.

Ecol. Forests up to 1600 m. Among limestone
outcrops (New Ireland), on coral limestone (New He-
brides), on red clay-loam with occasional basic vol-
canic boulders (New Britain). F/. March, July, Sept.,
Dec., fr. Jan., March, Aug., Oct.

Vern. NE. New Guinea: dodadud, Bilia, aikel,
Amele, boak, Dumpu.

Notes. The leaves are fragrant when crushed
(Cuew RSNH 345) and the flowers are strongly
scented (RAYNAL& GILLISON RSNH 16425). This spe-
cies is notable in the genus for the lack usually of
clearly differentiated petals and for the hard, bony,
dorsally verrucose endocarp. The fruit wrongly de-
scribed for this species in ForMAN, Kew Bull. 30

178

FLORA MALESIANA

[ser. I, vol. 102

(1975) 98 was part of a mixed collection and repre-
sents Macrococculus pomiferus BECC.

The alkaloids of this species were investigated by
LoperR & NEARN in a study of tumour-inhibitory
plants (Austr. J. Chem. 25, 1972, 2193).

Uncertain

Pycnarrhena montana BAcK. [Bekn. Fl. Java (em.
ed.) fam. 34 (1941) p. 7, descr. neerl.|, Blumea 5

(1945) 493; Back. & Baku. /f. Fl. Java 1 (1963) 154.

The plant represented by the above type-collection
is possibly referable to P. lucida (see there) except
that the flowers are much larger: the larger interior
sepals 2.5 mm long; petals 3—4 (some laterally
joined), oblate, 1 by c. 1.5 mm. The type-collection
(KOORDERS 26732) with male flowers made in 1897 is
the only one known. In absence of further material
I prefer to regard P. montana as a species of uncer-
tain status.

2. MACROCOCCULUS

Becc. Malesia 1 (1877) 160; Dries, Pfl. R. Heft 46 (1910) 55, f. 18; Bot. Jahrb.
52 (1915) 187; Nova Guinea 14 (1923) 80; FoRMAN, Kew Bull. 26 (1972) 418. —
Fig. 2i—k.

Liane. Older branchlets producing conspicuous swollen cups around the
bases of the petioles. Leaves with petioles swollen at the base and apex, lamina
not or only slightly peltate, + ovate, penninerved. /nflorescences cauliflorous,
fasciculate. — Male flowers long-pedicellate; sepals in 5—6 whorls of 3 (i.e.
15—18 all together), imbricate; the outer 3—4 whorls minute, the inner 2—3
whorls much larger; petals 6, thick and fleshy; stamens 9 (a ring of 6 stamens
slightly joined at the base surrounding 3 inner ones); anther-cells distinct, de-
hiscing by lateral, longitudinal slits; connective prolonged into a flattened
tongue, filaments flattened. — Female flowers unknown; young infructescences
indicate S—10 carpels. Drupe large with style-scar lateral (adaxial) and inconspi-
cuous, subglobose, narrowed at base into a stipe, radiating from margin of
disciform carpophore; endocarp thinly bony, dorsally bearing a coarse network
of ridges, ventrally smooth. Seed broadly ellipsoidal, covered with a reticulate
membrane; endosperm absent; embryo straight, cotyledons very thick, partly
rugulose, radicle short, thick.

Distr. Malesia: New Guinea (incl. New Britain, New Ireland). Monotypic.

Notes. Beccaristated that the cotyledons are fused together into a solid mass from which the radicle is
not differentiated, and that in these respects Macrococculus differs from Pycnarrhena. In the fruit of CLEMENS
631, however, the cotyledons are free, and when split apart the small radicle is discernible lying between the
cotyledons at one end. Although Macrococculus appears to be very closely allied to Pycnarrhena, it is desir-
able, I feel, to maintain it as a distinct genus which can be distinguished especially by the stamens with their
distinctive tongue-like prolongations and well developed, flattened filaments. The large fruits with a thick and
bony endocarp, the more or less ovate, penninerved leaves and numerous sepals are further diagnostic features
of Macrococculus.

1. Macrococculus pomiferus Becc. Malesia 1 (1877)
161; Bot. Jahrb. 52 (1915) 187; Nova Guinea 14
(1923) 80; ForMAN, Kew Bull. 26 (1972) 418. — Fig.
2i-k.

Branchlets glabrous becoming rather pale and
mottled. Leaves with glabrous petioles 3—10 cm, in-

serted at the basal margin of the lamina (or 1—2 mm
from it); lamina ovate, lanceolate-ovate or broadly
elliptic, base slightly cordate to obtuse, apex acumi-
nate, 8—25 by 5—13 cm, lateral nerves 5—6 pairs,
both surfaces usually densely reticulate and gla-
brous, stiffly chartaceous to subcoriaceous. /nflores-

1986]

cences fasciculate, borne on protuberances on old
wood. — Male flowers yellow on slender puberulous
pedicels 1—4 cm; outermost sepals minute, scarcely 1
mm long, puberulous, larger inner 2 (or 3) whorls +
broadly elliptic, 2.5—3.5 mm long; petals 6, rotund
to cuneate, 1.5 mm long, thick and fleshy; stamens
3 mm long, the outer 6 in a ring with their flattened
filaments slightly joined at the base surrounding the
3 inner free stamens, the connective-appendages in-
curved. — Female flowers unknown but young in-
fructescences indicate S—10 carpels, pedicel becom-
ing thickened, 2.5—3.5 cm. Drupes red, globose, or

MENISPERMACEAE (Forman)

179

smooth; endocarp 3—6 cm g, dorsally bearing 3—5
raised longitudinal ridges which are connected by
anastomosing ridges forming a loose network.

Distr. Malesia: New Guinea (incl. New Britain,
New Ireland).

Ecol. Forests, from sea-level to 1000 m.

Uses. BEccCARI stated that the fruits are often
eaten by cassowaries.

Note. The fruits and infructescences described
under Pycnarrhena ozantha Diets in FORMAN, Kew
Bull. 30 (1975) 98 belong to M. pomiferus, the speci-
men being a mixture with leaves of Pycnarrhena.

subpyriform, up to 10 by 8 cm (BEccar)), glabrous,

3. ALBERTISIA

Becc. Malesia | (1877) 161; Dries, Pfl. R. Heft 46 (1910) 97, f. 35; ForMaAN,
Kew Bull. 30 (1975) 82; ibid. 39 (1984) 113. — Fig. 2a—h.

Lianes. Branchlets bearing prominent discoid petiole-scars. Leaves with peti-
oles conspicuously swollen at both ends; lamina + elliptic, penninerved with
1 —2 pairs of subbasal nerves (in Mal. spp.), the lateral nerves running alongside
the midrib for some mm before departing from it. — Male inflorescences axil-
lary or ramiflorous, cymose or fasciculate. — Male flowers with (1 or) 2 outer
whorls of 3 minute triangular sepals, the inner 3 sepals connate into a fleshy,
corolliform tube with a minute apical opening; petals 3 or 6 (or O in extra-Mal.
spp.), minute, fleshy; stamens c. 18—24, connate into a conical synandrium, an-
thers 2-celled, transversely dehiscent. — Female inflorescences mostly reduced
to solitary flowers. — Female flowers (only known in A. papuana) with sepals
and petals as in the male; staminodes 6; carpels 4—6(—12 in extra-Mal. spp.),
elongate-ovoid attenuated into a subulate style. Drupes radiating from the mar-
gin of a swollen, tomentellous carpophore terminating the peduncle, + ellip-
soidal, subcompressed, style-scar close to base, tomentellous; mesocarp gran-
ular (when dry); endocarp + ellipsoidal, crustaceous or thinly woody, surface
sublaevigate or slightly wrinkled. Seed straight or only slightly curved, + ellip-
soidal, without endosperm; cotyledons very thick.

Distr. 12 spp. in tropical and subtropical Africa; 5 spp. in Indo-Malesia, 3 of which in Malesia.

Notes. By merging the African genus Epinetrum the size and range of the genus was distinctly enlarged.

In the key to the Malesian species given below it should be noted that there were relatively few male flowers
available for dissection; they had either 3 or 6 petals. With further material this difference may not prove to
be constant.

KEY TO THE SPECIES

1. Lamina drying bullate, broadly elliptic to subobovate, 24—41 by 9—18 cm. Male flowers with 6 petals.
Co Oe eR PP Ors ee PEN ePrints Pe ed 1. A. crassa

1. Lamina not drying bullate, narrower in shape and smaller than above. Petals 3.

2. Drupes 5—6 cm long. Lamina 9—13(—15) by 3.5—4.5(—6) cm. Male flowers in fascicles 2. A. megacarpa

2. Drupes 2.5—3.8 cm long. Lamina generally 19-32 by 7—14 cm. Male flowers in axillary or ramiflorous
DEACIOD: a5 a's dics spp vin pia © odd vm.) bibl ole CME L TURN IO EEE oie otk cote ee Cee ee 3. A. papuana

180 FLORA MALESIANA [ser. I, vol. 102

k

Fig. 2. Albertisia papuana Brcc.a. Habit with infructescence, x 2/3, b. male inflorescences, x 2/3, c. male

flower with half of inner calyx removed, x4, d. female flower with half of inner calyx removed, x4, e. drupe

on carpophore, f. TS of drupe, g. endocarp, h. seed, all nat. size. — Macrococculus pomiferus Brcc. i. Male

flower, x4, j. stamens, x 12, k. dorsal view of anther, x 30 (a, e—h ForMAN6A, b—c FoRMAN 413, d FORMAN
6B, i-k NGF 20760).

1986]

1. Albertisia crassa FORMAN, Kew Bull. 30 (1975) 85.

Liane. Branchlets at first minutely tomentellous,
later glabrescent, smooth. Leaves with petioles
strongly thickened and tomentellous at base and
apex, otherwise subglabrous, 5—8(—17) cm; lamina
broadly elliptic or subovate, base cuneate (or round-
ed), apex abruptly and obtusely or long and very
acutely acuminate, 24—41 by 9-18 cm, lateral nerves
3—4 pairs with the inner subbasal pair ascending be-
yond the middle of the lamina, nerves and reticula-
tion prominent on both surfaces, strongly bullate,
glabrous, thinly coriaceous. /nflorescences appar-
ently axillary, otherwise unknown. — Male flowers
?fascicled; sepals: outer 3(+ 3) to 1.5 mm, tomentel-
lous, inner calyx yellow, 10 mm, externally tomentel-
lous; petals 6, thick and fleshy, broadly ovate, lateral
lobes inflexed, 1 mm; synandrium 4—5 mm bearing
c. 20 anthers. — Female flowers unknown. Infruc-
tescence a solitary peduncle in the axil of a leaf-scar.
Drupe yellow, oblong-ellipsoidal, c. 4 by 2.7 cm,
peduncle 4 mm g, 1.5 cm long (incl. carpophore);
endocarp 2.7—3 cm long, crustaceous with wall
scarcely 1 mm thick.

Distr. Malesia: Malay Peninsula (Pahang, Se-
langor), 2 coll.

Ecol. In forests, 200 m.

Vern. Akar cherat, Temuan.

2. Albertisia megacarpa DiELsS ex FORMAN, Kew
Bull. 30 (1975) 85.

Large liane. Branchlets puberulous, glabrescent,
smooth. Leaves with petioles swollen and sometimes
puberulous at base and apex, otherwise subglabrous,
2.5—4.5 cm; lamina elliptic or elliptic-lanceolate,
base cuneate, apex acuminate, 9—13(—15) by
3.5—4.5(—6) cm, main lateral nerves 4—S5 pairs,
nerves and reticulation prominent on both surfaces,
glabrous but midrib sometimes puberulous below to-
wards base, thinly coriaceous. — Male flowers in
few-flowered fascicles in the axils of leaf-scars; pedi-
cels tomentellous, 4-6 mm; sepals tomentellous,
outer 3(? + 3) triangular, | mm, inner calyx 5 mm;
petals 3, unguiculate-ovate, recurved, | mm; synan-
drium 4 mm bearing c. 18 anthers. — Female flowers
unknown. Drupe ellipsoidal, 5—6 by 3.5—4 cm, ped-
uncles 1—1.5 cm long, 2—3 mm @ in the axils of leaf-
scars on older branches; endocarp 3.5—4.7 by
1.8—2.3 cm, slightly rugose, with wall fibrous-
woody, 2 mm thick.

Distr. Malesia: Malay Peninsula (Pahang: Cam-
eron Highlands), 2 coll.

Ecol. Hill forest, 1500—1600 m.

MENISPERMACEAE (Forman)

181

3. Albertisia papuana Becc. Malesia | (1877) 162;
BoERL. Cat. Hort. Bog. 1 (1899) 44, incl. var. banca-
na BoERL. ef var. buruensis BOERL.; DrELs, Pfl. R.
Heft 46 (1910) 97, f. 13c—d & 35; FoRMAN, Kew Bull.
30 (1975) 87. — Fig. 2a—h.

Large liane. Branchlets at first tomentellous, later
glabrescent, rather smooth, drying blackish. Leaves:
petioles with tomentellous swellings at both ends,
otherwise subglabrous, 2—10 cm; lamina elliptic-
oblong or elliptic, base cuneate to broadly rounded,
apex acuminate, (11—)19—32 by (4—)7—14 cm, 1—2
pairs of subbasal nerves apart from 3—4 pairs of lat-
eral nerves, nerves and reticulation raised on both
surfaces, glabrous, upper surface often shining, thin-
ly coriaceous. /nflorescences axillary and on older,
leafless stems. — Male flowers in fascicles: pedicels
4—6 mm, tomentellous; outer sepals 3+3, 1.5 mm
long, tomentellous; inner calyx creamy white to yel-
low, 10 mm long, externally tomentellous; petals 3,
fleshy, oblate, scarcely 1 mm long, lateral margins
inflexed; synandrium 5 mm long consisting of 24 sta-
mens in 6 vertical rows of 4. — Female flowers soli-
tary: pedicel and outer sepals similar to male, inner
calyx 6 mm long; petals 6, + obovate, 1 mm long,
glabrous; staminodes subulate, 2 mm; carpels 4—6,
elongate-ovoid, 2.5 mm long, tomentellous, attenu-
ated into a | mm long, subulate, glabrous style.
Drupes orange, oblong-ellipsoidal to obovoid-ellip-
soidal, 2.7—3.8 by 2—2.6 cm, on peduncle c. 4 mm
thick and 10 mm long; endocarp with a marginal
raised ridge, 3 by 1.7 cm, smooth, crustaceous.

Distr. Thailand (Peninsular); in Malesia: N. Su-
matra, Banka, W. Java, Borneo (Sarawak, ?Sabah),
S. Celebes, Moluccas (Buru), New Guinea (incl. Aru
Te):

Ecol. Primary rain-forest at low altitude.

Uses. According to CHIN 2760 the plant is widely
cultivated in Barak Distr., Sarawak for the leaves
which are used as a flavour-enhancer in cooking; also
the fruits are edible.

Vern. Sarawak: bekai, Kenyah.

Notes. Apart from complete material collected
in New Guinea, isolated fruiting or sterile specimens
from a wide geographical range appear to belong to
this species. Fuller material is needed for confirma-
tion.

Two collections from Sabah (Kioss 19175 &
19179) are possibly A. papuana, but in their hairy
stems and leaves resemble A. mecistophylla (MIERS)
ForMAN from Assam.

182 FLORA MALESIANA [ser. I, vol. 102

.
a

a
a
Es
‘ee
- 3% 7
54 4

ok A

Fig. 3. Carronia thyrsiflora (Brecc.) Diets. a. Leaf with male inflorescence on young terminal part of shoot,

x 2/3, b. male bud with valvate inner sepals, x 10, c. inner sepal, x 15, d. male flower with sepals removed,

e. stamen adaxial view, f. ibid. abaxial view, all x 20, g. part of infructescence with drupes, x 2/3, h. endo-

carp, x2, i. LS of endocarp (white) showing curved seed (white), x 2. — Haematocarpus subpeltatus MERR.

Jj. Habit, male plant, x 2/3, k. male bud, x8, /. petal with stamen, adaxial view, m. ibid. side view showing

adaxial projection, both x20, n. drupe on carpophore, o. LS of endocarp (white) showing curved seed
(white), both x 2/3 (a—f NGF 17701, g—i NGF 97719, j—m BS 33750, n—o Ktoss 18690).

1986] MENISPERMACEAE (Forman) 183

4. CARRONIA

F. v. M. Fragm. 9 (1875) 171; Diets, Pfl. R. Heft 46 (1910) 75, f. 26; FoRMAN,
Kew Bull. 30 (1975) 94. — Bania Brecc. Malesia | (1877) 161. — Fig. 3a—i.

Lianes. Branchlets often bearing prominent discoid petiole-scars. Leaves with
lamina + elliptic, penninerved. J/nflorescences axillary, terminal or rami-
florous, pseudo-spicate or thyrsoid (composed of pseudo-spikes), the flowers in
congested clusters. — Male flowers: sepals 9—30 in whorls of 3, the inner ones
larger, subcoriaceous, externally densely pilose, internally glabrous, the inner-
most whorl(s) valvate; petals 6, minute, + unguiculate; stamens 6, free, anthers
introrse with vertical or oblique slits. — Female flowers: sepals and petals as in
male; staminodes 0; carpels 6, densely pilose; styles erect, or recurved, subulate,
glabrous. Drupes narrowed at base into a stipe or subsessile on gynophore, sub-
obovoid or elongate-obovoid with style-scar near the base; endocarp with or
without a dorsal ridge, surface knobbly, slightly rugose or almost smooth, con-
dyle forming a longitudinal septum; seed horseshoe-shaped, endosperm absent,
cotyledons thick, semi-cylindrical.

Distr. 2 spp. in Australia (New South Wales, Queensland) and 1 sp. in Malesia: New Guinea.

1. Carronia thyrsiflora (BEcc.) Diets, Pfl. R. Heft
46 (1910) 76, f. 26A—H; Forman, Kew Bull. 30
(1975) 96. — Bania thyrsiflora Becc. Malesia 1
(1877) 161; Wars. Bot. Jahrb. 18 (1893) 193; K. Scu.
& Laut. Fl. Deut. Schutzgeb. Siidsee (1900) 315. —
Fig. 3a—i.

Stems tomentellous at first, later glabrescent.
Leaves with tomentellous to glabrous petioles 3—5
cm; lamina elliptic to ovate-elliptic, base rounded to
slightly cordate, apex acuminate, 14—30 by 7—14cm,
main lateral nerves 7—8 pairs, the lowest 1—2 pairs
arising at the base, upper surface glabrous with
nerves usually drying impressed, lower surface softly
pubescent to glabrous, stiffly papyraceous. — Male
inflorescences ramiflorous or terminal, narrowly
thyrsoid up to 22 cm with lateral branches up to 3.5
cm, greyish-brown tomentellous. — Male flowers:

cream-white sepals densely pale greyish-brown pilose
outside, c. 9; innermost sepals valvate, broadly ellip-
tic, concave 1.5 mm long; petals 6, unguiculate-
ovate, 0.5 mm long, + equal; stamens 6, 0.75 mm
long, anthers dehiscing vertically. — Female inflo-
rescences similar to male, lateral branches less than
2 cm long. — Female flowers with c. 12 outer smaller
sepals, inner sepals ovate, 1.5 mm long; petals as in
male; carpels 6, gibbous-ovoid, | mm long, densely
pilose; style subulate, recurved, glabrous. Drupes
creamy grey to red borne on tomentellous stipes 2—4
mm long, subobovoid, 13—15 mm long, drying
smooth without a prominent dorsal ridge, tomentel-
lous; endocarp rather smooth with only a very weak
dorsal ridge.

Distr. Malesia: New Guinea.

Ecol. Forests at low altitudes.

5. HAEMATOCARPUS

Miers, Ann. Mag. Nat. Hist. ser. 3, 13 (1864) 124; ibid. ser. 3, 19 (1867) 194;
Contr. Bot. 3 (1871) 323, t. 134; Diets, Pfl. R. Heft 46 (1910) 56; FoRMAN, Kew
Bull. 26 (1972) 419; ibid. 30 (1975) 81. — Baterium Miers, Ann. Mag. Nat. Hist.
ser. 3, 13 (1864) 124. — Fibraureopsis YAMAmoto, J. Soc. Trop. Agric. 16
(1944) 91, t. 1, p.p.; FORMAN, Kew Bull. 30 (1975) 81. — Fig. 3j—o.

Lianes. Leaves not peltate or very slightly peltate; lamina + elliptic, tripli-
nerved. Inflorescences axillary, terminal or cauliflorous; paniculate or race-
mose. — Male flowers: sepals and petals streaked with dark red lines and spots;

184 FLORA MALESIANA [ser. I, vol. 102

sepals 12—15 in whorls of 3, imbricate, composed of 2 large inner whorls and
2—3 much smaller outer whorls; petals 6, the inner 3 auriculate at the base and
clasping the opposite stamen; stamens 6, free, connective enlarged, projecting
inwards (adaxially). — Female flowers: sepals and petals as in male flowers; sta-
minodes 6, minute; carpels 6, style reflexed. Drupe large (c. 4 cm long), oblong-
ellipsoidal, narrowed at the base into a stalk, style-scar near base; endocarp
smooth with a thin inner longitudinal septum (/.e. condyle) around which the
seed is bent double; endosperm absent; cotyledons long and thick, radicle short.

Distr. SE Asia (Assam, Bangladesh, Peninsular Thailand, Andaman Is.); in Malesia: Sumatra, W. Java,
Borneo, Celebes and the Philippines.

Notes. The scanty material of this genus is inadequate to decide with certainty how many species should
be recognized. It is difficult to assess the significance of some of the rather marked differences between speci-
mens. In view of the nature of the material available I feel that the broader view of HooKER f. & THOMSON

of species is preferable and I therefore recognize only two species in the genus.

Baterium was first reduced by DrEzs (1910).

KEY TO THE SPECIES

[al eavesmotineltaters ccuue cht sare ceysid Posse doactsy teseseds
1. Leaves peltate (petiole inserted c. 1 mm from basal margin of lamina).............

1. Haematocarpus validus (Miers) BAKH. f. ex For-
MAN [nom. illegit. in Back. & Baku. f. Fl. Java 1
(1963) 154], Kew Bull. 26 (1972) 420. — Fibraurea
haematocarpus Hook.f. & TH. FI. Ind. 1 (1855) 204.
— Baterium validum Mters, Ann. Mag. Nat. Hist.
ser. 3, 13 (1864) 124. — H. comptus Mirrs, ibid. ser.
3, 19 (1867) 197, nom. illegit.; Contr. Bot. 3 (1871)
326, t. 134; Diets, Pfl. R. Heft 46 (1910) 58, f. 54;
Back. Bekn. Fl. Java (em. ed.) 3 (1941) fam. 34: p.
8; YaMAmorTo, J. Soc. Trop. Agric. 16 (1944) 34. —
H. thomsonii Miers, Ann. Mag. Nat. Hist. ser. 3, 19
(1967) 197; Contr. Bot. 3 (1871) 325, t. 134; Hook.
f. & Tu. Fl. Br. India 1 (1872) 106; Drets, Pfl. R.
Heft 46 (1910) 58; KanmaLt & Das, Fl. Assam 1
(1934) 57. — H. incusus Miers, Contr. Bot. 3 (1871)
S27:

Branchlets slightly puberulous or glabrous.
Leaves: petioles 1.2—3.5 cm, glabrous, attached at
the margin of the lamina; lamina elliptic, slightly
ovate-elliptic or slightly obovate-elliptic, rounded or
obtuse at the base, acuminate (or rounded) at the
apex, 7—16 by 3—8.5 cm, nervation mostly promi-
nent, especially below. /nflorescences axillary, cauli-
florous (or terminal) paniculate or subracemose,
slightly puberulous or glabrous, up to 50 cm long
with lateral branches up to 9 cm. — Male flowers on
pedicels 2—4 mm; sepals, the outer ones minute (less
than 1 mm long), ciliate, the inner ones larger, ellip-
tic, 3 mm long, glabrous; petals + broadly elliptic,
1.5—2 mm long; stamens 1 mm long with the connec-
tive much swollen on the adaxial side. — Female

flowers: sepals and petals as in male flowers; stami-
nodes rod-like, 0.5 mm; carpels ovoid-ellipsoidal, 1
mm long, style sharply reflexed, about half the
length of the carpel. Drupe red, obovoid-ellipsoidal,
glabrous, 4.5 by 2.5 cm; embryo 3.5 cm long (mea-
surements from material in alcohol).

Distr. SE. Asia; in Malesia: Sumatra (Atjeh, 1
coll.; Simalur I., 2 coll.; S. Palembang), W. Java
(Priangan: Tjadasmalang, | coll.).

Ecol. In forests, 400—1200 m.

Vern. W. Java: areuy katyamang, S; Simalur L.:
olor palinggam dotan, olor sigalinggam, M.

Note. The very few known specimens of this spe-
cies show a considerable range of characters, espe-
cially amongst the Assam specimens, three of which
Miers regarded as three distinct species.

2. Haematocarpus subpeltatus Merr. Philip. J. Sc.
14 (1919) 383; En. Philip. 2 (1923) 145; FoRMAN, Kew
Bull. 26 (1972) 421. — Fibraurea chloroleuca (non
Miers) Merr. Un. Cal. Publ. Bot. 15 (1929) 59. —
Fibraureopsis smilacifolia YAMAMOTO, J. Soc. Trop.
Agric. 16 (1944) 91, t. 1, p.p. — Fibraurea elliptica
YAMAMOTO, Trans. Nat. Hist. Soc. Taiwan 34 (1944)
229, f. 4. — Fig. 3j—o.

Branchlets sparsely puberulous, glabrescent. Lea-
ves: petioles 1.2—3 cm, glabrous or subglabrous, at-
tached at c. 1 mm from the base of the lamina; lami-
na elliptic, subobovate-elliptic or ovate-lanceolate,
base rounded to cordate, acuminate at the apex,
8—16 by 4—7 cm, nervation prominent on both sur-

1986]

faces. Inflorescences axillary or cauliflorous, 4—20
cm, either laxly paniculate with flowers mostly sub-
sessile or racemose and few-flowered with pedicels
5—11 mn, sparsely puberulous or glabrous. — Male
flowers mostly subsessile; sepals ciliate, especially
the outermost minute ones, inner larger sepals ellip-
tic, 3.5—4.5 mm; petals + broadly elliptic, 3—3.5

MENISPERMACEAE (Forman)

185

known. Drupe red, oblong-ellipsoidal, glabrous, 4
by 1.5 cm; seed 2.7 cm long (measurements from
dried specimen).

Distr. Malesia: Sarawak (Kuching, 1 coll.), E.
Borneo (W. Kutai: Belajan R., 1 coll.), N. Borneo
(Tawao, Sandakan, 2 coll.), Celebes (NE. 1 coll., SE.
1 coll.), Philippines (Luzon, 2 coll.).

mm; stamens | mm, connective produced on adaxial Ecol. Forests, from sea-level to 100 m.

side into a thin vertical wing. — Female flowers un-

6. TILIACORA

CoLesr. Trans. Linn. Soc. 13 (1821) 53; Miers, Ann. Mag. Nat. Hist. ser. 3,
14 (1864) 252; Contr. Bot. 3 (1871) 75, t. 104; Hook. f. & Tu. Fl. Ind. (1855)
186; Fl. Br. India 1 (1872) 99; Drezs, Pfl. R. Heft 46 (1910) 59, f. 20—22; Trov-
PIN, Monogr. Menisp. afric. (1962) 47; FORMAN, Kew Bull. 30 (1975) 89; ibid.
37 (1982) 369. — Fig. 9i—m.

Lianes. Branchlets bearing prominent discoid petiole-scars. Leaves petiolate;
lamina pinnately nerved, often with steeply ascending basal nerves, thus subpal-
mately nerved at base. /nflorescences axillary or cauliflorous, pseudo-racemose,
composed of few-flowered, peduncled cymes or solitary flowers especially in fe-
male inflorescences. — Male flowers: sepals 6—12, the outermost smallest, the
3 innermost much larger and valvate (or subimbricate in Australian sp.); petals
3 or 6, minute; stamens 3—9, free (in Asia), anthers dehiscing with oblique or
longitudinal slits, introrse. — Female flowers: sepals and petals as in male;
staminodes absent; carpels (3—)8—12(—30 in Africa) inserted on a gynophore;
style recurved or stigma sessile. Drupes subobovoid, stipitate (borne on
branches of carpophore), remains of style near base; endocarp subobovoid with
a straight groove running up the middle of each side, condyle forming a longitu-
dinal septum, ornamented dorsally with transverse, branched ridges or almost
smooth; seed horseshoe-shaped, endosperm deeply ruminate (in Asia); cotyle-
dons elongate, flattened.

Distr. 19 species in tropical Africa and 2 in SE. Asia of which 1 in Malesia (Malaya), and | endemic in

N. Australia.

1. Tiliacora triandra (CoLeBR.) Diets, Pfl. R. Heft 46
(1910) 62; Ript. Fl. Mal. Pen. 1 (1922) 110; Burk.
Dict. (1935) 2161; GaGnep. Suppl. Fl. Gén. I.-C.
(1938) 130; Martin, Introd. Ethnobot. Cambodge
(1971) 62; Forman, Kew Bull. 30 (1975) 92. — Coc-
culus triandrus Co.esr. Trans. Linn. Soc. 13 (1821)
64. — Menispermum triandrum Roxs. (Hort. Beng.
(1814) 72, nomen}, Fl. Ind. ed. Carey 3 (1832) 816.
— Limacia triandra (Covesr.) Hook. f. & Tu. FI.
Ind. (1855) 188; Fl. Br. India 1 (1872) 100; Miers,
* Contr. Bot. 3 (1871) 112; Mig. FI. Ind. Bat. 1, 2
(1858) 80; Kurz, Fl. Burma (1877) 55; Kina, J. As.
Soc. Beng. 58, ii (1889) 382; GaGnep. Fl. Gén. I.-C.
1 (1908) 146. — Fig. 9i—m.

Stems puberulous to glabrous, striate. Leaves with
puberulous to glabrous, rugulose petioles 0.5—2 cm;
lamina elliptic, lanceolate or sometimes subovate,
base cuneate to rounded (to subcordate), apex acute
to obtuse, often acuminate, 6.5—11(—17) by
2—4(—8.5) cm, with 3—5 subpalmate basal nerves
apart from 2—6 pairs of lateral nerves, main nerves
tending to link up towards the margin, midrib on
lower surface rugulose near the base, glabrous, stiff-
ly papyraceous. /nflorescences axillary or cauliflor-
ous, pubescent, 2—8 (—17) cm long bearing 1—few-
flowered peduncled cymes c. 0.5 cm long. — Male
flowers yellow; inner sepals broadly elliptic 2 mm
long, subglabrous; petals 3 or 6, cuneate, emar-

186 FLORA MALESIANA [ser. I, vol. 10?

Fig. 4. Chlaenandra ovata Mig. a. Leaf, x 1/2, b. male inflorescence, X 1/2, c. male flower, x 6, d. petal,

x 12, e. stamen, x12. — Tinospora glabra (Burm. f.) Merr. f. Leaf, x 1/2, g. domatia at leaf base, x4,

h. young male inflorescence arising from old stem, x 1/2, i. male inflorescence, x 1/2, j. male flower, k. pet-

al, /. stamen, all x 6. — Parabaena megalocarpa Merr. m. Unlobed leaf with male inflorescence, x L/24ns

lobed leaf, x 1/2, o. male flower with 2 large sepals removed, x 6, p. petal, side view, x12 @ NGF 47855,
b—e ScHoppE 4443, f—g, i—/ ForMAN 418, h Jacoss 4769, m SAN 1428, n—p S 36012).

1986]

ginate, 1 mm long, glabrous; stamens 3, clavate,
1.5—2 mm long. — Female flowers: inner sepals orbi-
cular, 2 mm long, externally puberulous; petals 6,
oblong-elliptic, 1 mm long; carpels c. 8—9, less than
1 mm long, borne on short branches of a glabrous gy-
nophore; stigma sessile. Drupes red, borne on a car-
pophore 3—4 mm long with branches 2—3 mm long,
subcompressed, obovoid, 7-10 by 6—7 mn, gla-
brous; endocarp transversely and irregularly ridged.

Distr. S. Burma (once), Assam (Khasya), Thai-
land, Cambodia, Laos, Vietnam; in Malesia: Malaya
(Kedah, Trengganu; also in Penang & Langkawi Is.),
4 coll.

Ecol. In Thailand: limestone hills, evergreen for-
est near sea and also by the side of streams in scrub

MENISPERMACEAE (Forman)

187

jungle at low altitudes up to 200 m; in Vietnam: on
rocky and clayey soils up to 800 m; in Malaya: Kedah
Peak at 1300 m alt.

Uses. Leafy shoots mixed with other plants are
used in Cambodia for the preparation of a medicine
for dysentery (MARTIN, 1971). Used for cordage in
Vietnam. Widely used in Thailand as a flavouring in
cooking.

Note. Tiliacora triandra was recorded from
Fraser Hill (BuRKILL & Hottrum 8620) and Bukit
Kutu (RIDLEY s.n.) in Malaya by BurRKILL & HoLt-
TuM (Gard. Bull. S. S. 3, 1923, 34). These records
were based, however, on misidentified specimens of
Cyclea elegans KiNG in the Singapore Herbarium.

7. CHLAENANDRA

Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1863) 83; BEcc. Malesia | (1877) 144; DrELs,
Pfl. R. Heft 46 (1910) 131; FoRMAN, Kew Bull. 39 (1984) 114, f. IM—N. — Po-
rotheca K. ScH.in K. Sch. & Laut., Nachtr. Fl. Deut. Schutzgeb. Stidsee (1905)
263. — Fig. 4a—e, 5m—n.

Large woody climber. Leaves with petiole swollen and geniculate at base,
swollen at apex; lamina + ovate, entire, palmatinerved at base. /nflorescences
borne on old, leafless stems, paniculate. — Male flowers: sepals 12 in 4 whorls
of 3, the outermost whorls minute, the inner 2 whorls much larger and subequal;
petals 6; stamens 6 with broad flattened filaments much broader than the small
anthers. — Female inflorescences and flowers unknown. /nfructescences pani-
culate, borne on old leafless stems, carpophores very short. Drupes large; endo-
carp bony, densely covered with branched spines partly joined in longitudinal
rows; condyle deeply intrusive into the seed-cavity and containing a hollow
chamber subdivided by a thin median wall; seed deeply concave, curved around
the intrusive condyle, with abundant endosperm, cotyledons very broad, radicle
very short.

Distr. Malesia: New Guinea. Monotypic.

1. Chlaenandra ovata Mig. Ann. Mus. Bot. Lugd.-
Bat. 4 (1863) 84; Becc. Malesia 1 (1877) 144; Drezs,
Pfl. R. Heft 46 (1910) 131, f. 48; Renpve, J. Bot. 61,
Suppl. (1923) 4; Forman, Kew Bull. 39 (1984) 115, f.
IM-—N. — Tinospora megalobotrys Laut. & K. Scu.
Fl. Deut. Schutzgeb. Siidsee (1900) 311. — Porothe-
ca petiolata K. Scu. in K. Sch. & Laut., Nachtr. FI.
Deut. Schutzgeb. Siidsee (1905) 263. — Fig. 4a—e,
5m—n.

Large woody climber, entirely glabrous. Young
stems finely striate, bearing prominent discoid
petiole-scars; old stems covered with rough, ridged
bark. Leaves with petioles 5—9 cm, strongly thick-

ened and geniculate in basal 1—1.5 cm, also shortly
thickened at apex; lamina ovate, 12—19 by 7—12 cm,
base rounded or slightly cordate, apex acuminate,
palmately 3—5-nerved at base with 3—4 pairs of distal
lateral nerves which run alongside midrib for several
mm before departing from it, reticulation fine and
prominent on both surfaces, thinly coriaceous. —
Male inflorescences borne on old, leafless stems,
paniculate, up to c. 50 cm long with spreading lateral
branches up to 15 em. — Male flowers on pedicels up
to 5mm: sepals yellowish, 6 minute outermost sepals
less than | mm long, 6 main sepals broadly obovate
to oblate, concave, 2—2.5 mm long; petals 6, broadly

188

FLORA MALESIANA

[ser. I, vol. 102

elliptic, 1.25 mm long; stamens 6, 2 mm long, fila-
ments flattened, elliptic, much broader than the an-
thers. — Female inflorescences and flowers un-
known. I/nfructescences c. 50—60 cm long, lateral
branches up to 20 cm, fruit-peduncles 2—3 cm,
carpophores subdiscoid, 1—2 mm long. Drupes red-
dish with glaucous bloom, ovoid to globose, 5—6 cm
@ when fresh, 4—4.5 cm @ when dry; endocarp sub-
globose, c. 4 cm @, densely covered with branched
spines c. 5 mm long partly joined in longitudinal
rows, the hollow intrusive condyle with a ventral ex-
ternal opening bordered with spines, the seed-cavity

forming to the seed-cavity, ventrally deeply concave,
(according to BEccAr)) the dorsal surface irregular,
with copious uniform endosperm.

Distr. Malesia: New Guinea.

Ecol. Lowland swamp- and rain-forest, and
periodically flooded riverine forest, on sandy clay or
clay, up to 1200 m. Beccari stated that the fruits,
like the large fruits of Macrococculus pomiferus
Becc., are avidly eaten by cassowaries. F/. March—
April, Sept.; fr. July—Jan.

Vern. W. New Guinea: jebwoneer, Kebar; SE.
New Guinea: pfagabuba, Kutubu.

a hollow subhemisphere (/.e. cupular). Seed con-

8. TINOSPORA

Miers, Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 35; Hook. f. & TH. Fl. Ind. (1855)
182; Miers, Ann. Mag. Nat. Hist. ser. 3, 13 (1864) 315; Contr. Bot. 3 (1871)
30; Hook. f. & Tu. Fl. Br. India 1 (1872) 96; Diets, Pfl. R. Heft 46 (1910) 133,
f. 49; TRouPIN, Monogr. Menisp. afric. (1962) 191; Back. & BAKH./f. Fl. Java
1 (1963) 157; FORMAN, Kew Bull. 36 (1981) 379; ibid. 39 (1984) 112. — Hypsipo-
des Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 82. — Fawcettia F. v. M. Fragm.
10 (1877) 93; F.M. Batey, Queensl. Fl. 1 (1899) 29; Diets, Pfl. R. Heft 46
(1910) 133. — Fig. 4f-1, 6, 7.

Woody climbers. Stems with bark often becoming detached on drying, some-
times pergamentaceous. Leaves with petiole swollen and geniculate at base, la-
mina often + cordate, margin usually entire, occasionally dentate, rarely
3-lobed, palmatinerved mostly with 3—5 basal nerves and | —3 pairs of distal lat-
eral nerves, sometimes with domatia or glandular patches present in the basal
nerve-axils. Inflorescences thyrsoid, pseudopaniculate, pseudoracemose or
pseudospicate, in some species not coetaneous with the leaves. — Male flowers:
sepals usually free, rarely joined at base, 6 (occasionally 1—3 additional minute
outer ones present), outer 3 usually smaller, sometimes subequal, sepals + ellip-
tic, often membranous; petals 6, occasionally 3, often broadly cuneate-ovate
with the lateral edges inrolled, usually fleshy and often glandular-papillose ex-
ternally towards the unguiculate base; stamens 6 and free (in Asia). — Female
flowers: sepals and petals similar to male except petals often narrower; stami-
nodes 6, subulate; carpels 3, curved-ellipsoidal, stigma reflexed with short,
pointed lobes. Drupes borne ona short or columnar carpophore, style-scar ter-
minal; endocarp bony, dorsally convex and often verrucose or tuberculate, ven-
trally with central aperture leading to a cavity (the condyle) or with shallow
longitudinal groove; seed with endosperm usually ruminate.

Distr. An Old World genus of 32 species: 7 in tropical Africa, 2 in Madagascar, 23 in Asia to Australia
and the Pacific, throughout Malesia.

Ecol. Animportant study by Dr. H. BANziGER on fruit-piercing moths in Thailand (Mitt. Schweiz. Ento-
mol. Ges. 55, 1982, 213—240) has demonstrated the important role played by species of Tinospora in the bio-

1986] MENISPERMACEAE (Forman) 189

Fig. 5. Endocarps of Tinosporeae. a—b. Aspidocarya uvifera Hook.f. & Tu. a. dorsal and ventral views, b.

in CS; c-e. Parabaena megalocarpa Merk. c. ventral view, d. lateral view, e. in CS; f. P. sagittata Miers

ex Hook.f. & Tu. dorsal and ventral views; g. P. elmeri Duets, dorsal and ventral views; h. P. denudata Dies,

dorsal and ventral views; /—k. P. tuberculata Becc. j. dorsal and ventral views, k. in CS; /. P. echinocarpa

Deis, dorsal and ventral views; m—n. Chlaenandra ovata Mig. m. lateral abaxial view, n. in CS, All x3 (a—b

GamaB_e 9682, c—e Clemens 26222, f Gamace 9712, g Loner 1975, A BS 43053, /—~k NGBF 1065, / Etmer
14224, m—n Scunoppe 2428). Drawn by Mrs. M. Church, Courtesy Kew Bulletin.

190 FLORA MALESIANA [ser. I, vol. 102

logical chain which leads to extensive damage to certain fruit crops in Thailand. Longan (Dimocarpus longan
Lour.) and citrus including mandarin are the main crops damaged by noctuid Lepidoptera which feed on the
sap by piercing the skin of the fruits. In Thailand the moth mainly responsible is Othreis fullonia. Dr. BANz1-
GER has demonstrated that the chief host plants for the larval stage are 7. sinensis and T. crispa throughout
Thailand and 7. baenzigeri in Central, S. and NE. Thailand. Other species of Menispermaceae also act as
hosts for the larvae, especially in periods when these species of Tinospora are leafless during the dry season,
which is more prolonged in the north. These three species of 7inospora in Thailand are typical components
of secondary vegetation, where forest has been destroyed or disturbed. The exceptional capacity of these
climbers to survive damage and to regenerate from detached lengths of stem encourages their spread in these
disturbed habitats. It is therefore apparent that destruction of forests in Thailand leads to the spread of Tino-
spora, which in turn promotes the increase of noctuid moths, which damage the fruit crops.

Morph. There are various types of inflorescence in the genus and the flowers also vary appreciably. The
flowers are basically arranged in cymes, but these are sometimes reduced to single flowers, which can be ses-
sile. The inflorescences can be apparently paniculate, racemose or spiciform. A thyrse, consisting of a raceme
of lax cymes, occurs in 7. trilobata. A pseudopaniculate inflorescence occurs in 7. dentata and T. dissitiflora,
and apparently also in 7. arfakiana and T. hirsuta, where only infructescences are known.

The sepals of T. trilobata are unusual in being connate at the base, while in the other species they are com-
pletely free. Most species have unequal sepals, the outer whorl of three being smaller, but in 7. dentata, T.
homosepala, T. sumatrana, T. trilobata and sometimes in T. sagittata they are subequal. In T. tinosporoides,
T. trilobata (and occasionally in T. cordifolia and T. merrilliana) there are up to three minute sepals in an
additional outer whorl. The petals are generally 6 in number, opposite to the equal number of stamens, but
in 7. crispa only the outer whorl of three petals usually develops. Although minute in size, the petals do vary
in form between species but fortunately we do not have to rely on them in order to distinguish the species.

Phytochemistry. Bisset (Kew Bull. 36, 1981, 377; ibid. 39, 1984, 100) discussed the fairly numerous
researches on alkaloids and bitter substances found in the species, and provided a bibliography.

Uses. Various species are used for medicinal purpose; see for example under T. crispa.

Notes. Tinospora has proved particularly difficult to revise owing to the incompleteness of the material
of several species and the fact that some species flower when the plants are leafless.

If complete material were known for all the species, it should not be difficult to provide separate keys for
male flowering, female flowering and fruiting plants. Male flowers, female flowers or fruits are unknown for
some species, and it is therefore impossible at present to provide a key using only morphological characters
which would work for every single specimen, whether flowering or fruiting.

KEY 20 THE SPECIES

I eeIsCAVESIGeCpVeS=LODECE rae. he ee rele Romer ae tema to) GMs Bate eeteeus ohoinne Mates 1. T. trilobata
1. Leaves not deeply lobed, margin entire, toothed or only slightly lobed.
2 ILevismmencyin Meal ENANy Toyo nso (GMINVEIN)) eo peo oa uddsuondc dono pdddododoodoondss.c5- T. dentata
2. Leaf-margin entire or slightly lobed.

Sh Peay esnvery nanow.10)4—2scm Drodgda(Auisthalia)ieaciserite.s 2 sass. cle oie ae Sele ce omnes T. angusta

3. Leaves broader.
4. Leaves hairy (sometimes sparsely) beneath.
5. Weavesisapiitatestosnastate (Chinas, Vietiann)ie sacri: © scale ele ics «se seen eee T. sagittata
5. Leaves + ovate.
6. Leaves puberulous above, tomentellous or rather densely puberulous beneath (SE. Asia, S. China,
Hongkong phaina nm) toa nears s Sux cus G chine oe aidan the gs, oe ay ee Oe oie te ore eee T. sinensis
6. Leaves glabrous above.
7. Fine reticulation not or scarcely visible on both surfaces, which dry minutely wrinkled; uncinate
Hains absentetnommainunenvessbelledthiieir tatty eee ene ater -0 es 2. T. merrilliana
7. Fine reticulation raised on both surfaces; some uncinate hairs present on main nerves beneath
3. T. hirsuta
4. Leaves glabrous beneath.
8. Leaves narrowly elliptic to elliptic.
Dmikeat-bascisavittateyncncis.e wea. os eee ee eee Oo ant ele eee 4. T. celebica
85 Weatbase rounded iorsubtruticate:..).caswis heirs b Peni ea las eS Se a OE 5. T. glandulosa
8. Leaves + ovate, rotund or triangular.

1986]

MENISPERMACEAE (Forman)

191

10. Inflorescences appearing when plant is leafless.
11. Stems strongly tuberculate (tubercles may be little-developed on young stems). Leaves lacking hol-
low domatia in basal nerve-axils. Petals usually 3. Endocarp 11—13 mm long..... 6. T. crispa

11. Stems not tuberculate; petals 6.

12. Leaves with a pair of hollow domatia on lower surface in basal nerve-axils. Petals usually 6. Endo-

CA 1) MMIONE”. ws. See one ees ees

oan Se oa ate Can aes nen seine ne 7. T. baenzigeri

12. Leaves with glandular patches on lower surface in basal nerve-axils, + triangular with sides
straight or sometimes concave towards base. Endocarp shortly pointed at both ends (Australia)

T. smilacina

10. Inflorescences appearing together with leaves.
13. Leaves 18—28 by 11—23 cm, thinly coriaceous, lacking domatia or glandular patches in basal nerve-
axils. Endocarp 1.7—2.4 cm long, surface very smooth, whitish.............. 8. T. arfakiana
13. Leaves smaller and thinner, sometimes with domatia or glandular patches in basal nerve-axils.

Endocarps not as above.

14. Leaves drying matt and minutely wrinkled, fine reticulation not or scarcely visible on either sur-
face, main nerves usually impressed on upper surface. Petals unguiculate, apically subreniform
and markedly thickened. Endocarp thinly crustaceous, broadly elliptic in outline, 7-8 mm long,

surface smooth or obscurely tuberculate

14. Characters not combined as above.

Fog ahs Susy Sey REA Us teenth Se hs Be 2. T. merrilliana

15. Inflorescences with lateral branches, at least towards base. Leaves with very fine, raised reticula-
tion, glandular patches present in basal nerve-axils. Flower-pedicels 5S—10 mm. Endocarps 10—12

mm long, strongly tuberculate.........

15. Inflorescences unbranched.

Sere, a sista sie Og suerelers auers) sip se: Soot ereiane 9. T. dissitiflora

16. Flowers in sessile clusters; inflorescences sometimes zigzag. Endocarps 3.5—4.5 cm long.

Pie sepals subequal 2 72. Th. f eae se sass

PT Oe En On eT 10. T. sumatrana

17. Sepals unequal, the outer 3 much smaller than the inner 3 ........... 11. T. macrocarpa
16. Flowers pedicellate; inflorescences always straight. Endocarps up to 2.5 cm long.
18. Leaves with a pair of domatia in basal nerve-axils. Endocarp 2—2.5 cm long, the surface

bearing scattered spinules ..........

Pee ies aorta ASE 12. T. teijsmannii

18. Leaves with glandular patches or domatia (rarely neither) in basal nerve-axils. Endocarp 6—8

mm long, pointed at both ends.

19. Leaves with domatia usually present in basal nerve-axils (replaced by glandular patches in
New Guinea). Flower-pedicels 8—12 mm. Outer sepals much smaller than inner sepals

13. T. glabra

19. Leaves with glandular patches in basal nerve-axils. Flower-pedicels 1-5 mm.

20. Male inflorescences 4—7 cm long; male flower-pedicels 1—2.5 mm. Drupes borne on a +
subglobose «1 shortly 3-branched carpophore, 1.5—2 mm long (Australia). T. smilacina

20. Male inflorescences (5—)7—15 cm long; male flower-pedicels 4-5 mm. Drupes borne on a

columnar carpophore, 4—5 mm long

1. Tinospora trilobata Drets, Pfl. R. Heft 46 (1910)
144; WInKLER, Bot. Jahrb. 49 (1913) 369; Merr. En.
Born. (1921) 249; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 96; ForMAN, Kew Bull. 36 (1981) 381, f.
1A.

Slender climber. Stems drying striate, bearing
short to long rigid hairs. Leaves: petioles 3—12 cm,
hispid to hispidulous; lamina deeply 3-lobed, lobes
subelliptic to subtriangular and acuminate at the
apex, base deeply cordate, 9-18 by 10—16 cm, both
surfaces hispid or hispidulous, papyraceous. — Male
inflorescences supra-axillary, thyrsoid consisting of
a raceme of lax cymes, hispidulous, 15-22 cm long,
lateral branches 3—4.5 cm. — Male flowers on slen-
der pedicels c. 5 mm; sepals yellow, joined at the
base, glabrous or sparsely hispidulous, 1—2 addition-
al minute outermost sepals present, outer 3 narrowly

PAE ed PO Ue ha Cart eet 14. T. subcordata

elliptic, 2.5—3 mm long, inner 3 broadly elliptic,
2—2.5 mm long; petals 6, cuneate, fleshy with lateral
edges incurved, | mm long and broad; stamens 6,
very short and thick, 1—1.5 mm long, filaments
thickened apically and adaxially with the anthers
horizontal and extrorse. — Female flowers and fruits
unknown.

Distr. Malesia: Borneo; 6 coll.

Ecol. Primary forest, 100-1000 m, in Sarawak
recorded on limestone.

Note. A very distinctive species in the genus on
account of its deeply lobed leaves, sepals joined at
the base and form of the stamens. It is unfortunate
that the fruits have not yet been collected; they are
needed to confirm the generic position of the species.

2. Tinospora merrilliana Die.s, Pfl. R. Heft 46

192 FLORA MALESIANA [ser. I, vol. 102

Fig. 6. Tinospora dissitiflora (Laut. & K. Scu.) Diets. a. Endocarp, dorsal and ventral views, x 2. — T. tino-
sporoides (F. v. M.) ForMAn. b. Leaf, x 2/3, c. endocarp, dorsal and ventral views, d. in TS, all x 2. — T.
macrocarpa DIELS. e. Endocarp, dorsal and ventral views, f. in TS, all x 2. — T. teijsmannii BoERL. g. Endo-
carp, dorsal and ventral views, h. in TS, all x 2. — T. glabra (BuRM. f.) MERR. /j. Leaf, x 2/3, k. leaf base
with domatia, «4, /—m. endocarp, dorsal and ventral views, x2. — T. subcordata (MiqQ.) Diets. n. Leaf,
x 2/3, p. endocarp, dorsal and ventral views, x 2 (a NGF 44584, b NSW 90151, c—d FawcetrTs.n., e—f KL
2190, g—h Hort. Bogor., /—k FoRMAN 418, /—m Jacoss 4758, n ZIPPELIUSS.n., p. KocHs.n.). Drawn by Mrs.
M. Church. Courtesy Kew Bulletin.

1986]

MENISPERMACEAE (Forman)

193

(1910) 137; in Elmer, Leafl. Philip. Bot. 4 (1911)
1164; Merr. En. Philip. 2 (1923) 146; ForRMAN, Kew
Bull. 36 (1981) 389, f. IH—K. — T. negrotica DIELs,
Pfl. R. Heft 46 (1910) 137; in Elmer, Leafl. Philip.
Bot. 4 (1911) 1164; Merr. En. Philip. 2 (1923) 146;
YAMAMOTO, Trans. Nat. Hist. Soc. Taiwan 34 (1944)
307, incl. var. monticola YAMAMOTO. — T. havilan-
dii Diets, Pfl. R. Heft 46 (1910) 138; Merr. En.
Born. (1921) 249; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 217. — Parabaena amplifolia Diets, Pfl.
R. Heft 46 (1910) 146; Merr. En. Born. (1921) 249;
YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 96. — T.
hastata Eimer, Leafl. Philip. Bot. 10 (1939) 3760. —
Fawcettia merrilliana (Diets) YAMAMOTO, J. Soc.
Trop. Agric. 16 (1944) 230.

Slender woody climber. Stems up to 1 cm 9; young
stems 2—3 mm @ drying striate, puberulous or gla-
brous; older stems covered with raised corky lenti-
cels. Leaves: petioles 2—9 cm, puberulous (some-
times hispidulous) or glabrous, geniculate and slight-
ly swollen at base; lamina narrowly ovate to ovate (or
broadly ovate) or triangular, base truncate to deeply
cordate or hastate with acute basal lobes, apex acu-
minate, 7—22 by 3—16.5 cm, palmately 5—7-nerved
at base, main nerves usually impressed on upper sur-
face and prominent below, both surfaces glabrous or
sometimes puberulous (or hispidulous) along nerves
especially below, both surfaces drying matt and mi-
nutely wrinkled, texture stiffly papyraceous, doma-
tia absent. — Male inflorescences axillary or arising
from older leafless stems, pseudo-racemose (or nar-
rowly ‘paniculate’), solitary or several arising togeth-
er, 5—16 cm long, very slender, puberulous, mostly
without flowers in lower half. — Male flowers on
pedicels 1—5 mm; sepals pale green, very thin, gla-
brous or sparsely puberulous, (sometimes with 1—2
additional outermost oblong sepals 0.5 mm long),
outer 3 + ovate, 0.5—1.5 mm long, inner 3 obovate
to spathulate, 1.5—2.5 mm long; petals 6, unguicu-
late with subreniform limb, 1.5 mm long, apically
fleshy; stamens 6, narrowly clavate, 2 mm long. —
Female flowers and inflorescences unknown. Infruc-
tescences narrowly pseudo-paniculate, up to 40 cm,
pendent with long slender peduncle up to 23 cm, pu-
berulous. Drupes pinkish white or white, radiating
from a subglobose carpophore 1—2 mm @ on a ped-
uncle 4—6 mm; pericarp glabrous, drying close to
endocarp, the endocarp thinly crustaceous, usually
pale, broadly elliptic in outline, 7—8 mm long, dor-
sally with an obscure medium ridge, surface smooth
or obscurely tuberculate, ventrally flattish with ellip-
tic aperture leading to deeply intrusive condyle.

Distr. Malesia: Borneo, Philippines (Palawan,
Luzon, Catanduanes, Panay, Mindanao) and NE.
Celebes (Minahassa).

» Ecol. Forests up to 1800 mand in Luzon in thick-
ets on limestone cliffs, in Palawan on limestone hill.

Fl. Nov.—May, fr. Febr.—Oct.

Vern. Philippines: cangogang, pisok, Luzon;
balang-batang, Negros; columpangi, kari, laganat,
Mindanao.

Note. This species varies considerably in leaf-
shape. Most specimens from Mt Kinabalu have small
ovate leaves which are truncate or only moderately
cordate at the base. From central and eastern Borneo
the leaves may be large and deeply cordate. Some
specimens from the Philippines and Celebes are
hastate with acute basal lobes, as exemplified by the
synonym 7. hastata ELMER.

3. Tinospora hirsuta (BECC.) FORMAN, Kew Bull. 36
(1981) 391, f. 2A—D. — Aspidocarya hirsuta BEcc.
Malesia | (1877) 136. — Parabaena hirsuta (BECC.)
Drets, Pfl. R. Heft 46 (1910) 147, p.p. quoad typum.

Woody climber. Stems puberulous, sometimes
soon glabrescent, drying striate and developing a
subnitidous bark with scattered small lenticels.
Leaves: petioles 8—10 cm, puberulous; lamina ovate
to broadly ovate, base cordate, apex long-acuminate,
14—18 by 8.5—12 cm, main nerves impressed above,
prominent below, reticulation very fine and promi-
nent on both surfaces, upper surface glabrous, on
lower surface main nerves hispidulous with some
hairs uncinate at the tip, puberulous along finer
nerves, papyraceous, domatia absent. — Male
flowers and inflorescences unknown. — Female
flowers unknown. Infructescence arising from older
leafless stems, narrowly paniculate, 18—25 cm, pu-
berulous, lateral branches c. 1—1.5 cm, upper
branches reduced to single pedicel 0.5 cm. Drupes
white, borne on a subglobose carpophore 2 mm 9;
endocarp cream-whitish, crustaceous, subrotund or
broadly elliptic in outline, slightly keeled at the apical
end, 8—11 by 8 mm, the surface covered with moder-
ately scattered very small pointed tubercles, ventrally
flattish with elliptic aperture leading to a globose
condyle deeply intrusive in the seed-cavity.

Distr. Malesia: Borneo (Sarawak; G. Kenepai); 3
coll.

Ecol. Alluvial forest in valley floor at c. 60 m.

4. Tinospora celebica Diets, Pfl. R. Heft 46 (1910)
143; ForMAN, Kew Bull. 36 (1981) 392, f. 2E—F.
Slender woody climber, entirely glabrous. Stems
drying substriate. Leaves: petioles 4.5(-7, (este
Diets) cm, geniculate near base; lamina narrowly el-
liptic, base sagittate with small acute lobes, apex acu-
minate, 10-12 by 3.5—4.5 cm; reticulation fine,
raised on both surfaces, stiffly papyraceous, domatia
present beneath in basal nerve axils. — Male and fe-
male flowers and inflorescences unknown, Drupes
(pericarp unknown) with endocarp bony, rather
oblong in outline with squarish corners but shortly
pointed at base and apex, 7 by 5 mm, whole surface

194

FLORA MALESIANA

granular-rugulose, dorsal side also slightly and irre-
gularly tuberculate, ventrally with shallow, small
central depression, condyle only slightly intrusive in-
to seed-cavity.

Distr. Malesia: N. Celebes (Gorontalo); 1 coll.

Notes. A unicate from Sarawak may belong to
this species (S 36766), at c. 530 m in poor forest with
mostly small trees and numerous climbers on igneous
derived brownish-yellow sandy soil. Its leaves have
the very characteristic shape of those of 7. celebica
except that the short lobes of the hastate base are
bluntly pointed and not acute; the petioles are short-
er, being about 3 cm. The single detached pseudo-
racemose infructescence (unknown in 7. celebica) is
19 cm long; the fruits are borne on slender pedicels
5—8 cm, each terminating in a narrow clavate carpo-
phore 4 mm long. The drupes have slightly larger en-
docarps, 10 by 6 mm, and are more prominently or-
namented with thin ridges irregularly interrupted and
sometimes divided into sharp points; the surface of
the endocarp is generally rough.

With so very little material known of both the Ce-
lebes and Sarawak plants, it is not possible to be sure
of the significance of the differences and whether or
not these two specimens are conspecific.

5. Tinospora glandulosa Mere. J. Str. Br. R. As.
Soc. n. 85 (1922) 171; Yamamoto, J. Soc. Trop.
Agric. 16 (1944) 96; ForMAN, Kew Bull. 36 (1981)
394, f. 2G—H.

Woody climber, entirely glabrous. Stems drying
striate when young, with scattered raised lenticels,
developing a papery later subcoriaceous bark.
Leaves: petioles 4—5 cm; lamina elliptic, base round-
ed or subtruncate, apex long-acuminate, 10—13 by
4—6 cm, 3(—5)-nerved at the base and with 4—6 pairs
of distal lateral nerves, reticulation raised on both
surfaces, stiffly papyraceous, domatia with distinct
apertures present beneath in the main nerve-axils. —
Flowers unknown. Infructescence arising from
older, leafless region of stem, 25 to over 60 cm, sub-
racemose with one or two pedicels 7—10 mm arising
together in the axil of a 1—2 mm long bract, some-
times short lateral branches 2—4 cm long present to-
wards the base. Drupes 10 mm long, drying irregular-
ly very wrinkled and angled borne on columnar car-
pophores 3—4 mm long; endocarp + broadly elliptic
in outline, abruptly pointed at base and apex, 9 by 6
mm, surface granular with irregular scattered point-
ed protuberances, ventrally flattish with a central el-
liptic aperture.

Distr. Malesia: N. Borneo (Sabah: Sandakan)
and NE. Celebes (Minahassa); 3 coll.

Ecol. Climbing bamboo forest at low altitude.

Note. This species is distinctive in its elliptic
leaves and long infructescences sometimes with short
lateral branches in the lower part. The reticulate

[ser. I, vol. 102

venation of the leaves and the domatia each with a
distinct aperture are very much as in 7. glabra and T.
celebica, to which T. glandulosa appears closely al-
lied.

6. Tinospora crispa (L.) Hook. f. & Tu. Fl. Ind. 1
(1855) 183; Mrers, Contr. Bot. 3 (1871) 34; GAGNEP.
Fl. Gén. I.-C. 1 (1908) 132; KirtrKAR & Basu, Indian
Medic. Pl. 1 (1918) 48, t. 34; Crevost & PETELOT,
Bull. Econ. Indoch. n.s. 1-1929, n. 199 (1929) 30 with
accompanying plate and figure; Merr. J. Arn. Arb.
19 (1938) 341; Back. & Baku. f. Fl. Java 1 (1963)
158; Lien, Acta Phytotax. Sin. 13 (1975) 37; For-
MAN, Kew Bull. 36 (1981) 394, f. 3A—C; ibid. 39
(1984) 113. — Funis felleus RumpH. Herb. Amb. 5
(1747) 44, f. 1. — Menispermum crispum LINNE, Sp.
Pl. ed. 2 (1763) 1468. — Menispermum tuberculatum
LAMK, Encycl. Méth. 4 (1797) 96. — JT. rumphii
BoeERL. Cat. Hort. Bog. (1901) 116, nom. illegit.;
Drts, Pfl. R. Heft 46 (1910) 135; Merr. Int.
Rumph. (1917) 220; Sp. Blanc. (1918) 145; En. Phi-
lip. 2 (1923) 146; Santos, Philip. J. Sc. 35 (1928) 187,
t. 1; Quis. Medic. Pl. Philip. (1951) 300, p.p. — T.
tuberculata (LAMK) BEUMEE ex HEYNE, Nutt. Pl. 1
(1927) 69; Burk. Dict. (1935) 2164. — Fig. 7a—c.
For a complete synonymy, see FORMAN (1981).
Woody climber up to c. 15 m, entirely glabrous.
Stems drying striate when young but later becoming
very prominently tuberculate, containing an exceed-
ingly bitter milky sap, producing very long filiform
aerial roots. Leaves: petioles S—15(—30) cm; lamina
broadly ovate to orbicular, base deeply to shallowly
cordate, apex usually long-acuminate, 7—14(—25) by
6—12(—24) cm, palmately 5—7-nerved at the base,
very thinly papyraceous, domatia usually absent al-
though a flat pocket sometimes present in axil of ba-
sal nerves beneath. Inflorescences not coetaneous
with the leaves. — Male inflorescences arising from
the older, leafless stems, often a few together,
pseudo-racemose, very slender, (S—)9—20 cm long,
flowers in 1—3-flowered fascicles. — Male flowers
on filiform pedicels 2—4 mm; sepals pale green, outer
3 + ovate, thickened at base, 1—1.5 mm long, inner
3 obovate, unguiculate or acute at base, 3—4 mm
long; petals 3, only the outer whorl usually developed
(sometimes 1—3 reduced inner petals present), nar-
rowly oblanceolate, flat, lacking papillae, 2 mm
long; stamens 6, 2 mm long. — Female inflores-
cences similar to male but shorter, 2—6 cm, with
flowers mostly arising singly along the axis. — Fe-
male flowers: sepals and petals as in male; stami-
nodes 6, subulate, scarcely 1 mm long; carpels 3, el-
lipsoidal, 2 mm long, stigma very shortly lobed. Jn-
fructescences (from Assam and Burma specimens)
bearing lateral peduncles 1.5—2 cm terminating in a
subpyramidal 2—3 mm long carpophore below which
usually persist reflexed ovate sepals 2 mm long.

1986]

Drupes orange, ellipsoidal, 2 cm long (when fresh),
with whitish endocarp, + ellipsoidal, 11—13 by 7—9
mm, surface obscurely rugulose or almost smooth,
with a conspicuous dorsal ridge and with a small el-
liptic ventral aperture, condyle deeply intrusive into
seed-cavity.

Distr. Bengal, Assam, Burma, Cambodia, Yun-
nan; in Malesia: Malaya (incl. Singapore I.), W. Java
(incl. Christmas I.), Lesser Sunda Is. (Sumbawa),
and the Philippines (Luzon, Mindoro, Mindanao).

In Christmas I. (Indian Ocean) it may have well
been introduced in the past by immigrant workers.

Ecol. In Thailand in mixed deciduous forest and
village hedgerows up to 900 m, also cultivated. In the
Philippines recorded from primary forest (Minda-
nao) and at 1000 m (Mindoro: Mt Yagaw). Also cul-
tivated as a medicinal plant in Ceylon and India.

Uses. Burxmt (1935, under 7. tuberculata) lists
the many medicinal uses of this plant. The Malays
drink an infusion of the stem as a vermifuge and of
the whole plant to treat cholera.

According to CreEvost & PETELOT (1929, /.c.) the
species was introduced into northern Vietnam (Ton-
kin) by the Sisters of St. Vincent de Paul under the
name of liane-quinine (= ddy ki nin) and it was culti-
vated by various Christian communities; but it was
also known elsewhere in the region. It is used by local
people to treat fevers and jaundice. The stem is cut
into small pieces and scraped, then it is infused in
boiling water, which after cooling is drunk. The
stems can also be dried and pounded into a powder,
which is used as quinine. This powder mixed with
fodder is used to fatten horses and cattle by stimulat-
ing their appetite; a similar use is reported from N.
Thailand by BANZIGER.

MERRILL (1918, under 7. rumphii) stated that this
is perhaps the most generally used medicinal plant in
the Philippines. It contains an extremely bitter prin-
ciple and it is known in the Philippines together with
the more common 7. glabra as makabuhay, but T.
crispa is more effective in use. The bitter principle of
makabuhay has been investigated by MARANON (Phi-
lip. J. Sc. 33, 1927, 357), who found it to be glucosi-
dal. QuisumBING’s account of the species (as T.
rumphii) and its uses in his Medic. Pl. Philip. (1951)
300 deals in part with 7. glabra.

According to THORNBER (Phytochem. 9, 1970,
167), berberine has been reported in 7. crispa, but
this could be based on misidentified material of 7.
glabra.

The anatomy of the stem and leaf has been de-
scribed by Santos (Philip. J. Sc. 35, 1928, 187).

Vern. Java: akar pahat, andawali, brotowali, pu-
trowali; Sarawak: daun akar wali; Philippines: ma-
kabuhay, meliburigan, Mindanao.

» Notes. Female flowers and fruit were described
from extra-Mal. specimens as they are as yet un-

MENISPERMACEAE (Forman)

195

known from Malesia. Even in continental Asia fruits
are rare, at least rarely collected.

Writing at the end of the 17th century, RUMPHTUS
gave a long and detailed account of this species ac-
companied by an illustration showing the charac-
teristic broad, deeply cordate and long-acuminate
leaves, together with the stem densely covered in
raised tubercles, which the artist had incorrectly ar-
ranged in longitudinal lines. RuMpuHtus stated that
this climber was brought to Amboina around 1690
and it flowered, when leafless, in Nov. 1691. He
mentioned its bitter sap and explained that the Ja-
vanese and Balinese names meant ‘bitter rope’, and
therefore he gave it the Latin name Funis felleus. He
also described the medicinal uses of the plant in Java
and Bali.

The confusion about the application of the name
Menispermum crispum L., lasting for two centuries,
originated from LINNE, who cited the correct plate in
RumMPuHIus, but the wrong name, ‘Funis quadrangu-
laris’, which is a Cissus (Vitaceae).

As a result of intensive searches in Thailand, Dr.
BANZIGER finally succeeded in collecting the fruits of
T. crispa, which proved to be clearly different from
those of T. baenzigeri.

The stems have a remarkable capacity when cut in-
to pieces to remain succulent and alive for a long pe-
riod: the dried sap effectively seals the cut ends.
RUMPHIUs stated that when originally brought to
Amboina about 1690, the coiled stems had been ina
closed box for some months, and when planted they
soon produced shoots. In confirmation of this prop-
erty, several portions of stem some 15 cm long were
received at Kew in Oct. 1977, collected by Dr. BANz1-
GER in Thailand some 10 to 12 months previously, yet
some were still green and succulent, the tissue appar-
ently still living.

In Thailand, according to BANzIGER, leaves are
present during the rainy season April-May to
Nov.—Dec. or later if growing in a humid place.
Plants flower late Jan.—March; the flowers are
scented. Fruits were collected in April and May.

The typical number of petals in this species is 3,
only the outer whorl developing, contrasting with 6
in the closely allied 7. baenzigeri. There are, how-
ever, specimens which have in addition | to 3 petals
of the inner whorl (usually reduced) together with the
warty stems characteristic of 7. crispa. It could be
that there has been some hybridisation between the
two species, whose areas of distribution overlap in
Central Thailand.

7. Tinospora baenzigers ForMAN, Kew Bull. 36
(1981) 399, f. 3D—G; ibid. 39 (1984) 113. — Fig.
7d—g.

Allied to 7. crispa, but differing by: Old stems up
to 6 cm @, bearing scattered pustular lenticels but

196 FLORA MALESIANA [ser. I, vol. 102

Fig. 7. Tinospora crispa (L.) Hook.f. & Tu.a. Leaf, x 2/3, b. stem, nat. size, c. endocarp, dorsal and ventral

views, x 2. — T. baenzigeri FoRMAN. d. Leaf, x 2/3, e. leaf base showing pocket-domatia, x4, f. stem, nat.

size, g. endocarp, dorsal and ventral views, x 2. — T. smilacina Bru. h—j. Leaves, x 2/3, k. leaf base showing

glandular patches, x 4, /. endocarp, dorsal and ventral views, x 2 (a BANZIGER 71-6, b BANZIGER 71-21, c BAN-

ZIGER 86, d—e BANZIGER 30-10, f BANZIGER 30-14, g BANZIGER 30-17, A ScHuLTz711,/ PARKER 477, k LAZARI-
DEs 6531, / Must 1289). Drawn by Mrs. M. Church. Courtesy Kew Bulletin.

1986]

MENISPERMACEAE (Forman)

lacking prominent tubercles. Leaves with a pair of
hollow domatia present in axils of basal nerves on
lower surface. Flowers with 6 petals. Drupes yellow,
radiating from a subglobose carpophore | mm long
on peduncle 5—7 mm; pericarp drying very thin;
endocarp thinly bony, blackish grey, 7—9 by 5—6
mm, broadly elliptic in outline, rounded at base,
slightly keeled at apex, with a weak dorsal ridge, sur-
face papillose or almost smooth.

Distr. Thailand, S. Vietnam; in Malesia: SW. Ja-
va (Christmas I., Indian Ocean).

Ecol. In Thailand, according to Dr. Banziger,
leaves form at the beginning of the rainy season
(April—May) and persist until the end of the rainy
season (Oct.—Nov.). Flowering begins mid-Dec. and
lasts until mid-Febr., the individual plants remaining
in flower for about one month. The flowers have a
strong but pleasantly fragrant scent. Fruits appear
from mid-Jan.

In Christmas I. (see below) male flowers and leaves
were collected in Dec. 1980 and between June and
Sept. 1981.

According to Dr. BANZIGER it is most common in
Central Thailand in areas with a prolonged dry sea-
son of 4—6 months, although it also occurs in parts
of S. Thailand with a rather wet climate; it often oc-
curs in open areas, sometimes on an isolated tree, at
altitudes up to about 400 m. The species is apparently
absent from the northern parts of Thailand where
low temperatures occur. KERR collected specimens
from scrub-land, scrambling over bushes, and from
wasteland around Bangkok.

Vern. A few specimens from Thailand collected
by Kerr and Marcan bear the same vernacular
names ching cha li and chincha chali as are used for
T. crispa, but T. baenzigeri does not appear to be in
general use for medicinal purposes. This may be con-
nected with the fact that the bitter substances present
in the stems of 7. baenzigeri are different from those
in 7. crispa.

Notes. Stem material was phytochemically anal-
ysed by LacHaty & SCHNEIDER (Arch. Pharm.,
Weinheim, 314, 1981, 251—256) under the erroneous
identification 7. cordifolia.

A most surprising extension of the range is the oc-
currence in Christmas I. (Indian Ocean), where it was
found in 3 localities (1980) and where it grows as a
climber enveloping small trees on the shore terraces,
and covering limestone pinnacles, in one place to-
gether with 7. crispa. This raises the possibility that
it may have been introduced together with that spe-
cies. On the other hand, 7. baenzigeri may in the fu-
ture be found to occur in other localities, e.g. Java,
Sumatra, and thus prove to have a wider distribution
than is now realized. Tinospora baenzigeri, with its
inconspicuous flowers appearing when the plant is
leafless, may yet be eluding collection in unsuspected
territory.

197

A noticeable feature of the Christmas I. material
is that the flower pedicels can be as long as 13 mm
compared with a maximum of c. 4 mm in specimens
from Thailand. Otherwise all the distinctive charac-
ters of T. baenzigeri are present in the Christmas I.
specimens.

8. Tinospora arfakiana Becc. Malesia | (1877) 140,
excl. infloresc.; FORMAN, Kew Bull. 36 (1981) 407, f.
4F—H; ibid. 89 (1984) 114. — Tinomiscium arfakia-
num (Becc.) Dtets, Pfl. R. Heft 46 (1910) 116, excl.
infloresc. — Parabaena scytophylla Diets, Bot.
Jahrb. 52 (1915) 189.

Woody climber, entirely glabrous. Stems rather
smooth without conspicuous lenticels when young,
bark on old stems with raised elongate lenticels, c.
5—10 mm. Leaves: petioles 10—15 cm; lamina ovate,
cordate or slightly so at the base, acuminate to
broadly acuminate at the apex, 18—28 by 11—23 cm,
palmately 5—7-nerved at the base, with a series of
short tertiary nerves running at right angles to the
midrib, nervation very prominent below, less so abo-
ve, thinly coriaceous. — Male and female flowers un-
known. /nfructescences cauliflorous, paniculate to-
wards the base with lateral branches up to 12 cm,
pseudo-racemose towards apex, 18—70 cm. Drupes
red, 1(—3) on peduncles 8—15 mm, drying smooth,
broadly ellipsoidal, ventrally flattened and slightly
concave, 17—24 by 14—18 mn, pericarp drying very
thin; endocarp very smooth and whitish, wall 1 mm
thick with ventral elongate groove divided by a longi-
tudinal septum, condyle intruding into the seed-
cavity. Seed ellipsoidal, ventrally concave; embryo
with the broad flat, slightly overlapping cotyledons
enclosed in entire endosperm, radicle median, cylin-
drical.

Distr. Malesia: New Guinea; 4 coll.

Ecol. Primary forest, in the Vogelkop at only 30
m, in E. New Guinea at 600—700 m and in montane
rain-forest at 1500 m.

9. Tinospora dissitiflora (LAuT. & K. Scu.) DIELS,
Pfl. R. Heft 46 (1910) 144; ForMAN, Kew Bull. 36
(1981) 408. — Aspidocarya dissitiflora Laut. & K.
Scu. Fl. Deut. Schutzgeb. Siidsee (1900) 312. — As-
pidocarya stenothyrsus K. Scu. Nachtr. Fl. Deut.
Schutzgeb. Siidsee (1905) 264. — 7. peekelii Diets,
Bot. Jahrb. 52 (1915) 188. — Fig. 6a.

Slender woody climber, entirely glabrous. Stems
striate when young, lenticellate, later developing a
pergamentaceous subnitidous bark, drying wrinkled.
Leaves: petioles 6—13 cm; lamina ovate to broadly
ovate, base slightly cordate or truncate, apex abrupt-
ly acuminate, 11—16 by 7.5—12 cm, reticulation very
fine and raised on both surfaces, stiffly papyraceous,
glandular patches present in basal nerve axils be-
neath. — Male inflorescences axillary or arising from

198

FLORA MALESIANA

[ser. I, vol. 102

older, leafless stems, pseudo-paniculate, 13—20 cm
long, the lower lateral branches up to 3.5 cm. —
Male flowers on very slender pedicels S—10 mm; sep-
als pale green, outer 3 ovate, 1.5—1.8 mm long, inner
3 broadly elliptic, 4.5—5.5 mm long; petals 6, broad-
ly spathulate to obovate-cuneate, externally papil-
lose in basal region, 2.5—3 mm long; stamens 6, nar-
rowly clavate, 3—S mm long. — Female inflores-
cences pseudo-paniculate towards the base, pseudo-
racemose towards the apex with the flowers in fasci-
cles, c. 30—40 cm long. — Female flowers (known
only from buds): sepals and petals similar to male;
staminodes 6, subulate, 0.7 mm long; carpels 3, ellip-
soidal, 1.3 mm long including spreading slightly
lobed stigma. Drupes red, usually only one develop-
ing on each 3 mm long columnar carpophore, on
peduncles 1.3—2.5 cm, very knobbly when dry with
pericarp drying close to endocarp; endocarp bony,
strongly and irregularly tuberculate, rather oblong in
outline with squarish corners but pointed at base and
apex, 10—12 by 7—8 mm, ventrally flattish with a
large elliptic cavity (7.e. the condyle).

Distr. Malesia: New Guinea (incl. New Britain
and New Ireland; rare in W.).

Ecol. Lowland rain-forest and swamp-forest at
low altitudes up to 300 m, also on coral shores.

Note. This species is easily recognizable by its
drupes and inflorescences and also by its leaves
which show (when dried) a very fine raised reticula-
tion on both surfaces.

10. Tinospora sumatrana (SCHEFF.) BEcc. Malesia 1
(1877) 139; Borer. Cat. Hort. Bog. 1 (1899) 36, p.p.;
ibid. 2 (1901) 118; Drets, Pfl. R. Heft 46 (1910) 144;
FORMAN, Kew Bull. 36 (1981) 411, 421. — Limacia
sumatrana SCHEFF. Obs. Phytogr. pt 3 (1872) 76, t.
9; Nat. Tijd. N. I. 32 (1873) 398.

Woody climber, entirely glabrous. Stems striate
with pustular lenticels, later covered with a smooth
pergamentaceous bark. Leaves: petioles 3.5—8 cm;
lamina broadly ovate to ovate, base cordate to trun-
cate, apex acuminate, 8—12 by 4.5—8 cm, papillose
patches sometimes (esp. in young leaves) obscurely
visible in basal nerve-axils on lower surface, surfaces
drying matt with reticulation obscure, papyraceous.
— Male inflorescences: a few arising together from
the older, leafless stems, unbranched, very slender,
zigzag or not, 7-15 cm long, flowers in spaced re-
trorse fascicles of c. 3, each fascicle subtended by a
retrorse bract 1 mm long. — Male flowers: minute,
sessile; outer 3 sepals elliptic, 2 mm long, inner 3 sep-
als elliptic, concave, 1.5 mm long; petals 6, oblong
with lateral edges incurved, 1 mm long; stamens 6, 1
mm long. — Female inflorescences and flowers un-
known. Jnfructescences unbranched, 20 cm. Drupes
3 on stout peduncles 1.5 cm, subnavicular-ellipsoid,
4.5 by 1.5—2 cm, pericarp (dried) very thin, endocarp

thinly bony, 1 mm or less thick, slightly verrucose
with a prominent apical carina and a shallow ventral
groove scarcely intruding into the large seed-cavity;
seed subhemicylindrical, ventrally flattened and with
a median groove, endosperm ventrally transversely
ruminate; embryo (BECCARI) with divaricate broad
thinly foliaceous cotyledons, radicle terete, superior.

Distr. Malesia: S. Sumatra (Lampong Distr.),
Billiton I.; 2 coll.

Notes. This species is very closely related to T.
macrocarpa from Malaya, with which it shares its
major distinctive features. The inner and outer sep-
als, however, are subequal in 7. sumatrana, and on
this basis the two species are regarded as distinct.

T. sumatrana var. hanadae YAMAmoToO, J. Soc.
Trop. Agric. 16 (1944) 94. The type (Sarawak, Ku-
ching, HANADA BOIS) has not been traced: the status
of this varietal name based on a sterile specimen must
remain uncertain.

11. Tinospora macrocarpa Diets, Pfl. R. Heft 46
(1910) 141; Rmx. Fl. Mal. Pen. 1 (1922) 103; For-
MAN, Kew Bull. 36 (1981) 412, f. SE—F. — T. uligino-
sa (non Miers) Hook.f. & TH. Fl. Br. India 1 (1872)
97. — Fig. 6e—-f.

Scandent shrub, entirely glabrous. Stems striate
with pustular lenticels, later covered with a smooth
pergamentaceous bark. Leaves: petioles (2—)5—10
(—16) cm; lamina broadly ovate to ovate (or elliptic-
ovate), base cordate to truncate (or rounded), apex
acuminate, 6—13(—21) by (3.5—)6—10(—17) cm, sur-
face drying matt with reticulation rather obscure, pa-
pillose patches sometimes present in basal nerve-axils
beneath, very thinly to stiffly papyraceous. — Male
inflorescences: a few arising together from the older,
leafless stems, unbranched, very slender, sometimes
slightly zigzag, 7-20 cm long; flowers in spaced
fascicles (sometimes retrorse) of c. 3 flowers; each
fascicle subtended by a retrorse bract 0.5—1 mm
long. — Male flowers: minute, subsessile (pedicels
up to 0.5 mm); outer 3 sepals triangular-ovate, 0.8
mm long, inner 3 sepals broadly elliptic, concave,
1.5—2 mm long; petals 6, oblong with lateral edges
incurved, | mm long; stamens 6, 0.8 mm long. — Fe-
male inflorescences and flowers unknown. Infructes-
cences rather slender and unbranched, 19—27 cm
long, bearing very prominent discoid scars. Drupes 3
on stout peduncles 1—2 cm, orange-yellow, ellipsoid-
al, 3.5—4.5 by 1.5—2 cm, style subterminal; pericarp
(dried) very thin; endocarp thinly bony, mostly less
than 1 mm thick, papillose-tuberculate or almost
smooth with a dorsal carina more pronounced to-
wards the apex, and with an elongate ventral groove
which only slightly intrudes into the large seed-
cavity. Seed ellipsoidal, ventrally grooved, contain-
ing copious, ventrally ruminate endosperm; embryo
with divergent, thin, foliaceous cotyledons with fine-

1986]

ly lobed margins, radicle cylindrical, superior.

Distr. Malesia: Malaya (Wellesley, Selangor,
Malacca, also in Penang and Singapore Is.), 7Borneo
(Sabah); 7 coll.

Ecol. Presumably in forest, in Wellesley at 150
m.
Vern. Akar kepayang, buah pelay tedong, Te-
muan.

Notes. Very closely related to 7. sumatrana; the
latter has subequal sepals, while in 7. macrocarpa
the outer sepals are much smaller than the inner ones.

I gave a lengthy discussion on the very limited and
incomplete specimens, which presented certain prob-
lems (FORMAN, 1981).

One collection in fruit from Sabah is probably T.
macrocarpa (or T. sumatrana).

12. Tinospora teijsmannii BoerL. Cat. Hort. Bog.
(1901) 117; Diets, Pfl. R. Heft 46 (1910) 141; Merr.
En. Born. (1921) 249; Yamamoto, J. Soc. Trop.
Agric. 16 (1944) 96; FoRMAN, Kew Bull. 36 (1981)
414, f. 5G—H; ibid. 39 (1984) 114. — Fig. 6g—h.

Slender woody climber, entirely glabrous. Stems
drying striate, later developing scattered raised lenti-
cels. Leaves: petioles 4—10 cm; lamina broadly cor-
date, often obtuse at insertion of petiole within the
broad basal sinus, apex acuminate, 10—13 by 9—12
cm, one pair of domatia present in basal nerve-axils
beneath, very thinly papyraceous. — Male inflores-
cences and flowers unknown. — Female inflorescen-
ces a few arising together from older leafless stems,
pseudo-racemose, lax, 25—30 cm long, the flowers
arising 1—2(—3) together. — Female flowers with
pedicels 9—12 mm; outer 3 sepals ovate, 2 mm long,
inner 3 sepals elliptic, 3 mm long; petals 6, narrowly
obovate, 0.8 mm long; staminodes 6, subulate, 1 mm
long; carpels 3, 1.5 mm long including shortly divid-
ed stigma, borne on a pyramido-globose gynophore.
Drupes \1—3 on peduncles 12—20 mn, arising from
main axis of infructescence, ellipsoidal, 20—25 by c.
15 mm, pericarp drying thin in loose folds around
endocarp; endocarp thinly bony, 16—18 by 9-10
mm, surface bearing sparsely scattered very short
pointed tubercles, otherwise smooth, with a dorsal
carina more pronounced towards the apex, and with
an elongate ventral groove intruding about 1/3 way
into seed-cavity. Seed ellipsoidal, ventrally grooved.

Distr. Malesia: Borneo, only known from the
type, cultivated in Hortus Bogoriensis, collected by
TEUSMANN.

13. Tinospora glabra (BurM./.) Merr. J. Arn. Arb.
19 (1938) 340; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 95; Back. & Baku. /. Fl. Java 1 (1963) 157;
ForMAN, Kew Bull. 36 (1981) 414, f. 5J-M. — Meni-
spermum glabrum Burma. f. Fl. Ind. (1768) 216
(‘316’), excl. syn. Rueepe. — Cocculus coriaceus BL.

MENISPERMACEAE (Forman)

199

Bijdr. (1825) 25. — Cocculus bantamensis BL. I.c.
26. — Dioscorea spiculata Bi. En. Pl. Jav. 1
(1827/28) 22, excl. syn. RuMpPH.; HALL. f. Meded.
Rijksherb. 1 (1911) 41. — Dioscorea aculeata (non
L.) Zoti. & Mor. Syst. Verz. (1845/46) 92. — Coc-
culus crispus [non (L.) Dc.] Hassx. Pl. Jav. Rar.
(1848) 166. — T. uliginosa Miers, Ann. Mag. Nat.
Hist. ser. 3, 13 (1864) 321; Contr. Bot. 3 (1871) 35;
Hook.f. & Tu. Fl. Br. India 1 (1872) 97; BEcc. Ma-
lesia 1 (1877) 139; Kina, J. As. Soc. Beng. 58, ii
(1889) 378; Back. Fl. Bat. 1 (1907) 34; Voorl. (1908)
8. — T. reticulata Miers, Ann. Mag. Nat. Hist. ser.
3, 13 (1864) 321; Contr. Bot. 3 (1871) 36; Diets, Pfl.
R. Heft 46 (1910) 143; in Elmer, Leafl. Philip. Bot.
4 (1911) 1164; Merr. Fl. Manila (1912) 204; En. Phi-
lip. 2 (1923) 146; Philip. J. Sc. 29 (1926) 368; SANTos,
ibid. 35 (1928) 198, t. 5; YAMamorTo, J. Soc. Trop.
Agric. 16 (1944) 95; Harus. Mem. Fac. Agric. Kago-
shima Un. 5 (1966) 30; PANCHO, Vasc. Fl. Mt Maki-
ling 1 (1983) 279, f. 83. — 7. pseudo-crispa BOERL.
ex Back. FI. Bat. 1 (1907) 35, nomen in syn. — T.
crispa [non (L.) Hook. f. & TH.], saepe p.p., incl.
Diets, Pfl. R. Heft 46 (1910) 143, pro maj. parte, t.
49D-—O. — T. andamanica Drets, Pfl. R. Heft 46
(1910) 141. — 7. coriacea (BL.) BEUMEE ex HEYNE,
Nutt. Pl. 1 (1927) 619; Back. Onkr. Suiker. (1930)
248, t. 259 (1936). — Fig. 4f-1, 6j—m.

Woody climber, entirely glabrous. Stems striate
when young, becoming warty (with raised lenticels)
and later developing a smooth thin papery bark
which often becomes detached on drying. Leaves:
petioles 4—8(—12) cm; lamina oblong-ovate or nar-
rowly to broadly ovate, base cordate to truncate,
with basal lobes rounded or obtusely pointed, apex
acuminate, 7—12(—15) by 5—9(—13) cm reticulation
raised on both surfaces, papyraceous, domatia with
distinct apertures usually present beneath in basal
nerve-axils, the floor of the domatia carpeted with
glands, occasionally domatia absent but glandular
patches present (e.g. in New Guinea). — Male inflo-
rescences axillary or arising from older, leafless
stems, pseudo-racemose, slender, lax, 10—20 cm, not
or sparsely flowered in the lower 1/3 to 1/2, flowers
solitary or in fascicles of 2—5 in the axil of a subulate
bract 1 mm long. — Male flowers on very slender
pedicles 8—12 mm; sepals yellow, greenish (or white),
outer 3 narrowly ovate 1 mm long, inner 3 broadly el-
liptic, concave 4-5 mm long; petals 6, broadly
cuneate-obovate with lateral edges incurved, exter-
nally papillose at base, 2—3 mm long; stamens 6, cla-
vate, 3—5 mm long. — Female inflorescences similar
to male but up to 35 cm. — Female flowers: sepals
and petals as in male but inner sepals 3 mm long; sta-
minodes 6, subulate, 0.5—1 mm; carpels 3, ellipsoid-
al, 1.5 mm, including reflexed lobed stigma, borne
on a subglobose gynophore |—1.5 mm long. Drupes
red, radiating from unbranched short to columnar

200

FLORA MALESIANA

[ser. I, vol. 102

carpophore 2—4 mm long on peduncle 4—10 mm
arising from main axis of infructescence; pericarp
drying thin and close to endocarp; endocarp thinly
bony, 6—8 by 4—5 mm, subrotund or subelliptic in
outline, pointed at base, keeled at apex, dorsally con-
vex with a median ridge and irregularly tuberculate,
ventrally flattish with a small elliptic aperture to con-
dyle.

Distr. From Hainan and S. Andaman to the So-
lomon Is. (Reef and Rennell Is.); in Malesia: Malaya,
Sumatra (incl. Krakatoa in Sunda Strait), Java (incl.
Nusa Barung and Madura Is.), Lesser Sunda Is. (all),
Borneo, Philippines (all islands), S. Moluccas (Te-
nimber Is.), E. New Guinea (incl. Normanby I. and
New Britain).

Ecol. In a great number of situations in littoral
rain-forest and Casuarina equisetifolia forest, man-
grove and on sandy beaches, often inland in dis-
turbed forest and shrubberies, secondary growths,
very frequently on limestone (Java, Sumba, Timor)
and on black soils under seasonal conditions; up to
500 m.

Morph. The anatomical structure of the stem
and leaf of 7. glabra was investigated by SANTOS
(1928, /.c.).

As a rule the outer sepals are much smaller than
the inner ones, but exceptionally (PNH 17161) the
sepals vary from subequal in some flowers to very
unequal in others. This approaches the closely allied
species 7. homosepala Diets, where the sepals are
equal.

The inflorescences of the species are characteristi-
cally unbranched, but in the anomalous PNH 9142
the infructescences have a few lateral branches up to
4 cm, rather similar to those of 7. glandulosa MERR.

Specimens from New Guinea differ in certain re-
spects from the rest of the material. Domatia are
lacking on the lower surface of the leaves, where they
are normally present in the basal nerve-axils. There
occur instead, in these positions, flat glandular-
papillose patches similar to the glandular areas that
are found within domatia. The female inflorescences
are only 8—10(—15) cm, which is shorter than in ma-
terial from west of New Guinea. In Lag 52539 and
CLEMENS 11066 the endocarps have a larger ventral
aperture than is usual; the carpophore of Lar 52539
is shortly and divaricately branched, one branch be-
low each fruit.

Uses. In the Philippines this species is used for a
variety of medicinal purposes, e.g. burnt leaves used
to treat pinworms; ground bark is applied to sore
breasts of nursing mothers. Together with 7. crispa
this species is known in the Philippines as makabu-
hay, but apparently 7. crispa is medicinally more ef-
fective. The account of makabuhay in QuUISUMBING
(Medic. Pl. Philip. 1951, 300) is given under the
name 7. rumphii BoERL., a synonym of T. crispa,

but the description in part refers to 7. glabra. Qut-
SUMBING mentions a number of medicinal uses as well
as reports of alkaloids: some of these may refer to 7.
glabra. The species is also used in the Philippines for
baiting wild pigs by mixing sliced roots with Jpomoea
batatas.

The alkaloid berberine has been reported in 7.
crispa (THORNBER, Phytochem. 9, 1970, 167). The
material tested, however, may well have been 7. g/a-
bra since the correctness of its identification is uncer-
tain.

Vern. Flores: wasé wages; Philippines: papaitan,
Palawan, makabuhay, tabin tabin, Mindoro, maka-
buhay, Luzon, manongal, Panay, agmamali, ca-
sopo, glingu melibutigan, sangawnaw, Mindanao;
nono, New Britain.

Notes. In his original description of Menisper-
mum glabrum, BURMAN incorrectly cited as a syno-
nym ‘Cit-amerdu’ of RHEEDE, Hort. Malab. 7 (1688)
39, t. 21. RHEEDE’s description, however, clearly im-
plies that his plant had hairy stems and leaves; it is,
in fact, part of the basis of the later name Menisper-
mum malabaricum LAMK, which is now a synonym
of Tinospora sinensis (LOUR.) MERR.

The type of Menispermum glabrum Burm. f. is a
specimen in the DELESSERT Herbarium at Geneva,
which was acquired by BURMAN from the herbarium
of PRYON.

14. Tinospora subcordata (MiQ.) Drets, Pfl. R. Heft
46 (1910) 136; ForRMAN, Kew Bull. 36 (1981) 419. —
Hypsipodes subcordatus Mig. Ann. Mus. Bot.
Lugd.-Bat. 4 (1868) 82; VaLet. Bull. Dép. Agric.
Ind. Néerl. 10 (1907) 11. — T. polygonoides DiE1s,
Pfl. R. Heft 46 (1910) 136; J. Arn. Arb. 20 (1939) 73.
— Fig. 6n, p.

Small woody climber, entirely glabrous. Stems
drying striate when young, later becoming minutely
verruculose and bearing scattered raised lenticels.
Leaves: petioles 2.5—9 cm; lamina triangular to
broadly triangular, base broadly cordate to truncate
with rounded, sometimes subhastate, basal lobes,
apex acute, 6—10 by 4—9 cm, reticulation raised on
both surfaces, papyraceous, glandular patches pres-
ent on lower surface in axils of main nerves. — Male
inflorescences axillary, pseudo-racemose, (5—)7—15
cm long, the lower half without flowers, arising sing-
ly or 2—3 directly from the leaf-axils, or sometimes
2—3 arising from very short 1—1.6 cm long axillary
shoot, flowers mostly in fascicles of 3—4. — Male
flowers on slender pedicels 4—5 mm; sepals white,
outer 3 + ovate, 1—1.5 mm long, inner 3 elliptic, 4
mm long; petals 6, obovate-cuneate, 1.5 mm long,
fleshy, externally minutely papillose-glandular near
base; stamens 6, narrowly clavate with filament
broadened apically, 3.5—4.5 mm long. — Female in-
florescences pseudo-racemose, the flowers arising

1986]

MENISPERMACEAE (Forman)

201

singly. — Female flowers on pedicels 4—6 mm; sepals
and petals similar to male but slightly smaller and
petals thin; staminodes 6, oblong, 0.5 mm long;
gynophore columnar, | mm long; carpels 3, gibbose-
ellipsoidal, 1 mm long, stigma flat and expanded
with margin minutely lobed. /nfructescences race-
mose, 7—9 cm. Drupes red, radiating from columnar
gynophore 4—5 mm long on peduncles 5—8 mm;
pericarp drying thin; endocarp bony, + elliptic in

outline pointed) at apex, 7 by 4 mm, dorsally with a
median ridge, surface coarsely and irregularly tuber-
culate and also minutely rugulose, ventrally with el-
liptic aperture to shallow ventral cavity.

Distr. Malesia: Lesser Sunda Is. (Timor), S. Mo-
luccas (Tenimber Is., Selaru I.), S. New Guinea (Me-
rauke, Mabaduan, Wassi Kussa: Tarara).

Ecol. Largely in lowland vegetation subject to a
distinct dry season.

outline, shortly pointed at base, strongly keeled (in

9. PARABAENA

Miers, Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 35, 39; Hoox. f. & Tu. FI. Ind.
(1855) 181; BENTH. in B. & H. Gen. Pl. 1 (1862) 34; Miers, Ann. Mag. Nat.
Hist. ser. 3, 14 (1864) 51; Contr. Bot. 3 (1871) 57; Hoox. f. & Tu. Fl. Br. India
1 (1872) 95; Diets, Pfl. R. Heft 46 (1910) 145; Forman, Kew Bull. 39 (1984) 103.
— Fig. 4m—p, 5c—I.

Slender woody climbers. Leaves with petiole swollen and geniculate at base,
lamina often cordiform or hastate, rarely deeply 3—5-lobed, margin entire or
repand-dentate, palmatinerved at base, papyraceous. /nflorescences axillary,
cymose or thyrsoid (a raceme or panicle of cymes). — Male flowers: sepals 6,
free, equal or the inner 3 broader; petals 6, often minute, sometimes with a pair
of thickenings or projecting lobes on basal inner surface; synandrium usually
peltate with the anthers situated around the margin usually dehiscing transver-
sally. — Female flowers occasionally functionally hermaphrodite: sepals and
petals as in male; staminodes 6, minute but these occasionally developing into
claviform polliniferous stamens; carpels 3, stigma reflexed, lobed or laciniate.
Drupes 3, borne on subglobose carpophore; endocarp bony, dorsal and lateral
surfaces variously ridged or spiny, sometimes with prominent dorsal ridge as
well as basal and apical keels, condyle represented by a ventral concavity some-
times bordered by incurved spines or developed into an inflated ventral cham-
ber, seed-cavity flattened. Seed with copious endosperm; embryo with very thin
divaricate broad cotyledons and prominent radicle.

Distr. Continental SE. Asia (1); Malesia: N. Borneo (1), Philippines (3) and New Guinea & Solomons (1),
in all 6 spp.

KEY TO THE SPECIES
Based on male flowering material
N.B.; female inflorescences usually shorter and less branched

1. Inflorescences usually repeatedly dichotomously branched; leaves often sagittate with pointed basal lobes
and margin often repand-dentate (Nepal to Yunnan, NE. India, Andaman Is., Indochina) P. sagittata
1. Inflorescences with a distinct main axis bearing lateral branches; leaves mostly ovate or cordiform with
entire or subentire margins (rarely 3—5-lobed).
2. Inflorescences glabrous, 5—18 cm long; synandrium with apical conical appendage. Leaves cordiform to
broadly cordiform, glabrous (or sparsely hispidulous below) .........6. 660000005 5. P. echinocarpa

202

FLORA MALESIANA [ser. I, vol. 102

2. Inflorescences pubescent or, if subglabrous, then 18—35 cm long; synandrium flat or domed at apex.
3. Inflorescences elongate, mostly 23—36 cm long, narrow with + equal lateral branches 2—6 cm long; an-
thersiwithsverticaleslits wise seis ae Rite «rete Vets eal Abe ey epee ae 1. P. megalocarpa
3. Inflorescences shorter or, if over 20 cm long, then with lowermost lateral branches longest and decreasing
upwards; anthers with transverse slits.
4. Lamina ovate to narrowly ovate, rather sparsely pubescent below; sepals equal .... 3. P. denudata
4. Lamina cordiform to broadly cordiform or broadly ovate, usually very pubescent below, inner sepals
broader.
5. Inflorescences 7—15 cm long, narrow, rather densely pubescent.................... 2. P. elmeri
5. Inflorescences 18—35 cm long with spreading lateral branches up to 12 cm long, patent-hispidulous
tOVSUD glabrous sec atae arse fee eV aerators ciekaie wcrw od Gis ares ermo eens NG 4. P. tuberculata

KEY TO THE SPECIES
Based on fruiting material (dried fruits)

IRquitsssubellipsordy 292 —2- SiCmalon pw nispideaeyy see. eic sijsrta sole eitrcrs oie > «ye eboney ean Sere
1. Fruits irregular in shape and much smaller, glabrous.

2. Infructescences usually repeatedly dichotomously branched; fruits 5-6 mm long with conspicuous apical
keel and dorsal ridge of endocarp visible; leaves often sagittate with pointed basal lobes and margin often
repand-dentate (Nepal to Yunnan, NE. India, Andaman Is., Indochina) ............... P. sagittata

2. Infructescences with a distinct main axis bearing lateral branches; fruits 7-11 mm long; leaves mostly
ovate or cordiform with entire or subentire margins.

3. Infructescences rather densely pubescent. Leaves pubescent below ...................
3. Infructescences with scattered hairs or glabrous.

1. P. megalocarpa

2. P. elmeri

3. P. denudata

AMMAN ALOVAtCHLOMMATNOWLY OVALE Milani tin ccc sist ele dere eue see e ae cele a thinner

4. Lamina cordiform to broadly cordiform.

5. Fruits strongly ridged, the endocarp bearing a prominent dorsal ridge produced at the ends into basal
and apical keels and also bearing prominent thin lateral wings; leaves usually pubescent below

4. P. tuberculata

5. Fruits lacking clear dorsal and lateral ridges, not or scarcely pointed at the ends, dorsal surface of
endocarp densely covered with slender patent spines; leaves glabrous (or sparsely hispidulous below)

1. Parabaena megalocarpa Merr. Un. Calif. Publ.
Bot. 15 (1929) 59; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 97; ForMaAN, Kew Bull. 36 (1981) 392; ibid.
39 (1984) 105, f. 1C—E. — P. hirsuta (non BEcc.)
Drs, Pfl. R. Heft 46 (1910) 147, p.p. quoad Havi-
LAND 2833. — Fig. 4m—p, 5c—e.

Young stems hispid to hispidulous or puberulous.
Leaves with hispid to shortly pubescent petioles 5—14
cm; lamina ovate to broadly ovate or deeply
3—5-lobed with narrow sinuses, base cordate, apex
acuminate, 12—25 by 8—18.5 cm, margin entire, both
surfaces hispid to hispidulous. — Male inflorescen-
ces supra-axillary (or terminal), elongate, a raceme
of cymes, (12—)23—36 cm, the lateral branches 2—6
cm, puberulous. — Male flowers on pedicels 1—1.5
mm: sepals yellow or cream, papillose or puberu-
lous, main outer sepals narrowly elliptic, 2—2.5 mm
long, inner sepals broader, 2—2.5 mm long; petals ro-
tund, 0.5—0.75 mm long with 2 basal inwardly pro-
jecting lobes, or these joined together; synandrium
cylindrical, thick, 1.25 mm long; anthers with verti-
cal slits. — Female inflorescences pseudo-racemose
or with lateral branches cymose or racemose, 9—15

5. P. echinocarpa

cm. — Female flowers: sepals and petals similar to
male; staminodes 1 mm; carpels 1.5 mm. Drupes yel-
low turning red, subellipsoidal, hispid with fine
spines of endocarp projecting through pericarp when
dry, 2.2—2.8 cm long, peduncles c. 5 mm; endocarp
resembling a hedgehog, densely covered with long
thin patent spines, apart from the closed inflated
ventral chamber, c. 2 cm long, seed-chamber dorsi-
ventrally very compressed.

Distr. Malesia: Borneo (Sarawak, Sabah).

Ecol. Forests, including riparian forests, up to
900 m. Fl. fr. Jan.—Dec.

Uses. The fruit is stated to be edible and sour.

Vern. Sabah: paruka-paruka; Sarawak: akar
pelir-udok, \ban.

Note. This species shows a number of strong re-
semblances to Tinospora trilobata Dies, especially
when the leaves are deeply lobed and hispid. Both
species have similar supra-axillary, very long thyr-
soid inflorescences.

2. Parabaena elmeri Diets, Pfl. R. Heft 46 (1910)
147; in Elmer, Leafl. Philip. Bot. 4 (1911) 1164;

1986]

Merr. En Philip. 2 (1923) 147; YAMaAmorTo, Trans.
Nat. Hist. Soc. Taiwan 34 (1944) 308, incl. var. phi-
lippinensis (MERR.) YAMAMOTO; PANCHO, Vasc. FI.
Mt Makiling 1 (1983) 277; FoRMAN, Kew Bull. 39
(1984) 107, f. 1G. — P. philippinensis MERR. ex
Drts, Pfl. R. Heft 46 (1910) 148; in Elmer, Leafl.
Philip. Bot. 4 (1911) 1164; Merr. En. Philip. 2
(1923) 147. — Fig. 5g.

Stems moderately to lightly pubescent, glabres-
cent. Leaves with pubescent petioles S—11 cm; lami-
na broadly ovate or deltoid-ovate, base cordate or
Sagittate-cordate (or subtruncate), apex acuminate,
10—16(—23) by 8—12(—23) cm, margin entire or re-
motely dentate, both surfaces pubescent, usually
more densely beneath. /nflorescences axillary, com-
posed of a raceme or narrow panicle of cymes, slen-
der, 7—15 cm long, usually densely pubescent. —
Male flowers on pedicels 1—2 mm: sepals 2 mm long,
glabrous, outer ones + elliptic, inner ones broadly
elliptic; petals oblong, 0.75 mm long, flat; synan-
drium | mm long, anthers with transverse slits. — Fe-
male flowers: sepals subequal, 2 mm long; petals lan-
ceolate, 0.75 mm long; staminodes oblong, 0.5 mm
long; carpels ovoid-ellipsoidal, 1.25 mm long with
recurved, lobed stigma. Drupes drying irregularly
ridged, 8-10 mm long, on pubescent pedicels c. 2
mm, glabrous; endocarp broadly elliptic in outline
with conspicuous apical keel, dorsally bearing subad-
pressed spinules and fimbriate or toothed diagonal
ridges with the margin encircled by a skirt-like
toothed ridge, the large ventral cavity bordered by a
thin incurved toothed margin.

Distr. Malesia: Philippines (Mindoro, Luzon).

Ecol. Thickets and forests at low to medium alti-
tudes. Fi. Febr., May—July, fr. Febr., July—Oct.

Vern. Mindoro: bugbog-puro.

3. Parabaena denudata Diets, Pfl. R. Heft 46 (1910)
147; in Elmer, Leafl. Philip. Bot. 4 (1911) 1164;
Mere. En. Philip. 2 (1923) 147; Forman, Kew Bull.
39 (1984) 108, f. IH. — Fig. 5h.

Stems up to c. 1.3 cm @, sparsely pubescent when
young. Leaves with petioles pubescent to subgla-
brous, 4—10 cm; lamina ovate to narrowly ovate with
base truncate to cordate, or sagittate, apex mostly
long-acuminate, 8—15 by 3—9 cm, margin entire or
repand-denticulate, upper surface subglabrous,
lower surface lightly pubescent with prominent fine
reticulation. Inflorescences axillary, a raceme or
panicle of cymes, very slender with delicate ultimate
branching, 9-20 cm long, shortly pubescent. —
Male flowers on pedicels c. 1 mm: sepals greenish
white, equal, elliptic to obovate, 2 mm long, gla-
brous; petals broadly obovate-cuneate, 0.5 mm long;
synandrium | mm long, anthers with transverse slits.
— Female flowers not seen. Drupes drying spinulate-
muricate with conspicuous apical keel, 7-8 mm

MENISPERMACEAE (Forman)

203

long, glabrous on puberulous pedicels c. 5 mm;
endocarp 7 by 6 mm, rotund in outline (excluding
apical keel), dorsally bearing a median double row of
short spines which lead into a prominent apical keel,
rest of dorsal surface bearing short spines with the
surface between them rather rough, ventral cavity
bordered by radial incurved spines surrounding a
central aperture.

Distr. Malesia: Philippines (Luzon, Samar, Ne-
gros, Bohol, Mindanao, Basilan).

Ecol. In Negros along rivers and in thickets in
damp ravines at 700—750 m. Fi. Febr.—May, /r.
Aug.

4. Parabaena tuberculata Becc. Malesia 1 (1877)
137, p.p. excl. fruct.; Diets, Pfl. R. Heft 46 (1910)
148; p.p.; C.T. Wuire, J. Arn. Arb. 10 (1929) 208,
212; FORMAN, Kew Bull. 39 (1984) 108, f. 1J-K. —
Stephania cincinnans K.Scu.in K. Sch. & Hollr., Fl.
Kaiser Wilh. Land (1889) 44. — P. myriantha K.
Scu. in K. Sch. & Laut., Nachtr. Fl. Deut. Schutz-
geb. Stidsee (1905) 264 (‘myriaditha’, sphalm.);
Dre.s, Pfl. R. Heft 46 (1910) 149; RENDLE, J. Bot. 61
Suppl. (1923) 4. — P. cincinnans (K. Scu.) DIELs,
Pfl. R. Heft 46 (1910) 149. — P. psilophylla Diets,
l.c. 148; KANEH. & Hatus. Bot. Mag. Tokyo 56
(1942) 472. — Fig. 5j—k.

Young stems pubescent, sometimes sparsely so.
Leaves with pubescent (or subglabrous) petioles
5—13 cm; lamina broadly cordiform to rotund, base
cordate with rounded or sometimes obtusely angled
basal lobes or truncate, apex abruptly acuminate,
10—22 by 9-22 cm, margin entire or sometimes
repand-denticulate, both surfaces pubescent (or sub-
glabrous), fine reticulation prominent on lower sur-
face. Inflorescences axillary, apparently paniculate,
18—35 cm long with spreading to retrorse lateral
branches, the lower ones up to 12 cm, patent-hispid-
ulous to subglabrous. — Male flowers on pedicels
1—2 mm: sepals white or yellow, 2.5 mm, externally
sparsely pubescent to glabrous, the outer 3 + ellip-
tic, the inner 3 broader and concave; petals + rhom-
boid, 1 mm long, fleshy with lateral thickenings; syn-
andrium c. 1 mm long, sometimes domed at apex,
anthers with transverse slits. — Female flowers on
pedicels 3—4 mm: sepals and petals similar to male;
staminodes minute, peg-like c. 0.25 mm long but
sometimes (as in lectotype) stamen-like, c. 0.75 mm
long; carpels | mm long, stigma laciniate with 3-6
reflexed lobes. Plants with all flowers hermaphrodite
sometimes occurring, these with 6, free stamens c.
0.75 mm long. Drupes white (or red) radiating from
subglobose carpophore |—1.5 mm @ terminating a
glabrous pedicel 3—4 mm, drying strongly ridged, +
broadly elliptic in outline and abruptly pointed at
both ends, 9-11 mm long, glabrous; endocarp
broadly elliptic in outline, with a prominent median

204

FLORA MALESIANA

[ser. I, vol. 102

dorsal ridge (composed of a double row of closely set
spines) produced at both ends into basal and apical
keels, and two thin prominent lateral wings which are
continuous or composed of separate spines, with fur-
ther small spines present over dorsal surface, ventral
surface with a ring of radially incurved spines bor-
dering a ventral chamber and leaving a central oval
aperture.
Distr. Solomon Is.; in Malesia: New Guinea.
Ecol. Secondary rain-forest, Pometia-Intsia for-
est with much regrowth, river-side and cane-brakes
in floodbed, up to 1000 m. Fi. fr. Jan.—Dec.
Vern. W. New Guinea: kakoep, Biak; NE. New
Guinea: simuganbang, Waskuk, jehmu, Wagu.
Note. For a full discussion of this species see
FORMAN (1984). Some elements must be discarded
from the original material; a lectotype was selected;
it does not occur in the Aru Is. and in Timor; the bi-
sexual flowers BECCARI mentioned are anomalous in
his specimen.

5. Parabaena echinocarpa Drets, Philip. J. Sc. 8
(1913) Bot. 157; MerRrR. En. Philip. 2 (1923) 147; Ya-
MAMOTO, Trans. Nat. Hist. Soc. Taiwan 34 (1944)
307, incl. var. pubescens YAMAMOTO; FORMAN, Kew
Bull. 39 (1984) 110, f. 1L. — Fig. 51.

Stems glabrous. Leaves with glabrous (or subgla-
brous) petioles S—18 cm; lamina cordiform to broad-
ly cordiform or broadly triangular-ovate, base deep-
ly to shallowly cordate or truncate, apex acuminate,
9—15(—25) by 7—13(—20) cm, margin entire, both
surfaces glabrous (or sparsely hispidulous below).
Inflorescence axillary, a panicle of cymes, slender,
5—18 cm, glabrous. — Male flowers on pedicels 1—2
mm: sepals subequal, elliptic to oblanceolate,
2.25—3 mm long, glabrous, petals elliptic, 1 mm
long; synandrium 1.5 mm long with apical conical
appendage, anthers with transverse slits. — Female
flowers not seen. Drupes whitish, drying irregularly
rugose, 8—9 mm long, glabrous, on glabrous pedicels
4—5 mm; endocarp broadly elliptic to subrotund in
outline, dorsally densely covered with slender patent
spines, apical keel moderately or little developed,
ventral cavity covered by a thin wall with the central
aperture bordered by a radially fimbriate incurved
margin or by flattened incurved spines.

Distr. Malesia: Philippines (Luzon,
Leyte, Mindanao, Camiguin de Misamis).

Ecol. Thickets and forests at altitudes up to c.
500 m. Fi. April—June, Nov., fr. April, Nov.—Dec.

Vern. Polillo I.: baya-bayatian, Dum.

Polillo,

10. TINOMISCIUM

Miers [Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 44, nomen] ex Hook.f. & Tu. FI.
Ind. (1855) 205; Miers, Ann. Mag. Nat. Hist. ser. 3, 13 (1864) 489; Contr. Bot.
3 (1871) 44; DreELs, Pfl. R. Heft 46 (1910) 115; FoRMAN, Kew Bull. 40 (1985) 542,
f. 1. — Fig. 8.

Woody climbers containing white latex in a laticiferous system which extends
throughout the plant. Stems ferrugineous-pubescent when young. Leaves +
ovate to elliptic, base 3—5S-nerved, petioles usually long, bent and swollen at
base and sometimes at apex, upper surface when dry revealing the laticiferous
system as a fine, dense network of + parallel ridges. Inflorescences racemose
arising from old, leafless stems, usually ferrugineous-tomentose. — Male
flowers: sepals 9 in 3 whorls of 3, the outermost whorl much smaller than the
inner ones; petals 6 with the lateral edges incurved; stamens 6, variable, some-
times apiculate, the anthers sometimes immersed in the thickened connective,
dehiscence longitudinal to transverse. — Female flowers: sepals and petals as in
male; staminodes 6, very narrow; carpels 3, stigma lobed. Drupes 3, radiating
from discoid carpophore, style-scar terminal; endocarp compressed, elliptic to
subovate in outline; seed flat, endosperm present, cotyledons thin, flat, imbri-
cate, radicle small.

Distr. Assam, Burma, Nicobar Is., Thailand, Yunnan, Indochina; almost throughout Malesia, except the
Lesser Sunda Is. Monotypic.

1986]

1. Tinomiscium petiolare Hoox. f. & Tu. FI. Ind.
(1855) 205; Mia. Fl. Ind. Bat. 1, 2 (1858) 87; Miers,
Contr. Bot. 3 (1871) 45, t. 94; Hook.f. & Tu. Fl. Br.
India 1 (1872) 97; Kina, J. As. Soc. Beng. 58, ii
(1889) 379; Riot. J. Str. Br. R. As. Soc. n. 33 (1900)
42; GaGnepP, Fl. Gén. I.—C. 1 (1908) 127, t. 14/1-
10; Drets, Pfl. R. Heft 46 (1910) 33, f. 16A & B, 118;
Rpt. J. Str. Br. R. As. Soc. nm. 54 (1910) 14; MERR.
En. Born. (1921) 248; Rpt. Fl. Mal. Pen. 1 (1922)
105, t. 9; NORMAN, J. Bot. 62, Suppl. (1924) 5; Burk.
& HENpD. Gard. Bull. S. S. 3 (1925) 344; HEYNE,
Nutt. Pl. (1927) 618; Henp. Gard. Bull. S. S. 4
(1928) 219; Burk. Dict. (1935) 2163; YAmMamorTo, J.
Soc. Trop. Agric. 16 (1944) 41; Back. & Baku. /f. FI.
Java 1 (1963) 157; THoTHATHRI ef a/. Bull. Bot. Surv.
India 15 (1976) 15; ForMAN, Kew Bull. 40 (1985) 542,
f. 1. — T. phytocrenoides Kurz ex TEUSM. & BINN.
Nat. Tijd. N. I. 27 (1864) 36; ScHeErFF. ibid. 32 (1873)
394, t. 3; BoeRL. Cat. Hort. Bog. 1 (1899) 37; DreELs,
Pfl. R. Heft 46 (1910) 117, f. 42; Back. Schoolfl.
(1911) 42; Koorp. Exk. Fl. Java 2 (1912) 233; Hey-
NE, Nutt. Pl. (1927) 618; Burk. Dict. (1935) 2136;
Yamamoto, J. Soc. Trop. Agric. 16 (1944) 40;
STEENIS-KRUSEMAN, Bull. Org. Sc. Res. Indon. 18
(1953) 35; Back. & Baku. f/f. Fl. Java 1 (1963) 157.
— T. pyrrhobotryum Mig. Ann. Mus. Bot. Lugd.-
Bat. 4 (1868) 81. — 7. javanicum Miers [Ann. Mag.
Nat. Hist. ser. 3, 13 (1864) 490, nomen], Contr. Bot.
3 (1871) 45. — T. elasticum Becc. Malesia 1 (1877)
141; Drets, Pfl. R. Heft 46 (1910) 116; KANEH. & Ha-
Tus. Bot. Mag. Tokyo 56 (1942) 474. — T. philippi-
nense Diets, Pfl. R. Heft 46 (1910) 116; in Elmer,
Leafl. Philip. Bot. 4 (1911) 1164; W.H. Brown, Mi-
nor Prod. Philip. For. 3 (1921) 186; Merr. En. Phi-
lip. 2 (1923) 146; Burk. Dict. (1935) 2163; Quis.
Medic. PI. Philip. (1951) 299; PANcHo, Vasc. Fl. Mt
Makiling 1 (1983) 279, f. 83. — T. molle Diets, Phi-
lip. J. Sc. 8 (1913) Bot. 157; Merr. En. Philip. 2
(1923) 146. — Fig. 8.

Large woody climber to 30 m, exuding white latex
when cut. Old stems coarsely striate, glabrous, up to
c. 2.5 cm @; young leafy stems conspicuously striate,
ferrugineous-pubescent at first, later glabrescent.
Leaves: petioles 6—21 cm, often rather slender, pu-
berulous to glabrous, bent and slightly swollen at
base and sometimes also at apex; lamina ovate to
broadly ovate or elliptic, base truncate to cordate or
obtuse, apex acuminate, 11—25(—29) by 4.5—20 cm,
base 3—5-nerved with 2—3(—4) pairs of distal lateral
nerves, nervation prominent on lower surface, upper
surface (when dry) covered with a dense network of
+ parallel hair-like ridges, glabrous, lower surface
puberulous to glabrous, stiffly papyraceous. /nflo-
rescences arising several together from protuber-
ances on old stems, racemose, (5—)8—28(—35) cm,
férrugineous-tomentose, rarely glabrous. — Male
flowers on puberulous pedicels 2.5—5 mm; 3 outer

MENISPERMACEAE (Forman)

205

sepals triangular-ovate to narrowly triangular, 1—2
mm long, lightly puberulous to glabrous; 6 main in-
ner sepals white to yellow, elliptic, 4-5 mm long, re-
flexed at anthesis, glabrous or externally lightly pu-
berulous (rarely tomentellous); petals broadly ellip-
tic, 2.5—3.5 mm long, erect and contiguous at anthe-
sis, concave with lateral edges incurved, glabrous;
stamens with or without apiculus, 2—2.5 mm, an-
thers either immersed in the thickened connective or
rather prominent, dehiscence longitudinal to trans-
verse. — Female flowers: sepals and petals as in
male; staminodes linear-oblong, acute, 3 mm; car-
pels curved-ellipsoidal, 2 mm long, stigma shortly
multi-lobed. Drupes at first green with white spots,
later white to yellow (or orange), containing white la-
tex, radiating from discoid carpophore terminating
pedicels 1-2 cm, drying + compressed-ellipsoidal,
glabrous, base narrowed into a short stalk; endocarp
compressed, narrowly to broadly elliptic or subovate
in outline, 2—3.5 by 1—2 cm, base rounded to obtuse,
apex obtuse to sharply acute, surface obscurely to
strongly rugose or rugulose.

Distr. India (incl. Nicobar Is.), Burma, Thai-
land, Yunnan, Vietnam; in Malesia: Sumatra, Ma-
laya, W. Java, Borneo (incl. Natuna Is.), Philippines
(Luzon, Mindanao), New Guinea.

Ecol. In forests up to 1400 m, but most collec-
tions from below 500 m, on a variety of soils includ-
ing limestone (Java). Fl. fr. Jan.—Dec. The flowers
are fragrant; PomLANE 29891 (Vietnam) indicates that
the odour is like that of methyl-salicylate (oil of win-
tergreen).

Uses. The milky exudate is used against dental
caries (Vietnam), to alleviate sprue and fever (VAN
STEENIS-KRUSEMAN, /.c.), and diluted it is used as an
eyewash (Philippines; see BRown, /.c., under 7. phi-
lippinense). The fruits are used as fish-poison (Phi-
lippines; BuRKILL, /.c., under 7. philippinense), yet
Wuirtmoke in his field-notes for Fri 3381 stated that
the seeds are edible and sweet. The plant (? fruits) is
used as a rat-poison in S. Sumatra (BURKILL, /.c., un-
der 7. phytocrenoides), while the roots and stems are
used medicinally in Java.

Vern. Thailand: parai hortawng; Malaya: akar
mumbulu, akar pelis kura, lumpaung; Java: (akar)
ki konéng, S, kuniran, ojod konéng, ojod tjatjing,
seriawan susun, susun, susun sapi, J; W. Borneo:
kunyet utan, Natuna Is.; SW. Borneo: hujan panas,
Karimata Arch.; Philippines: /agtang, maglabtang,
Luzon, calumpangi, Mindanao.

Note. In his monograph Dtets (1910) recognized
7 species although he commented that these were
very close and that their number should perhaps be
reduced. The characters used by Diets to distinguish
the species were: shape and indumentum of leaves,
and sizes of inflorescences, flower-pedicels and sep-
als. With the abundant material of the genus now

206 FLORA MALESIANA [ser. I, vol. 102

asst as are ee

aa
VY

PNET yn
{

Wp,

ye

Prue. *
| /

ao

Z : Se Te SS
9s ; Ox ZS
as ye \
f ov, §
'‘* Es.
. mes Hee meer
2 SS 2 Ri i
vind eal a Ze
£) ei) Ke
3 ,
we
\ 4

Fig. 8. Tinomiscium petiolare Hoox. f. & TH. a. Habit, x 2/3, b. detail of lower leaf surface, x20, c. male

inflorescences, x 2/3, d. male flower, front petals removed, x4, e—f. varying stamens, x15, g. female

flower, all petals and front staminodes removed, x 4, 4. drupes, ventral view, nat. size, i. TS of drupe contain-

ing compressed seed, x 1 1/2, j—/. varying endocarps, dorsal views, nat. size (a—b, h—j S 36494, c—d KOsTER-
MANS 19354, e—f Unesco 238, g FRI 3381, k PNH 4850, / DE WILDE 16507).

1986]

available, it is clear that there is much variation in the
characters and that they can no longer be used to
draw taxonomic distinctions. The stamens and endo-
carps have also proved to be very variable. The sta-
mens can be conspicuously apiculate or the apiculus
can be completely lacking; both conditions can occur

MENISPERMACEAE (Forman)

207

prominent, /.e. projecting from the connective, with
vertical slits to being immersed in the connective with
horizontal slits; intermediate examples also occur.
The endocarps are very variable in shape and surface
ornamentation and also vary in the degree of flatten-
ing.

in the same flower. The anthers vary from being

11. FIBRAUREA

Lour. Fl. Coch. (1790) 626; Miers, Ann. Mag. Nat. Hist. ser. 3, 13 (1864) 487;
Contr. Bot. 3 (1871) 41; Drets, Pfl. R. Heft 46 (1910) 119; FoRMAN, Kew Bull.
40 (1985) 546. — Fig. 9a—h.

Woody climbers with yellow wood, entirely glabrous. Older stems with grey-
ish buff bark, irregularly and coarsely striate; young stems smoothly and finely
striate. Leaves + elliptic to ovate, base 3(—5)-nerved with the main basal later-
als running alongside the midrib for several (— 15) mm before curving outwards,
with 2—4 pairs of distal lateral nerves. /nflorescences: lax panicles, often rami-
florous. — Male flowers: 6 main sepals with 2—3 minute outer ones; petals 0;
stamens 3 or 6, the filament thick with a prominent collar around the base of
the anthers, dehiscence longitudinal to oblique. — Female flowers: sepals as in
male; petals 0; staminodes 6, subulate; carpels 3, stigma cleft-like. Drupes ra-
diating from a small knob-like carpophore, drying coarsely wrinkled; endocarp
subellipsoidal with ventral narrow longitudinal groove; seed subellipsoidal, with
narrow longitudinal groove, endosperm abundant around the embryo, cotyle-
dons thin, foliaceous.

Distr. Two species, one wide-spread in the Nicobar Is., Burma, Indochina, S. China, and Malesia: Suma-
tra and Malaya to Celebes and the Philippines, the other one endemic in S. China and Indochina.

Ecol. Both species have yellow wood but field-notes of F. tinctoria also mention the presence of a white
latex or sticky sap. Although not mentioned in the field-notes for F. recisa, it may well occur in that species
also.

Note. There has been some confusion about the identity of LouREIRO’s type species, a matter discussed
by Forman, /.c.

KEY TO THE SPECIES

1. Stamens 3. Wall of endocarp very thin, firmly crustaceous, less than 0.5 mm thick. S. China, Indochina
F. recisa

1. Stamens 6. Wall of endocarp much thicker, hard and rigid, c. 1 mm thick ............ 1. F. tinetoria

1. Fibraurea tinctoria Lour. Fl. Coch. (1790) 626;
Hook. f. & Tu. Fl. Ind. (1855) 204; Mig. Fl. Ind.
Bat. 1, 2 (1858) 87; Ann. Mus. Bot. Lugd.-Bat. 4
(1868) 81; Miers, Contr. Bot. 3 (1871) 41; Hook. /.
Fl. Br. India 1 (1872) 98; Scnerr. Obs. Phyt. 3 (1872)
73,1. 4; Kurz, J. As. Soc. Beng. 43, ii (1874) 60; For.
Fl. Burma | (1877) 53; Becc. Malesia | (1877) 142;
Gaonep. Fl. Gén. I.-C. 1 (1908) 135; Merr. Trans.

Am. Phil. Soc. n.s. 24 (1935) 157; FoRMAN, Kew
Bull. 40 (1985) 549. — Cocculus rimosus Bu. Bijdr.
(1825) 24. — F. chloroleuca Miers [Ann. Mag. Nat.
Hist. ser. 3, 13 (1864) 489, nomen], Contr. Bot. 3
(1871) 42, t. 93; Kina, J. As. Soc. Beng. 58, ii (1889)
380; Rip. Trans. Linn. Soc. Bot. Il, 3 (1893) 274;
Boer. Cat. Hort. Bog. 1 (1899) 37; Rint. J. Str. Br.
R. As. Soc. n. 33 (1900) 42; Diets, Pfl. R. Heft 46

208 FLORA MALESIANA [ser. I, vol. 102

f ||
yoy
UW fo af!

yO \

WW 4 1, W, *
Ay

ih

all g

' ea

yy

Fig. 9. Fibraurea tinctoria Lour. a. Leaf, b. male inflorescence, both x 2/3, c. male flower, front sepals re-

moved, d. stamen, side view, both x 10, e. female flower, front sepals and staminode removed, x6 (from

spirit material), f. drupe, g. endocarp, both x 2/3, A. TS of endocarp showing section of seed, mainly endo-

sperm, with 2 separated thin cotyledons, x 1 1/2. — Tiliacora triandra (COLEBR.) DiELs. i. Habit, male plant,

x 2/3, j. male flower with valvate inner sepals, x 10, k. male flower with front inner sepal removed, x 15,

/. drupes on branched carpophore, x 2, m. endocarp, <3 (a, e FORMAN 76, b—d FORMAN 77, f—h SAN 26487,
i—k Curtis 447, /—m Co.t.ins 119).

1986]

(1910) 120; Back. Schoolfl. (1911) 42; Koorp. Exk.
Fl. Java 2 (1912) 233; MerRR. En. Born. (1921) 248;
Row. Fl. Mal. Pen. 1 (1922) 105; Merr. En. Philip.
2 (1923) 146; Burk. & HEND. Gard. Bull. S. S. 3
(1925) 344; Norman, J. Bot. 64, Suppl. (1926) 142;
Drets & HACKENB. Bot. Jahrb. 60 (1926) 307; HEy-
NE, Nutt. Pl. (1927) 619; Burk. Dict. (1935) 1000;
YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 41; BACK.
& Baku. f. Fl. Java 1 (1963) 157. — F. fasciculata
Miers, Contr. Bot. 3 (1871) 43. — F. laxa MIERs,
Contr. Bot. 3 (1871) 43; Diets, Pfl. R. Heft 46 (1910)
120; Merr. En. Born. (1921) 249; Yamamoto, J.
Soc. Trop. Agric. 16 (1944) 42. — Fig. 9a—h.
Large woody climber up to c. 40 m. Stem up to 5
cm @, containing white latex, young shoot-tips ten-
drilliform. Leaves: petioles (2—)4—13 cm, often dry-
ing blackish at least at the swollen base; lamina ellip-
tic, elliptic-ovate to ovate or oblong-elliptic, base
sometimes subpeltate, usually rounded, apex acumi-
nate, often shortly so, (9—)11—21(—28) by (3.5—)
5—14 cm, upper surface often drying greyish and
smooth with reticulation obscure, thinly coriaceous.
Inflorescences axillary or ramiflorous, c. 10-38 cm
with lateral branches up to 12 cm. — Male flowers
sweetly scented, on pedicels up to 5 mm or sessile;
main sepals white or yellow, broadly elliptic, con-
cave, 2.5—4 mm long; stamens 6, 2—2.5 mm long, fil-
ament thickly columnar, incurved, anthers rather
elongate and narrowly rounded at apex with lateral
longitudinal slits. — Female flowers: sepals and pet-
als as in male; staminodes subulate, 2 mm; carpels el-
lipsoidal, 1.75 mm long, stigma cleft-like, small. Jn-
fructescences often ramiflorous up to c. 55 cm. Dru-
pes yellow to orange on pedicels 6—15 mm; endocarp
2—2.5 cm long, wall c. 1 mm thick, hard and rigid.
Distr. NE. India (Manipur, Nicobar Is.), Bur-
ma, Thailand, Indochina; in Malesia: Malaya, Su-

MENISPERMACEAE (Forman)

209

matra (incl. Enggano I.), Java, Borneo, NE. Cele-
bes, Philippines (Dinagat I., N. of Mindanao).

Ecol. In Thailand recorded up to c. 100 m, local-
ly common in dry evergreen forest, also in bamboo
forest and scrub; in Vietnam at c. 1200 m on clayey
soil in forest with undergrowth of tea bushes. In Ma-
laya at low altitudes in primary forest, dense bamboo
forest, lowland secondary and disturbed forest,
along riverside and on rocky coast. In Sumatra at
1000 m. In Sarawak locally abundant in peat swamp
forest, also in primary lowland forest and in second-
ary forest at 1000 m. In Brunei on yellow sandy loam
in secondary forest and in mixed peat swamp forest.
In Sabah up to 900 m in primary and secondary for-
est on ultrabasic, sandstone and stony blackish soil,
in logged forest and along river banks. F/. Jan.—
Oct., fr. Jan.—Dec.

Uses. The stems are used medicinally in Malaya,
as a stomach medicine in Sarawak, also for dyeing
and for cordage. HEYNE (1927) and BurKILL (1935)
recorded various uses, against dysentery, diabetes
and headache; alkaloids occur; the yellow dye is pos-
sibly berberine, but see BIssET in FORMAN, /.c. 540.

Vern. Peninsular Thailand: kamin krua, kumin
kua, man miet; Malaya: sekunyit, Johore; Sumatra:
akar kunyit, akar stupai, olor labai, Simalur 1.; Ban-
ka: akar mangkédun, M; Java: areuj gember, areuj
ki konéng, S, peron, J; Sarawak: akar badi, akar
kunyit, akar penawar, \ban, war birar, Murut.

Notes. Inhis key, Drets (/.c. 1910) distinguished
F. laxa by its tricomposite male inflorescences up to
50 cm compared with F. chloroleuca with bicompo-
site male inflorescences up to 20 cm. This distinction
is not valid when all the material now available is
considered: indeed, the type of F. /axa, the only
specimen of that species cited by Drexs, has a single
inflorescence no longer than c. 38 cm.

12. ARCANGELISIA

Becc. Malesia | (1877) 145; Drets, Pfl. R. Heft 46 (1910) 103; FoRMAN, Kew
Bull. 32 (1978) 333; Fercuson, /.c. 341; WILKINSON, /.c. 350.

Lianes. Leaves palmately nerved at base with small papillose region on upper
surface above insertion of petiole; hollow domatia with the aperture sometimes
fringed with hairs present in the axils of the nerves and main veins, lamina other-
wise glabrous. /nflorescences axillary or cauliflorous, paniculate with lateral
branches spicate or subspicate. — Male flowers sessile or subsessile; sepals
9—10, glabrous, the outermost 3—4 minute, larger inner sepals 3 + 3; petals 0;
synandrium a sessile, globose cluster of 9—12 anthers. — Female flowers (DIELS,
l.c.) with sepals + as in male; petals 0; staminodes present (? number); carpels
3, stigma broad. Infructescence with club-shaped unbranched carpophores.
Drupes transversely subovoid or subglobose with style-scar lateral, large; endo-

210 FLORA MALESIANA [ser. I, vol. 102

carp not sculptured but bearing a layer (continuous or broken) of radially ar-
ranged fibres; condyle inconspicuous or absent; seed broadly ellipsoidal; endo-
sperm deeply ruminate; cotyledons apparently divergent and much folded (ac-

cording to BEccARI and MAINGAY).

Distr. Two species in Hainan, S. Thailand, Indochina; in Malesia: Malaya to New Guinea.

Palyn. Pollen of Arcangelisia was described by FERGUSON (Kew Bull. 32, 1978, 341).

Notes. There are clearly two species: the wide-spread, small-fruited A. flava and the large-fruited New
Guinea endemic A. tympanopoda, which can unfortunately only be identified in fruiting condition. Miss Wi-
KINSON (/.c.) found that there are stomatal-anatomical differences in the leaf; FERGUSON (/.c.) found no differ-

ence in the pollen.

There is no indication that A. tympanopoda occurs outside New Guinea.

KEY TO THE SPECIES

1. Fruits transversely subovoid, 2.2—3 cm long, 2.5—3.3 cm broad (long axis); endocarp covered with a dense

matonnadiallysarranged fibres. 22--25..2...2+-.

Dash Bee eh eA Oe ee 1. A. flava

1. Fruits subglobose, 4.5—5.5 cm @; endocarp bearing an interrupted layer of radially arranged fibres, which

form a dense dorsal ridge as well as thin lateral transverse plates ...............

1. Arcangelisia flava (L.) Merr. Int. Rumph. (1917)
222; W.H. Brown, Minor Prod. Philip. For. 2
(1921) 388; ibid. 3 (1921) 67, 185; Merr. En. Born.
(1921) 248; En. Philip. 2 (1923) 145; HeEyNe, Nutt.
Pl. (1927) 621; Hottu. & Lam, Blumea 5 (1942) 180;
YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 36; HEINE
in Fedde, Rep. 54 (1951) 226; Back. & Baku. f/f. FI.
Java 1 (1963) 155; ForMAN, Kew Bull. 32 (1978) 334;
PANCHO, Vasc. Fl. Mt Makiling 1 (1983) 281, f. 85.
— Tuba flava Rumen. Herb. Amb. 5 (1747) 38, t. 24.
— Menispermum flavum LINNnE, Herb. Amb. (subm.
Stickman) (1754) 18; Syst. ed. 10 (1759) 992. — Me-
nispermum flavescens LAMK, Encycl. Méth. 4 (1797)
98. — Cocculus flavescens (LAMK) DC. Syst. 1 (1817)
520; Prod. 1 (1824) 97. — Anamirta flavescens
(LAMK) Mig. FI. Ind. Bat. 1, 2 (1858) 79. — Anamir-
ta lemniscata Miers [Ann. Mag. Nat. Hist. ser. 3, 14
(1864) 51, nomen], Contr. Bot. 3 (1871) 54, t.
97/1—6. — Anamirta luctuosa Miers, Contr. Bot. 3
(1871) 55. — A. lemniscata (MtERs) BEcc. Malesia 1
(1877) 147; Bogert. Cat. Hort. Bog. 1 (1899) 38;
Drets, Pfl. R. Heft 46 (1910) 106, f. 38; Back.
Schoolfl. (1911) 40; Drexs in Elmer, Leafl. Philip.
Bot. 4 (1911) 1163; Nova Guinea 8 (1912) 869;
Koorp. Exk. Fl. Java 2 (1912) 232. — A. inclyta
Becc. Malesia 1 (1877) 147. — Anamirta loureiri
PiERRE, For. Fl. Coch. (1885) t. 110; Kina, J. As.
Soc. Beng. 58, ii (1889) 379. — Mirtana loureiri
(PIERRE) PreRRE, Bull. Soc. Bot. Fr. 52 (1905) 490.
— A. loureiri (P1ERRE) DrELs, Pfl. R. Heft 46 (1910)
104; Rpt. Fl. Mal. Pen. 1 (1922) 107; Burk. &
HEND. Gard. Bull. S. S. 3 (1925) 344. — Tinospora
havilandii Diets, Pfl. R. Heft 46 (1910) 138, p.p.,
quoad folia.

2. A. tympanopoda

Plant glabrous apart from leaf-domatia. Stems
with yellow wood and exuding yellow sap when cut,
bearing prominent cup-like petiole-scars. Leaves:
petioles (4—)7—15(—20) cm, swollen at both ends,
geniculate at base; lamina usually ovate, elliptic-
ovate or broadly ovate, base usually rounded, trun-
cate or slightly cordate, apex abruptly acuminate,
(10—)12—25 by (5.5—)8—19 cm, palmately 5-nerved
at the base and with 1—3 pairs of lateral nerves usual-
ly arising from above halfway along the midrib, main
nerves prominent, especially below, both surfaces
usually drying matt with a rather obscure reticulum,
coriaceous; hollow domatia present on lower surface
in the axils of main nerves, with the aperture fre-
quently puberulous. /nflorescences axillary or cauli-
florous, paniculate, slender, 10-50 cm, lateral
branches spicate or subspicate, 1-5 cm. — Male
flowers sessile or subsessile subtended by an ovate
bracteole c. 1 mm long which is strongly thickened at
the base; 3—4 minute outer sepals less than 1 mm
long, 3 +3 larger inner sepals elliptic, ovate or nar-
rowly obovate, 1.5—2.5 mm long; synandrium 0.5—1
mm long. — Female flowers (DiELs) with 6 main sep-
als narrowly oblong with the apex becoming re-
flexed, 2.5—4 mm long; staminodes minute, scale-
like; carpels 3, 1.5 mm long, stigma broad, sessile,
papillose. Jnfructescences cauliflorous, usually
branched, (S5—)7—30(—45) cm, with thickened axis
and branches, 3—6 mm g, the fruits plus carpophores
borne on the lateral branches; 1—3 borne together on
a club-shaped, unbranched carpophore swollen at
the apex, up to 4 cm. Drupes yellow, slightly laterally
compressed, transversely subovoid, 2.2—3 by
2.5—3.3 cm (long axis), 2—2.5 cm thick, drying finely

1986]

MENISPERMACEAE (Forman)

211

rugulose, glabrous; endocarp woody, surface cov-
ered with a dense mat of radially arranged fibres.

Distr. Hainan, Indochina, S. Peninsular Thai-
land; in Malesia: N. & Central Sumatra, Malaya (in-
cl. Langkawi), Central Java, throughout Borneo and
Philippines, Central & N. Celebes, N. Moluccas (Ta-
laud Is., Halmahera), New Guinea.

Ecol. Forests at altitudes up to 1000 m, some-
times near river banks. On limestone hill, low alti-
tudes in N. Celebes.

Vern. Thailand: hap; Sabah: takop.

Notes. Alkaloids found in this species are ber-
berine, columbamine, jatrorrhizine and palmatine,
according to THORNBER (Phytochem. 9, 1970, 160).
JEWERS ef al. (/.c. 663) reported the same alkaloids,
but not columbamine, from extracts of stems and
roots.

Details of the primary xylem elements were given
by Zamora (Philip. Agric. 50, 1966, 439, f.
899-912).

The fruits are eaten and dispersed by orang-utans,
gibbons and macaques in E. Borneo (information
from Dr. M. LEIGHTON, Harvard Univ.).

2. Arcangelisia tympanopoda (LAuT. & K. Scu.)
Diets, Pfl. R. Heft 46 (1910) 106, f. 39; FoRMAN,
Kew Bull. 32 (1978) 337. — Macrococculus tympa-

flowers

nopodus Laut. & K. Scu. Fl. Deut. Schutzgeb. Std-
see (1900) 314.

Incompletely known. Leaves apparently indistin-
guishable from those of A. flava, except for stomata
(see notes under the genus). — Male inflorescences
and flowers apparently as in A. flava. — Female
unknown. Infructescence cauliflorous,
23—60 cm, either slender, 2—3 cm 9, unbranched,
terminating in a carpophore, or branched with main
axis to 5mm; carpophore thick, claviform, to 4 cm
long, 2 cm @ at apex. Drupes yellow, subglobose,
slightly laterally compressed with a faint longitudinal
dorsal ridge running all round, 4.5—5.5 cm, surface
drying finely granular, glabrous; endocarp woody,
surface bearing an interrupted layer of radially ar-
ranged fibres, these forming a dense dorsal ridge as
well as thin lateral, transverse plates.

Distr. Malesia: New Guinea.

Ecol. Secondary forest on clay soil, well drained
alluvial soil, from forest edge overhanging water, up
to 350 m.

Note. The difference between the two forms of
infructescence is remarkable. Since they both bear
similar large-sized fruits, it must be concluded that
they both belong to this species. The position on the
plant, e.g. whether on older or younger stems, may
determine the form of the infructescence.

13. ANAMIRTA

Co.esr. Trans. Linn. Soc. 13 (1821) 52; BENTH. in B. & H., Gen. PI. 1 (1862)
35; Miers, Ann. Mag. Nat. Hist. ser. 3, 14 (1864) 49, t. 97; Contr. Bot. 3 (1871)
49; Hook. f. Fl. Br. India 1 (1872) 98; Diets, Pfl. R. Heft 46 (1910) 108, f. 10,
40; ForMAN, Kew Bull. 32 (1978) 329. — Fig. 10h—j, 11.

Large lianes, almost entirely glabrous. Leaves palmately nerved at the base,
hairy domatia present in the axils of the nerves and main veins. /nflorescences
usually cauliflorous (male sometimes axillary), paniculate. — Male flowers
pedicellate: usually 2 minute outer sepals with 3 + 3 concave, imbricate larger in-
ner sepals, becoming reflexed; petals 0; synandrium consisting of a shortly
stalked globular cluster of c. 30—35 transversely dehiscing anthers. — Female
flowers: sepals as in male; petals 0; staminodes 6, minute; carpels 3(—4) arising
laterally from a central conical gynophore which greatly lengthens in fruit, stig-
ma recurved. Drupes subreniform-globose, style-scar sublateral, borne on short
terminal divergent branches of a thickened columnar carpophore continuous
with peduncle (i.e. flower-pedicel); pericarp very thin; endocarp woody with re-
ticulate surface, bearing 2 small sublateral perforations on the ventral (concave)
side, with a deeply intrusive bilobed condyle around which the subhemispherical
seed lies, each lobe of the condyle being hollow and leading to one of the exter-
nal perforations; endosperm copious; embryo with divaricate, thin, foliaceous
cotyledons much larger than the radicle.

D2 FLORA MALESIANA [ser. I, vol. 102

Fig. 10. Coscinium fenestratum (GAERTN.) CoLeBR. da. Habit, male plant, x 2/3, b. male flower, x 8, c. inner

sepal, inner view, x 10, d. stamens, x 20, e. infructescence, x 2/3, f. LS of drupe, x 2/3. — C. blumeanum

Miers ex Hook.f. & TH. g. Female flower, front sepals removed, x6. — Anamirta cocculus (L.) WiGcuHtT &

Arn. h. Part of infructescence with branched carpophore, nat. size, i. LS of drupe, x 1 1/2, /. half of drupe

without seed showing one lobe of bilobed hollow condyle, x1 1/2 (a vAN Batcooy 2426, b—d DE WILDE
12588, e—f KosTERMANS 13932, g MainGay 118/2, h—j Forses 3391).

1986]

MENISPERMACEAE (Forman)

213

Distr. Ceylon and India to Indochina; throughout Malesia. Monotypic.
Palyn. Pollen of Anamirta (cocculus) was described by FERGUSON (Kew Bull. 32, 1978, 340).

1. Anamirta cocculus (L.) WiGHT & Arn. Prod. |
(1834) 446; Arn. Ann. Sci. Nat. ii, 2 (1834) 65, t. 3;
in Hook. f. & Th., Fl. Ind. (1855) 185; Mig. Fl. Ind.
Bat. 1, 2 (1858) 78; Ann. Mus. Bot. Lugd.-Bat. 4
(1868) 80; Hook.f. Fl. Br. India 1 (1872) 98; SCHEFF.
Nat. Tijd. N. I. 32 (1873) 395; Becc. Malesia 1 (1877)
143; Hemst. Rep. Chall. Bot. 1 (1885) 118; ENGL.
Bot. Jahrb. 7 (1886) 455; K. Sco. & Hoitr. FI. Kai-
ser Wilh. Land (1889) 44; K. Scu. Notizbl. Berl.-
Dahl. 2 (1898) 116; BoeRL. Cat. Hort. Bog. 1 (1899)
37; Drets, Pfl. R. Heft 46 (1910) 108, f. 10, 40; Back.
Schoolfl. (1911) 41; Diets in Elmer, Leafl. Philip.
Bot. 4 (1911) 1165; Koorp. Exk. Fl. Java 2 (1912)
232; Merr. Int. Rumph. (1917) 221; Sp. Blanc.
(1918) 145; W.H. Brown, Minor Prod. Philip. For.
1 (1920) 375; ibid. 3 (1921) 79, 185; Merr. En. Phi-
lip. 2 (1923) 145; Heyne, Nutt. Pl. (1927) 620; Ka-
NEH. & Hatus. Bot. Mag. Tokyo 56 (1942) 471; Ya-
MAMOTO, J. Soc. Trop. Agric. 16 (1944) 37; Back. &
Baku. f. Fl. Java 1 (1963) 156; FoRMAN, Kew Bull.
32 (1978) 329; PANcHOo, Vasc. Fl. Mt Makiling 1
(1983) 281. — Natsiatum RHEEDE, Hort. Mal. 7
(1688) t. 1. — Tuba baccifera RumpH. Herb. Amb.
5 (1747) 35, t. 22. — Menispermum cocculus LINNE,
Sp. Pl. (1753) 340; GaERTN. Fruct. 1 (1788) 219, t.
70; Roxs. Fl. Ind. ed. Carey 3 (1832) 807; BLANCO,
Fl. Filip. ed. 1 (1837) 809. — Menispermum lacuno-
sum LAMK, Encycl. Méth. 4 (1797) 98. — Cocculus
populifolius Dc. Syst. 1 (1817) 519; Prod. 1 (1824)
97; Decne, Nouv. Ann. Mus. Paris 3 (1834) 423;
Span. Linnaea 15 (1841) 163. — A. jucunda MIERS,
Contr. Bot. (1871) 55. — Fig. 10h—j, 11.

For complete synonymy, see FORMAN (1978).

Young stems and petioles usually drying pale grey-
ish straw-coloured, striate, glabrous. Leaves with
glabrous petioles 6—18(—26) cm, swollen at both
ends, geniculate at the base; lamina ovate to broadly
ovate, base cordate to truncate (or broadly obtuse),
apex usually acuminate, 16—28 by 10—24 cm, pal-
mately 3—5(—7)-nerved at base and with 4—S pairs of
lateral nerves running + parallel with the main, /.e.
distal, pair of basal nerves, the lateral nerves linked
together with scalariform veins; lower surface with
reticulum clearly visible and slightly raised, drying
pale straw-coloured against a pale grey-brown back-
ground, midrib and nerves very prominent; upper
surface drying slightly darker, subnitidous, midrib
prominent; glabrous on both surfaces apart from
pockets of hairs in the axils of the nerves and main
veins, thinly coriaceous. /nflorescences 16—40 cm
with lateral branches 2—5 cm, glabrous, bracteoles c.
0.5 mm long. — Male flowers with glabrous pedicels
becoming up to 2—3 mm; sepals white, yellow or pale
green, outer sepals 2, scarcely | mm long, inner sep-

als 6, broadly elliptic, 2.5—3 by 2 mm, glabrous,
apart from often minutely papillose margin; synan-
drium 1.5—2 mm long. — Female flowers: pedicels
and sepals as in male; staminodes 6, minute, scarcely
0.25 mm; carpels 3(—4), curved-ellipsoidal, 1.5—2
mm; stigma thick, recurved. Infructescences wholly
glabrous, lateral branches up to 15 cm; carpophore
(3—)6—16 mm, continuous with pedicel 8—20 mm.
Drupes white, 9-11 mm long, glabrous; endocarp
subreniform-globose, surface reticulate-rugulose
with a weak dorsal groove; cotyledons + narrowly
elliptic 5-7 mm long, 2 mm broad.

Distr. Ceylon, India, Thailand, Indochina; in
Malesia: N. Sumatra (once), E. Java (twice), Lesser
Sunda Is. (Sumba, Flores, Alor, Timor, Wetar, Da-
mar, Babar), Moluccas (Tenimber, Kei, Ceram, Sula
Is., Halmahera), Philippines (Luzon, Mindoro, Ba-
silan, Mindanao), New Guinea (incl. Aru Is.).

Ecol. Lowland, in a variety of conditions, on
banks of rivers and streams, coastal forest, savan-
nahs, on basalt, limestone and sandy soil, both in
rain-forest conditions and in seasonal climates, but
judging from the scanty occurrence in Java and Su-
matra and absence in Malaya and Borneo, with a dis-
tinct preference for seasonal conditions, it accounts
for the high frequency in the Lesser Sunda Is.

Uses. The stem produces bast-fibres. The fruits
are used as a fish-poison and are also used to kill lice
in the hair. They are a source of picrotoxin, which
has proved to be a mixed crystallizate of picrotoxi-
nin, which is a violent convulsant poison, and picro-
tin, which is very much less toxic. Picrotoxin has
been used in the treatment of schizophrenia and is an
effective antidote for barbiturate and morphine pois-
oning. A review of the chemical constituents and
pharmacological properties is given by QUISUMBING
(Medic. Pl. Philip. 1951, 290, 1030) and in Wealth of
India, Raw Materials 1 (1948) 75.

According to FLUckiGeR & HANBURY (Pharmaco-
graphia ed. 2, 1879, 31-33) the fruits have been
known in Europe at least since the 16th century when
they were being imported via Alexandria and other
centres in the Middle East. They are well figured in
Gerarve’s Herbal of 1597. In the 1633 edition, p.
1548 he stated that they were ‘well known in shoppes
by the name of Cocculus Indicus, some call them
Cocci Orientales . . . They are used with good suc-
cess to kill lice in children’s heads... . In England
we use the fruit called Cocculus Indi in pouder mixed
with flower, hony, and crummes of bread to catch
fish with, it being a numming, soporiferous, or sleep-
ing medicine, causeth the fish to turn up their bellies,
as being senceless for a time.’ In 1635 the fruits were
subject in England to an import duty of 2s. per

[ser. I, vol. 102

FLORA MALESIANA

Fig. 11. Anamirta cocculus (L.) Wicut & ARN. in flower (male). Central Thailand, Saraburi (Photogr. H.

BANZIGER).

1986]

MENISPERMACEAE (Forman)

215

pound. Hooker/. & THOMSON (1855) reported of the
fruits that ‘in England they are extensively used in the
adulteration of beer.’

Vern. Sumatra: waran pisang, Alas; Lesser Sun-
da Is.: Kruppe, Sumba; Philippines: array, Minda-
nao, /agtang, ligtang, Luzon.

Notes. The anomalous stem-structure was de-

scribed by SANTOS (Philip. J. Sc. 44, 1931, 385—407).
Details of the primary xylem elements were given by
ZAMORA (Philip. Agric. 50, 1966, 437—440, f.
961—973).

According to BRown (1920, /.c.) the flowers are
fragrant; the field notes on some Philippine speci-
mens describe the odour as unpleasant.

14. COSCINIUM

Coepsr. Trans. Linn. Soc. 13 (1821) 51; Hook. f. & Tu. Fl. Ind. 1 (1855) 177;
BENTH. in B. & H. Gen. PI. 1 (1862) 35; Miers, Contr. Bot. 3 (1871) 19; HooKk.
f. Fl. Br. India 1 (1872) 98; Drets, Pfl. R. Heft 46 (1910) 110; ForMAN, Kew
Bull. 32 (1978) 324. — Fig. 10a—g.

Large lianes. Leaves often peltate, palmately nerved, lamina tomentellous,
often whitish below. /nflorescence supra-axillary or ramiflorous, composed of
a raceme of peduncled + globose heads of flowers. — Male flowers: sepals 9,
imbricate in 3 whorls, externally sericeous; petals 0; stamens 6, the outer 3 free
with 1-locular introrse anthers, the inner 3 with connate filaments and with
2-locular latrorse anthers. — Female flowers: sepals as in male; petals 0; stami-
nodes 6; carpels 3; densely pilose, style filiform recurved. Jnfructescence with
globose carpophore. Drupes (only known in C. fenestratum) subglobose, to-
mentellous, style-scar sublateral, endocarp covered with anastomosing fibrous
ridges, condyle deeply intrusive, thickly clavate and containing 2 ducts, each
linking the seed-cavity with a pore on the basal surface of the endocarp; seed
subglobose, hollow, enveloping the condyle, endosperm surrounding the divari-
cate, folded and divided cotyledons.

Distr. Two species: Ceylon, India, Thailand, Indochina; in W. Malesia: Malaya, Sumatra, W. Java,
Borneo.
Palyn. Pollen of Coscinium was described by FerGusoN (Kew Bull. 32, 1978, 342).

KEY TO THE SPECIES

1. Lamina + broadly ovate, less than 1 1/2 times as long as broad, peltate or not; upper surface drying fairly
smooth. Male flowers in several-flowered heads c. 7 MM © ......-.-. 50 eee eeeeees 1. C. fenestratum
1. Lamina elongate, more than 1 1/2 times as long as broad, peltate with petiole inserted 1.5 to 5 cm from
margin; upper surface often drying rugose with main nerves markedly impressed. Male flowers in many-
nen heads e119. sii Oi) iid, 0.20 RG.) . . Pode ae es ba 2. C. blumeanum

1. Coscinium fenestratum (GaerTN.) CoLepr. 42; Diets, Pfl. R. Heft 46 (1910) 113, f. 41H—N;

Trans. Linn. Soc. 13 (1821) 65; Hook. Bot. Mag.
(1852) t. 4658; Hook. f. & Tu. Fl. Ind. (1855) 178;
Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 80; Miers,
Contr. Bot. 3 (1871) 22, t. 88; Hook.f. & Tu. Fl. Br.
India | (1872) 99; Scnerr. Nat. Tijd. N. I. 32 (1873)
395, t. 5; Becc. Malesia | (1877) 144; Kina, J. As.
Soc. Beng. 58, ii (1889) 381; Boer. Cat. Hort. Bog.
1 (1899) 38; Ror. J. Str. Br. R. As. Soc. n. 33 (1900)

Back. Schoolfl. (1911) 41; Merr. En. Born. (1921)
248; Philip. J. Sc. 29 (1926) 367; Yamamoto, J. Soc.
Trop. Agric. 16 (1944) 38, incl. var. macrophyllum
Yamamoro et var. ovalifolium YAMAMOTO, l.c. 39;
Back. & Baku. f. Fl. Java 1 (1963) 156; FORMAN,
Kew Bull. 32 (1978) 325. — Menispermum fenestra-
tum GAERTN. Fruct. | (1788) 219, t. 45, f. 5. —C.
wallichianum Miers [Ann. Mag. Nat. Hist. ser. 2, 7

216

FLORA MALESIANA

[ser. I, vol. 102

(1851) 37, nomen], Contr. Bot. 3 (1871) 23; DIELs,
Pfl. R. Heft 46 (1910) 112; Rrpv. Fl. Mal. Pen. 1
(1922) 106; Burk. Dict. (1935) 669. — C. blumea-
num (non Miers ex Hook. f. & Tu.) Mia. Fl. Ind.
Bat. 1, 2 (1858) 77, p.p.; Hook.f. & TH. Fl. Br. India
1 (1872) 99, p.p.; ScHEFF. Nat. Tijd. N. I. 32 (1873)
396, t. 6; BeEcc. Malesia | (1877) 144; Kina, J. As.
Soc. Beng. 58, ii (1889) 381, p.p.; BoeRL. Cat. Hort.
Bog. 1 (1899) 38, incl. var. epeltatum BOERL.; RIDL.
Fl. Mal. Pen. 1 (1922) 106, p.p.; HEYNE, Nutt. Pl.
(1927) 621; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 38. — C. maingayi Pierre, Fl. Coch. (1885)
sub t. 112 (‘mangayi’, sphalm.). — C. miosepalum
Diets, Pfl. R. Heft 46 (1910) 113; Back. Schoolfl.
(1911) 41; Koorp. Exk. Fl. Java (1912) 232; Back.
& Baku. f. Fl. Java 1 (1963) 156. — C. peltatum
Merr. Un. Cal. Publ. Bot. 51 (1929) 59. — Fig.
10a—f.

Large liane with yellow wood and sap. Branchlets
terete, obscurely ridged or smooth, brownish tomen-
tose at first, later glabrescent, becoming whitish,
bearing prominent disciform petiole-scars. Leaves:
petioles at first brownish tomentose, 3—16 cm, often
conspicuously swollen at both ends, geniculate at
base, inserted up to 0.8(—2.7) cm from basal margin
of lamina; lamina usually broadly ovate or ovate,
rarely subpanduriform with basal, lateral lobes, base
broadly rounded, truncate or shallowly cordate,
rarely broadly obtuse, apex acuminate, 11—33 by
8—23 cm; upper surface glabrescent, usually drying
smooth, midrib and other main nerves sunken, lower
surface often whitish tomentellous with fine reticula-
tion visible, palmately 5—7-nerved at base and also
usually two pairs of distal lateral nerves, thinly coria-
ceous. Inflorescences: flowers in several-flowered
globose heads 6—7 mm @ on peduncles 10—30 mm,
arranged in a raceme 5—11 cm, supra-axillary or
from older, leafless stems; inflorescences arising
singly or a few together, axis and branches slender,
brown tomentose or tomentellous, bracts subulate,
4—5 mm long. — Male flowers sessile or with pedi-
cels up to 1 mm; sepals densely sericeous-pilose exter-
nally, glabrous within, broadly elliptic to obovate,
the inner 3—6 spreading, yellow, 1.5—2 mm long;
outermost sepals smaller, 1—1.5 mm long, inserted
lower; stamens 6, 1 mm long. — Female flowers: sep-
als as in male flowers; staminodes 6, claviform, 1 mm
long; carpels 3, curved-ellipsoidal, 2 mm long, dense-
ly pilose; style filiform, recurved. Infructescence
with carpophore globose, tomentellous, 7-8 mm @,
bearing 1—3 drupes. Drupes subglobose, tomentel-
lous, brown to orange or yellowish, 2.8—3 cm g; peri-
carp drying woody, c. 1 mm thick; endocarp bony,
2.2—2.5 cm 9; wall 3 mm thick covered with anasto-
mosing fibrous ridges; condyle deeply intrusive,
thickly clavate. Seed whitish, subglobose, enveloping
the condyle; endosperm present and within this are

immersed the divaricate, much folded and divided
cotyledons.

Distr. Ceylon, S. India, Cambodia, Vietnam; in
Malesia: Malaya, Sumatra (incl. Banka), W. Java,
Borneo.

Ecol. Primary lowland forest; soils include gra-
nitic sand (Banka) and sandstone (Sabah).

Uses. The wood produces a yellow dye (used to-
gether with Curcuma in Cambodia). A decoction of
the stem and leaves is used medicinally. It has been
known in Europe as False calumba, being a substi-
tute for Calumba (Jateorhiza). The plant has alleged
antiseptic properties and is used in Malaya to dress
wounds and ulcers. The species is used as an ingre-
dient for arrow poisons in Malaya (Bisset & Woops,
Lloydia 29, 1966, 194). According to the notes on S
32149 and 33332 from Sarawak, the effects of intoxi-
cation can be avoided if the roots are chewed and the
juices swallowed before drinking.

GRESHOFF (Meded. Lands Pl. Tuin 25, 1898, 22)
recorded that the leaves contain picrotoxin-like bitter
substances. Like most previous authors, GRESHOFF
wrongly applied the name C. b/umeanum to the pres-
ent species. Alkaloids found by JEwerset al. (Phyto-
chem. 9, 1970, 663) were: palmatine, berberine and
jatrorrhizine. The record of the same three alkaloids
by THORNBER (Phytochem. 9, 1970, 163) under C.
blumeanum probably refers to this species. Further
alkaloids were reported by Stwon et al. (Planta Me-
dica 38, 1980, 24).

Vern. Malaya: akar kuning, (kunyit-kunyit) ba-
bi, kopak, kupak, tol; Banka: akar kunyit; Java:
akar kuning; Borneo: abang asuh, Sabah, binap ko-
kop, upak-upak, E. Borneo, perawan, dipang, Sara-
wak.

Note. The fruits are eaten and dispersed by
orang-utans, gibbons and macaques in E. Borneo
(information from Dr. M. LeEicHton, Harvard
Univ.).

2. Coscinium blumeanum Miers [Ann. Mag. Nat.
Hist. ser. 2, 7 (1851) 37, nomen] ex Hoox. f. & Tu.
FI. Ind. (1855) 179; Mia. FI. Ind. Bat. 1, 2 (1858) 77,
p.p.; Miers, Contr. Bot. 3 (1871) 23; Hook.f. & Tu.
Fl. Br. India 1 (1872) 99, p.p.; Kine, J. As. Soc.
Beng. 58, ii (1889) 381, p.p.; Diets, Pfl. R. Heft 46
(1910) 113, f. 41A—G; Rit. Fl. Mal. Pen. 1 (1922)
106, p.p.; FORMAN, Kew Bull. 32 (1978) 328. — Fig.
10g.

Large liane. Branchlets terete, obscurely ridged or
smooth, pale fawn to brownish at first, later glabres-
cent and whitish, bearing prominent disciform
petiole-scars. Leaves: petioles whitish tomentose,
6—20 cm, conspicuously swollen at both ends, insert-
ed 1.5—5 cm from basal margin of lamina; lamina
oblong, lanceolate-oblong or narrowly ovate, occa-
sionally subpanduriform, base broadly rounded or

1986]

truncate, apex acuminate, 15—35 by 6—20 cm, above
glabrous, often drying + bullate, beneath whitish to-
mentellous with fine reticulation visible, palmately
7—11-nerved at base with 2—3 pairs of distal lateral
nerves, thinly coriaceous. /nflorescences: flowers in
globose (or slightly elongate) densely and ~-flowered
heads 10-13 mm @ on peduncles 10—25 mm ar-
ranged in araceme 12—14 mn, supra-axillary or aris-
ing from older leafless stems, axis and branches of in-
florescence stout and brown tomentose, bracts in-
conspicuous, scale-like, 1—2 mm long. — Male flow-
ers with pedicels 1.5—2 mm; sepals densely sericeous-
pilose externally, glabrous within, inner 3—6 spread-
ing at anthesis, broadly elliptic to spathulate-
obovate, 2.5—3 mm long, the outermost 3 elliptic,
1.5—2 mm long, inserted lower; stamens 6, 1 mm
long. — Female flowers: 6 larger inner sepals oblong

MENISPERMACEAE (Forman)

247

to oblanceolate, 4—4.5 mm long, staminodes 6, car-
pels 3 as in C. fenestratum. Drupes unknown.

Distr. Peninsular Thailand; in Malesia: Malaya
(Penang and Pangkor I.).

Ecol. In Peninsular Thailand recorded from
evergreen forest at c. 300 m, and on Terutao I. at 15
m.

Notes. This distinctive species has a restricted
distribution. The name C. blumeanum has often
been wrongly applied to specimens of the more wide-
ly distributed C. fenestratum, thus resulting in a
long-standing confusion between the two species.

There is no indication in the available field-notes
that the wood is yellow as in C. fenestratum.

Mature fruits have apparently never been collect-
ed. It would be most interesting to know how they
compared with those of the other species.

15. PACHYGONE

Miers, Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 37; Hook.f. & Tu. Fl. Ind. 1 (1855)
202; Miers, Contr. Bot. 3 (1871) 328; DieEzs, Pfl. R. Heft 46 (1910) 241, f. 80;
ForMAN, Kew Bull. 12 (1958) 457; ibid. 22 (1968) 374. — Tristichocalyx F. v.
M. Fragm. 4 (1863) 27. — Fig. 12g-i.

Woody climbers. Leaves + ovate, base 3- to S5-nerved. Inflorescences axil-
lary, pseudo-racemose. — Male flowers: sepals 6(— 12), inner ones larger, imbri-
cate; petals 6, auriculate towards base; stamens 6. — Female flowers: sepals and
petals similar to male, staminodes 6, carpels 3, glabrous, style reflexed, stigma
entire. Drupe curved with style-scar near base, subcompressed-obovoid; endo-
carp rather smooth, with a dorsal median groove and on each lateral face a small
central sublunate perforation leading to the central hollow condyle. Seed
strongly curved; endosperm absent; cotyledons large, thick.

Distr. China, SE. Asia, Malesia, Australia and Polynesia. In Malesia | sp.; 11 more have been described

from surrounding regions, but this number will probably have to be reduced.

1. Pachygone ovata (Porr.) Hook.f. & Tu. Fl. Ind.
1 (1855) 203; Mig. FI. Ind. Bat. 1, 2 (1858) 86; Miers,
Ann. Mag. Nat. Hist. ser. 3, 19 (1867) 321; Mia.
Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 87, incl. var. ro-
tundifolia Mig. et var. dasyphylla Miq.; MIERS,
Contr. Bot. 3 (1871) 331, t. 135; Hook. f. & Tu. FI.
Br. India | (1872) 105; Becc. Malesia | (1887) 158;
Bogert. Cat. Hort. Bog. | (1899) 43; K. Scu.& Laut.
Fl. Deut. Schutzgeb. Siidsee (1901) 315; VALETON,
Bull. Dép. Agr. Ind. Néerl. 10 (1907) 12; Diets, Pfl.
R. Heft 46 (1910) 243, f. 80; ForMAN, Kew Bull. 12
(1958) 457; ibid. 22 (1968) 374, with a full synonymy.
— Cissampelos ovata Poir. Encycl. Méth. Bot. 5
(1804) 10; Dc. Syst. 1 (1817) 537; Prod. 1 (1824) 102.
=- Cocculus brachystachyus De. Syst. 1 (1817) 528;
Prod. | (1824) 99; Decne, Ann. Nouv. Mus. Paris 3

(1834) 424; Span. Linnaea 15 (1841) 163. — Coccu-
lus leptostachyus De. Syst. 1 (1817) 528; Prod. 1
(1824) 99; Decne, Ann. Nouv. Mus. Paris 3 (1834)
424; Span. Linnaea 15 (1841) 163. — P. pubescens
Bentu. Fl. Austr. 1 (1863) 58; Drets, Nova Guinea
8 (1910) 283; Pfl. R. Heft 46 (1910) 245; WINKLER,
Bot. Jahrb. 49 (1913) 369; Merr. En. Born. (1921)
250; YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 103.
— P. brachystachyua (Dc.) Miers, Ann. Mag. Nat.
Hist. ser. 3, 19 (1867) 321; Contr. Bot. 3 (1871) 334.
— P. leptostachya (Dc.) Miers, Ann. Mag. Nat.
Hist. ser. 3, 19 (1867) 321; Contr. Bot. 3 (1871) 335;
Diets, Pfl. R. Heft 46 (1910) 244. — P. hebephylla
Miers [Ann, Mag. Nat. Mlist. ser. 3, 19 (1867) 321,
nomen\, Contr. Bot. 3 (1871) 333; Diets, Pfl. R.
Heft 46 (1910) 245; Back. Schoolfl. (1911) 45;

218

FLORA MALESIANA

[ser. I, vol. 102

Koorp. Exk. Fl. Java 2 (1912) 235; YAMAmoTo, J.
Soc. Trop. Agric. 16 (1944) 103; Back. & BAKuH. f.
Fl. Java 1 (1963) 159. — Limacia nativitatis Rw. J.
Str. Br. R. As. Soc. n. 45 (1906) 170. — P. zeylanica
SANT. & WAGH, Bull. Bot. Surv. India 5 (1963) 107,
nom. illeg., cf. FORMAN, Kew Bull. 22 (1968) 374. —
Fig. 12g-i.

For complete synonymy, see FORMAN (1958, 1968).

Woody climber up to 15 m or more high. Branch-
lets puberulous to pubescent with yellowish hairs, la-
ter glabrescent. Leaves: petioles 2.5—4 cm, yellowish
pubescent to glabrous; lamina ovate to ovate-lanceo-
late or broadly ovate, (S—)7—11 by 3—7.5(—12) cm,
apex obtuse to acutely acuminate, mucronate, base
truncate, rounded or very obtuse (rarely subcordate),
3—5-nerved; lateral nerves 1—2 pairs; both surfaces
softly pubescent to glabrous; papyraceous. /nflores-
cences axillary, solitary or 2—3 arising together,
pseudo-racemose with flowers often in small clusters
of about 3—5 along main axis, 4—17 cm long, yellow-
ish tomentose to pubescent. — Male flowers yellow
on pedicels 1—3(—7) mm; sepals 6(—12), outer 3(—6)
bracteoliform, + elliptic, 1—2 mm long, puberulous
outside, inner 3(—6) elliptic to rotund, 1.5—2.5 by
1.25—1.5 mm, glabrous or sparsely hairy outside;

petals 6, + oblong, with basal auricles clasping op-
posite stamen, 1—1.5 mm long, glabrous; stamens 6,
filaments slender, 1.25—1.5 mm. — Female flowers:
sepals and petals similar to male, carpels 3, + ovoid,
0.75 mm long, style flattened, staminodes 6, minute.
Drupes on pedicels c. 3—5 mm long, subcompressed,
+ obovoid, 7—8 by 6 mm, 5 mm thick, glabrous,
rather smooth.

Distr. Ceylon, India, through Malesia to NE.
Australia; in Malesia: N. Borneo, Java (incl. Christ-
mas I.), Lesser Sunda Is. (Timor), S. Celebes, Mo-
luccas (Sula Is.: Mangoli), New Guinea.

Ecol. On seashores and in lowland forests, up to
100 m, also in seasonal regions.

Vern. Java: geureung ikan; N. Borneo: luod, Ba-
jau dial.

Notes. Dries maintained as distinct 4 of the spe-
cies listed above, viz. P. ovata, P. pubescens, P.
leptostachya and P. hebephylla.

It appears that there is, in fact, only one species of
Pachygone in Malesia with a variable leaf shape, be-
coming broader eastwards, and with considerable
variation in the degree of hairiness on the leaves. The
inflorescence, floral and fruit characters are rather
constant.

16. HYPSERPA

Miers [Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 40, nomen], ibid. ser. 3, 14 (1864)
363; Contr. Bot. 3 (1871) 100; Dies, Pfl. R. Heft 46 (1910) 205; Forman, Kew
Bull. 12 (1958) 451. — Selwynia F. v. M. Fragm. 4 (1864) 153. — Fig. 12a—f.

Scandent shrubs or woody climbers. Stems with young growing tips some-
times tendrilliform. Leaves + ovate to elliptic, base 3—7-nerved with the side
nerves sometimes supra-basal. Inflorescence axillary or supra-axillary, cymose
or thyrsoid. — Male flowers: sepals 7—12, spirally arranged, glabrous or sub-
glabrous, outer ones minute and bracteoliform, inner ones larger, imbricate;
petals 5—9, fleshy; stamens 9—40, free or connate. — Female flowers: sepals
and petals similar to male; staminodes 0—several; carpels 2—3, stigma entire, re-
flexed. Drupes curved, subcompressed-obovoid to -globose with style-scar near
base; endocarp laterally convex with 2 lateral cavities each with an external aper-
ture, dorsally rugose to rugulose. Seed horseshoe-shaped, narrow, embedded in
endosperm.

Distr. From China and tropical SE. Asia (incl. Ceylon) to Australia and Polynesia; throughout Malesia.
In all 6 spp.
KEY TO THE SPECIES

APH Orescences ela bLOus) Wace Gos eee clots. . e URE Bde shag OR eee ee 1. H. laurina
1. Inflorescences hairy.

1986]

MENISPERMACEAE (Forman)

219

2. Leaf-base mostly 3-nerved. Inflorescences 0.5—1.5(—2.5) cm broad. Stamens 9—10(—14), free for most of

USGI? [EN TEU Teas eet de ee eo oe

2. H. nitida

2. Leaf-base mostly 5—7-nerved. Inflorescences 1.5—5 cm broad. Stamens about 20—40, connate

1. Hypserpa laurina (F. v. M.) Diets, Pfl. R. Heft 46
(1910) 209, f. 72; ForMAN, Kew Bull. 12 (1958) 452.
— Selwynia laurina F. v. M. Fragm. 4 (1864) 153. —
H. selwynii F. v. M. Fragm. 9 (1875) 82. — Limacia
selwynii (F. v. M.) BAILEy, Queensl. Fl. 1 (1899) 30.
— H. parvifolia KANEH. & Hatus. Bot. Mag. Tokyo
56 (1942) 471.

Woody climbers, entirely glabrous (bracts and
axillary buds sometimes puberulous). Leaves: peti-
oles 1—1.5(—3) cm; lamina elliptic to oblong-elliptic,
7-12 by 3—4.5 cm, apex obtuse, often minutely
emarginate, base broadly cuneate to rounded, +
5-nerved; lateral nerves 2—3 pairs; stiffly papyra-
ceous. — Male inflorescences thyrsoid, very lax,
4—15(—20) by 1—4(—9) cm. — Male flowers green or
white on pedicels 1-5 mm long; sepals 7—9, outer
2—3 about | mm long, inner, 5—6 + rotund, 1.5—2
mm Q; petals (rarely S—)7—9, + obtriangular 0.75—1
mm long; stamens 11—15, free (but with occasional
connate pairs). — Female inflorescences and flowers
unknown. [/nfructescence 6—8 cm long, teste DIELs.]
Drupes red on pedicels about 5 mm, broadly obovate
in outline, 12 by 9 mm, 8 mm thick. Endocarp dor-
sally obscurely rugose, laterally convex and smooth.

Distr. Queensland and E. Malesia (SW. Papua).

Ecol. Rain-forest and /mperata fields.

2. Hypserpa nitida Miers in Hook., Kew J. Bot. 3
(1851) 258; Contr. Bot. 3 (1871) 102; Drets, Pfl. R.
Heft 46 (1910) 210; FormMaAN, Kew Bull. 12 (1958)
453; ibid. 22 (1968) 361. — [Cocculus cuspidatus
WALL. Cat. n. 4960 (1831/32), nomen.] — Limacia
cuspidata Hook. f. & Tu. Fl. Ind. 1 (1855) 189; FI.
Br. India | (1872) 100; ScHeFF. Nat. Tijd. N. I. 32
(1873) 397, t. 8. — Limacia microphylla Mig. FI.
Ind. Bat., Suppl. 1 (1861) 386. — H. cuspidata
(Hook. f. & TH.) Miers, Ann. Mag. Nat. Hist. ser.
3, 14 (1864) 365; Contr. Bot. 3 (1871) 102, t. 108;
Becc. Malesia | (1877) 149; Boer. Cat. Hort. Bog.
(1899) 39, incl. var. microphylla (MiQ.) BOERL.;
Diets, Pfl. R. Heft 46 (1910) 207; in Elmer, Leafl.
Philip. Bot. 4 (1911) 1165; Merr. En. Born. (1921)
249; Rip. Fl. Mal. Pen. 1 (1922) 111; Merr. En.
Philip. 2 (1923) 147; Burk. Dict. (1935) 1219; Yama-
moto, J. Soc. Trop. Agric. 16 (1944) 97. — Limacia
borneensis Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868)
83. — H. heteromera Miers [Ann. Mag. Nat. Hist.
ser. 3, 14 (1864) 365, nomen], Contr. Bot. 3 (1871)
104; Diets, Pfl. R. Heft 46 (1910) 210; Merr. En.
Born. (1921) 249; Yamamoro, J. Soc. Trop. Agric.
16 (1944) 98. — H. praevaricata Miers [Ann. Mag.
Nat. Hist. ser. 3, 14 (1864) 365, nomen], Contr. Bot.

3. H. polyandra

3 (1871) 103. — H. propensa Miers [/.c. 365, no-
men}, /.c. 105; BEcc. Malesia | (1877) 149. — [H. tri-
flora (non (Dc.) Miers) Miers, Ann. Mag. Nat.
Hist. ser. 3, 14 (1864) 365; Contr. Bot. 3 (1871) 106,
p.p. non quoad Cocculus triflorus Dc.: KiNG, J. As.
Soc. Beng. 58, ii (1889) 384; Ripi. J. Str. Br. R. As.
Soc. n. 33 (1900) 42.] — H. borneensis (MiQ.) BEcc.
Malesia | (1877) 149. — AH. jagorii Diets, Pfl. R.
Heft 46 (1910) 211; in Elmer, Leafl. Philip. Bot. 4
(1911) 1165; MerrR. En. Philip. 2 (1923) 148. — Phy-
tocrene loheri MERR. Philip. J. Sc. 7 (1912) Bot. 293;
En. Philip. 2 (1923) 492, p.p., leaves only. — H. nan-
dinifolia YAMAMOTO, Trans. Nat. Hist. Soc. Taiwan
34 (1944) 310, f. 5. — Fig. 12a—-f.

Scandent shrub. Branchlets sparsely to densely pu-
bescent with yellowish hairs when young, later gla-
brescent. Leaves: petioles 0.8—2 cm, sparsely to
densely pubescent; lamina very variable in shape,
lanceolate, lanceolate-elliptic, broadly elliptic or
ovate (rarely narrowly oblong-elliptic), 4—12 by
1.5—7 cm, apex obtusely (rarely acutely) acuminate
or obtuse, usually mucronulate, base obtuse to
rounded or truncate (rarely acute), usually 3-nerved
with the side nerves supra-basal; main lateral nerves
c. 2 pairs, often indistinct; both surfaces usually gla-
brous, sometimes pubescent (especially in Celebes
and the Philippines); papyraceous to subcoriaceous.
Inflorescences cymose to paniculate, 1—4(—12) by
0.5—1.5(—2.5) cm, yellowish pubescent. — Male
flowers yellow on pubescent pedicels 1—2 mm; sepals
7—11, outer ones minute and bracteoliform, subtri-
angular, scarcely 1 mm long, puberulous outside, in-
ner 4—5 + ovate-rotund, 2.5 by 1—1.75 mn, gla-
brous; petals 5, obovate, c. 1 mm long; stamens 9—10
(—14), connate at the base only, free above, 1—1.75
mm long. — Female flowers: sepals and petals as in
male; carpels 2, + ellipsoid, scarcely 1 mm long; stig-
ma plate-like. Drupes yellow to red on pedicels, 1—4
mm, subglobose, (S—)6—8 mm @. Endocarp rugu-
lose, sometimes perforate.

Distr. Ceylon, India, Assam, Lower Burma,
Thailand, Indochina; in Malesia: S. Sumatra (incl.
Banka and Riouw), Malaya (incl. Langkawi, Penang
and Singapore), Borneo, Philippines (throughout)
and Celebes.

Ecol. Forest, from sea-level to 2000 m.

Vern. Banka: akar sentjaw; Malaya: akar min-
jak; N. Borneo: akar suganda; Philippines: /alapau,
mamana, Sub.

Note. The size, shape and degree of hairiness of
the leaves of this species vary considerably. In gener-
al the leaves tend to be more hairy in Celebes and

220 FLORA MALESIANA [ser. I, vol. 102

Fig. 12. Hypserpa nitida Mrers.a. Habit, male plant, x 2/3, b. male flower, x 8, c. female flower, front sep-
als and petal removed, x 8, d. drupe, e. endocarp, f. CS of endocarp showing the central 2-chambered condyle
and the seed (white) cut through twice, all x 3. — Pachygone ovata (Por.) Hook.f. & TH. g. Drupe, h. endo-
carp, 7. LS of endocarp showing the seed curved around the hollow condyle, all «3. — Limacia scandens
Lour./j. Habit, male plant, x 2/3, k. leaf, «2/3, /. bud, x8, m. male flower, x 8, n. petal, x 15, o. stamen,
x 15, p. drupe, g. endocarp, lateral and median views, r. LS of endocarp, all nat. size (a SAN A492, bSAN
32243, c CLEMENS 40880, d—f SAN 22153, g—i BRANDERHORST 136, j, /—o RmpDLEY 10182, & SF F732, p—r
CHEW WEE LEK 246).

1986]

Philippines, while further west they are usually gla-
brous. The inflorescences are usually short (1—4 cm)
but they can become very long (up to 12 cm) in Phi-
lippine specimens and in those from Celebes. There
are all kinds of intermediates between the small, gla-
brous leaved form with short inflorescences and the
larger, glabrous or hairy leaved form with long inflo-
rescences. It has therefore not been possible to main-
tain more than one species, nor has it been possible
to recognise distinct infraspecific taxa.

3. Hypserpa polyandra Becc. Malesia | (1877) 148;
Wars. Bot. Jahrb. 13 (1891) 315; K. Sco. & Laut.
Fl. Deut. Schutzgeb. Siidsee (1901) 315; Diets, Pfl.
R. Heft 46 (1910) 213; Wuite, J. Arn. Arb. 10 (1929)
212; ForMAN, Kew Bull. 12 (1958) 455; ibid. 22
(1968) 360. — H. selebica Becc. Malesia 1 (1877)
148; Diets, Pfl. R. Heft 46 (1910) 212. — Limacia
monilifera Burk. in Hook., Ic. Pl. (1899) 2585. —
H. monilifera (BuRK.) Diets, Pfl. R. Heft 46 (1910)
209. — H. raapii Dre1s, /.c. 212; YAMAMoToO, J. Soc.
Trop. Agric. 16 (1944) 98. — H. latifolia Mia. ex
Drets, Pfl. R. Heft 46 (1910) 213.

Scandent shrub or woody climber up to c. 40 m.
Branchlets minutely yellowish puberulous to glabres-
cent or tomentose. Leaves: petioles 2—4.5(—6) cm,
puberulous or tomentose; lamina ovate to ovate-el-
liptic, 6-17 by 4—11 cm, apex shortly and often ab-
ruptly acuminate, or obtuse, sometimes mucronu-
late, base obtuse to rounded and sometimes abruptly
cuneate, 5—7-nerved; lateral nerves 1—3 pairs; both
surfaces sparsely puberulous to glabrous or tomen-
tose; stiffly papyraceous to coriaceous. /nflores-
cences pseudo-paniculate, 2.5—11 by 1.5—5 cm, +
triangular in outline, yellowish puberulous or tomen-
tose. — Male flowers yellow, subsessile or with pedi-
cels up to 1 mm; sepals 7—12, outer 2—5 + triangular
to rotund, 0.5—1 mm long, puberulous outside, inner
(4—)5—7 + rotund, 2—3 mm @, margin sometimes
minutely ciliolate; petals (S—)7—8, very variable in

MENISPERMACEAE (Forman)

221

shape, 0.75—1 mm long, margin often undulate; sta-
mens c. 20—40, connate, 1.5—2 mm long. — Female
flowers: sepals and petals similar to male; carpels 3,
+ ellipsoid, 0.75 mm long. Drupes white or red on
pedicels up to 2 mm long or subsessile, subrotund to
obovate in outline, 7—8 mm long and broad, 4—6
mm thick. Endocarp rugulose, perforate.

Distr. Australia (Queensland), New Hebrides,
Solomons, Carolines; in Malesia: New Guinea (incl.
Aru Is.), Moluccas (Ternate), SE. Celebes, Lesser
Sunda Is. (Flores, Timor), W. Sumatra (Batu Is.).

Ecol. Mangrove swamps and lowland mixed
rain-forest, up to 1200 m. Fr. July, Nov., f/. March,
May, July, Nov.

Uses. Bark after being pounded and powdered is
applied to the head as a treatment for headaches in
Bougainville I.

Notes. Hypserpa raapii was distinguished by
Diets from H. polyandra by the number of inner sep-
als being 4 in the former and 6 in the latter. H. raapii
is still only represented by the solitary, male type spe-
cimen (RAAP 607) which does indeed usually have 4
inner sepals, but 5 also occur. Since (4—)5—7 inner
sepals are found in H. polyandra, and RaAap 607
agrees in all other respects with that species it is not
possible to maintain H. raapii as a distinct species.

This results in a rather curious distribution for H.
polyandra, since RAAP 607 was collected in the Batu
Is. west off Sumatra while H. polyandra is only
known otherwise in S. and E. Malesia.

var. tomentosa FORMAN, Kew Bull. 22 (1968) 360.
Differs from the type variety in having tomentose
branches, leaves (mainly beneath) and inflorescen-
ces, these being in the type variety glabrescent or mi-
nutely puberulous.
Distr. East New Guinea (Morobe Distr.).
Ecol. Regrowths and near rivers and lakes,
70—1200 m.
Vern. Brewa, Herzog Ra., Buang dial.

17. LIMACIA

Lour. Fl. Cochinch. (1790) 620; Hook. f. & Tu. Fl. Ind. 1 (1855) 187, p.p.;
Miers, Contr. Bot. 3 (1871) 108; Diets, Pfl. R. Heft 46 (1910) 213; ForMAN,
Kew Bull. 12 (1958) 447. — Fig. 12j-r, 13.

Woody climbers. Stems with young growing tips sometimes tendrilliform.
Leaves with base 3- or 5-nerved. Inflorescences axillary or supra-axillary, cy-
mose or pseudo-paniculate. — Male flowers: sepals 6 (or 9), in whorls of 3, to-
mentose, inner whorl valvate and larger than outer whorl(s); petals 6, concave;
stamens 6, free; rudimentary carpels 3 or 0. — Female flowers: sepals and petals
similar to male; staminodes 6 or 0; carpels 3, tomentose, style reflexed, stigma
entire. Drupes with style-scar near base, subcompressed-obovoid, abruptly nar-

222 FLORA MALESIANA [ser. I, vol. 102

‘ «gh ’ F “ -

Fig. 13. Habit of Limacia blumei (BoERL.) DiELs in the Botanic Gardens, Bogor (X VI.D.7) (Photogr. L.L.
FORMAN).

rowed at base into a short stipe; endocarp laterally convex containing 2 large lat-
eral cavities each with a large external aperture and separated internally by a sep-
tum with a small central hole, dorsally bearing a raised longitudinal band, sur-
face smooth or slightly rugose. Seed horseshoe-shaped, narrow, embedded in
endosperm.

Distr. Tropical SE. Asia (Lower Burma, Thailand, Indochina); in Malesia: Sumatra, Malaya, Borneo
(W., Sarawak, Sabah), Java, Lesser Sunda Is. (Sumba, Timor), SE. Celebes (Buton I.), Philippines (Minda-
nao); 3 spp.

Note. Hooker f. & THOMSON considered Hypserpa as a synonym of Limacia, but nearly all subsequent
authors have retained the two genera as distinct. Limacia differs from Hypserpa in having the sepals in distinct
whorls of three, those of the inner whorl being valvate. In Hypserpa the sepals are all imbricate, irregular in
number and not arranged in whorls.

1986 |

MENISPERMACEAE (Forman)

223

KEY TO THE SPECIES

1. Leaf-base 5-nerved, petals tomentose ............

1. Leaf-base 3-nerved, petals glabrous.

1. L. blumei

2. Young stems velvety tomentellous. Inflorescence a lax pseudo-panicle, male up to 16 cm long. Drupes

i eerninlong | fet! 65859 Sas ee eee. .

AFOUL AT CORE AE 2. L. oblonga

2. Young stems tomentose. Inflorescence a peduncled congested cyme, less than 3 cm long. Drupes 20—25

TTT? a A Rie dae de Me ek ae ae

1. Limacia blumei (BoERL.) Diets, Pfl. R. Heft 46
(1910) 215; Back. Schoolfl. (1911) 43; MerRR. Philip.
J. Sc. 7 (1912) Bot. 266; En. Philip. 2 (1923) 148; Ya-
MAMOTO, J. Soc. Trop. Agric. 16 (1944) 99; FoRMAN,
Kew Bull. 12 (1958) 448; ibid. 22 (1968) 362. — Coc-
culus blumei BoERL. Cat. Hort. Bog. 1 (1899) 40. —
Fig. 13.

Woody climber up to 15 m or more high. Branch-
lets densely puberulous at first, later glabrescent.
Leaves: petioles 2—5 cm, puberulous; lamina ovate
to elliptic-ovate, 10—26 by 5—16 cm, apex acute with
long mucronate acumen, base cunate to rounded,
5-nerved; lateral nerves 2—3 pairs, sparsely to moder-
ately puberulous and very finely reticulate on both
surfaces; papyraceous. /nflorescences cymose, few
flowered, 0.75—1.5 cm long, puberulous. — Male
JSlowers cream on pedicels 1—2 mm; sepals 6 or 9, to-
mentose, outer 3 or 6 minute up to 0.75 mm long, in-
ner 3 elliptic, 2 by 1.25 mm; petals 6, tomentose,
broadly elliptic, 1 mm long; stamens 6, sublinear,
glabrous, 1.5 mm; rudimentary carpels 3. — Female
flowers (from Ramos & EDANO 49144): sepals and
petals larger than in male; carpels 3, obliquely subel-
lipsoid, 2 mm long, style elongate, grooved, stami-
nodes 6, linear, 1 mm. Drupes unknown.

Distr. Thailand (Lower) and Malesia: Borneo
(Sabah), Java, Lesser Sunda Is. (Sumba, Timor),
SE. Celebes (Buton I.), Philippines (Mindanao: Da-
vao).

Ecol. Thickets and forests at low altitude.

Note. This rare species was described in 1899
from plants cultivated in the Botanic Gardens at Bo-
gor, all said to have come from Java. One of these
plants, n. XVI.D.7, was still alive and strongly grow-
ing in 1956, when in May I collected male flowering
material from it. No other collections are known
from Java. The records from Thailand, Borneo,
Sumba and Celebes are based on single sterile collec-
tions.

2. Limacia oblonga Hook. /. & Tu. Fl. Ind. 1 (1855)
189; Mio. FI. Ind. Bat. 1, 2 (1858) 80; Miers, Contr.
Bot. 3 (1871) 109; Hook. f. & Tu. Fl. Br. India 1
(1872) 100; Kina, J. As. Soc. Beng. 58, ii (1889) 382;
Diets, Pfl. R. Heft 46 (1910) 214; Rip. Fl. Mal.
Pen. | (1922) 109; Burk. & Henn. Gard. Bull. S.S.
3 (1925) 344; Henn. ibid. 4 (1928) 220; Burk. Dict.
2 (1935) 1343; Forman, Kew Bull. 12 (1958) 448. —

» OL A ee ee ES Eee 3. L. scandens

L. distincta Miers, Contr. Bot. 3 (1871) 111, t. 109.
— L. inornata Mters, l.c. — L. velutina Hook. f. &
Tu. var. glabrescens KinG, J. As. Soc. Beng. 58, ii
(1889) 383.

Woody climber up to about 10 m. Branchlets, peti-
oles and inflorescences covered with a very short,
velvety, + yellow-brown indumentum. Leaves: peti-
oles 1.5—4 cm; lamina elliptic, oblong-elliptic,
broadly elliptic or elliptic-obovate, 9-25 by 3—10
cm, apex usually acutely acuminate, mucronulate,
base acute to rounded, 3-nerved; lateral nerves 4
pairs, prominent on lower surface; both surfaces gla-
brous apart from puberulous midrib; papyraceous.
— Male inflorescences pseudo-paniculate, often
about 3 arising together, 5—16 cm long, with very
slender branches. — Male flowers greenish yellow,
subsessile: sepals 9, + ovate, tomentose, outer 3
minute, 0.5 mm long, middle 3, 0.75 mm long, inner
3, 1.5 mm long and broad; petals 6, obovate, 0.75
mm long, glabrous; stamens 6, claviform, 0.75 mm.
— Female inflorescences similar to male but shorter,
1—4 cm long, branches thickening when in fruit. —
Female flowers: sepals and petals + as in male; car-
pels 3, obliquely ellipsoidal, 1 mm long, stigma auri-
culate; staminodes linear. Drupes yellow, obliquely
obovate in outline, 14—20 by 12—16 mm, puberulous
to glabrescent.

Distr. Lower Thailand; in Malesia: W. Malaya
(from Penang and Perak to Singapore), Sumatra (E.
Coast), Borneo (Sarawak).

Ecol. Primary and secondary forests up to about
350 m, on deep brown sandy loam in Sarawak. Fi.
Sept., Nov., fr. June, Aug., Oct.

Uses. Root extract applied to sore eyes; possibly
contains berberine (BURKILL, 1935). Fruits edible,
sweet (VAN BALGooy 2155, Malaya).

Vern. Malaya: akar china, akar kuning, akar
kunyit-kunyit.

3. Limacia scandens Lour. Fl. Cochinch. (1790) 620;
Hook. f. & Tu. Fl. Ind. (1855) 189; Miers, Contr.
Bot. 3 (1871) 109; Diets, Pfl. R. Heft 46 (1910) 214;
Merr. Trans. Am. Phil. Soc. n.s. 24 (1935) 157;
ForMAN, Kew Bull. 12 (1958) 449. — Cocculus lima-
cia De. Syst. 1 (1817) 526. — Menispermum limacia
(Dc.) SPRENG. Syst. 2 (1825) 155. — [Cocculus velu-
tinus WALL. Cat. n. 4970 (1831/32), nomen.| — L.
velutina Hook. f. & Tu. Fl. Ind. 1 (1855) 189; Mig.

224

FLORA MALESIANA

[ser. I, vol. 102

Fl. Ind. Bat. 1, 2 (1858) 80; Mrers, Contr. Bot. 3
(1871) 110; Hook. f. & Tu. Fl. Br. India 1 (1872) 100;
Becc. Malesia | (1877) 151; Kurz, For. Fl. Burma 1
(1877) 55; Rpt. J. Str. Br. R. As. Soc. n. 33 (1900)
42; GaGneP. Fl. Gén. I.-C. 1 (1908) 144; Diets, Pfl.
R. Heft 46 (1910) 215; in Elmer, Leafl. Philip. Bot.
4 (1911) 1165; Koorp. Exk. Fl. Java 2 (1912) 234;
Ript. Fl. Mal. Pen. 1 (1922) 109; Merr. En. Philip.
2 (1923) 148; Norman, J. Bot. 62 (1928) Suppl. 5;
HEND. Gard. Bull. S. S. 4 (1928) 220; Yamamoto, J.
Soc. Trop. Agric. 16 (1944) 98; HENpD. Mal. Nat. J.
6 (1951) 417, t. 378. — L. cerasifera BEcc. Malesia
1 (1877) 150; Diets, Pfl. R. Heft 46 (1910) 215;
Merr. En. Born. (1921) 249 (‘cerasifolia’); YAMA-
moto, J. Soc. Trop. Agric. 16 (1944) 98. — Fig.
12j-r.

Woody climber. Young branchlets, petioles and
inflorescences yellowish to golden-brown (or rusty)
tomentose, branchlets later glabrescent, young
shoots sometimes tendrilliform. Leaves: petioles 1—3
cm; lamina ovate-elliptic, elliptic or obovate-elliptic,
7—17 by 3—8 cm, apex usually broad and abruptly
cuspidate, or obtusely apiculate (occasionally long
acuminate), base cuneate to rounded, 3-nerved; lat-
eral nerves 4—6 pairs; both surfaces at first yellowish
to golden-brown pubescent or tomentose, especially

on lower surface along the nerves, later glabrescent;
stiffly papyraceous. Jnflorescences up to 2.5 cm,
composed of about 2—6 densely flowered cymes, 4—5
mm across with peduncles (3—)10—20 mm. — Male

flowers green to white, subsessile; sepals 9, ovate, to-

mentose, outer 3 1 by 0.75 mm, middle 3 1.25 by 1
mm, inner 3 2.5—3.25 by 1.75—2.5 mm, thick; petals
6, obovate, unguiculate, | by 0.75—1 mm, glabrous;
stamens 6, claviform, | mm, filaments usually
sparsely pilose adaxially. — Female flowers: sepals
and petals + as in male, petals clasping linear stami-
nodes; carpels 3, + obliquely ellipsoidal, 1.5 by 1.25
mm, stigma auriculate. Drupes obliquely obovate in
outline, 20—25 by 16—20 mm, pubescent to glabres-
cent.

Distr. SE. continental Asia (Lower Burma,
Lower Thailand, Annam, S. Indochina); in Malesia:
Central & S. SUMATRA (INCL. LINGGA), W. MALAYA
(PENANG TO SINGAPORE), W. BORNEO (SW. SARA-
WAK, BRUNEI), ? JAVA, PHILIPPINES (DIELS, /.c.).

Ecol. Secondary growths and open habitats, at
low altitude; common in Singapore and parts of Ma-
laya; in Sumatra also in primary swamp forest.

Note. For discussion concerning the identity of
LOUREIRO’s type see FORMAN (1958).

18. SARCOPETALUM

Fry. MM. Pl. Vict: 1 (1860) 26, t. suppl. 3; Diets, Pfl. R. Heft. 46 (910) 23258
85; FORMAN, Kew Bull. 22 (1968) 361. — Fig. 15m—p.

Woody climbers. Leaves subpeltate or peltate. Inflorescences axillary or aris-
ing from old, leafless stems, pseudoracemes. — Male flowers: sepals (2—)3—5,
minute; petals 3—5, thick and fleshy, larger than sepals; stamens with the fila-
ments connate in a column; anthers 3—4, free, arising horizontally from the top
of column. — Female flowers: sepals and petals as in male; staminodes equal
in number to the petals, free; carpels 3—6, stigma recurved, divided at apex into
2—3 subulate points. Drupe curved with style-scar near base; endocarp subsemi-
circular in outline with the base and apex separated by a short, + straight edge,
dorsally spinulose and/or ridged, laterally concave. Seed semi-annular. Embryo
subterete embedded in endosperm; cotyiedons elongate, flattened.

Distr. Monotypic. E. Australia (Victoria, New South Wales, Queensland); in Malesia: S. New Guinea.

1. Sarcopetalum harveyanum F. v. M. Pl. Vict. 1
(1860) 27, t. suppl. 3; Moore, Handb. Fl. N.S.W.
(1893) 20; BatLey, Queensl. Fl. 1 (1899) 32; Diets,
Pfl. R. Heft 46 (1910) 252, f. 85; J. Arn. Arb. 20
(1939) 73; BEADLE et al. Handb. Vasc. Pl. Sydney
(1962) 137, f. 29/B; FormMaAN, Kew Bull. 22 (1968)
361. — Fig. 15m—p.

Leaves with petioles 4— 10 cm long inserted almost

at or up to 3 mm from the basal margin of the la-
mina; lamina broadly ovate or deltoid-ovate, base
deeply or shallowly cordate, apex acuminate or ob-
tuse, apiculate at the tip, 9-15 by 7—12 cm, reticula-
tion prominent on both surfaces, glabrous, papyra-
ceous. — Male inflorescences c. 4 cm long, minutely
puberulous, bearing minute, narrowly lanceolate
bracteoles, 2 mm long. — Male flowers with pedicels

1986 |

2—3 mm long; sepals + triangular, c. 1 mm long,
margin irregularly dentate; petals very fleshy, +
broadly cuneate, 2 mm long, 1.5 mm thick; synan-
drium 1.5 mm long. — Female inflorescences 4—16
cm long. — Female flowers: sepals and petals as in
male; staminodes minute, 0.5 mm long; carpels semi-
ovoid, 1.5 mm long. Drupe red, glabrous; endocarp
6 by 5 mm, dorsally covered with numerous, + scat-
tered sharp points, these sometimes partly arranged
in transverse rows.

MENISPERMACEAE (Forman)

225

lections.

Ecol. Whereas the Papuan collections have been
found at low altitude, the Queensland ones are found
montane (1000—1200 m). All are recorded from
‘rain-forest’, but the Papuan localities are subject to
a seasonal climate. Fr. Sept., Jan.

Notes. The endocarps from Papua differ slightly
in ornamentation from the Australian specimens but
seem to represent merely a regional variation.

Although suspected to be toxic to livestock in Aus-

Distr. E. Australia; in Malesia: S. New Guinea
(Lake Daviumbu, Middle Fly; Wassi Kussa), 2 col-

tralia, feeding tests have proved negative (EVERIST,
Poison. Pl. Austr. rev. ed. 1981, 527).

19. LEGNEPHORA

Miers, Ann. Mag. Nat. Hist. ser. 3, 19 (1867) 89; Contr. Bot. 3 (1871) 287;
Diets, Pfl. R. Heft 46 (1910) 222, f. 76; FORMAN, Kew Bull. 22 (1968) 369; ibid.
27 (1972) 275, f. 1. — Fig. 14, 15g—h.

Woody climbers. Leaves palmately 3—7-nerved at the base, + broadly ovate.
Inflorescences: pedunculate cymes, one to few supra-axillary or racemosely ar-
ranged. — Male flowers: sepals 6, outer 3 usually narrower than inner 3; petals
6 with sides folded inwards around the opposite stamen, glabrous; stamens 6,
free; anthers dehiscing transversely, + introrse, the cells separated abaxially by
the thickened connective. — Female flowers: sepals 6—9; petals 0; staminodes
6, claviform; carpels 3; stigmas recurved, flattened. Drupe curved with style-
scar near base; endocarp rotund to obovate in outline with entire dorsal wing
and prominent lateral horseshoe-shaped or cucullate crests. Seed curved; em-
bryo imbedded in endosperm, narrow with radicle slightly longer than cotyle-
dons.

Distr. E. Australia, Solomon Is.; in Malesia: New Guinea, Lesser Sunda Is. (Timor), Moluccas, Philip-
pines. In all 5 spp.

Note. Single collections confirm the genus in the Lesser Sunda Is. (Timor) and Solomons (Santa Cruz),
but in the absence of fruits the species are uncertain.

KEY TO THE SPECIES

1. Leaves long and finely acute at the apex, entirely glabrous. Endocarp 8—9 mm long with a dorsal wing 1
mim: broad and:lateral cucullatecrests)2—3 mm broad 359 i065 os celal aplele srs store outset 1. L. acuta
1. Leaves rounded or broadly pointed at the apex, hairy, at least on the petioles.
2. Lateral crests on endocarp with fimbriate margins. Endocarps 9-11 by 10-11 mm. Australia L. moorei
2. Lateral crests on endocarp with margins entire or bearing a few teeth.
3. Endocarps up to 6 mm long.
4. Endocarps 4—5 mm long with lateral crests scarcely | mm broad, bearing a few teeth along the margins.
Senals tomentellais:s > (5 sca Mea eae ie clade pala a Mk aeet eee eals 2. L. philippinensis
4. Endocarps 6 mm long with lateral cucullate crests 3 mm broad, entire at the margins. Sepals sparsely
mnai CL OUIB S Siyh:c. 5's: ive wine e 9d Hie oR RDG a 8 Matra ee alay uv Oey Onna ven eReT 3. L. microcarpa
3. Endocarps 10—18 mm long, with a dorsal 3—4 mm broad wing, and lateral 4—5 mm broad cucullate crests
4. L. minutiflora

226

FLORA MALESIANA

102

[ser. I, vol.

Fig. 14. Endocarps of Legnephora Miers. — L. acuta FoRMAN, a. side view, a’. top view. — L. moorei (F.

v. M.) Miers, b. side view, b’. top view. — L. philippinensis FoRMAN, c. side view, c'. top view. — L. micro-

carpa ForMAN, d. side view, d'. top view. — L. minutiflora (K. Sc.) DiELs, e. side view, e'. top view. All

x5 (a MANNER & STREET 344, b CLEMENS 43871, c. PNH 10020, d NGF 21169, e CLEMENS 8682). Courtesy
Kew Bulletin.

1. Legnephora acuta ForMAN, Kew Bull. 27 (1972)
276, f. 1A. — Fig. 14a.

Stems glabrous apart from pubescent leaf-axils.
Leaves: petioles glabrous, 3—4.5 cm; lamina elliptic-
ovate, apex long and finely acute, base broadly ob-
tuse, 7.5—10 by 3.5—5.5 cm, glabrous, reticulation
prominent on both surfaces, 3-nerved at the base,
thinly coriaceous. — Male flowers unknown. — Fe-
male inflorescences composed of a few cymes ar-
ranged in a raceme, 1—2 per axil, 4.5—6 cm, subgla-
brous. — Female flowers pedicellate; outermost sep-
als 1—2, less than 1 mm, slightly puberulous, main
sepals 6, elliptic, 2.5 mm, glabrous; staminodes 5—6
less than 1 mm, apex bilobed; carpels 1 mm, stigma
reflexed. Drupe obovoid, 8—9 mm, glabrous; endo-
carp 8 mm, bearing a narrow dorsal wing 1 mm
broad and lateral cucullate wings 2—3 mm long, sur-
face smooth.

Distr. Malesia: E. New Guinea (Western High-
lands); 1 coll.

Ecol. Primary and old secondary forests on
shale, 1750 m.

Vern. Gawa kun, Maring.

2. Legnephora philippinensis FORMAN, Kew Bull. 27
(1972) 278, f. 1C. — Fig. 14¢.

Young stems both yellowish-hispid and yellowish-
puberulous (indumentum mixed). Leaves: petioles
hispid, sometimes also puberulous, 7—16 cm; lamina
broadly ovate, apex acute with a pubescent mucro,
base slightly cordate or truncate, 8—17 by 6.5—14.5
cm, both surfaces sparsely to moderately pubescent,
7-nerved at the base, stiffly papyraceous. — Male
flowers unknown. — Female inflorescences supra-
axillary, cymose, 3—8 cm on peduncles 1.2—5.5 cm,
branches tomentellous. — Female flowers shortly

1986 |

pedicellate; sepals 9 in 3 whorls, tomentellous, outer-
most 3 very narrowly oblong, 1.5 mm, middle 3 nar-
rowly elliptic, 2.5 mm, inner 3 broadly elliptic, 2.5
mm; staminodes 6, 1.25 mm, narrowly obtriangular,
puberulous; carpels 1 mm, densely pilose, stigma
broadly infundibular and recurved. Drupe (? young)
pubescent; endocarp subrotund in outline, 4-5 mm
9, c. 2mm thick, dorsal wing c. 1 mm broad, lateral
crests scarcely 1 mm broad with toothed margins.

Distr. Malesia: Philippines (Mindanao, Mt Ka-
tanglad), 1 coll.

3. Legnephora microcarpa FoRMAN, Kew Bull. 27
(1972) 278, f. 1D. — Fig. 14d.

Young stems yellow-puberulous. Leaves: petioles
yellow-tomentellous to -puberulous, 2—6 cm; lamina
broadly ovate to broadly elliptic, apex rounded or
obtuse sometimes with a fine mucro, base cordate,
truncate or rotund, (6—)8—11.5 by (4.5—)6—11 cm,
yellow-tomentellous below, sparsely puberulous or
subglabrous above, reticulation prominent and
dense above, 3—5-nerved at the base, stiffly papyra-
ceous. — Male inflorescences: cymes 2—3.5 cm, ped-
uncles 1—2 cm, puberulous. — Male flowers shortly
pedicellate; sepals + equal, elliptic, 1.5 mm, sparsely
puberulous outside; petals broadly rhomboid, 0.75
mm, glabrous; stamens 0.75—1 mm. — Female
flowers unknown. Infructescence c. 6 cm long with
peduncle c. 3 cm, puberulous. Drupe 6 mm 9, gla-
brous; endocarp 5—6 mm with a narrow dorsal ridge
less than 1 mm high, bearing extended lateral cucul-
late crests 3 mm long, surface smooth.

Distr. Malesia: E. New Guinea (Morobe Distr.:
Bulolo), 2 coll.

Ecol. Forests, 800—1200 m.

MENISPERMACEAE (Forman)

227

4. Legnephora minutiflora (K. ScH.) Diets, Pfl. R.
Heft 46 (1910) 222; ForMAN, Kew Bull. 27 (1972)
279, f. 1E. — Tinospora minutiflora K. ScH. &
Laut. Nachtr. Fl. Deut. Schutzgeb. Stidsee (1905)
262. — L. nyctericarpa Dre1s, Bot. Jahrb. 52 (1915)
189. — Phytocrene malacothrix SLEUMER, Notizbl.
Berl.-Dahl. 15 (1941) 361; Blumea 17 (1969) 237; FI.
Mal. I, 7 (1971) 80, 83. — Fig. 14e, 15g—h.

Stems and petioles covered with a fulvous indu-
mentum of either short hairs or long, straight hairs,
or both mixed together. Leaves: petioles (2.5—)4.5—8
cm; lamina broadly ovate, suborbicular or broadly
elliptic, apex broadly pointed to rounded, often with
a puberulous mucro, base cordate, truncate or
rounded, sometimes subpeltate, (6—)8—20 by
(5—)7—20 cm; beneath usually puberulous (some-
times subtomentose or subglabrous), above glabres-
cent, 3—S-nerved at the base, stiffly papyraceous. —
Male inflorescences subumbelliform, 2.5—6 cm long
with peduncles 1—4 cm, puberulous. — Male flowers
green or white on slender pedicels up to 4 mm; sepals
6, outer 3 narrowly elliptic, 2 mm long, inner 3
broadly elliptic, 2 mm long, tomentellous; petals 6,
0.5 mm long; stamens 6, 0.75 mm long, anthers
dehiscing transversely. — Female inflorescences sim-
ilar to male. — Female flowers: sepals as in male
flowers; staminodes 6, narrowly oblong 1 mm long,
glabrous; carpels 1 mm long, densely pilose, stigma
recurved. Drupe when dry similar to endocarp in size
and shape, densely fulvous-pilose when young, indu-
mentum becoming sparser; endocarp rotund in out-
line, 1—1.8 cm g, dorsally bearing a very thin wing,
3—4 mm broad, laterally bearing cuculliform crests
projecting 4—5 mm and with entire margins.

Distr. Malesia: S. Moluccas (Tenimber Is.: Jam-
dena) and New Guinea (incl. New Ireland).

Ecol. Rain-forest, from sea-level to 1800 m.

20. PERICAMPYLUS

Miers Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 36; ibid. ser. 3, 14 (1864) 369;
Contr. Bot. 3 (1871) 116; Diets, Pfl. R. Heft 46 (1910) 216, f. 74 & 75; FORMAN,
Kew Bull. 22 (1968) 365. — Fig. 15a—f.

Woody climbers. Leaves not peltate (in Malesia), palmately nerved. /nflores-
cences axillary, cymose and subumbelliform, pedunculate, cymes solitary or
fasciculate. Flowers: sepals 9, the outermost 3 narrow and minute, the inner 6
imbricate and concave, the innermost 3 broader than the others; petals 6, cu-
neate. — Male flowers with stamens 6, free (in Malesia); anthers dehiscing
longitudinally. — Female flowers with 6 filamentose staminodes; carpels 3; stig-
ma deeply bifid, recurved. Drupe curved with style-scar near base; endocarp ro-
tund in outline, dorsally covered with short pointed processes, laterally concave,
condyle septiform, imperforate; seed horseshoe-shaped; embryo enclosed in

228 FLORA MALESIANA [ser. I, vol. 102

Fig. 15. Pericampylus glaucus (LAMK) M_ErR. a. Habit, male plant, x2/3, b. male flower, one sepal removed,
x 14, c. stamen with petal, «30, d. female flower, front sepal, petal and staminode removed, x12, e. drupe,
x4, f. endocarp, x4. — Legnephora minutiflora (K. ScuH.) Diets. g. Stamen with petal, adaxial view, h. sta-

men, abaxial view, both x36. — Diploclisia kunstleri (KING) Drets. i. Male inflorescence, 1/2, j. male

flower, one inner sepal removed, x9, k. drupes on carpophore, x2/3, /. endocarp, nat. size. — Sarcopetalum
harveyanum F. v. M. m. Male flower, <9, n. stamens, X15, 0. drupe, x3, p. endocarp, x3 (a—c BACKER

17053, d DE VOGEL 3736, e—f CLEMENS 27416, g—h PLEYTE 71, i—j HAvILAND 1814, kK—/ STONE FSC 330, m—n

FLETCHER 19042, o—p Brass 7714).

a

as a ae

1986 |

MENISPERMACEAE (Forman)

229

endosperm, elongate and narrow, terete, radicle much longer than the subterete

cotyledons.

Distr. About 2—3 spp. in tropical and subtropical Asia; one throughout Malesia.

1. Pericampylus glaucus (LAMK) Merr. Int. Rumph.
(1917) 219; W.H. Brown, Minor Prod. Philip. For.
1 (1920) 375; Merr. En. Born. (1921) 250; En. Phi-
lip. 2 (1923) 148; Philip. J. Sc. 29 (1926) 368; Burk.
Dict. 2 (1935) 1693; HottH. & Lam, Blumea 5 (1942)
180; Yamamoto, J. Soc. Trop. Agric. 16 (1944) 99;
HEND. Mal. Nat. J. 6 (1951) 416, t. 376; Back. &
BakH. f. Fl. Java 1 (1963) 158; FoRMAN, Kew Bull.
22 (1968) 366; PANcHo, Vasc. Fl. Mt Makiling 1
(1983) 283, f. 86. — Menispermum glaucum LAMK,
Encycl. Méth. 4 (1797) 100, based on ‘Folium luna-
tum minus’ RumpH. Herb. Amb. 5 (1747) 40, t. 25,
f. 1. — Cocculus glaucus (LAMK) Dc. Syst. 1 (1817)
521; Mia. FI. Ind. Bat. 1, 2 (1858) 82. — Cocculus in-
canus CoLesr. Trans. Linn. Soc. Lond. 13 (1822) 57;
ScuerF. Nat. Tijd. N. I. 32 (1873) 398, t. 10. — Coc-
culus corymbosus BL. Bijdr. (1825) 24. — Cocculus
lanuginosus BL. |.c. — Clypea tomentosa BL. I.c. 27.
— Stephania tomentosa (Bu.) SPRENG. Syst. Veg. ed.
16, 4 (Cur. Post.) (1827) 316; Hassk. Pl. Jav. Rar.
(1848) 170. — Cocculus cinereus ZOLL. & Mor. Syst.
Verz. (1846) 38. — P. incanus (CoLEBR.) Hook. f. &
Tu. FI. Ind. (1855) 194; Mia. Fl. Ind. Bat. 1, 2 (1858)
83; Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 85; Miers,
Contr. Bot. 3 (1871) 118; Hook. f. & Tu. FI. Br. In-
dia 1 (1872) 102; Becc. Malesia 1 (1877) 151; KING,
J. As. Soc. Beng. 58 (1889) 385; K. Sco. & HOLtrR.
Fl. Kais. Wilh. Land (1889) 44; Boer. Cat. Hort.
Bog. 1 (1899) 41; K. ScuH. & Laut. Fl. Deut.
Schutzgeb. Stidsee (1901) 311; Back. Fl. Bat. 1
(1907) 37; Voorl. Schoolfl. (1908) 9; Drets, Pfl. R.
Heft 46 (1910) 217, f. 74; Back. Schoolfl. Java
(1911) 43; Drets in Elmer, Leafl. Philip. Bot. 4(1911)
1165; Koorp. Exk. Fl. Java 2 (1912) 234; KiRTIKAR
& Basu, Ind. Medic. Pl. 1 (1918) 58; Rip. Fl. Mal.
Pen. 1 (1922) 112; Burk. & HeNpb. Gard. Bull. S. S.
3 (1925) 344; HEND. ibid. 4 (1928) 220. — P. lanugi-
nosus (Bi.) Mig. Fl. Ind. Bat. 1, 2 (1858) 83. — P.
membranaceus Miers, Contr. Bot. 3 (1871) 122;
Diets, Pfl. R. Heft 46 (1910) 220; Riot. Fl. Mal.
Pen. 1 (1922) 112; Yamamoto, J. Soc. Trop. Agr. 16
(1944) 100. — Cissampelos pareira (non L.) Rw. FI.
Mal. Pen. | (1922) 114; Henn. Gard. Bull. S. S. 4

(1928) 220. — Fig. 15a—f.

Woody climber. Young stems yellowish-tomen-
tose, older stems glabrescent, longitudinally ridged.
Leaves with yellowish-tomentose petioles 3—7 cm;
lamina broadly triangular-ovate, margin usually
broadly and shallowly crenate, apex apiculate, usual-
ly broadly rounded, or obtuse, sometimes acute,
base shallowly cordate or truncate, sometimes ob-
tuse, S—10 cm long and broad; palmately S-nerved,
lower surface tomentose or softly pubescent, upper
surface sparsely pubescent; margin shallowly cren-
ate; papyraceous. /nflorescences yellowish-tomen-
tose, 2—4 cm long. — Male flowers white or yellow
with pedicels c. 1 mm; sepals 9, hairy outside, outer
3 narrow, minute c. 0.5 mm long, middle 3 oblanceo-
late, inner 3 obovate, all 1 mm long; petals 6, obcu-
neate, 0.5 mm long, glabrous; stamens 6, 0.75 mm
long. — Female flowers: sepals and petals as in male
flowers; staminodes 6, filamentose; carpels 0.5 mm
long; stigma bifid recurved. Drupe purple to black,
glabrous; endocarp rotund in outline, 5 mm 9, dor-
sally bearing 2 rows of c. 15 pointed projections,
each of which is linked by a ridge to a laterally point-
ing projection, the latter being arranged in 2 lateral
rows.

Distr. Throughout
throughout Malesia.

Ecol. Primary and secondary forests and thick-
ets, up to 1700 m.

Uses. Stems used for tying and basketry. Sap
used as an eye medicine.

Vern. (Only names taken from specimens ex-
amined are listed). Malaya: akar mumpanang, akar
tali gasing; Java: areuj geureung, S.; Sumbawa: se-
kompalai; Talaud 1.: talimba-as; Halmahera: gou-
male, Tidorese; Ternate: ginato; Philippines: bug-
bug, Mindoro, malabawugan, silong-pugo, Luzon;
maripari, Biliran; gapus nibid, Mindanao; further
sec. MERRILL (1923): botang-botang, C. Bis., gapisi-
lid, Sub., hah-tin, Yak., Sul., /agauat, Bag., pama-
20, Bik., pisok, lg., silong-pugo, tugian-tugian, Tag.

Note. The field-notes for RSNB 2615 from Mt
Kinabalu record a tuber 30 cm broad.

the generic range and

21. DIPLOCLISIA

Miers, Ann. Mag. Nat. Hist. ser. 2, 7 (1851) 37; Contr. Bot. 3 (1871) 280; Diets,
Pfl. R. Heft 46 (1910) 224, f. 77; FoRMAN, Kew Bull. 22 (1968) 362. — Fig.

15i—I.

230 FLORA MALESIANA [ser. I, vol. 102

Woody climbers. Leaves sometimes peltate. Inflorescences supra-axillary and
cymose (but not in Malesia), or cauliflorous and composed of a raceme of
cymes. — Male flowers: sepals 6, outer 3 narrower than inner 3; petals 6 with
sides folded inwards around the opposite stamen; stamens 6, free; anthers de-
hiscing with a transverse slit. — Female flowers: sepals and petals as in male
flowers; staminodes 6, filamentose with rudimentary anthers; carpels 3; stigmas
recurved, flattened with margins dentate. Drupe laterally compressed, obovate
(extra-Mal.) or narrowly obovate and curved in outline with style-scar close to
base; endocarp dorsally bearing many transverse ridges, with an elongate curved
depression on each lateral face (Mal. spp.). Seed (Mal. spp.) narrowly horse-
shoe-shaped, sharply curved around the deeply intrusive narrow condyle; em-
bryo narrow, radicle much shorter than the flat cotyledons; endosperm scanty.

Distr. 3 spp. in tropical continental SE. Asia and throughout Malesia as far as W. New Guinea, not yet
recorded from the Lesser Sunda Is. The third species in S. China.

Note. The only constant difference between the two Malesian species is in the fruits. No floral or inflor-
escence differences were found.

KEY TO THE SPECIES

1. Endocarp 14—20 by 8—11 mm, lacking a median dorsal ridge. Leaves usually not peltate 1. D. glaucescens

1. Endocarp (19—)25—30 by 16-17 mm with a pronounced median dorsal ridge. Leaves peltate

1. Diploclisia glaucescens (BL.) Diets, Pfl. R. Heft 46
(1910) 225, f. 77A—L; Back. Schoolfl. Java (1911)
44; Diets in Elmer, Leafl. Philip. Bot. 4 (1911) 1166;
Koorp. Exk. Fl. Java 2 (1912) 235; Merr. En. Phi-
lip. 2 (1923) 148; Back. & BAKH. f. Fl. Java 1 (1963)
158; FORMAN, Kew Bull. 22 (1968) 363, p.p. (excl. D.
kunstleri), with full discussion and synonymy. —
Cocculus glaucescens Bu. Bijdr. (1825) 25; Mia. FI.
Ind. Bat. 1, 2 (1858) 82; Ann. Mus. Bot. Lugd.-Bat.
4 (1868) 84; ScHEFF. Nat. Tijd. N.1. 32 (1873) 399. —
Cocculus macrocarpus W. & A. Prod. (1834) 13. —
D. macrocarpa (W. & A.) Miers, Contr. Bot. 3
(1871) 280, t. 127/20—28; Becc. Malesia 1 (1877)
152; BoeRL. Cat. Hort. Bog. 1 (1899) 41.

Woody climber recorded up to 30 m long, totally
glabrous. Stem up to 5 cm @, finely striate when
young. Leaves usually not peltate with petioles 5—12
cm long; lamina broadly ovate to suborbicular, apex
rounded to acute, base rounded to cordate, 6—11 by
6—11 cm; fine reticulation more obvious on the
sometimes glaucous lower surface, margin some-
times broadly and shallowly crenate, chartaceous.
Inflorescences cauliflorous, up to 52 cm long and 7
cm wide. — Male flowers pale yellow with pedicels
2—4 mm long; sepals 6, strongly marked (in sicco) by
a dark brown reticulum, outer 3 elliptic, 2.5 mm
long, inner 3 broadly elliptic, 2.5 mm long; petals 6,
ovate-rhombic, 1 mm long, apex acute or emargi-

2. D. kunstleri

nate; stamens 6, 2 mm long, anthers dehiscing with
a transverse slit. — Female flowers: sepals and petals
as in male flowers; staminodes 6, filamentose; car-
pels 2 mm long. Drupe yellow to orange; endocarp
elongate, + narrowly obovate in outline, slightly
curved, 14—20 mm long, 8—11 mm broad, dorsally
ornamented with many transverse ridges, lacking a
median dorsal ridge.

Distr. S. China, India, Ceylon, Burma, Thai-
land, Indochina; in Malesia: Sumatra, Java, Cele-
bes, Philippines, 7Moluccas (Halmahera, Sula Is.,
Ceram), ?W. Guinea.

Ecol. In forests, up to 600 m. Fi. Feb., June,
Ochi. }

Vern. Java: areuj geureung, geureung, S, sluru-
rut, tjlurutan, J.

Notes. The leaves are occasionally peltate. Col-
lections from the Moluccas with peltate leaves and
from W. Guinea with both peltate and non-peltate
leaves are possibly D. glaucescens, but fruits are
needed from these regions for confirmation.

The only fruiting collections known from Malesia
are two from the Philippines: from Mindanao with
peltate leaves (teste Diets) and from Sulu Is. with
non-peltate leaves.

2. Diploclisia kunstleri (KrNG) Diets, Pfl. R. Heft 46
(1910) 227, f. 77M—N; Merk. En. Born. (1921) 250;

1986 |

Ripi. Fl. Mal. Pen. 1 (1922) 107; MAsAMuNE, En.
Phan. Born. (1942) 274. — Cocculus kunstleri KING,
J. As. Soc. Beng. 58, ii (1889) 384. — D. glaucescens
(non (KING) DrELs) sensu FORMAN, Kew Bull. 22
(1968) 363, p.p. — Fig. 15i-l.

Characters as for D. glaucescens except: Leaves
peltate with petioles inserted (1—)10—28 mm from
basal margin. Drupe glaucous: endocarp (19—)
25—30 by 16—17 mm with a pronounced median dor-

MENISPERMACEAE (Forman)

231

sal ridge.

Distr. Malesia: Malaya, Borneo.

Ecol. In primary and secondary forests, some-
times by streams, up to 1500 m. F/. April—Oct.; fr.
May—Nov.

Vern. Sabah: bakauk, takob; Sarawak: akar tu-
ak, Kenyah.

Note. According to S 34725, the stem produces a
yellowish exudate and the flowers are fragrant.

22. COCCULUS

DC. Syst. Veg. 1 (1817) 515, nom. cons.; Mrers, Contr. Bot. 3 (1871) 253;
Drts, Pfl. R. Heft 46 (1910) 227, f. 78 & 79; FORMAN, Kew Bull. 15 (1962) 479;
ibid. 22 (1968) 372; ibid. 29 (1974) 477, t. 14 & 15. — Nephroia Lour. FI. Co-
chinch. (1790) 565. — Holopeira Miers, Ann. Mag. Nat. Hist. ser. 2, 7 (1851)
42, nomen; ibid. ser. 3, 19 (1867) 27; Contr. Bot. 3 (1871) 270, t. 126. — Fig. 16.

See for a complete synonymy FORMAN (1962).

Slender woody climbers, erect shrubs or small trees. Leaves not peltate, rarely
lobed (in Mal. spp.). Inflorescences cymose or thyrsoid. — Male flowers: sepals
6—9 in 2—3 whorls, the outer sepals smallest; petals 6, bifid or emarginate at
apex, with basal inflexed auricles clasping the opposite stamen; stamens 6, free.
— Female flowers: sepals and petals as in male; staminodes 6 (in Mal. spp.); car-
pels 3 or 6, styles subulate, reflexed. Drupes curved with style-scar near base,
obovate or rotund in outline; endocarp dorsally verruculose or ridged and with
a curved aperture on both lateral faces. Seed curved almost into a ring, broad,
dorsiventrally flattened; endosperm very thin; embryo with liguliform cotyle-
dons.

Distr. Central & North America, Africa, SE.—E. Asia, Malesia (Malaya, Sumatra, Java, Philippines) to
Polynesia. In all 8 spp.

Notes. I made a special study (1974) of the ornamentation patterns of the endocarp of 7 species which
show a remarkable variety, even allowing them to be identified by a key and from Sem photographs.

The leaf-epidermal characters of the genus were investigated by D.K. Ferguson in Kew Bull. 29 (1974)
483-492, tt. 16-21.

KEY TO THE SPECIES IN MALESIA

1. Slender climber. Lamina usually broadest below the middle, not acute at both apex and base; basal pair
of nerves about equally prominent as the main lateral nerves, usually not running parallel to the margin
and usually becoming indistinct at or below the middle of the lamina. Carpels 6 .... 1. C. orbiculatus

1. Erect shrub or small tree. Lamina broadest at or slightly above the middle, both apex and base acute; basal
pair of nerves much stronger than the lateral nerves, running + parallel to margin and continuing promi-
nently beyond the middle of the lamina. Carpels 3 .......... 00. cc cee cee cence eeeees 2. C. laurifolius

1. Cocculus orbiculatus (L.) DC. Syst. 1 (1817) 523;
Prod. | (1824) 98; FormAN, Kew Bull. 22 (1968) 374;
ibid. 29 (1974) 479, t. ISA—G. — Menispermum
orbiculatum L. Sp. P1. (1753) 341. — Menispermum

trilobum Tuuns. Fl. Jap. (1784) 194. — C. trilobus
(THuns.) De. Syst. 1 (1817) 522; Driers, Pfl. R. Heft
46 (1910) 232, f. 78A—L; in Elmer, Leafl. Philip.
Bot. 4 (1911) 1166; Spracue, Bot. Mag. 139 (1913)

232 FLORA MALESIANA [ser. I, vol. 102

Fig. 16. Cocculus orbiculatus (L.) DC. a. Habit, male plant from Malaya, 2/3, b. leaf shape occurring in
Java, 2/3, c. bud showing the sepals, d. male flower, e. petal with stamen, f. female flower, g. petal with
staminode, all x10, h. 4 of 6 drupes (dried) from one flower, x2, i. drupe (dried), x2, /. endocarp, x6.
— C. laurifolius DC. k. Habit, male plant, «2/3, /. male flower, m. petal with stamen, n. female flower,
all «10, o. 2 of 3 drupes (dried) from one flower, 2, p. endocarp, x6 (a, c—e FRI 5038, b COMMERSON
s.n., f—g BS 27231, h—j Poore 1140, kK—m Wintr 97 (Thailand), n FORMAN 36, o—p EvRrArD 535 (Vietnam)).

1986]

t. 8489; Merr. En. Philip. 2 (1923) 149; ForMAN,
Kew Bull. 15 (1962) 480, f. 1; BAcK. & Baku. f. FI.
Java 1 (1963) 159. — Nephroia sarmentosa Lour. FI.
Coch. 2 (1790) 565. — Menispermum ovalifolium
VAHL ex Pers. Syn. Pl. Ench. Bot. 2 (1807) 628. —
C. ovalifolius (V AHL ex PERS.) Dc. Syst. 1 (1817) 526;
BL. Bijdr. (1825) 25; Hassk. Tijd. Nat. Gesch. Phy-
siol. 10 (1843) 132; Marer, Nat. Tijd. N. I. 3 (1852)
465; Mia. FI. Ind. Bat. 1, 2 (1858) 81; Ann. Mus.
Bot. Lugd.-Bat. 4 (1868) 84; ScHEFF. Nat. Tijd. N.
I. 32 (1873) 399, t. 11; BeEcc. Malesia 1 (1877) 151;
BoerL. Cat. Hort. Bog. 1 (1899) 39; Back. Fl. Bat.
1 (1907) 36; Voorl. Schoolfl. (1908) 8; Rui. Fl. Mal.
Pen. 1 (1922) 110. — C. triflorus Dc. Syst. 1 (1817)
529. — C. cynanchoides PRESL, Rel. Haenk. 2 (1830)
79. — Limacia kunstleri KiNG, J. As. Soc. Beng. 58,
ii (1889) 383; Rm. Trans. Linn. Soc. II, Bot. 3
(1893) 274; J. Str. Br. R. As. Soc. n. 33 (1900) 42. —
C. sarmentosus (Lour.) Drets, Pfl. R. Heft 46 (1910)
233; Back. Schoolfl. Java (1911) 44; Drets in Elmer,
Leafl. Philip. Bot. 4 (1911) 1166; Koorpb. Exk. FI.
Java 2 (1912) 231; Merr. Philip. J. Sc. 13 (1918) Bot.
10, incl. var. stenophyllus MerrR.; En. Philip. 2
(1923) 149; BEumMEE, Hand. N. I. Nat. Congr. (1925)
175; Merr. Trans. Amer. Phil. Soc. n.s. 24 (1935)
156; YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 102;
HEND. Mal. Nat. J. 6 (1951) 417, t. 377. — Nephroia
elegans Riwv. J. Str. Br. R. As. Soc. n. 54 (1910) 15.
— C. elegans (Rw .) Ript. Fl. Mal. Pen. 1 (1922)
111. — Fig. 16a—j.

For the full synonymy of this species see FORMAN
(1962).

Slender climber. Stems herbaceous or slightly
woody, 1—1.5 mm g, in leaf-bearing parts, puberu-
lous to subglabrous. Leaves with puberulous petioles
5—10(—20) mm; lamina variable in shape, in Malaya
and Sumatra usually elliptic to ovate-elliptic, in Java
usually ovate to triangular-ovate, in the Philippines
+ narrowly elliptic to very broadly elliptic, apex
usually rounded, sometimes acute or emarginate,
with a mucronate tip, base obtuse to truncate, rarely
acute, 3—10 by 1—5.5 cm; base 3- (or 5-)nerved; both
surfaces thinly puberulous to glabrous; papyraceous.
— Male inflorescences axillary and terminal, the
flowers in peduncled cymes 0.8—1.5 cm long which
are often solitary when axillary, or arranged in a nar-
row thyrse up to 7 cm, puberulous throughout with
yellowish hairs. — Male flowers white, glabrous
throughout, borne on pedicels 1—2 mm; sepals in 3
whorls of 3, outermost whorl minute, often with one
or more parts lacking, middle whorl up to | mm
long, inner whorl with sepals obovate to rotund
1—2.5 mm long; petals 6, + oblong with the apex di-
vided into 2 acute lobes, with 2 incurved lobes shortly
above the base clasping the filament of the stamen
Opposite, |—1.5 mm long; stamens 6, 1 mm long. —
Female inflorescences axillary and terminal, much

MENISPERMACEAE (Forman)

233

fewer-flowered than in the male, appearing race-
mose, up to 5cm long. — Female flowers on bracteo-
late ‘pedicels’ c. 5 mm long (i.e. glabrous pedicel +
puberulous inflorescence-branch); sepals and petals
as in male; staminodes 6, minute, 0.3 mm long, gla-
brous; carpels 6, glabrous, 0.75 mm long; style slen-
der, curved outwards. Drupes dark blue, rotund in
outline, 4.—5 mm 9, glabrous; endocarp with a small
curved aperture on both sides, dorsally ornamented
with branched ridges.

Distr. Eastern Himalayas, east to China, Japan,
Taiwan and Hawaii; in Malesia: NE. Sumatra, Ma-
laya (also Penang & Singapore), W.—Central Java,
Philippines (Luzon, Mindanao, Batan Is.). In Réu-
nion and Mauritius possibly as an introduction.

Ecol. Often climbing over trees and shrubs by the
sea-shore, inland sometimes on limestone terraces
and found associated with other coastal plants (cf.
BEUMEE, 1925).

Notes. The extensive synonymy of this species is
largely the result of combining four species which
have previously been recognized as distinct, viz., C.
trilobus (THUNB.) DC., C. sarmentosus (LourR.)
DrEts, C. mollis Hook. f. & Tu., and C. ferrandia-
nus GAUDICH.

The material now available is considerably more
than that available to Diets (1910). The specimens I
have examined display great variability in the leaves
with regard to shape and degree of hairiness, which
completely obscures the distinctions set out by Diets
in that part of his key which separates the above-
mentioned four species. It now seems apparent that
the epithets ‘trilobus’, ‘sarmentosus’, ‘mollis’ and
‘ferrandianus’ are applicable only to forms each of
which is more dominant in a different part of the to-
tal area of the species, but which are linked together
by so many intermediates that the recognition of
infraspecific taxa is impracticable.

I have discussed in detail (1962, under C. trilobus)
the great variation in shape and indumentum of the
leaves in relation to the geographical distribution of
the species. This variation has resulted in the exten-
sive synonymy which I gave in full (/.c.).

Riw.ey has twice recorded the stems as being latici-
ferous, under Limacia kunstleri KiNG, Trans. Linn.
Soc. I], Bot. 3 (1893) 274, and under C. elegans
(Ripi.) Rpt. Fl. Mal. Pen. 1 (1922) 111.

2. Cocculus laurifolius DC. Syst. 1 (1817) 530; De-
Less. Ic. Pl. 1 (1820) 25, t. 97; Cocepr. Trans. Linn.
Soc. 13 (1822) 65; Hook. f. & Tu. FI. Ind. (1855) 191;
Mig. Fl. Ind. Bat. 1, 2 (1858) 81; Ann. Mus. Bot.
Lugd.-Bat. 4 (1868) 84; Hook. /. & Tu. Fl. Br. India
1 (1872) 101; Scuerr. Nat. Tijd. N. I. 32 (1873) 393,
t. 12; Boert. Cat. Hort. Bog. 1 (1899) 39, incl. var.
angustifolius (Hassk.) Boeri. et var. triplinervis
Boert. /.c. 40; K. & V. Bijdr. 9 (1903) 95; Diets, Pfl.

234

FLORA MALESIANA

[ser. I, vol. 102

R. Heft 46 (1910) 239, f. 79; Back. Schoolfl. Java
(1911) 45; Koorp. Exk. Fl. Java 2 (1912) 231; MERR.
En. Philip. 2 (1923) 149; Burk. Dict. 1 (1935) 594;
YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 102; For-
MAN, Kew Bull. 15 (1962) 485; Back. & BAKH. /. FI.
Java 1 (1963) 159; ForMAN, Kew Bull. 29 (1974) 479,
t. ISA—G. — C. angustifolius Hassk. Cat. Hort.
Bog. (1844) 172; Wap. Rep. 5 (1846) 17; Hassk. PI.
Jav. Rar. (1848) 167. — Holopeira laurifolia (Dc.)
Miers, Ann. Mag. Nat. Hist. ser. 3, 19 (1867) 29;
Miers, Contr. Bot. 3 (1871) 276. — Holopeira
australis Miers [Ann. Mag. Nat. Hist. ser. 3, 19
(1867) 29, nomen], Contr. Bot. 3 (1871) 277. — Fig.
16k—p.

Erect shrub or tree up to 6 m, with main stem up
to 12 cm go. Branchlets glabrous. Leaves: petioles
3—6(—10) mm; lamina elliptic or oblanceolate-
elliptic, apex acute, often rather attenuate, base usu-
ally acute, occasionally cuneate, 7—11(—15) by 3—S.5
cm, base strongly 3-nerved with the basal nerves run-
ning almost parallel to the margin beyond the middle
of the lamina; both surfaces glabrous, stiffly papyra-
ceous. — Male inflorescences axillary, composed of
cymes arranged in a thyrse, or sometimes consisting
of a single cyme, 0.5—4 cm long, glabrous or sparsely
puberulous; bracts subulate, 1 mm long, frequently
puberulous. — Male flowers yellow, glabrous
throughout, borne on pedicels up to 1 mm; sepals in
2 whorls of 3, all + broadly elliptic, outer sepals
0.5—0.75 mm long, inner sepals 1—1.25 mm long;
petals 6, cuneate, apex emarginate or divided into 2
rounded lobes, also with 2 incurved lateral lobes
clasping the stamen opposite, 0.5 mm long; stamens

6, 0.75 mm long. — Female inflorescences similar to
male but fewer-flowered thus appearing subrace-
mose. — Female flowers on pedicels up to 5 mm, sep-
als and petals as in male flowers; staminodes 6, mi-
nute; carpels 3, each with slender reflexed style.
Drupes rotund in outline, c. 4mm @, glabrous; endo-
carp with a small curved aperture on both sides, fine-
ly ridged over the dorsal surface.

Distr. India to Nepal, Burma, S. China, Japan,
Taiwan, Indochina and Thailand; in Malesia: Suma-
tra, throughout Java, Philippines (Palawan, Luzon,
Mindoro).

Ecol. Open woodland, thickets, grassland, river-
banks, teak and banana plantations up to 1500 m.

Uses. BurkKILL (1935) records the presence of an
alkaloid in the bark which has an action similar to
that of curare, and is known as cocculine or coclau-
rine.

Vern. Java: ki patjar, tedjan, S.

Notes. In contrast to C. trilobus, C. laurifolius
displays remarkably little variation in its leaf-shape.
This may be connected with the fact that this species
is not a climber, but an erect shrub or small tree, one
of the rare examples of this life-form in the family.
Judging from dried material, the fruits appear to be
much less fleshy than those of C. trilobus, but notes
on their size and colour in the fresh state are lacking.

Although the species occurs in Thailand and Indo-
china and again in Sumatra and Java, it apparently
does not occur in the Malay Peninsula.

Cultivated as an ornamental in milder parts of
Europe and North America.

23. CISSAMPELOS

Linn, Sp) P1..(1753).1031; DC. Syst. 1.(1817).531,,p.p:; _Prod.4,.(1824) 4100;
p.p.; WALP. Rep. 1 (1842) 96, p.p.; MiERS, Contr. Bot. 3 (1871) 127; DiE.Ls, Pfl.
R. Heft 46 (1910) 283, f. 91; FoRMAN, Kew Bull. 22 (1968) 355. — Fig. 17.

Scandent shrubs or lianes. Leaves peltate or not peltate. — Male inflorescen-
ces: flowers in axillary, peduncled, corymbose cymes, these solitary or fascicled
(and sometimes borne along an axillary shoot bearing reduced leaves, but not
normally so in Malesia). — Male flowers: sepals 4, obovate; petals connate into
a cupuliform corolla (rarely free in extra-Malesian spp.); stamens connate into
a peltate synandrium, anther-cells 4 (in Malesian sp.). — Female inflorescences
axillary, thyrsoid, elongate, composed of fascicles arising in the axils of ac-
crescent + orbicular bracts (these not accrescent in some American spp.). — Fe-
male flowers: sepal 1; petal 1 (rarely 2—3 in extra-Malesian spp.); staminodes
0; carpel 1. Drupe curved with style-scar near base, pubescent; endocarp bearing
dorsally 2 rows of transverse ridges; seed horseshoe-shaped; embryo embedded
in endosperm, elongate, narrow, terete, cotyledons flattened, radicle as long as
or longer than the cotyledons.

1986] MENISPERMACEAE (Forman) 235

Fig. 17. Cissampelos pareira L. var. hirsuta (DC.) FormMaN. a. Habit, male plant, 2/3, b. male flower, c.

sepal, d. male flower with 2 front sepals and front half of corolla removed, e. corolla and stamens, all «13,

Se habit, female plant, «2/3, g. part of female inflorescence, cymule and bracts, x7, A. female flower, x13,
i. LS of carpel, x13, j. endocarp, x3. Courtesy Flora of Tropical East Africa.

236

FLORA MALESIANA

[ser. I, vol. 102

Distr. Pantropical, with 20—25 species. Only one species with one variety in Malesia.

1. Cissampelos pareira LINNE var. hirsuta (BUCH. ex
DC.) FORMAN, Kew Bull. 22 (1968) 356; PANCHo,
Vasc. Fl. Mt. Makiling 1 (1968) 286, f. 87. — C. pa-
reira LINNE, Sp. Pl. (1753) 1031, p.p. quoad ‘B’;
Hook. f. & Tu. Fl. Ind. (1855) 199; Mig. Ann. Mus.
Bot. Lugd.-Bat. 4 (1868) 85, incl. var. orbiculata
(Dc.) Miqg.; Hook. f. & Tu. Fl. Br. India 1 (1872)
103; ScHEFF. Nat. Tijd. N. I. 32 (1873) 401, incl. var.
peltata SCHEFF.; BEcc. Malesia | (1877) 156; KING,
J. As. Soc. Beng. 58, ii (1889) 386; BoERL. Cat. Hort.
Bog. 1 (1899) 43; GaGnep. Fl. Gén. I.-C. 1 (1908)
149; Drets, Pfl. R. Heft 46 (1910) 286, f. 91, incl.
var. typica Diets; in Elmer, Leafl. Philip. Bot. 4
(1911) 1167; Merr. Fl. Manila (1912) 204; Sp. Blanc.
(1918) 144; W.H. Brown, Minor Prod. Philip. For.
3 (1921) 186; Merr. En. Born. (1921) 251; En. Phi-
lip. 2 (1923) 149; Burk. Dict. 1 (1935) 559; KANEH.
& Hatus. Bot. Mag. Tokyo 56 (1942) 471; YAmMa-
MoTO, J. Soc. Trop. Agric. 16 (1944) 144; TrRouPIN,
Bull. Jard. Bot. Brux. 25 (1955) 140; Monogr. Me-
nisp. Afric. (1962) 274. — C. convolvulacea WILLD.
Sp. Pl. 4 (1805) 862; Dc. Syst. 1 (1817) 536; Prod. 1
(1824) 101; Miers, Contr. Bot. 3 (1871) 196; Hassk.
Pl. Jav. Rar. (1848) 171, incl. var. hirsuta (Dc.)
Hassk. — C. discolor De. Syst. 1 (1817) 534; Prod.
1 (1824) 101; A. Gray, Bot. U.S. Expl. Exped.
(1854) 38, incl. var. cardiophylla A. Gray. — A. cu-
mingiana TuRCZ. Bull. Soc. Imp. Nat. Moscou 27 (2)
(1855) 283. — Fig. 17.

For further synonyms see DIELs (1910: 288), TRou-
PIN (1962: 274) and FORMAN (1968: 356).

Scandent shrub. Old stems woody. Leafy stems
slender, densely to sparsely pubescent (or puberu-
lous) to glabrous. Leaves with densely to sparsely pu-
bescent or puberulous petioles, 2—9 cm, inserted
1—18 mm from basal margin of lamina; lamina +
broadly ovate, base rounded, truncate or cordate,
apex mostly acuminate (in Mal.), sometimes very ob-
tuse, mucronate at the tip, 4.5—11 by 4.5—12 cm,
lower surface tomentose to pubescent or puberulous,
upper surface sparsely pubescent or puberulous, so-
metimes tomentose; palmately 5—7-nerved, charta-
ceous. — Male inflorescences: flowers in subcorym-
bose, peduncled cymes, 2—4 cm long, pubescent, sol-
itary or a few arising in a fascicle. — Male flowers
green to yellowish on pedicels 1—2 mm; sepals 4,
obovate, 1.25—1.5 mm long, pilose outside; corolla
cupuliform, c. 0.5 mm long, puberulous outside;
synandrium c. 0.75 mm long. — Female inflo-
rescences thyrsoid, narrow, up to 18 cm, composed
of a pseudoraceme of fascicles, each fascicle in the
axil of an accrescent, + suborbicular bract, up to 1.5
cm long, puberulous to tomentose. — Female flo-
wers on pedicels 1—1.5 mm; sepal 1, broadly obo-
vate, 1.5 mm long; petal 1, broadly cuneate-obovate,
0.75 mm; long; ovary scarcely 0.5 mm long, pilose,

about equal in length to the thick, glabrous style;
stigma divaricately 3-lobed. Drupe orange or red,
pubescent, with endocarp obovate in outline, 5 mm
long, dorsally bearing 2 rows of 9—11 very promi-
nent, transverse ridges, lateral faces of condyle bor-
dered by a horseshoe-shaped ridge.

Distr. Pantropical, through continental South-
east Asia, Australia (Queensland); in Malesia: N.
Borneo, Philippines (throughout), Celebes (incl. Ka-
baena I.), Lesser Sunda Is. (Lombok, Sumbawa,
Flores, Wetar, Timor), Moluccas (Halmahera, Ce-
ram, Tenimber Is.), New Guinea (incl. Aru Is.). Not
known from Sumatra, Malaya, and Java.

Ecol. Recorded from primary and secondary for-
ests and thickets, climbing over trees or river-banks,
up to 1300 m.

Uses. According to W.H. Brown (1921) the root
when brewed is considered diuretic, lithotriptic, pec-
toral and febrifugal; pounded leaves are used for
snake-bites and as an antiscabious remedy.

Vern. Philippines: abobo, C. Bis., bangbdngau,
kalaad, kalkalaad, kuskusipa, \\k., batang-bdtang,
gulagulamdnan, kalakalamayan, makabo-o, sansdu,
sinchdo-sinchaduan, sinsau-sinsauan, TVag., hampa-
padre, himpapara, sampare, Bis., kauas, Sub., mala-
ruto, Ibn., samang, Bon.; Moluccas: mangaloke,
Ceram.

Notes. Cissampelos pareira L. has been erro-
neously recorded from Malaya and Java in several
publications. Records for Malaya were given by Ripb-
LEY (Fl. Mal. Pen. 1, 1922, 114) and by HENDERSON
(Gard. Bull. S. S. 4, 1928, 220). From an examina-
tion of the relevant specimens in the Singapore Her-
barium, it is clear that these records are based on mis-
identified specimens of Pericampylus glaucus
(LAMK) MERR.

The origin of the erroneous records for Java was
MiIQuEL (FI. Ind. Bat. 1, 2, 1859, 85), where, under
C. pareira, MiQugEL listed as a synonym Stephania ca-
pitata (BL.) SPRENGEL together with the synonyms of
the latter species.. Clearly, therefore, MIQUEL was
misidentifying Stephania capitata with C. pareira, as
indeed he later admitted (Ann. Mus. Bot. Lugd.-Bat.
4, 1868, 86). The Sundanese vernacular name ‘ojat
tjam-tjoear’ given by MIQUEL resembles ‘tjamtjau’,
which commonly refers to Stephania capitata and
also to Cyclea barbata Miers. As a result of MI-
QUEL’s misidentification, the name C. pareira still
persists in publications on the flora of Java, although
the occurrence there of the species is sometimes men-
tioned as being doubtful. The following works on
Javanese plants record C. pareira: BACKER,
Schoolfl. Java (1908) 47; Koorpers, Exk. Fl. Java 2
(1912) 237; Backer, Bekn. Fl. Java (em. ed.) 3 (fam.
34) (1941) 19; BAcKER & BAKH. f. Fl. Java 1 (1963)
160, with doubt.

1986] MENISPERMACEAE (Forman) 237

24. CYCLEA

ARNOTT ex WIGHT, Ill. Ind. Bot. 1 (1840) 22; Miers, Contr. Bot. 3 (1871) 234;
Drets, Pfl. R. Heft 46 (1910) 309, f. 93; ForMAN, Kew Bull. 14 (1960) 68; ibid.
34 (1980) 565. — Fig. 18.

Slender woody climbers. Leaves often peltate, palmately nerved. Jnflorescen-
ces axillary, terminal or cauliflorous, pseudoracemose or thyrsoid. — Male
flowers: sepals 4(—5), free or connate into a 4(—5)-lobed calyx; petals 4, free or
connate into a + cup-shaped corolla or rarely 0; stamens connate into a peltate
synandrium, anthers 4—5 dehiscing transversely. — Female flowers: sepals and
petals 1 (extra-Mal.), 2—3, rarely petals 0; staminodes 0; carpel 1, stigma
3—5-fid. Drupes curved with style-scar near base, obovate to rotund in outline,
sometimes pubescent; endocarp bony with the condyle a central cavity around
which the seed is curved, perforate ventrally between style-scar and base, often
perforate laterally, dorsally ornamented with 3—6 rows of tubercles. Seed horse-
shoe-shaped; embryo narrow, terete, embedded in endosperm.

Distr. India to Central and S. China, Thailand, Indochina; in Malesia: Sumatra, Malaya, Java, Borneo,
Philippines. About 29 spp., of which 9 in Malesia.

Note. Species with free sepals are now included in Cyclea; the calyx of the male flowers is distinguished
from that of Stephania by the single whorl of sepals in contrast to the two whorls in Stephania. The inflores-
cences of Cyclea and Stephania can easily be distinguished. Cyc/ea has thyrses (in some species the lateral
branches are reduced to sessile clusters along the main axis), whereas in all Asiatic species of Stephania the
inflorescence is umbelliform or composed of umbelliform parts, which are sometimes reduced to peduncled,
disciform capitula. It is noteworthy that the fruits of Stephania are always glabrous although they are some-
times hairy in Cyclea.

KEY TO THE SPECIES

1. Inflorescences unbranched; flowers densely crowded in spaced, sessile clusters .... 1. C. kinabaluensis
1. Inflorescences thyrsoid.
2. Lamina lanceolate to ovate-lanceolate, usually glabrous below ....................5. 2. C. elegans

2. Lamina broader, mostly broadly ovate, pubescent below.
3. Young stems bearing short hairs interspersed or not with long, straight spreading hairs; sepals of male
flowers only shortly connate.
pememumaenot peltate 2.20000. Ew. lle > Ceiba gerne es ld aee tee 3. C. atjehensis
4. Lamina peltate.
5. Petiole inserted 1—5 mm from basal margin of lamina.
6. Lower surface of leaves tomentose; upper surface with a very dense, raised reticulation 4. C. merrillii
6. Lower surface sparsely hairy; upper surface with a very lax, scarcely raised reticulation 5. C. insularis
5. Petiole inserted more than 6 mm from basal margin of lamina.
7. Lamina 15—21 cm long. Male inflorescence with rhachis and branches stiff and straight, the flowers
in congested peduncled cymes. Endocarp bearing 2 rows of c. 8 transverse ridges. 6. C. robusta
7. Lamina 7.5—12.5 cm long. Male inflorescence very slender and flexuose, the flowers in lax cymose
clusters without obvious peduncles. Endocarp bearing 4 rows of c. 14 papilliform tubercles
7. C. cauliflora
3. Young stems bearing only long, straight, spreading hairs; sepals of male flowers united for at least half
their length.
8. Flowers male.
oi Calyx glabrous or subglabrous; petals free.) 70.798 0 OR APN 8. C. laxiflora
* 9, Calyx puberulous.
RO Petals frees 55 ie VPN EOF LH OR IE ROPER BIR 9. C. peregrina

238 FLORA MALESIANA [ser. I, vol. 102

Fig. 18. Cyclea atjehensis FoRMAN. a. Habit with female inflorescences, <1/2, b. portion of stem, x4, c.

male inflorescence, x2/3, d. male flower (in spirit), X10, e. corolla and synandrium of male flower, X10,

f. female flower, abaxial view, x16, g. sepal and petal of female flower, x20, A. fruit (in spirit), x2 2/3,

j. endocarp, x4 (a—b BANzIGER 65-11, c—e BANZIGER 65-15, f—g BANZIGER 65-10, h—j BANZIGER 65-8).
Drawn by Mrs. A. Davies. Courtesy Kew Bulletin.

1986]

POs Petalsiconnatete: #2.) 2, eRe eee

8. Flowers female.
11. Flowers clearly spaced in lax clusters.

12. Young carpels glabrous or subglabrous....
12. Young carpels puberulous ...............
11. Flowers tightly crowded in subglobose heads; carpels tomentose ...............

1. Cyclea kinabaluensis FORMAN, Kew Bull. 14 (1960)
69.

Slender woody climber. Leaves with petioles 3—6
cm long inserted c. 1 mm above basal margin; lamina
deltoid-ovate to triangular-ovate, apex long and fine-
ly acuminate, base very obtuse, truncate or cordate,
palmately 5—7-nerved, 7—12.5 by 4.5—10 cm, both
surfaces sparsely puberulous or glabrous with a fine
and prominent reticulation, stiffly papyraceous. —
Male inflorescences axillary, unbranched, 4—7 cm
long with the flowers in sessile clusters 0.5—1 cm
apart. — Male flowers green, sessile; sepals 4, basally
connate or almost free, broadly obovate 1.5—2 mm
long; synandrium subdiscoid, slightly lobed at the
margin, 0.5—1 mm long, fleshy, glabrous. — Female
inflorescences similar to male, up to 13 cm long. —
Female flowers: sepals 2 or 3, free, obovate or subre-
niform, 1 mm long, externally puberulous; petals 2
or 3, free or basally connate, + cuneate to reniform,
0.75 mm long, glabrous; carpel subellipsoidal, 1 mm
long, stigma 3—S-lobed. Drupes sessile, rotund in
outline, 7 mm 9, glabrous or subglabrous. Endocarp
bearing 4 dorsal rows of tubercles.

Distr. Malesia: North Borneo (Mt Kinabalu).

Ecol. In forest, 1700—2800 m.

var. hispida FORMAN, Kew Bull. 14 (1960) 70.
Differs from var. kinabaluensis in its glabrous in-
florescences and flowers, and hispid branchlets. In
the type variety inflorescences, stems and petioles are
golden-brown puberulous.
Known only from one specimen.

2. Cyclea elegans Kinc, J. As. Soc. Beng. 58, ii
(1889) 387; Diets, Pfl. R. Heft 46 (1910) 311; Rit.
Fl. Mal. Pen. 1 (1922) 115; Burk. & HENbD. Gard.
Bull. S. S. 3 (1925) 344; Airy SHAw, Kew Bull. 1939
(1940) 538; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 145; ForMan, Kew Bull. 14 (1960) 71. — C.
caudata Merr. J. Str. Br. As. Soc. n. 85 (1922) 172;
Yamamoto, J. Soc. Trop. Agric. 16 (1944) 145. — C.
acuminatissima Merr. Pap. Mich. Ac. Sc. 24 (1939)
68. — [C. tonkinensis (non GAGNEP.): YAMAMOTO,
J. Soc. Trop. Agric. 16 (1944) 145.] — C. scyphigera
Suessenc. & Herne, Mitt. Bot. Staatssamml.
Miinch. 2 (1950) 59, incl. f. angustifolia SUESSENG. &
Heine; Heine in Fedde, Rep. 54 (1951) 227.
Slender climber, up to c. 4m. Stems woody, 1.5—4
mm @, puberulous to glabrous. Leaves: petioles 2—3
cm, minutely puberulous to glabrous, inserted at, or

MENISPERMACEAE (Forman)

259

10. C. barbata

8. C. laxiflora
9. C. peregrina
10. C. barbata

up to 5(—12) mm from, basal margin; lamina lanceo-
late or ovate-lanceolate, apex usually attenuately
acuminate to caudate, sometimes abruptly acumi-
nate, base rounded, truncate or cordate, 6—15 by
(2—) 3—4(—S.5) cm; palmately 3—5-nerved; both sur-
faces usually glabrous but sometimes lightly pube-
scent below, papyraceous. /nflorescences axillary,
narrowly thyrsoid, 7-17 cm, puberulous, female
sometimes a compound thyrse up to c. 40 cm termi-
nating a lateral shoot. — Male flowers yellow, borne
on puberulous pedicels up to 1.5 mm; calyx subcam-
panulate, shortly 4-lobed, 1 mm long, 1.25 mm
broad, glabrous; corolla entire, broadly cyathiform,
0.75 by 1 mm, glabrous; synandrium not exserted, 1
mm long. — Female flowers greenish borne on pube-
rulous pedicels at first 0.5 mm and later up to 1.5
mm; petals and sepals early caducous; sepals 2, +
broadly elliptic, 1 by 0.5 mm, minutely puberulous
outside; petals 2, subrotund, 0.5 mm long, glabrous;
carpel obliquely ovoid 0.75 mm long, glabrous, stig-
ma obscurely and bluntly lobed. Drupes on pedicels
1—2 mm, obliquely obovate in outline, 6—7 by 5—6
mm, 3 mm thick, glabrous; endocarp bearing 3 rows
of papilliform tubercles on both faces.
Distr. Malesia: Sumatra, Malaya, Borneo.
Ecol. Forests at 300—1800 m.

3. Cyclea atjehensis FORMAN, Kew Bull. 14 (1960) 72;
ibid. 34 (1980) 565, f. 3. — C. barbata (non Miers):
Cras, Fl. Siam. En. 1 (1925) 71. — Fig. 18.
Slender woody climber, 2—5 m. Stems densely cov-
ered with a mixture of yellow hairs c. 2 mm long to-
gether with much shorter hairs, eventually glabres-
cent. Leaves with petioles 6—8.5 cm long covered with
both long and short hairs; lamina not peltate, deltoid
ovate, apex acuminate, base + truncate (in Mal.) and
laterally sublobed, palmately S-nerved, 11—14 by
9—10 cm, upper surface softly pubescent especially
along the nerves, lower surface subtomentose, thinly
papyraceous. — Male inflorescences axillary or on
older, leafless stems, narrowly thyrsoid, 4.5 cm long,
0.8 cm broad, yellowish pubescent, flowers in short
lateral cymes. — Male flowers on 1—2 mm pedicels;
sepals 4, shortly connate at the base, oblong, 2 mm
long, externally pilose; corolla reduced to a fleshy
disc 0.25 mm long; synandrium | mm long. — Female
inflorescences similar to male, but larger. — Female
Slowers on pedicels 0.5 mm long; sepals 2, suboppo-
site, elliptic, 1 mm long, externally pilose; petals 2,
episepalous, broadly obovate, 0.25 mm long, fleshy.

240

FLORA MALESIANA

[ser. I, vol. 102

Carpel subellipsoidal, 1 mm long, puberulous be-
coming hispid, stigma with 4—5 spreading, subulate
lobes. Infructescences on older, leafless stems up to
19 cm by 2.5 cm, with the drupes crowded on short
lateral branches. Drupes very shortly pedicellate or
sessile, covered with both long and short hairs, ro-
tund in outline, 5 mm @. Endocarp bearing 6 dorsal
rows of tubercles.

Distr. Thailand; in Malesia: N. Sumatra (Atjeh).

Ecol. Young secondary growth in newly felled
forests at 700—1260 m.

Note. The female flowers are described from
specimens from Thailand.

4. Cyclea merrillii Drets, Pfl. R. Heft 46 (1910) 312,
f. 93F; in Elmer, Leafl. Philip. Bot. 4 (1911) 1167;
Merk. En. Philip. 2 (1923) 151, p.p.; FORMAN, Kew
Bull. 14 (1960) 72; PANcHo, Vasc. Fl. Mt. Makiling
1 (1968) 286.

Slender climber. Stems herbaceous or slightly
woody, 1—2 mm g, puberulous and also bearing
long, spreading hairs. Leaves with tomentose or gla-
brescent petioles 2—5 cm, inserted 1—5 mm above the
basal margin; lamina broadly ovate to triangular-
ovate, apex acuminate, terminating in a long, fine
mucro, base rounded, truncate or subcordate, 7—14
by 5—12 cm, palmately nerved with 5 main ascending
nerves; upper surface pubescent at first, later gla-
brescent, very finely reticulate; lower surface tomen-
tose; papyraceous. — Male inflorescences axillary (?
also cauliflorous), thyrsoid, 6—12(—23) cm long, yel-
lowish pubescent to tomentose; flowers borne in
small cymose clusters on lateral branches up to 1(—4)
cm. — Male flowers with pedicels c. 1 mm; sepals 4,
free, + oblanceolate-elliptic, 1.5 by 0.75 mm, pu-
bescent outside; corolla campanulate, margin suben-
tire, 1 mm long, glabrous; synandrium c. 1 mm long.
— Female inflorescences similar to male but also ter-
minal. — Female flowers on pedicels 1—2 mm; sepals
2, obovate, 1—1.25 by 0.75 mm, densely pilose out-
side; petals 2, obovate, 1 by 0.5 mm, sparsely pilose
outside; carpel + ellipsoidal, very densely pilose, 1
mm long, style short, glabrous, stigma laciniately
4—5-lobed. Drupes unknown.

Distr. Malesia: Philippines (Luzon, Panay).

Ecol. Thickets and forests at low and medium al-
titudes.

Vern.
Negr.

Philippines: abiab, P. Bis., malatuigi,

5. Cyclea insularis (MAKINO) Hatus. Mem. Fac.
Agric. Kagoshima Un. 5, 3 (1966) 29, incl. var. luxu-
rians Hatus. — Cissampelos insularis MAKINO, Bot.
Mag. Tokyo 24 (1910) 227.

Slender climber. Stems sparingly long-pubescent
with short hairs also present, at least on young parts.
Leaves with petioles 3—11 cm inserted 1—2 mm from

basal margin; lamina deltoid-ovate to reniform,
6.5—10 by 6—12 cm, palmately 7—8-nerved, base
broadly to deeply cordate, apex broadly acuminate
to broadly obtuse, upper surface sparsely hairy with
lax reticulation, lower surface rather sparsely long-
pubescent, hairs more dense along the nerves, thinly
papyraceous. /nflorescences axillary, narrowly thyr-
soid up to 18 cm long (female shorter) with lower lat-
eral branches up to 2 cm long, puberulous. — Male
flowers with pedicels 0.5 mm long; sepals 4—S,
joined at base, ovate, 2mm long, sparsely pubescent;
petals united into a cup-shaped corolla 1 mm long;
synandrium 1.5 mm long. — Female flowers: sepals
2, elliptic, 0.5 mm long; petals 2, rotund, 0.25 mm
long; carpel glabrous or sparsely hispid, scarcely 1
mm long, short style with divaricately 3-lobed stig-
ma. Drupes pink, subrotund in outline, 5 mm diam.,
sparsely hispid; endocarp dorsally and laterally set
with 6 rows of short blunt angular points, c. 18—20
per row, with a small slit-like aperture on both lateral
faces.

Distr. Japan; in Malesia: Philippines (Batan I.,
Mt Iraya).

Ecol. In thickets and forest slopes at 100—300 m.
Fl. May, Nov.; fr. May.

6. Cyclea robusta Becc. Malesia | (1877) 157; Diets,
Pfl. R. Heft 46 (1910) 316; Merr. En. Born. (1921)
251; YAMAMOTO, J. Soc. Trop. Agric. 16 (1944) 145;
FORMAN, Kew Bull. 14 (1960) 73; ibid. 22 (1968) 360.

Climber. Stems 2—6 mm g, puberulous and pat-
ently hispid when young, later glabrescent and woo-
dy. Leaves with puberulous (later glabrescent) peti-
oles 6—11 cm, inserted 2—4 cm from the base of lami-
na; lamina triangular-ovate to broadly ovate, apex
gradually acuminate and terminating in a mucro,
base truncate (or slightly emarginate), 15—21 by
9—17 cm; palmately 11—14-nerved; above glabrous,
subnitidous, beneath subtomentose, very prominent-
ly and finely reticulate, stiffly papyraceous. — Male
inflorescences cauliflorous, thyrsoid, 30—35(—55)
cm long, yellowish pubescent, lower branches up to
8 cm, flowers borne in peduncled congested cymose
clusters. — Male flowers with pedicels about 1 mm;
sepals 4, almost free, elliptic to oblanceolate,
puberulous outside, 1.5 by 0.5—0.75 mm; corolla
turbinate, margin crenulate, glabrous, scarcely 1 mm
long; synandrium 1.5 mm long. — Female flowers
unknown. /nfructescence cauliflorous and thyrsoid,
similar to the male inflorescence, and 25 cm long.
Drupes on pedicels 4.5 mm, rotund in outline, 6—7
mm @, sparingly puberulous; endocarp bearing 2
rows of c. 8 transverse + bilobed ridges, condyle
slightly inflated.

Distr. Malesia: Borneo (West, Sarawak, and Sa-
bah); 5 coll.

Ecol. Once recorded from forests at 300 m.

1986 |

7. Cyclea cauliflora MERR. Philip. J. Sc. 26 (1925)
452; En. Philip. 4 (1926) 248; ForMaNn, Kew Bull. 14
(1960) 74.

Slender climber. Stems 1—6 mm @, woody, puber-
ulous or sparsely hispid when young, later glabres-
cent. Leaves: petioles puberulous, 3—7 cm, inserted
11—22 mm above the basal margin; lamina broadly
ovate to suborbicular, apex abruptly acuminate to
subcaudate, acumen finely mucronate, base truncate
to cordate, 7.5—12.5 by 6.5—11 cm; palmately
11—13-nerved; glabrous or subglabrous above, sub-
tomentose below; submembranous. — Male
inflorescences axillary or cauliflorous (sometimes
having the appearance of a terminal or cauliflorous
flowering stem from which the leaves have fallen),
15—120 cm, laxly and repeatedly branched, puberu-
lous, flowers in cymose clusters, bracts linear, to-
mentose, up to 5 mm. — Male flowers white, borne
on pedicels 1—1.5 mm; sepals 4, free, elliptic, 1.5—2
by 0.5 mm, pubescent outside; corolla campanulate,
margin entire, 1 mm long, glabrous, synandrium |
mm long. — Female inflorescences ?cauliflorous, c.
30 cm, with lower lateral branches up to 3(—5) cm,
puberulous; flowers in rather dense clusters arranged
along lateral branches, bracts as in male inflorescen-
ces. — Female flowers borne on pedicels 1—1.25 mm;
sepals 2, variable in shape, + rotund, 0.75 mm long,
pilose outside; petals ?none; carpel obliquely ellip-
soidal, 1 mm long, densely pilose; style 0.25 mm, gla-
brous, lacinately 4- or 5-lobed, lobes 0.75 mm. Dru-
pes on pedicels 3—5 mm, rotund in outline, 5 mmo,
sparsely puberulous; endocarp bearing 4 longitudi-
nal rows of c. 14 papilliform tubercles, both faces
smooth with slight depression at centre and bordered
by prominent ridge.

Distr. Malesia: Philippines (Luzon, Mindoro,
Catenduanes, Leyte, Negros).

Ecol. Once recorded from a forested ridge at 460
m.
Notes. In the specimens of Ramos & EDANO
28836 and 75112 which I have examined, the long
male inflorescences have been coiled round with
leafy stems in such a manner that it is extremely diffi-
cult to follow the course of the inflorescences and
their branches. It seems probable that the apparently
long (up to 120 cm) flowering shoots really represent
a succession of axillary panicles from which the sub-
tending leaves have fallen. There are several un-
doubtedly axillary inflorescences about 15 cm long.

In the two specimens of Ramos 41103 which I have
seen, the female inflorescences are completely de-
tached, and their position on the plant is uncertain.

Cyclea apoensis Yamamoto, Trans. Nat. Hist.
Soc. Taiwan 34 (1944) 312 is probably a synonym of
this species. The type (Etmer 11272, Mindanao) with
very young fruits has narrower leaves.

MENISPERMACEAE (Forman)

241

8. Cyclea laxiflora Miers [ Ann. Mag. Nat. Hist. ser.
3, 18 (1866) 19, nomen], Contr. Bot. 3 (1871) 241;
Diets, Pfl. R. Heft 46 (1910) 317; Rm. Fl. Mal.
Pen. 1 (1922) 114; Burk. & HEND. Gard. Bull. S. S.
3 (1925) 344; Burk. & Hantrr, ibid. 6 (1930) 171;
Burk. Dict. 1 (1935) 722; HEND. Mal. Nat. J. 6
(1951) 415, t. 375; FoRMAN, Kew Bull. 14 (1960) 75.
— C. peltata {non (LaMk) Hook. f. & TH.]: BEcc.
Malesia | (1877) 157, p.p.; Ri. J. Str. Br. R. As.
Soc. n. 33 (1900) 43, ‘var. arnotti Miers’. — C. kort-
halsii Diets ex NorMAN, J. Bot. 62 (1924) Suppl. 5,
nomen.

Slender climber up to c. 15 m. Stems hispid to gla-
brous, herbaceous or slightly woody. Leaves with
pubescent, usually hispid, petioles (1.3—)4—6.2 cm,
inserted 8—22 mm above the basal margin; lamina
triangular-ovate, deltoid-ovate or broadly ovate,
apex acutely acuminate with acumen finely mucro-
nate, base truncate to slightly emarginate, 9-16 by
S—13.5 cm, margin usually hispid; palmately
9—11-nerved; sparsely hispid to glabrescent above,
fairly densely hispid to puberulous or subtomentose
below; papyraceous. — Male inflorescences axillary,
flowers borne in clusters on a lax thyrse 15—55 by
10—20 cm, puberulous. — Male flowers with pedicels
1—2 mm; calyx white or cream, turbinate, glabrous
or subglabrous, 1—1.5 mm long, lobes 4, broadly tri-
angular, about half the length of tube; petals 4, free,
0.5 mm long, glabrous; synandrium c. 1 mm long,
shortly exserted. — Female inflorescences cauliflo-
rous (? always), similar to male, 10—35 by 2.5—10
cm. — Female flowers on pedicels c. 0.5 mm; petals
and sepals unknown (? early caducous); carpel
curved-ellipsoidal, 1 mm long, pilose or glabrous,
stigma with 3 filiform, divaricate lobes. Drupes
white, obliquely obovate to rotund in outline, 5—8 by
4—6 mm, sparsely pubescent; endocarp bearing 3
rows of papilliform tubercles on both faces.

Distr. Extreme Lower Thailand (Pattani); in
Malesia: Sumatra (incl. Banka), Malay Peninsula
(common) and Anambas Is. (Siantan).

Ecol. Hedges, cultivated land, scrub, coastal and
secondary forest; 0O—1200 m.

Uses. According to BURKILL a decoction made
from the roots is used medicinally for fever, piles,
following childbirth and as a vermifuge for children.

Vern. Malaya: akar gasing bukit, a. pahit, a.
rempenang, a. terong kemang (kemar), chawan, me-
timum tikus.

Note. Although Diets described the inflores-
cences as cauliflorous, the male inflorescences are
clearly axillary, as can be seen even in some of the
specimens he cited. In a few examples, the leaf sub-
tending the inflorescence has fallen.

9. Cyclea peregrina Miers [Ann. Mag. Nat. Hist.
ser. 3, 18 (1866) 20, nomen|, Contr. Bot. 3 (1871)

242

FLORA MALESIANA

[ser. I, vol. 102

242; Diets, Pfl. R. Heft 46 (1910) 316; Merr. En.
Born. (1921) 251; Yamamoto, J. Soc. Trop. Agric.
16 (1944) 145; ForMAN, Kew Bull. 14 (1960) 76.

Slender climber. Stem 1.5—4 mm @, hispid when
young, later glabrescent and woody. Leaves with pu-
berulous to glabrescent petioles 2—3 cm inserted
9—16 mm above the basal margin; lamina broadly
ovate, apex acutely acuminate with acumen finely
mucronate, base slightly emarginate, 7.5—10.5 by
5.5—9 cm; palmately 11—12-nerved; subglabrous
above, puberulous below; papyraceous. — Male in-
florescences cauliflorous, 14 by 5 cm, puberulous,
flowers crowded in dense clusters on lateral bran-
ches. — Male flowers subsessile or with puberulous
pedicels up to 0.5 mm; calyx turbinate, sparsely pu-
berulous, 1.25 mm long, lobes 4, broadly triangular,
almost equal to length of tube; petals 4, free, 0.25
mm long, glabrous; synandrium | mm long, scarcely
exserted. — Female inflorescences similar to male.
— Female flowers sessile; sepals and petals un-
known; carpel curved-ellipsoidal, 1 mm long, puber-
ulous, stigma 3-laciniate. Drupes subrotund in out-
line, 5 by 5 mm, puberulous; endocarp bearing 3
rows of papilliform tubercles on both faces.

Distr. Malesia: SE. Borneo.

Note. This species, known only from two speci-
mens collected by MorLey at Bandjarmasin about a
century ago, is intermediate between C. barbata and
C. laxiflora, having the hairy calyx and dense inflo-
rescence of the former together with the minute, free
petals of the latter. It is significant that these two spe-
cies are not known from Borneo. If more material of
C. peregrina becomes available it may prove to be
conspecific with C. Jaxiflora. The dense inflo-
rescence of MorLey 684, although bearing open
flowers, may possibly be immature with the buds
having opened on drying. Furthermore, sparsely
hairy calyces do occasionally occur in C. laxiflora.

10. Cyclea barbata Miers [Ann. Mag. Nat. Hist. ser.
3, 18 (1866) 19, nomen], Contr. Bot. 3 (1871) 237;
Drts, Pfl. R. Heft 46 (1910) 314; Back. Schoolfl.
(1911) 47; Koorp. Exk. Fl. Java (1912) 237; Cras,
Fl. Siam. En. 1 (1925) 70; Heyne, Nutt. Pl. (1927)
618; Burk. Dict. 1 (1935) 721; GaGNep. Suppl. FI.
Gén. I.-C. 1 (1938) 139, p.p.; YAMAMOTO, J. Soc.
Trop. Agric. 16 (1944) 144; ForMAN, Kew Bull. 14
(1960) 77; Back. & Baku. f. Fl. Java 1 (1963) 161.
— C. peltata [non (LaMk) Hook. f. & Tu.]: Mia. Fl.
Ind. Bat. 1, 2 (1858) 86, p.p.; SCHEFF. Nat. Tijd. N.
I. 32 (1873) 393, t. 15; BoERL. Cat. Hort. Bog. 1
(1899) 43; BoorsMA, Teysmannia 11 (1900) 515;
Back. FI. Bat. 1 (1907) 40; Voorl. Schoolfl. (1908) 9.
— C. wallichii Diets, Pfl. R. Heft 46 (1910) 315. —
C. ciliata CratB, Kew Bull. 1922 (1922) 230; Fl. Si-
am. En. 1 (1925) 71.

Slender climber, up to c. 5 m. Roots tuberous.

Stems herbaceous or woody, hispid when young, lat-
er glabrescent. Leaves with hispid petioles 3—6.5 cm,
inserted 6—25 mm above the basal margin; lamina
ovate, deltoid-ovate or broadly ovate, apex acutely
acuminate to obtuse with a finely mucronate acu-
men, base slightly emarginate, truncate or rounded,
6—17.5 by 4—12.5 cm, margin often hispid; palmate-
ly 9—12-nerved; hispid to subglabrous above, puber-
ulous to tomentose below, occasionally hispid along
nerves; papyraceous. — Male inflorescences axillary
or cauliflorous, 7—12(—30) by 0.5—4(—12) cm, pu-
berulous; flowers in dense, subcapitate, hairy
clusters interruptedly borne on short lateral branches
1—4(—7) cm. — Male flowers with pedicels 1—2 mm,
calyx greenish, turbinate, puberulous, 1.5—2 mm
long, lobes 4 (or 5), triangular, about half the length
of tube; corolla gamopetalous, turbinate, margin
truncate or obtusely lobed, 0.75 mm long, glabrous;
synandrium 1.5—2 mm long, usually exserted. — Fe-
male inflorescences similar to male but usually broa-
der, 11—19 by 3—7 cm. — Female flowers sessile in
dense heads; sepals 2, rhomboid to obovate, 0.5 mm
long and broad, pilose on outer surface; petals 2, +
reniform, opposite to and much broader than sepals,
0.5 by 0.75—-1 mm, glabrous; ovary curved-
ellipsoidal, 1 mm long, densely pilose, stigma 3-la-
ciniate. Drupes obliquely obovate to rotund in out-
line, 5—7 mm long, 4—5.5 mm broad, puberulous;
endocarp bearing 3 rows of papilliform tubercles on
both faces.

Distr. Assam, Burma, Thailand, Cochinchina;
in Malesia: NW. Sumatra (Simalur I.), P. Sebesi and
Krakatoa Is. in Sunda Strait, W.—E. Java.

Ecol. In forests (including teak and bamboo for-
est) and in alang? fields, O—1000 m.

Uses. The leaves are commonly used in Java to
prepare a refreshment in the form of a jelly called
‘tjintjau’. This is usually eaten together with a sweet
syrup. The leaves are crushed in water and the mix-
ture left to set. The jelly is also regarded as a stom-
ach-medicine. According to HEYNE, the roots attain
considerable size and when dried are used medicinal-
ly: a brew prepared from them is used as a prophylac-
tic against fever. The very bitter taste is due to the al-
kaloid ‘cycleine’. The starch content of the roots is
high, and there is about 10% fat content.

Vern. Simalur I.: olor kalimenang, o. labana
uding; Java: areuj tarawulu, tjamtjau, S, djudju,
kepleng, krotok, J, kelemaju telor, terung kemau,
tjintjau, Md.

Notes. Three collections by Koorpers differ in.
some respects from the rest of the material. These are
KoorDERS 27853, 34037 (both from Central Java)
and 38535 (E. Java). DieLs annotated the Bogor spe-
cimens of these numbers as ‘Cyclea korthalsii DrELs’,
a name later published as a nomen nudum by Nor-
MAN (/.c.), where it was applied to ForBes 2622a

1986]

from Sumatra, which belongs to C. /axiflora MIERs.
In the index to collectors’ numbers in Diets’ mono-
graph these three KOORDERS numbers are all referred
to ‘Cyclea tomentosa’, a name which I have not been
able to find elsewhere. However, Diets cited the
Same numbers under his general treatment of C.
barbata.

In these specimens the lamina is smaller and nar-
rower (5—8 cm long, 2.5—5 cm broad) than typical C.
barbata and the petiole is inserted only 1—2 mm from
the basal margin of the lamina. The stems differ in
being puberulous and not hispid. The numbers 34037

MENISPERMACEAE (Forman)

243

male buds and very few open flowers, of which two
were dissected and were both found to have one petal
completely free from the rest of the gamopetalous
corolla. | am uncertain of the taxonomic value of the
above differences. These collections may be provi-
sionally regarded as representing aberrant forms of
C. barbata.

Drets placed C. barbata in his key under ‘Synan-
drium inclusum’, although the synandrium is usually
exserted. Cyclea wallichii, which DIcLs originally de-
scribed and placed under ‘Synandrium exsertum’ in
his key cannot be distinguished from it and is accord-

and 38535 are in fruit. Number 27853 bears young __ ingly reduced.

25. STEPHANIA

Lour. Fl. Coch. 2 (1790) 608; Miers, Contr. Bot. 3 (1871) 210; Drets, Pfl. R.
Heft 46 (1910) 259; ForMAN, Kew Bull. 11 (1956) 43; ibid. 22 (1968) 352. — Cly-
pea BL. Bijdr. (1825) 26; Miers, Contr. Bot. 3 (1871) 205. — Fig. 19, 20.
Climbers, mostly slender. Stems woody or herbaceous. Roots sometimes tu-
berous. Leaves with petioles usually geniculate at base; lamina peltate, usually
+ ovate to suborbicular, palmately 8—13-nerved. Inflorescences axillary or
arising from old, leafless stems, usually composed of peduncled umbelliform
cymes which are solitary or racemosely arranged, at least the 1st(—2nd) orders
of branching umbellate (in Mal. spp.), the ultimate branching sometimes irregu-
lar, or sometimes the cymes condensed to disciform capitula. — Male flowers
symmetrical: sepals free, imbricate, 6 or 8 in two equal or unequal whorls, or
only 2—3 in S. capitata, usually + obovate; petals free, 3 or 4 or 2—3 in S. capi-
tata, usually + broadly obovate with lateral margins often involute; stamens
connate into a peltate synandrium, anther-cells 4—8 dehiscing transversely. —
Female flowers symmetrical or asymmetrical: sepals 1—8, petals 2—4, both simi-
lar to male; carpel 1, style very short or absent, stigma shortly lobed or divari-
cately laciniate. Drupes obovoid with style-scar near base, glabrous; endocarp
bony, dorsally bearing a horseshoe-shaped band of 2 or 4 longitudinal rows of
processes or transverse ridges, condyle often perforate. Seed horseshoe-shaped.
Embryo with cotyledons + equalling the radicle, surrounded by endosperm.

Distr. About 35 spp. in the warmer parts, mostly tropics, of the three continents in the Old World;
throughout Malesia.

Notes. The Malesian species of Stephania fall into Diets’ sections Thamnothyrsa and Eustephania. These
sections are not maintained in the following account since the distinctions between them, which were based
on the position and form of the inflorescence, are unreliable.

As to life-form, the species differ: S. japonica, S. psilophylla and S. venosa seem to have annual or seasonal
stems, while S. corymbosa and S. zippeliana have perennial woody stems.

The form of the inflorescence is specifically important in Stephania; in all the Malesian species its organisa-
tion is based on one or more peduncled, umbelliform cymes, which show varying degrees of condensation
in different species. In S. montana and S. corymbosa the cymes are lax and bear pedicellate flowers. In S.
Japonica the flowers are sessile in subcapitate globose clusters. Extreme condensation of the cyme-branches
occurs in S. capitata and S, dictyoneura resulting in a solid, disciform receptacle on which the flowers are

244 FLORA MALESIANA [ser. I, vol. 102

very densely crowded. The composition of the inflorescence is usually either a solitary umbelliform cyme as
in S. japonica and S. venosa or a racemose arrangement of umbelliform cymes as in the other species. The
former composition is a character used by Diets for his section Eustephania and the latter for his section
Thamnothyrsa. \n S. psilophylla, however, a racemose arrangement of cymes is usual but solitary cymes also
occur.

Externally, the fruits of different species are similar; the differences are to be found internally. The remark-
able ornamentation of the endocarp is usually distinct for each species. There are 2 or 4 rows of processes
in the form of ridges, hooks, rods or plates, the rows running longitudinally along the dorsal surface of the
endocarp. Each species has a characteristic range in the number and form of these processes.

KEY TO MALE PLANTS

(Note: The male of S. dictyoneura is not known but from female inflorescences it is clear that the species be-
longs next to S. capitata in this key.)

1. Inflorescences composed of umbelliform cymes.
Dablowersisessile on subsessilemm denserchustensiget racks oe sere wie. 2 titctet oe tote! sl dels ons ol veneeds 1. S. japonica
2. Flowers pedicellate.
3. Inflorescences verruculose; peduncles of cymes arising retrorsely from main axis. Leaf-margin with wide,
shallow crenations; petiole inserted’ 2—3 mm from base .....................-.-- 2. S. grandiflora
3. Inflorescences glabrous, papillose or puberulous; peduncles of cymes not retrorse along main axis. Leaf-
margin not widely crenate; petiole inserted more than (6—)10 mm from base.
4. Leaves submembranous, margin often slightly lobed, reticulation scarcely raised and usually drying red-
dish brown. Inflorescence usually a solitary axillary subumbel ..................... 3. S. venosa
4. Leaves papyraceous to coriaceous.
5. Inflorescences less than 1.5 cm long. Reticulation on lower surface of leaf very fine and raised
4. S. reticulata
5. Inflorescences 3 cm or more long.
6. Sepals unequal. Petals papillose-puberulous within. Upper surface of leaf often minutely subpapil-
losen(lanceyepicenmalsCells)icea metus qrae ee «teal Ge ve epreuaebaing. oouckols eu. ale pales 5. S. corymbosa
6. Sepals + equal. Petals glabrous. Upper surface of leaf not subpapillose.
7. Leaves with lax reticulation.
8. Leaves triangular-orbicular to orbicular, apex rounded to very broadly obtuse 6. S. salomonum
8. Leaves triangular to ovate, apex acute to obtuse.
9. Umbelliform cymes 2—5 cm with primary rays less than 1 cm. Leaves ovate to ovate-triangular,

APCRAODUISE Mr a Meera sc ae eae ow ciw Sikes ce sear anlereialna te ole. c leucts steroiee 7. S. moluccana
9. Umbelliform cymes 4—14 cm with primary rays more than 1 cm. Leaves triangular to narrowly
tian cilarerapexVeLVeAC leit .15 caraway Gh eae ces atin Marte Manteo ee 8. S. montana

7. Leaves with very fine reticulation.
10. Leaves 5—9 by 4—8 cm. Inflorescences axillary, composed of very slender umbelliform cymes

PAS ONCIIPLO MORN ee eee ee hee oad ORNS RLS COL 9. S. psilophylla
10. Leaves usually more than 16 by 10 cm. Inflorescences very large (over 30 cm long) usually on old,
leafless stems, composed of stout umbelliform cymes over 10 cm long ..... 10. S. zippeliana

1. Inflorescences composed of peduncled disciform capitula.
11. Leaves with a lax and rather obscure reticulation, apex often caudate; petiole inserted more than 10 mm
AORN ASC Fee ee NONI ar erat s clea BARE RESPIR OID, Sheet cae 11. S. capitata
11. Leaves with a close, raised reticulation, apex shortly acuminate; petiole inserted 3—7 mm from base
12. S. dictyoneura

KEY TO FEMALE PLANTS
1. Inflorescences composed of umbelliform cymes.

2. Inflorescences verruculose. Leaf-margin with wide, shallow crenations; petiole inserted 2-3 mm from
DASE treaty rats. ia Sab seeders GM aed - Ae eke BR A Ree ee 2. S. grandiflora

1986 |

MENISPERMACEAE (Forman)

245

2. Inflorescences glabrous, papillose or puberulous. Leaf-margin not widely crenate; petiole inserted more

than (6—)10 mm from base.

3. Leaves submembranous, margin often slightly lobed, reticulation scarcely raised and usually drying red-
dish brown. Inflorescence usually a solitary axillary cyme. Flowers asymmetrical: sepal 1, petals 2

3. Leaves papyraceous to coriaceous.

3. S. venosa

4. Drupes sessile or subsessile (excluding ultimate infructescence branchlets).
5. Leaves with a rather lax reticulation, often obscure on upper surface which is often minutely papillose.
Endocarp generally suborbicular in outline with 4 dorsal rows of (10—)12—19 hooks. Female flowers

PEMeRHESHIGAL~.; 5.002 vs. seins. Le eee

5. S. corymbosa

5. Leaves with a fine reticulation usually visible on both surfaces. Endocarp obovate in outline with 2
or 4 dorsal rows of ridges or lamelliform projections. Female flowers symmetrical . 1. S. japonica
4. Drupes pedicellate, apart from occasional sessile ones.
6. Leaves with very fine reticulation visible on one or both surfaces.
7. Drupes 4—5 mm long. Leaves 5—9 by 4—8 cm. Endocarp with 2 or 4 dorsal rows of 7—9 ridges or pa-

pilliform processes. Inflorescences axillary

9. S. psilophylla

7. Drupes 7—10 mm long. Inflorescences usually on old, leafless stems.
8. Leaves 7—13 by 5—10 cm. Endocarp with 4 dorsal rows of 9—11 very prominent projections, the

outer 2 rows lamelliform, dorsal surface very scabrid...................00005

4. S. reticulata

8. Leaves usually more than 16 by 10 cm. Endocarp with 4 dorsal rows of 9—14 short papilliform pro-
cesses or low ridges (sometimes T-shaped), dorsal surface slightly rough or smooth 10. S. zippeliana
6. Leaves with a lax reticulation visible on both surfaces.
9. Leaves triangular-orbicular to orbicular, apex rounded to very broadly obtuse . 6. S. salomonum
9. Leaves triangular to ovate, apex acute to obtuse.

10. Leaves ovate to ovate-triangular, apex obtuse .................2022eeeeee
10. Leaves triangular to narrowly triangular, apex very acute....................

7. S. moluccana
8. S. montana

1. Inflorescences composed of peduncled, disciform capitula.
11. Leaves with a lax and rather obscure reticulation, apex often caudate; petiole inserted more than 10 mm

IORI Saat 15s 2 ere a te AL Seas

11. S. capitata

11. Leaves with a close, raised reticulation, apex shortly acuminate; petiole inserted 3—7 mm from base

1. Stephania japonica (THUNB.) Miers, Ann. Mag.
Nat. Hist. ser. 3, 18 (1866) 14, p.p., excl. CUMING
1160; Contr. Bot. 3 (1871) 213, p.p., excl. CUMING
1160; Diets, Pfl. R. Heft 46 (1910) 277; in Elmer,
Leafl. Philip. Bot. 4 (1911) 1166; JuEL, Pl. Thunb.
(1918) 245; W.H. Brown, Minor Prod. Philip. For.
3 (1921) 186; GaGnep. Suppl. Fl. Gén. I.-C. 1 (1938)
134; YAMAMOTO, Taiwania | (1948) 46; FoRMAN,
Kew Bull. 11 (1956) 49. — Menispermum japonicum
Tuuns. Fl. Jap. (1784) 193; LamK, Encycl. Méth. 4
(1797) 96; WiLtp. Sp. PI. 4 (1806) 827. — Cissampe-
los hernandiifolia WiLp. Sp. Pl. 4 (1806) 861 (‘her-
nandifolia’); Dc. Syst. 1 (1817) 533; Prod. 1 (1824)
100; Roxs. Fl. Ind. ed. Carey 3 (1832) 842. — Coccu-
lus japonicus (THuNB.) De. Syst. 1 (1817) 516; Prod.
1 (1824) 96, incl. var. timoriensis Dc. — Cocculus
Jforsteri Dc. Syst. 1 (1817) 517; Prod. 1 (1824) 96;
Gutttemin, Ann. Sci. Nat. II, 7 (1837) 370. — Cly-
pea discolor Bu. Bijdr. (1825) 26. — S. discolor (BL.)
SprenGc. Syst. Veg. 4 (1827) 316; Hassk. Pl. Jav.
Rar. (1848) 168; Miers, Contr. Bot. 3 (1871) 224;
Bogert. Cat. Hort. Bog. 1 (1899) 42, incl. var. her-
nandiifolia (WiLLD.) BOERL.; BACK. Fl. Bat. 1 (1907)
39; Voorl. Schoolfl. (1908) 9; Rint. J. Str. Br. R. As.
Soc. n. 87 (1923) 52. — Clypea glaucescens Decne,

12. S. dictyoneura

Nouv. Ann. Mus. Paris 3 (1834) 423, t. 18; SPAN.
Linnaea 15 (1841) 164. — S. glaucescens (DECNE)
Wa Lp. Rep. | (1842) 96; Miers, Contr. Bot. 3 (1871)
214. — S. hernandiifolia (WiLLD.) WALP. Rep. 1
(1842) 96; Hook. f. & Tu. Fl. Ind. 1 (1855) 196, p.p.;
Mia. Fl. Ind. Bat. 1, 2 (1858) 83; BENTH. FI. Austr.
1 (1863) 57, p.p.; Kurz, Nat. Tijd. N. I. 27 (1864)
177; Miers, Contr. Bot. 3 (1871) 222; Mig. Ann.
Mus. Bot. Lugd.-Bat. 4 (1868) 85, incl. var. genuina
seu glabra et var. discolor (Bu.) Mig.; Hook. f. & TH.
Fl. Br. India 1 (1872) 103; ScHErFF. Nat. Tijd. N. I.
32 (1873) 401; Kurz, J. As. Soc. Beng. 43, ii (1874)
61; Becc. Malesia 1 (1877) 154; Kina, J. As. Soc.
Beng. 58, ii (1889) 385; Wars. Bot. Jahrb. 13 (1891)
314; Rit. Trans. Linn. Soc. I], 3 (1893) 274; K. Scu.
Notizbl. Berl.-Dahl. 2 (1898) 116; BamLey, Queensl.
Fl. 1 (1899) 33, p.p.; K. Scu. & Laut. Fl. Deut.
Schutzgeb. Siidsee (1901) 311; BRANbis, Indian Trees
(1906) 23; GaGnep. Fl. Gén. I.-C. 1 (1908) 147;
Diets, Pfl. R. Heft 46 (1910) 279; Back. Schoolfl.
Java (1911) 46; Koorp. Exk. Fl. Java 2 (1912) 335;
Merr. En. Born. (1921) 250; Ript. Fl. Mal. Pen. 1
(1922) 113; Burk. & Henpb. Gard. Bull. S. S. 3 (1925)
344; Crass, Fl, Siam. En. 1 (1925) 69; Heyne, Nutt.
Pl. 1 (1927) 617; Back. Onkr. Suiker. 2 (1930) 247;

246 FLORA MALESIANA [ser. I, vol. 102

Fig. 19. Stephania japonica (THUNB.) Miers. a—d, f—h. var. timoriensis (DC.) FORMAN, e. var. discolor (BL.)

FORMAN, g—i. var. japonica. a. Habit, 2/3, b. male flower, 20, c. sepal, x15, d. petal, inner view, x15,

e. female flower, x20, f. drupe, x2, g. endocarp, 4 with h. detail of ornamentation viewed along the me-

dian dorsal line, x10, 7. variant of ornamentation, x10. — S. venosa (BL.) SPRENG. j. Female flower with

1 sepal and 2 petals, x20. — S. corymbosa (BL.) Wap. k. Male flower, x10, /. petal, glandular inner face,

x15, m. drupe, <2, n. detail of ornamentation on endocarp, x10 (a—d ELBERT 3998, e S 40966, f—h RoBIn-
son 487, i BS 80201, 7 KoorDERS 25871, k—/ BS 80080, m—n Koss 19126).

1986 |

Atlas (1936) t. 258; Burk. Dict. 2 (1935) 2076; MERR.
J. Arn. Arb. 19 (1938) 340; Masam. En. Phan. Born.
(1942) 276; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 138; Taiwania 1 (1948) 48. — S. forsteri (Dc.)
A. Gray, Bot. U.S. Expl. Exped. | (1854) 36; DrELs,
Pfl. R. Heft 46 (1910) 278; Nova Guinea 8 (1910)
283; Back. Schoolfl. Java (1911) 46; Koorp. Exk.
Fl. Java 2 (1912) 336. — S. concinna Mrers [Ann.
Mag. Nat. Hist. ser. 3, 18 (1866), 15, nomen] Contr.
Bot. 3 (1871) 226. — S. exigua Miers [Ann. Mag.
Nat. Hist. ser. 3, 18 (1866) 16, nomen] Contr. Bot.
3 (1871) 228. — S. hallieri Diets, Pfl. R. Heft 46
(1910) 281; Merr. En. Born. (1921) 250; Masa. En.
Phan. Born. (1942) 276; Yamamoto, J. Soc. Trop.
Agric. 16 (1944) 140. — Fig. 19a—i.

For further names based on extra-Malesian types,
see FORMAN (1965: 55).

Slender climber, 2—10 m. Root tuberous. Stem
herbaceous or thinly woody, glabrous or crispately
puberulous. Leaves with glabrous or crispately pu-
berulous petioles 3-12 cm; lamina _ broadly
triangular-ovate to ovate, (4—)6—12(-17) by
4—10(—14) cm, apex usually + acuminate with the
acumen usually obtuse and mucronulate but some-
times very acute or very obtuse, base broadly round-
ed to slightly cordate; fine reticulation usually visible
on both surfaces, upper surface glabrous; lower sur-
face glabrous or crispately puberulous and also
sometimes glaucous (due to very minute whitish pa-
pillae);} + papyraceous. — Male inflorescence an
axillary compound umbelliform cyme 4—9 cm, bear-
ing flowers in dense, subcapitate, cymose clusters,
glabrous or puberulous, usually solitary, but occa-
sionally paired or a few borne on an axillary shoot.
— Male flowers green, white or yellow, sessile or sub-
sessile: sepals 6 or 8, glabrous or puberulous outside,
oblanceolate to spathulate, 0.75—1.25 mm; petals 3
or 4, glabrous, + obdeltoid to suborbicular, 0.5—1
mm; synandrium 0.5—1 mm long, exsert or not. —
Female inflorescence similar to male. — Female
flowers with petals and sepals as in male but number
of sepals often reduced; carpel + ovoid, 0.75—1 mm.
Drupe red, sessile or subsessile (occasionally with
pedicels up to 2 mm long), + obovate to suborbicu-
lar in outline, 4—8 by 4—6 mm. Endocarp usually
perforate, dorsally bearing 2 longitudinal rows of
about 8—10 transverse ridges which are often +
2-lobed thus forming 4 distinct rows of processes, +
papilliform in the 2 inner rows and broader and often
hooked in the 2 outer rows; surface between and
around tubercles usually smooth, rarely rough.

Distr. Japan, S. China, Indochina, Thailand, In-
dia, throughout Malesia, N. & E. Australia,
Polynesia.

Ecol. In hedges, thickets, river-banks, secondary
growth and forests, from sea-level to 2000 m. Fi. fr.
Jan.—Dec. (at least, in Java).

MENISPERMACEAE (Forman)

247

Uses. In var. discolor the tuberous root is bitter
and very poisonous due to its picrotoxin content. It
is used medicinally for fever, diarrhoea, urinary dis-
eases and stomach-ache. Crushed leaves in water
form a slightly gelatinous mass which is applied to
breast infections. Although reported to be poisonous
to livestock in Australia (WHITE, Queensl. Agric. J.
n.s. 8, 1917, 230), feeding tests have proved negative
(EvERIsT, Pois. Pl. Austr. rev. ed., 1981, 528).

Vern. Java: areuj gurrung, djadjo, djundju lala-
keh, kepleng, ojod minjak, ondjo ondjo, seloro, slu-
ru, tjiluru, tjimtjau minjak; Moluccas: ginato bobu-
do, gurnali babudo, Ternate; New Guinea: bidililim,
West; worarugan, East, Hagen-Chimbu.

Notes. The very extensive synonymy of this spe-
cies is the result of combining the synonyms quoted
by Diets for three species, which he recognized as
distinct, viz. S. forsteri(DC.) A. Gray, S. hernandii-
folia (WiLLD.) Wap. and S. japonica (THUNB.)
Mrers. The only differences between these taxa con-
cern the presence or absence of a puberulous indu-
mentum on the lower surfaces of the leaves and on
the inflorescences, as indicated in the key below. The
degree of hairiness varies considerably but the great
majority of the specimens I have examined can be
placed in one of these three taxa, which occupy dif-
ferent but overlapping geographical areas. I there-
fore consider S. forsteri, S. hernandiifolia and S. ja-
ponica to be no more than varieties of one wide-
spread species.

For a detailed discussion on the types see FORMAN
(1956).

The endocarp of S. japonica was described by
Diets as imperforate. It is, however, usually per-
forate in all three varieties of this species.

KEY TO THE VARIETIES

1. Leaves glabrous below (region around insertion of
petiole sometimes puberulous).

2. Inflorescences glabrous ...... a. var. japonica

2. Inflorescences puberulous.. b. var. timoriensis

1. Leaves puberulous below (apart from region
around insertion of petiole .... ¢. var. discolor

a. var. japonica

For synonymy, see FORMAN (1956: 49).

Plant entirely glabrous; the bracts and sepals
(outer surface) sometimes papillose.

Distr. Ceylon, SW. Deccan, E. China, Korea,
Japan, Taiwan, Tonga & Society Is.; in Malesia: Les-
ser Sunda Is. (Timor), Moluccas (Ceram), Philip-
pines (Batan Is., Luzon, Mindoro, Negros, Pala-
wan).

b. var. timoriensis (DC.) Forman, Kew Bull. 11

248

FLORA MALESIANA

[ser. I, vol. 102

(1956) 49. — Fig. 19a—d, f—h.

For synonymy, see FORMAN (/.c.).

Leaves glabrous (sometimes sparsely puberulous
on the lower surface around the insertion of the pet-
iole). Inflorescences crispately puberulous.

Distr. E. Bengal to N. Australia and Polynesia
(New Caledonia, Samoa, Tahiti); in Malesia: Central
& E. Java, Lesser Sunda Is., SW. Celebes, Moluccas
(Ceram, Ambon, Kei Is.), Papua New Guinea.

c. var. discolor (BL.) FORMAN, Kew Bull. 11 (1956)
49. — Fig. 19e.

For synonymy, see FORMAN (/.c.).

Leaves crispately puberulous below. Inflorescen-
ces crispately puberulous.

Distr. Tropical SE. Asia: S. China (Yunnan), In-
dia, Nepal, Burma, Thailand, Indochina to E. Aus-
tralia; in Malesia: throughout, except the Philip-
pines.

2. Stephania grandiflora FoRMAN, Kew Bull. 22
(1968) 354.

Woody climber. Young stems, petioles and inflo-
rescences verruculose. Leaves with petioles 3—5 cm
inserted 2—3 mm from the basal margin; lamina
triangular-ovate, 6—12 by 4.5—9 cm, base truncate,
apex acute or obtuse, margin broadly and slightly
crenate, reticulation dense, raised and very conspicu-
ous on both surfaces, glabrous, thinly coriaceous. —
Male inflorescences axillary, c. 22—35 cm long, com-
posed of umbelliform cymes 10—12 cm long with
peduncles 5—7 cm, arising retrorsely from the main
axis. — Male flowers on slender pedicels, c. 2 mm;
sepals 6 or 8, elliptic or broadly elliptic, 3-4 mm, gla-
brous; petals 3 or 4, cuneate-oblong, 3 mm with the
lateral margins inflexed, apex truncate, glabrous;
synandrium 3 mm long. — Female flowers un-
known. Drupe yellow on pedicel up to 2 mm long,
obovate in outline, 6 mm long. Endocarp imperfor-
ate with 4 dorsal rows of c. 13 processes, papilliform
in the median rows, but consisting of transverse rid-
ges, hooked on the outer margins in the lateral rows.

Distr. Malesia: East New Guinea (E., W.& S.
Highlands).

Ecol. Primary and seondary forest by streams
and in regrowth in logged areas, on peaty humic soil
and humic soil over orange-brown clay at 2300—2700
m. Fi. June, July; fr. Feb., June, Sept., Oct.

Uses. Local people make abdominal belts from
the stems.

Vern. Kepilam, kwik, Enga dial., kuip, Wabag,
kuru ponde, Hagen Distr.

Note. This is a most distinctive species of Stepha-
nia in several respects. The leaves differ from those
of other species in the wide and shallow crenations at
the margin of the lamina, which is densely reticulate
and comparatively thick in texture. The flowers are

the largest known in the genus. The inflorescences
are remarkable in that the main branches, /.e. the
peduncles of the umbelliform cymes, are directed
backwards towards the base of the inflorescence,
suggesting that the inflorescences are pendulous with
the retrorse branches directed upwards.

3. Stephania venosa (BL.) SPRENG. Syst. Veg. 4
(1827) 316; Wap. Rep. 1 (1842) 96; Diets, Pfl. R.
Heft 46 (1910) 271; Koorp. Exk. Fl. Java 2 (1912)
336; MasaM. En. Phan. Born. (1942) 276; YAMA-
moto, J. Soc. Trop. Agric. 16 (1944) 141; ForMAN,
Kew Bull. 11 (1956) 58; Back. & Baku. f. Fl. Java
1 (1963) 160 (‘venenosa’); FoRMAN, Kew Bull. 22
(1968) 352. — Clypea venosa BL. Bijdr. (1825) 27. —
S. prapatensis YAMAMOTO, J. Soc. Trop. Agric. 16
(1944) 143. — S. rotunda (non Lour.) Mig. Ann.
Mus. Bot. Lugd.-Bat. 4 (1868) 86; Scuerr. Nat. Tijd.
N. I. 32 (1873) 401, t. 14; Becc. Malesia 1 (1877) 155;
Rw i. J. Str. Br. R. As. Soc. n. 54 (1910) 15; ibid. n.
59 (1911) 67 (‘rotundifolia’); Fl. Mal. Pen. 1 (1922)
113. — S. hernandiifolia [non (WiLLD.) WALP.]
Rw. J. Str. Br. R. As. Soc. n. 59 (1911) 67. — Fig.
19}.

Slender climber up to c. 20 m, containing red sap.
Root tuberous. Stem herbaceous, glabrous. Leaves
with glabrous petioles (3—)6—22 cm; lamina broadly
triangular-ovate, margin often slightly lobed, 6—20
by 6—20 cm, apex obtusely mucronulate, base slight-
ly cordate; lower surface rather pale with a darker ve-
nation (in sicco usually reddish brown) very minutely
papillose, otherwise glabrous; upper surface usually
darker, glabrous; submembranous. — Male inflores-
cences an axillary, peduncled, umbelliform cyme,
6—15 cm, papillose (occasionally glabrous), usually
solitary. — Male flowers yellow to orange with pedi-
cels 1—2 mm; sepals 6, papillose outside, outer 3 ob-
lanceolate, 2—2.5 mm long; inner 3 obovate, ungui-
culate, 2—2.5 mm long; petals 3, obdeltoid-obovate,
1.25—1.5 mm long, glabrous; synandrium 1—1.75
mm long. — Female inflorescence similar to male. —
Female flowers with pedicels about 1 mm, asymmet-
rical; sepal 1, elliptic-oblong to obovate, 1—1.25 by
0.5 mm, papillose or glabrous outside; petals 2, +
suborbicular to obdeltoid, 0.5—0.75 mm, glabrous.
Sepal and petals on one side of the flower; carpel +
ellipsoidal, 1—1.5 by 0.5—0.75 mm. Drupe red with
pedicels up to 7 mm, obovate in outline, 6—9 by 5—6
mm. Endocarp perforate, dorsally bearing 4 longitu-
dinal rows of 12—16 papilliform processes.

Distr. Andaman Is., Thailand, Vietnam, S. Chi-
na (Yunnan); in Malesia: N. Sumatra, N. Malaya
(Perlis, Langkawi Is.), Java, N. Borneo (Mt Kinaba-
lu), Philippines (Luzon), and SW. Celebes.

Ecol. On hillsides, plains and mountains, scat-
tered in forests up to 1600 m. On limestone in Ma-
laya.

1986 |

Vern. Java: gorong bodas.

Note. Some male specimens from Langkawi Is.
have a fleshy scale on the inside of each petal which
may indicate a distinct taxon.

4. Stephania reticulata FORMAN, Kew Bull. 11 (1956)
65; ibid. 22 (1968) 352; ibid. 39 (1984) 113.

Woody climber. Leaves with petioles 4-12 cm,
glabrous or puberulous towards the apex; lamina tri-
angular-ovate, 7—13 by S—10 cm, apex usually atten-
uately acuminate, base truncate or broadly rounded,
lower surface with reticulation very fine and raised,
glabrous or sparsely puberulous towards the base of
the nerves, upper surface with laxer reticulation,
glabrous, papyraceous. — Male inflorescences axil-
lary, less than 1.5 cm long, a solitary umbelliform
cyme or a few cymes arising from a very short axis,
glabrous. — Male flowers on pedicels 1.25—1.5 mm
in dense subglobose clusters; sepals 6, elliptic or ob-
lanceolate, 1—1.25 mm long; petals 3, broadly ob-
ovate, the lateral margins inflexed, 0.5 mm long; syn-
andrium 0.75—1 mm long. — Female inflorescences
similar to male but much larger. /nfructescence axil-
lary or on older, leafless stems, 4—14 cm long, sub-
glabrous or puberulous. — Female flowers subsessile
or on pedicels c. 0.5 mm long; sepals and petals simi-
lar to male; carpel 1 mm long. Drupe yellow to red
on pedicel 3—5 mm, sometimes sessile, obovate to
suborbicular in outline, 7—9 by 7—8 mm. Endocarp
perforate with 4 dorsal rows of 9—11 projections irre-
gularly divided at the apices, the 2 median rows +
rod-like, the 2 outer rows lamelliform with the outer
margins hooked, the whole dorsal surface very sca-
brid.

Distr. Tenasserim and Peninsular Thailand (Pa-
talung; Pattani); in Malesia: W. Java and N. Borneo
(Sarawak; Mt Kinabalu & Crocker Range).

Ecol. In forests, in continental Asia at c.
300—400 m, in W. Java 500-1000 m, in Sabah
1000—1700 m.

Notes. This species is clearly distinguished by its
very small, axillary, male inflorescences bearing
shortly pedicellate flowers, and also by the pedicel-
late fruits containing remarkable endocarps which
dorsally bear 2 inner rows of + rod-like projections
and 2 outer rows of thin, transverse, plate-like pro-
jections, which are hooked on the outer margin. The
projections are irregularly divided at their tips and
the whole dorsal surface of the endocarp between the
projections is very rough.

Backer & Baku. /. (Fl. Java 1, 1963, 157) exclude
this species from W. Java, but I disagree (/.c. 1968).

5. Stephania corymbosa (Bi.) WALP. Rep. | (1842)
96; Mio. FI. Ind. Bat. 1, 2 (1858) 84; Ann. Mus. Bot.
Lugd.-Bat. 4 (1868) 86; Becc. Malesia | (1877) 155;
Diets, Pfl. R. Heft 46 (1910) 267; Back. Schoolfl.

MENISPERMACEAE (Forman)

249

Java (1911) 46; Koorp. Exk. Fl. Java 2 (1912) 336;
Back. Bull. Jard. Bot. Btzg II, 12 (1913) 31; Rwt. J.
Fed. Mal. Stat. Mus. 8 (1917) 14; Fl. Mal. Pen. 1
(1922) 113; Merr. En. Born. (1921) 250; Burk. &
HEND. Gard. Bull. S. S. 3 (1925) 344; Norman, J.
Bot. 64 (1926) Suppl. 142; Masam. En. Phan. Born.
(1942) 276; YAmMAmorTo, J. Soc. Trop. Agric. 16
(1944) 137; ForMAN, Kew Bull. 11 (1956) 60, f. 4;
Back. & Baku. f. Fl. Java 1 (1963) 160; PANcHo,
Vasc. Fl. Mt. Makiling 1 (1983) 285. — Clypea co-
rymbosa Bu. Bijdr. (1825) 27. — S. ramuliflora
Miers [Ann. Mag. Nat. Hist. ser. 3, 18 (1866) 16,
nomen], Contr. Bot. 3 (1871) 232. — ? S. cauliflora
Becc. Malesia | (1877) 155; Diets, Pfl. R. Heft 46
(1910) 267; HottH. & Lam, Blumea 5 (1942) 180. —
S. hernandiifolia {non (WitLLD.) WALP.] Koorp.
Meded. Lands Pl. Tuin 19 (1898) 340. — S. catose-
pala Diexs, Pfl. R. Heft 46 (1910) 268; in Elmer,
Leafl. Philip. Bot. 4 (1911) 1166; Merr. En. Philip.
2 (1923) 149, p.p., excl. MERRILL 828, 9705 et SANTOS
31806. — S. merrillii Diets, Pfl. R. Heft 46 (1910)
268; in Elmer, Leafl. Philip. Bot. 4 (1911) 1166;
Me_rr. En. Philip. 2 (1923) 150; YAMAmoro, J. Soc.
Trop. Agric. 16 (1944) 137. — S. ramosii DiELs, Phi-
lip. J. Sc. 7 (1912) Bot. 265; Merr. En. Philip. 2
(1923) 150. — S. menadonensis Diets ex Koorp.-
ScHum. Syst. Verz. (1914) 41, nomen. — ? S. bor-
neensis YAMAMOTO, J. Soc. Trop. Agric. 16 (1944)
137. — Fig. 19k—n.

Woody climber, about 5—12 m, main stem about
1—2 cma. Leaves with glabrous petioles 3—19 cm; la-
mina broadly triangular-ovate to suborbicular (often
narrowly triangular-ovate in Luzon), 3.5—13(—19)
by 3—11(—17) cm, apex acute and usually abruptly
acuminate, sometimes gradually narrowing or ob-
tuse, base broadly rounded to truncate; margin
sometimes revolute; reticulation rather lax, often ob-
scure On upper surface, which is often minutely sub-
papillose (large convex epidermal cells), both sur-
faces glabrous; papyraceous to subcoriaceous. —
Male inflorescence often borne on the older, leafless
parts of the stem, but axillary on young shoots,
3—10(—15) cm long, composed of peduncled umbel-
liform cymes 1.5—9 cm long, racemosely arranged,
papillose-puberulous to glabrous. — Male flowers
white, yellowish or orange with pedicels up to 3(—4)
mm long; sepals 6, glabrous or subglabrous outside,
outer 3 + narrowly oblong, 1.25—1.75 mm, apex ob-
tusely pointed, inner 3 obovate, 1.25—1.5 mm, apex
broadly rounded; petals 3, + broadly obtriangular,
c. 0.75 mm long, apical margin membranous, lateral
margins sometimes involute, papillose-puberulous
inside towards base; synandrium 0.75—1 mm long.
— Female inflorescence similar to male. /nfructes-
cence up to 20 cm or more. — Female flowers sessile,
asymmetrical; sepals 3, + elliptic, 0.5 mm long; pet-
als 1—2, broadly obtriangular, scarcely 0.5 mm long,

250

FLORA MALESIANA

[ser. I, vol. 102

papillose or glabrous within; carpel + ellipsoidal,
0.75 by 0.5 mm. Drupes red, sessile or subsessile (but
ultimate branchlets of inflorescence sometimes look
like pedicels), suborbicular to obovate in outline,
6—10 by 6-8 mm. Endocarp perforate, dorsally
bearing 4 longitudinal rows of (10—)12—19 trans-
versely hooked projections, the 2 inner rows directed
inwards and those of the 2 outer rows directed out-
wards (rarely the hooks scarcely or not developed).

Distr. Malesia: Sumatra (incl. Simalur I.), Ma-
laya (Maxwell’s Hill, Perak), W. Java, Borneo, Phi-
lippines, S., Central & NE. Celebes.

Ecol. Occurring in thickets and forests up to 1800
m. Fl. fr. Jan.—Dec.

Uses. Stems used for tying in Sabah.

Vern. Java: areuj geureung, geureng arent; Sa-
bah: penaki, puruut, Murut; Celebes: karokas sela.

Notes. In my revision (/.c. 1956: 62) I comment-
ed on the types of the synonyms, and discussed the
variability.

Drets (1910) distinguished between S. corymbosa
(Bt.) WaALP., S. cauliflora BEcc., S. catosepala DIELS
and S. merrillii DreLs by using the following charac-
ters in his key: relative length of lamina and petiole,
shape of sepals, leaf-texture, and thickness of synan-
drium-column. In the material I have examined, the
distinctions made by Drets are not reliable, nor do
there seem to be any other characters which could be
used to distinguish more than one species in this
group.

The endocarp of S. corymbosa is characterized by
the 4 dorsal longitudinal rows of transversely direct-
ed hooks, which are developed to various degrees of
prominence in specimens from different localities.
Only at the extreme eastern and western limits of the
geographical range of the species are the hooks not,
or only scarcely, developed, i.e. in Simalur I. and in
Celebes.

6. Stephania salomonum Drets in Rech., Bot. Zool.
Ergebn. Samoa-Salomons 5 (1913) 110, f. 21.
Woody climber, entirely glabrous. Leaves with
glabrous petioles (4—)7—10 cm inserted 2.5—6 cm
from the basal margin; lamina triangular-orbicular
to orbicular, apex rounded to very broadly obtuse,
mucronate, 9-19 by 9—19 cm, reticulation lax,
raised on both surfaces, glaucous below, margin
slightly crenate-undulate, papyraceous. — Male in-
florescences axillary or arising from leafless stems,
slender, lax, c. 20—23 cm long, composed of a ra-
zeme of umbelliform cymes 4—9 cm long on ped-
uncles 3—4 cm. — Male flowers on pedicels 1—2.5
mm; sepals 6, + equal, oblanceolate to obovate, 2
mm long, membranous; petals 3, broadly obdeltoid,
0.5—0.75 mm long, fleshy; synandrium 1 mm long.
— Female inflorescences and flowers not seen (ac-
cording to Diets: long-pedunculate, 40—45 cm long).

Infructescences 12(—?) cm long. Drupe red, on pedi-
cels 2-4 mm, obovate, pointed at style-scar which
projects below the base, 8—9 by 7 mm. Endocarp
perforate with 2 dorsal median rows of c. 12 erect
teeth with an equal number of short laterally directed
projections on both sides, the whole dorsal surface
papillose-rugulose.

Distr. Solomon Is. (Bougainville) and Malesia:
New Guinea (East: Long I.; Admiralty Is.: Manus).

Ecol. Open disturbed area in forest and regrowth
in deep gorge, up to 600 m. Fl. May; fr. Oct.

Note. Kom & SAND.-OLSEN 1555 from Mussau I.
has similar leaves and infructescences but the endo-
carps are different, suggesting a distinct species.

7. Stephania moluccana FoRMAN, Kew Bull. 11
(1956) 68.

Woody climber c. 6—8 m high, entirely glabrous.
Leaves with petioles 3—5 cm; lamina ovate or trian-
gular-ovate, 6—13 by 4.5—8 cm, apex shortly obtuse
or gradually narrowed to an obtuse tip, reticulation
lax on both surfaces, papyraceous. — Male inflores-
cences on leafless woody stems, composed of umbel-
liform cymes 2—5 cm long with primary rays less
than 1 cm and peduncle 1—4 cm arising from a slen-
der axis 10—20 cm long. — Male flowers on filiform
pedicels c. 2 mm; sepals 6, broadly oblanceolate or
spathulate, 1.5—2 mm long; petals 3, obdeltoid, 0.75
mm long, fleshy; synandrium 1 mm long. — Female
flowers unknown. Infructescences slender, c. 7—13
cm long. Drupe usually pedicellate up to 2 mm or ses-
sile, orbicular-obovate in outline, 5 mm long. Endo-
carp imperforate with 4 dorsal series of 10—13 papil-
liform processes.

Distr. Malesia: Moluccas (Morotai, Ternate).

Ecol. In forests, in Morotai at 30 m, in Ternate
at 1000 m.

Note. JAcoss 8682 (in fruit) from SE. New Gui-
nea may belong to this species.

8. Stephania montana Diets, Bot. Jahrb. 52 (1915)
189; FoRMAN, Kew Bull. 11 (1956) 69. — ? S. forma-
nii KUNDU & GuHA, Bot. Notis. 129 (1976) 259.
Slender climber. Stem glabrous. Leaves with gla-
brous petioles 3—6 cm; lamina triangular, narrowly
triangular or subtriangular, (6—)9—14 by 5—9 cm;
apex very acute, finely apiculate; base truncate to
rounded, sometimes apiculate at the basal corners;
reticulation lax, both surfaces glabrous, lower sur-
face sometimes strigulose along the nerves; papyra-
ceous. — Male inflorescences arising from leafless
parts of the stem, very lax and slender, glabrous or
papillose, composed of umbelliform cymes 4—14 cm
long with primary rays mostly more than 1 cm, ped-
uncles 3.5—8 cm, borne along a slender axis c. 40—50
cm. — Male flowers white or cream on pedicels up to
c. 3 mm; sepals 6, obovate, 1.5 mm long, glabrous;

1986 |

petals ?3, + broadly obovate, 0.75 mm long, gla-
brous. — Female flowers unknown. Infructescence
with peduncles of cymes 1—2 cm long. Drupes on
pedicels S—10 mm, rotund-obovate in outline, 5—6
by 4—5 mm. Endocarp ? perforate, dorsally bearing
4 longitudinal rows of 12—16 + papilliform pro-
cesses or short ridges.

Distr. Malesia: ?Philippines (Luzon), East New
Guinea.

Ecol. Mossy forests at 1300—2070 m.

Notes. The material I have seen of this species is
incomplete. The type specimen has only one leaf and
a few flower buds. Fruits are known only from CarR
14221 (BM), and these are possibly immature. Never-
theless, the species appears to be distinct on account
of its triangular leaves and long, slenderly branched
and lax inflorescences. Only on the leaf of the type
specimen are the basal corners of the lamina apicu-
late.

BS 23820 (in fruit) from Luzon may belong to this
species, although the processes on the endocarp are
transversely hooked.

Stephania formanii is represented by one collec-
tion with young male inflorescences. It probably be-
longs to this species.

9. Stephania psilophylla (PRESL) FORMAN, Kew Bull.
11 (1956) 63. — Cissampelos psilophylla Pres,
Reliq. Haenk. 2 (1835) 80; F.-VitL. Nov. App. (1880)
8. — S. corymbosa (non BL.) Turcz. Bull. Soc. Nat.
Hist. Mosc. 27, ii (1855) 281. — S. hernandiifolia
[non (WiLLD.) WaLpP.] VipaL, Phan. Cuming. Phi-
lip. (1885) 93, p.p., excl. CuminG 1533; Rev. PI.
Vasc. Filip. (1886) 45. — S. japonica [non (THUNB.)
Miers] Merr. En. Philip. 2 (1923) 149, p.p. — S.
catosepala (non Diets) Merr. En. Philip. 2 (1923)
149, p.p.

Slender climber. Stem herbaceous, up to 2 mm g@.
Leaves: petioles 2—9 cm, glabrous or sometimes
sparsely puberulous towards the top; lamina ovate to
broadly ovate, 5—9 by 4—8 cm, usually acuminate to-
wards the apex, base broadly rounded to very slightly
cordate; very fine and regular, reticulation raised on
both surfaces, glabrous, but lower surface sometimes
puberulous around insertion of petiole; papyra-
ceous. — Male inflorescence axillary (often slightly
supra-axillary) consisting of very slender, compound
umbelliform cymes 2.5—6 cm long, solitary or sever-
al along an axillary shoot, which is sometimes leafy;
usually glabrous, sometimes puberulous; frequently
with minute scales at the base. — Male flowers white,
entirely glabrous with pedicels |—2 mm, arranged in
clusters; sepals (6 or) 8, elliptic to (broadly) oblan-
ceolate, 1—1.5 mm, apex obtuse; petals (3 or) 4,
broadly obovate to obdeltoid, c. 0.5 mm long; synan-
drium about 0.5 mm long. — Female inflorescence
similar to male but cymes usually solitary. — Female

MENISPERMACEAE (Forman)

251

flowers with very short pedicels, c. 0.5 mm, or ses-
sile, entirely glabrous; sepals 3 or 4, + elliptic about
0.75 by 0.25 mm; petals as in male; carpel ellipsoidal,
1 mm long. Drupes red with filiform pedicels 2—4
mm, obovate to broadly obovate in outline, 4—5 by
3.5—4 mm. Endocarp perforate, dorsally bearing 2
longitudinal rows of 7—9 transverse ridges which are
sometimes transversely 2-lobed or divided, thus
forming 4 longitudinal rows of papilliform pro-
cesses: surface rugulose between ridges or processes.

Distr. Malesia: Philippines (Luzon).

Ecol. Primary forests and the edges of forests at
100—1300 m.

Note. This species has usually been confused
with S. japonica (THUNB.) MIERS var. japonica,
which also occurs in Luzon; but S. psilophylla can
easily be distinguished from it by the pedicellate male
flowers and fruits, and the very finely and regularly
reticulate leaves.

10. Stephania zippeliana Mig. Ann. Mus. Bot.
Lugd.-Bat. 4 (1868) 86, p.p., excl. folia; SCHEFF.
Ann. Jard. Bot. Btzg 1 (1876) 5; BEcc. Malesia 1
(1877) 153; Drets, Pfl. R. Heft 46 (1910) 266; Nova
Guinea 8 (1912) 869; Riot. Trans. Linn. Soc. II, 9
(1916) 16; Diets, Nova Guinea 14 (1923) 80; KANEH.
& Hatus. Bot. Mag. Tokyo 56 (1942) 474; FoRMAN,
Kew Bull. 11 (1956) 66; ibid. 22 (1968) 355. — S. flo-
rulenta Becc. Malesia 1 (1877) 153.

Large woody climber up to c. 20 m. Stem glabrous
with supra-axillary tufts of triangular to subulate
scales 1—3 mm long; main stems up to c. 1.5 cm @,
covered with flaky bark. Leaves with glabrous peti-
oles (6—)10—19 cm; lamina broadly ovate to suborbi-
cular (1 1—)16—22(—27) by (7—)10—17(—22) cm, apex
very acutely acuminate or obtuse, base rounded,
margin sometimes slightly repand; upper surface
smooth and slightly shining, usually with a raised,
very minute reticulation, sometimes + minutely
areolate; lower surface usually with minute, raised
reticulation (often with minute epidermal ridges run-
ning along and between the nervules); papyraceous
to coriaceous. — Male inflorescences very large (? up
to c. 100 cm) arising from old, leafless stems, occa-
sionally axillary, composed of compound umbelli-
form cymes 10—?30 cm long, with stout peduncles
arranged racemosely among and subumbellately at
the end of a long axis, minutely puberulous to gla-
brous. — Male flowers white with pedicels 1—4 mm;
sepals 6, + oblong-elliptic, 1.25—2.5 mm long, gla-
brous; petals 3, usually + broadly obovate, (0.25—)
0.75—1.5 mm long, glabrous; synandrium 0.5 mm
long. — Female inflorescence similar to male but
smaller. — Female flowers: sepals 3, petals 3, both
similar to male; carpel c. | mm. Drupe on pedicel
2-6 mm, broadly obovate to suborbicular in outline,
7-10 by 6-8 mm. Endocarp perforate or im-

D52

FLORA MALESIANA

[ser. I, vol. 102

perforate, surface either rough all over, and dorsally
bearing 4 longitudinal rows of 9—14 short papilli-
form processes (sometimes joined transversely) or
surface smooth, and dorsally bearing + longitudinal
rows of 9—10 T-shaped ridges.

Distr. Solomon Is. (San Cristobal, Guadalca-
nal), New Hebrides (Vanuata); in Malesia: New Gui-
nea (incl. Misool) and Moluccas (Ceram, Tenimber
Is.: Jamdena).

Ecol. In Ceram along the sea-shore, in New Gui-
nea in forests up to 1500 m.

Note. The appearance of the lower leaf surface
may vary from indistinct, fine reticulation to coarse
reticulate venation but this seems to be due to age
(METCALFE, Kew Bull. 11, 1956, 71).

11. Stephania capitata (BL.) SPRENG. Syst. Veg. 4
(1827) 316; Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1868)
86; SCHEFF. Nat. Tijd. N. I. 32 (1873) 401, t. 13;
Becc. Malesia | (1877) 156; Borer. Cat. Hort. Bog.
1 (1899) 42; Diets, Pfl. R. Heft 46 (1910) 281; Back.
Schoolfl. Java (1911) 46; Koorp. Exk. Fl. Java 2
(1912) 336; Merr. En. Born. (1921) 250; Riwi. FI.
Mal. Pen. | (1922) 113; Norman, J. Bot. 62 (1924)
Suppl. 5; Burk. & HEND. Gard. Bull. S. S. 3 (1925)
344; Heyne, Nutt. Pl. 1 (1927) 617; HEND. Gard.
Bull. S. S. 4 (1928) 220; Masam. En. Phan. Born.
(1942) 276; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 141; ForMAN, Kew Bull. 11 (1956) 46; STEEN.
Mt. Fl. Java (1972) pl. 33-1. — Clypea capitata BL.
Bijdr. (1825) 28. — Clypea acuminatissima BL. l.c.;
Miers, Contr. Bot. 3 (1871) 206. — S. acuminatis-
sima (BL.) SPRENG. Syst. 4 (1827) Cur. Post. 316
(‘acuminata’); WAP. Rep. 1 (1842) 96. — S. obvia
Miers [Ann. Mag. Nat. Hist. ser. 3, 18 (1866) 15,
nomen|, Contr. Bot. 3 (1871) 226. — S. longifolia
Becc. Malesia 1 (1877) 156. — S. truncata Y AMA-
MOTO, J. Soc. Trop. Agric. 16 (1944) 142. — Fig. 20.

Slender woody climber up to c. 15 m, entirely gla-
brous. Roots tuberous, fusiform. Leaves; petioles
(2—)3—5(—10) cm long; lamina lanceolate to broadly
ovate, 6—17 by 2.5—10.5(—14) cm, apex usually at-
tenuately acute or caudate, base broadly rounded to
slightly cordate; reticulation usually rather lax and
not prominent, + equally visible on both surfaces;
papyraceous. — Male inflorescences usually axillary
(sometimes borne on leafless parts of the stem) com-
posed of several, peduncled, disciform capitula aris-
ing in the axils of persistent, triangular to linear
bracts 1—2 mm along an axis usually less than 1 cm
long, occasionally to 14 cm on old stems; capitula
3—6(—8) mm in @ on filiform peduncles up to 20
(—35) mm long. — Male flowers sessile, yellow to
green, very densely crowded on a disciform recep-
tacle; sepals 2—3, + oblong-elliptic, 0.5—0.75 mm
long; petals 2—3, + obovate, 0.25 mm long; synan-
drium 1 mm long, exserted. — Female inflorescence

Fig. 20. Stephania capitata (BL.) SPRENG. Male inflo-

rescences, side view, <1 1/3; top view of capitulum

above, x4. Trang, Thailand (Photogr. H. BAN-
ZIGER).

as in male, but axis and peduncles thicker, especially
at fruiting, also peduncles longer, up to 55 mm. —
Female flowers with very short pedicels, about 1 mm,

1986]

sometimes partly united; sepals and petals as in male;
carpel + ovoid or ellipsoidal, c. 1 mm long. Drupe
red, with slender pedicels S—10 mm, obovate (to sub-
orbicular) in outline, sometimes attenuated towards
the base (6—)8—11 by (S—)6—8 mm. Endocarp per-
forate, dorsally bearing 4 longitudinal rows of 10—14
capitate projections divaricately lobed at their apices
(rarely the projections scarcely developed).

Distr. Thailand (Peninsular); in Malesia: Suma-
tra, Malaya, Java, Lesser Sunda Is. (Bali), N. & W.
Borneo.

Ecol. Scattered on plains and mountains and in
rain-forest from sea-level to 2000 m.

Uses. A substitute for Cyclea barbata (WALL.)
Miers in the preparation of ‘tjintjau’, a native pre-
paration used for abdominal diseases and fevers (?
leaves used).

Vern. Java: akar talur, areuj geureung, areuj
tiamtjau minjak, daun tjamtjau, gorong, sumpat
kendi, tjamtjau, tjintaun.

Notes. The axis of the inflorescence is usually
less than 1 cm long, but on LOrziNG 5509 from Su-
matra the axes are up to 14 cm and arise from an old
leafless stem.

12. Stephania dictyoneura Drets, Pfl. R. Heft 46
(1910) 281; Yamamoto, J. Soc. Trop. Agric. 16
(1944) 141; ForMAN, Kew Bull. 11 (1956) 48.
Slender woody climber, entirely glabrous. Leaves:
petioles 2.5—4 cm, inserted 3—7 mm from the basal
margin: lamina broadly triangular-ovate, 5—6.5 cm
long and broad, apex shortly acuminate, acumen ob-
tusely mucronulate, base truncate or slightly cordate;
reticulation close and raised, especially beneath;
stiffly papyraceous. — Male inflorescences and

MENISPERMACEAE (Forman)

253

flowers unknown. — Female inflorescence borne on

older, leafless stem, composed of several, peduncled,
disciform capitula, arising in the axils of subulate
bracts 1—2 mm long, along an axis about 5 cm long;
capitula c. 7 mm @ on peduncles c. 5 cm. — Female

flowers very densely crowded on a disciform recep-

tacle; sepals and petals + obovate, c. 0.5—0.75 mm

long; carpel ellipsoidal c. 0.75 mm. Fruits unknown.
Distr. Malesia: Central W. Sumatra (Mt Singa-

lang), only known from the type (BECCARI PS 8).

Ecol. Montane forest, 1600 m. Fi. June—July
(female).

Notes. The specimens at Kew and Leiden are ster-
ile but on one sheet of BEccaAR! PS 8 at Florence (FI)
there is an incomplete female inflorescence which
bears a single capitulum. This inflorescence arises
from a leafless, slightly woody stem 3—4 mm o.

This species is closely related to S. capitata (BL.)
SPRENG, which also has inflorescences composed of
dense capitula. Stephania dictyoneura DiE1s is easily
distinguished from S. capitata by its more broadly
shaped leaves, which show a prominent and close re-
ticulation, and by the insertion of the petiole which
is 3—7 mm from the base of the lamina.

Doubtful

Stephania neoguineensis KuNDU & GUHA, Bot.
Notis. 129 (1976) 257.

Only known from one collection from East New
Guinea, E. Highlands Distr. (Brass 32246). The
single large detached male inflorescence and the male
flowers agree with S. montana Diets, but not the
broadly triangular-ovate leaves.

Excluded

Aspidocarya kelidophylla Laut. & K. Scu. Fl. Deut. Schutzgeb. Siidsee (1901) 313, according to Diets,
Pfl. R. Heft 46 (1910) 320 = Cardiopteris moluccana Bu. (Cardiopteridaceae).

Cocculus flavicans WALL. Cat. n. 4976 (1831/32), nomen, from Penang I. is, according to BURKILL, Gard.
Bull. S. S. 4 (1929) 426 = Anisophyllaea gaudichaudiana BAL. (Rhizophoraceae).

Heckelia nymanii K. Scu. in K. Sch. & Laut., Nachtr. Fl. Deut. Schutzgeb. Stidsee (1905) 26 is, according
to Drets, Pfl. R. Heft 46 (1910) 320 = Rhipogonum album R.Br. (Liliaceae).

Juppia borneensis Mere. J. Str. Br. R. As. Soc. n. 85 (1922) 170 is, according to Harms, Notizbl. Berl.-
Dahl. 80 (1924) 717 = Zanonia indica L. (Cucurbitaceae).

Peripetasma polyanthum Ript. J. Bot. 58 (1920) 147; Fl. Mal. Pen. 1 (1922) 103 is, according to PRAIN
& Burkitt, Kew Bull. (1925) 66 = Dioscorea stenomeriflora PRain & BuRK. (Dioscoreaceae).

Tinospora curtisii Rr. J. Bot. 58 (1920) 148 is, according to FoRMAN, Kew Bull. 36 (1981) 420 = Zanonia

indica L. (Cucurbitaceae).

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|

MONIMIACEAE (W.R. Philipson, Christchurch)

Evergreen shrubs or trees, rarely lianes. Leaves decussate, or rarely in whorls
of three, exstipulate, simple, entire or dentate, with spherical oil cells in the la-
mina, bearing simple or stellate hairs or glabrous. /nflorescence terminal or axil-
lary (when in axils of reduced bracts appearing supra-axillary), sometimes cauli-
florous, cymose, paniculate, fasciculate or pleiochasial. Flowers unisexual or
bisexual, actinomorphic or very rarely (extra-Mal.) oblique, receptacle usually
well developed (perigynous), rarely reduced (hypogynous), + globose or urceo-
late to widely campanulate; tepals usually inconspicuous, sometimes larger and
petaloid, rarely distinct sepals and petals (extra-Mal.), decussate, radial or spi-
ral. — Male flowers with few to many stamens arranged in whorls or sometimes
spirally or disposed irregularly; filaments usually strap-shaped, short, occasion-
ally with 2 basal lobes; anthers 2—4 sporangiate, the loculi sometimes confluent
above (or rarely below) opening by slits or valves. — Female flowers with or
without staminodes; carpels few to many (rarely extra-Mal., only one), sessile
or stipitate, free or immersed in the receptacle, outer carpels of female flowers
sometimes sterile; ovule solitary, erect or pendulous, crassinucellar, bitegmic or
(extra-Mal.) unitegmic. Fruits of separate drupes or achenes, sometimes plu-
mose, frequently enclosed in the persistent receptacle or exposed by various
modes of splitting of the receptacle; endosperm copious, oily; embryo straight,
cotyledons appressed or divergent, sometimes with serrate margins.

Distribution. About 33 genera with an estimated 320 species, mainly in the warmer parts
of the southern hemisphere. There is a concentration of genera in Malesia (11 genera with 86 spp.)
with extensions south and east into Australia and the SW. Pacific; further concentrations occur
in the islands of the western Indian Ocean and in South America. The family is represented in
Africa only by two small aberrant genera and occurs on the Eurasian mainland only in the Malay
Peninsula, the Nicobar Islands and Peninsular Thailand.

The Malesian genera are either endemic or nearly so, with one or few species extending to the
Solomon Islands (Steganthera) or eastern Australia (Levieria, Palmeria, Steganthera, Kibara and
Dryadodaphne). Wilkiea has more species in Australia than in Malesia. Only Kibara extends
slightly westwards into the Nicobar Islands and Thailand. The concentration of genera in New
Guinea is striking: only Matthaea lies exclusively to the west of New Guinea, the family being re-
presented in western Malesia, otherwise by one species each of Steganthera, Levieria and Pal-
meria, and by four or five species of Kibara.

The Malesian genera fall within several subfamilies and each of these has a distinctive geograph-
ical relationship. Levieria is a member of the tribe Hedycaryeae with relationships in SW. Poly-
nesia; Dryadodaphne falls within the subfamily Atherospermatoideae, a subfamily that is pre-
dominantly Australian; while Pa/meria is most closely related to Monimia of the Mascarene Is-
lands. The remaining genera (Steganthera, Matthaea, Kairoa, Faika, Parakibara, Wilkiea and Ki-
bara form a closely knit group within the tribe Mol/linedieae which is characteristic of the Malesian
region.

Trimenia (Piptocalyx) is referred to the separate family Trimeniaceae.

The ratio of species to genera, in Malesia, is c. 8: 1, but if the largest genus, Kibara, is omitted
this reduces to c. 4.3 : 1. Five genera are represented by a single species. Fig. 1 & 2.

Fossils. MULLER (1981) pointed out that /naperturopollenites crispolensis (DoyLe et al. , 1977)
from the Lower Cretaceous is similar to that of Hedycarya pollen and that pollen of Stellatopollis

(255)

256 FLORA MALESIANA [ser. I, vol. 102

i ]

2 j s
1 1 :
1 7
2 RE 3
4
2 5 z =a 2 6 5 =.= |
3 | ee 76
| Fig.1 Seyi are are | | Fig.2 P Cried eee eds |) Se an
Fig. 1. Number of genera of Monimiaceae in Malesian islands; figures below the hyphen indicate endemic
genera. — Fig. 2. Number of species of Monimiaceae in the Malesian islands.

barghoornii (DoyLe et al., 1975) from the Middle Albian is comparable with that of Daphnandra
and Nemuaron (from New Caledonia).

SCHODDE (1969) has little doubt that the two fossilized woods A therospermoxylon (KRAUSEL,
1939) and Protoatherospermoxylon (MADEL, 1962) are correctly assigned to this family. He reject-
ed as doubtful all leaf records from the northern hemisphere, but accepts as probable fossil leaves
from Eocene-Oligocene and Early Miocene deposits in Argentina (Laurelia armarillana, BERRY,
1928, and Laurelia guinazui, BERRY, 1935) and from the Eocene-Oligocene of Seymour Island
(Laurelia insularis, DUsEN, 1908). Pollen grains of Laurelia are recorded in New Zealand from
Middle Oligocene to the present (COUPER, 1960).

References: BERRY, Proc. U.S. Nat. Mus. 73 (1928) 21; Bot. Gaz. 96 (1935) 752; CoupER, Pa-
laeont. Bull. 32, Geol. Surv. New Zeal. (1960) 1—87; Doyte ef a/., Pollen et Spores 17 (1975)
429—486; Bull. Centre Rech. Explor. Prod. Elf. Aquitaine 1 (1977) 451—473; Dusen, Wiss. Er-
geb. Schwed. Sudpolar-exped. 3 (1908); KRAusEL, Abh. Bayer. Akad. Wiss. math.-naturw. Abt.
NF 47 (1939) f. 3; MApDEL in Muller-Stoll & Madel, Trans. Geol. Soc. S. Afr. 65 (1962) 99; Mut-
LER, Bot. Rev. 47 (1981) 9; ScHoppE, A monograph of the Atherospermataceae R. Br., Thesis,
Univ. Adelaide (1969).

Ecology. Malesian Monimiaceae form an inconspicuous but remarkably consistent element
of the understorey of rain-forests in New Guinea and to a much lesser degree of the rest of Ma-
lesia. They are most frequently straggling shrubs or treelets, occasionally reaching the stature of
moderate trees, and more rarely are low to moderately high-climbing (c. 22 m) woody lianes. They
occur in forests from sea-level to the limit of trees, their greatest frequency being in lower montane
forest. They occur, though rarely, in coastal scrub on sandy soil or coral (e.g. Kibara rigidifolia)
and in the zone of scrub above tree-line (e.g. Kibara oligocarpella).

Flower biology. The majority of the Malesian genera are either monoecious or dioecious, the
only genus with bisexual flowers in Malesia being Dryadodaphne. Little is known of the floral
biology of this family. Sampson (1969a) found that Hedycarya arborea is wind-pollinated. The
greenish flowers lack nectar and no insects were observed to visit them. In Laurelia novae-
zelandiae the paired lobes on the filaments secrete nectar which accumulates on the floor of the
flower (SAMPSON, 1969b). Large numbers of bees and blowflies were observed visiting the flowers.
The nectar consists mainly of sucrose with possibly traces of raffinose. GOTTSBERGER (1977) re-
ported pollination of the neotropical genus Mollinedia by Thysanoptera. The female insects bore
holes in male and female flowerbuds and deposit their eggs in the receptacles. The larvae develop
in the buds, becoming adult as the flowers open. They emerge and transfer pollen to other flowers.
Similar infestations of flowerbuds have been observed in the Malesian genera Levieria and Kairoa
in which the male flowers open, as in Mollinedia, though no observations on pollination have been
made in these genera. ENDRESS (1980) recorded similar insect infestation in species of Wilkiea and
Steganthera, in which the male flowers have a restricted opening. The flowers of Steganthera are
frequently so deformed by gall insects that their function appears to be impaired (personal obser-
vation on S. ilicifolia and other species).

1986 | MOoONIMIACEAE (Philipson) 254

The reception and germination of pollen on a non-carpellary hyperstigma, long suspected in
Hennecartia (PERKINS & GiLG, 1901) has been confirmed and extended to three other genera,
Tambourissa, Wilkiea and Kibara (ENDREss, 1979, 1980). A zone in the narrow entrance to the
female receptacle secretes a mucilage which acts as a transmitting medium for the pollen tubes
and is continuous from the outer surface of the receptacle to the carpels. As the mucilage connects
all the stigmas in a receptacle it acts as an extragynoecial compitum.

The intensive field observations in the Malagasy region by LORENCE have considerably in-
creased information on the floral biology of the family (LORENCE, 1980; ENDREss & LORENCE,
1983). Seven species of Tambourissa were studied. Odour and possibly colour attract insects
which were rewarded in male flowers with pollen and in female flowers with a sugary secretion
or in some species by deceit. Some species are mainly dependent on flies and have short styles,
whereas others with longer styles are visited mainly by beetles. Different flowering times contri-
bute to reproductive isolation and hybridization was found to be extremely rare. LORENCE (1980,
1985) considered that Monimia ovalifolia, with a sweet odour, yellow to pink or orange recep-
tacles, shallow open white male flowers and fully exposed stamens and styles, appears well adapt-
ed to the small syrphid flies that visit it.

Dispersal. The fruit-clusters of most Malesian genera consist of few to many drupes, usually
black and shining when ripe, sessile or stipitate on receptacles, which are usually enlarged, fleshy
and bright yellow and orange. In Palmeria the drupes are exposed when mature by the splitting
of the receptacle. They are black or dark reddish brown and are borne on the inner side of the
receptacle which is bright red or pink. All these structures are clearly suitable for dispersal by birds
or animals, but no records of field observations of this in Malesia are known to me. LORENCE
(1980, 1985) records dispersal by birds in Malagasy species of Monimia and Tambourissa which
bear fruits of somewhat similar appearance to those of the Malesian genera. In Dryadodaphne
fruit structure and means of dispersal are very different, though again this has only been inferred
from their structure. The developing achenes are enclosed by the enlarged and indurated re-
ceptacle and this splits at maturity to release the ripe achenes. These are dry and spindle-shaped
with a persistent aculeate style which becomes markedly plumose. The structure appears well
adapted to wind dispersal.

Myrmecophily. The association of ants with the swollen nodes of Steganthera hospitans was
noted by Beccari (1877). BeEccaRi concluded that the entry holes were excavated in the distended
nodes by the ants (Hypoclinea scrutator). He also observed the presence of small homopterids
(Myzolecanium) inside the hollow nodes. Only adult female scale insects were seen, and since
these were too large to pass the entry holes, BEccaARI speculated on the possible life cycles of the
coccid and on their relationship to the ecology of the ants. Similar associations occur on other
species of Steganthera (S. royenii, S. moszkowskii and S. ledermannii) and also on species of Ki-
bara (K. ferox, K. latifolia, K. archboldiana and K. carrii). Only K. latifolia of the Moluccas oc-
curs outside New Guinea.

Beccari referred to analogous ant associations in the stems of Verbenaceae: Clerodendron;
Euphorbiaceae: Pimelodendron; Myristicaceae: Myristica and the rhizomes of the fern Polypo-
dium sinuosum WALL. and of species of the Rubiaceous epiphytes Myrmecodia and Hydnophy-
tum as well as some Melastomaceous epiphytes. In addition VAN STEENIS reported spindle-shaped
twigs inhabited by ants in several other genera, e.g. in Meliaceae: Aphanamixis, Chisocheton;
Elaeocarpaceae: Elaeocarpus; Euphorbiaceae: Drypetes, Glochidion, Homalanthus; Legumi-
nosae: Archidendron; Rubiaceae: Myrmeconauclea, Nauclea; Sapindaceae: Harpullia, and Sym-
plocaceae: Symplocos, and ForMAN added to these (in litt.) Myristicaceae: Myristica subalulata
and Proteaceae: Helicia macrostachya. They occur mostly in a single species of these genera, but
sometimes in several, e.g. in Nauclea.

References: Beccari, Malesia | (1877) 186-193; ENprRess, Experientia 35 (1979) 45; Pl. Syst.
Evol. 134 (1980) 79—120; ENpress & Lorence, PI. Syst. Evol. 143 (1983) 53—81; GorrsBERGER,
PI. Syst. Evol. Suppl. 1 (1977) 211-226; Lorence, A systematic and eco-evolutionary study of

258 FLORA MALESIANA [ser. I, vol. 102

the Monimiaceae in the Malagasy region, Thesis, Washington Univ., St. Louis (1980); Ann. Mo.
Bot. Gard. 72 (1985) 1—165; PERKINS & Gita, Pfl. R. Heft 4 (1901) 1—122; SAmMpson, (a) Austr.
J. Bot. 7 (1969) 403—424; (b) New Zeal. J. Bot. 7 (1969) 214—240.

Morphology. Little is known in detail of the shoot morphology of the trees, shrubs and
lianes of this family. In Laurelia the bole bears buttresses and the roots develop knee-pneumato-
phores. The bark is generally + smooth with only small fissures and flakes, an exception being
Kairoa with prominent corky ridges on the main stems. Throughout the family phyllotaxis is de-
cussate or rather rarely with the leaves in whorls of three (e.g. Kibara rigidifolia). Marked ani-
sophylly occurs in Glossocalyx. The stems are terete or + quadrangular, except near the nodes
which are somewhat flattened and dilated. The buds, both vegetative and reproductive, are usual-
ly enclosed in small scales and there may be more than one in an axil arranged either horizontally
or vertically (LORENCE, 1980, 1985), the central bud usually developing first. In spite of the variety
of leaf size, form, margin and indumentum, there is a family character which facilitates recogni-
tion in the field. This is due, principally, to the venation, which almost invariably is festooned
brachidodromous with the secondary veins arising from the midrib at regular intervals and at uni-
form angles. An exception noted by LORENCE (1980, 1985) is Ephippiandra. Leaves are usually
petiolate but may be amplexicaul (e.g. Kibara ferox). Leaf shape may vary with age. The juvenile
leaves of Steganthera hospitans are much narrower than those of adult trees and juvenile speci-
mens of Kibara ferox bear narrower and more dentate leaves than do adults. In Kibara ferox,
however, the narrow ultimate branches of adult plants bear much narrower leaves than those of
the basal parts of the shrub. Heterophylly also occurs in many Tambourissa species and in Hor-
tonia.

Inflorescences usually occur in the axils of foliage leaves. When several are grouped among the
terminal leaves of a shoot, apparently terminal leafy inflorescences are formed, and with the re-
duction of the leaves these may result in large panicles. It is doubtful, however, if the true terminal
bud is involved in these inflorescences, the terminal (vegetative) bud aborting. Inflorescences
often occur on basal parts of stems where the nodes bear much reduced scales. The upper parts
of these stems continue as indeterminate foliage shoots. The presence of multiple buds at nodes
which develop in succession allows inflorescences to persist on branches which have lost their fo-
liage and to become strikingly cauliflorous. The branching of the inflorescences is cymose, the
most simple element being a dichasium. However, several flowers or branches frequently arise
from a single node, or alternatively, several pairs of flowers occur along a simple axis (pleiocha-
sium — often referred to as racemose). As a result of combinations of these factors individual
inflorescences range from solitary flowers, through fascicles of flowers or branches to rather sim-
ple cymes and more diffuse and complex paniculate cymes.

References: LORENCE, A systematic and eco-evolutionary study of the Monimiaceae in the Ma-
lagasy region, Thesis, Washington Univ., St. Louis (1980); Ann. Mo. Bot. Gard. 72 (1985) 1-165.

Anatomy. General accounts of the vegetative anatomy are given by Money, BAILEY & SWAMY
(1950) and MeTcaLre & CHALK (1950). Of particular interest are 1) the universal presence of oil
cells (HoBEIN, 1889; PERKINS, 1898), 2) the unilacunar node, with simple strands or arcs of strands
entering the petiole, 3) the presence of hippocrepiform sclereids in the pericycle (but not in Siparu-
noideae) (MonEy, BAILEY & Swamy, 1950). Hairs may be simple unicellular, often in fascicles
from a common base, or stellate grading into peltate scales. Two-armed hairs have been reported
for some genera, including Matthaea.

Phloem plastids. Most members of the Monimiaceae have been found to contain P-type
plastids. Very consistently, the Atherospermatoideae have large protein crystals, protein fila-
ments and starch (BEHNKE, 1981). Except for the genera Monimia, Palmeria and Tambourissa,
the other subfamilies have been found to contain + small protein crystals besides the dominant
starch grains.

References: BEHNKE, Nordic J. Bot. 1 (1981) 381—400; Hosein, Bot. Jahrb. 10 (1889) 51—74;
METCALFE & CHALK, Anatomy of the Dicotyledons, Clarendon Press, Oxford (1950); Money,

1986 } MONIMIACEAE (Philipson) 259

Bamtey & Swamy, J. Arn. Arb. 31 (1950) 372—404; Perkins, Bot. Jahrb. 25 (1898) 547—577.

Wood anatomy. The characteristics of the secondary xylem have been described by the authors
listed below. A particularly full assessment is given by FOREMAN (1983). Certain features have
been found to characterize the subfamilies, for example the multiseriate rays of Atherospermatoi-
deae and Siparunoideae are narrower than those of the Monimioideae, and simple perforation
plates occur in Monimia and Palmeria. Peumus has the most distinctive xylem with spiral thicken-
ing in the vessels and simple perforation plates. The xylem of Hortonia has more primitive featu-
res than other genera.

References: BUTTERFIELD & MEYLAN, Austr. J. Bot. 20 (1972) 253—259; The structure of New
Zealand woods, Wellington (1973); DADSWELL & RECORD, Trop. Woods 48 (1936) 1—30; ForE-
MAN, The morphology and phylogeny of the Monimiaceae (sensu /ato) in Australia, Thesis, Univ.
of New England, Armidale (1983); GARRATT, Trop. Woods 39 (1934) 18—44; HoBEIN, Bot. Jahrb.
10 (1889) 51—74; KuCERA & PuiLipson, New Zeal. J. Bot. 15 (1977) 649—654; LEMESLE & PI-
CHARD, Rev. Gen. Bot. 61 (1954) 69—95; LorENCcE, A systematic and eco-evolutionary study of
the Monimiaceae in the Malagasy region, Thesis, Washington Univ., St. Louis (1980); Ann. Mo.
Bot. Gard. 72 (1985) 1—165; Matos ARAuJo & FiLHo, Brazil Florestal 4 (1973) 35—39, 41—45;
ibid. 5 (1974) 57—60; Rodriguesia 27 (1974) 153—162; Arq. Jard. Bot. Rio de Janeiro 20 (1977)
15—20; METCALFE & CHALK, Anatomy of the Dicotyledons, Clarendon Press, Oxford (1950); DEN
OUTER & VAN VEENENDAAL, Acta Bot. Neerl. 31 (1982) 265—274; PatEeL, New Zeal. J. Bot. 11
(1973) 587—598; ScHoppDE, A monograph of the family A therospermataceae R. Br., Thesis, Univ.
of Adelaide (1969); Taxon 19 (1970) 324—328; SOLEREDER, Systematic anatomy of the Dicotyle-
dons (transl. Boodle & Fritsch), Clarendon Press, Oxford (1908); Wetcu, J. Proc. R. Soc.
N.S.W. 62 (1929) 350—365.

Floral anatomy. Development and vascularization are discussed by MonEy, BAILEY & SWAMY
(1950), Sampson (1969a, b, c) and ENpREss (1972, 1980). The little then known of embryology
in the family was reviewed by Davis (1966) and BHANDARI (1971). Since then SAMPSON (1969a,
c)and Enpress (1972, 1980) have added much information. The ovule is bitegmic and crassinucel-
late (unitegmic in Siparuna), pendulous with an upwardly directed micropyle or lateral to basal
with a downwardly directed micropyle. The structure of the fruit and seed is described by ENDRESs
(1972), CoRNER (1976) and LoRENCE (1980, 1985). The fruits are either an achene, often plumose,
or they have a fleshy or leathery pericarp. The latter fruits are usually referred to as drupes be-
cause the endocarp consists of sclerotic cells. In some genera (e.g. Palmeria) the endocarp forms
a substantial stony layer, in Levieria it is thinner but still strongly sclerotic, while in others, not-
ably Kibara, the endocarp consists of a single papery layer. As these differences are of degree and
intermediate states occur, the same term (drupe) is used throughout, even though the fruit of spe-
cies with a thin endocarp might technically be better referred to as berries. In Kibara the mesocarp
of the species examined consists of uniform soft-walled tissue, whereas in Steganthera all species
examined had numerous nests of sclereids adjacent to the endocarp (possibly to be considered as
part of the endocarp) with many oil cells in the outer mesocarp. The cotyledons are divergent in
the Monimieae but in the Mollinedieae and Atherospermatoideae they are closely appressed. In
Hedycaryeae both conditions occur: Hedycarya appressed, Levieria divergent. The endosperm is
fleshy in most genera, but in Kairoa it is horny and in Levieria the central part of the endosperm
is milky. Rarely (extra-Mal.) a ‘stylar aril’ overlies the upper part of the exocarp.

The form of stamens is variable in respect of the presence or absence of a pair of glands on
the filaments; the number (2 or 4) of sporangia and the dehiscence by slits or valves. Stamen and
anther development is described by ENprRess (1980), FoREMAN (1983) and SAMPSON (1969b) and
the morphology of the glands discussed by Money, BAatLey & Swamy (1950), SAMPSON (1969c)
and Enpress (1980). The different types of cytokinesis in Monimioideae and Atherospermatoi-
deae, first described by SAMPSON (1969c), is extended by FoREMAN (1983) who discusses many as-
pects of androecial development.

- References; BUANDARI, J. Arn. Arb. 52 (1971) 1—39, 285—304; Corner, The seeds of Dicotyle-

260 FLORA MALESIANA [ser. I, vol. 102

dons 1 (1976) 194—197; Davis, Systematic embryology of the Angiosperms, New York (1966)
178—179; ENprREss, Bot. Jahrb. 92 (1972) 331—428; Pl. Syst. Evol. 133 (1980) 79—120, 199-221;
FOREMAN, The morphology and phylogeny of the Monimiaceae (sensu lato) in Australia, Thesis,
Univ. of New England, Armidale (1983); LoRENCE, A systematic and eco-evolutionary study of
the Monimiaceae in the Malagasy region, Thesis, Washington Univ., St. Louis (1980); Ann. Mo.
Bot. Gard. 72 (1985) 1—165; Money, BAILEY & Swamy, J. Arn. Arb. 31 (1950) 372—403; SAmp-
SON, (a) Austr. J. Bot. 17 (1969) 403—424; (b) /.c. 425—439; (c) New Zeal. J. Bot. 7 (1969)
214—240.

Palynology. Reviews of the morphology of pollen grains in the family are given by Money,
BamLtEy & Swamy (1950), ERDTMAN (1966), JEREMIE ef al. (1984), WALKER & DOYLE (1975), and
WALKER (1976a, b). Studies on the family include: on Hedycarya SAMPSON (1969a, 1977, 1982),
on Laurelia SAMPSON (1969b, 1975), on Peuwmus BARTH (1962), HEUSSER (1971), on Monimia and
other Malagasy genera LORENCE (1980) and LORENCE, ZENGER & VINAY (1984). Inaperturate
grains are typical of the family but the Atherospermatoideae have disulcate grains. Spinose grains
occur in Peumus, Monimia and Palmeria. The grains remain in tetrads in some species of Hedy-
carya and Kibaropsis.

References: BARTH, Mem. Inst. Oswaldo Cruz, Rio de Janeiro, 60 (1962) 405—420; ERDTMAN,
An introduction to palynology, 1, Pollen morphology and plant taxonomy, New York (1966);
HeusseR, Pollen and spores of Chile, Univ. Arizona Press, Tucsan (1971); JEREMIE ef a/., Pollen
et Spores 26 (1984) 161—180; LorENcE, A systematic and eco-evolutionary study of the Monimia-
ceae in the Malagasy region, Thesis, Washington Univ., St. Louis (1980); LORENCE, ZENGER &
Vinay, Grana 23 (1984) 11—22; Money, BaAILEy & Swamy, J. Arn. Arb. 31 (1950) 374—404; SAmp-
son, (a) Austr. J. Bot. 17 (1969) 403—424; (b) I.c. 425—439; Grana 15 (1975) 153—157; ibid. 16
(1977) 61—73; ibid. 21 (1982) 9-14; WALKER (a) in Beck (ed.), Origin and early evolution of the
Angiosperms, Columbia Univ. Press, New York (1976) 241 —299; (b) in Ferguson & Muller (eds.),
The evolutionary significance of the exine, Acad. Press, London (1976) 251—308; WALKER &
DoyLe, Ann. Mo. Bot. Gard. 62 (1975) 664—723.

Chromosomes. The only count known to me based on Malesian specimens is that for Kibara
(BoRGMAN, 1964). Counts of Australian species of Steganthera and Palmeria, together with a few
non-Malesian genera, will be found in the references below. From these counts it is considered
that the basic number for the Monimioideae (n= 19) differs from that for the Atherospermato-
ideae (n=22).

References: BORGMAN, Z. Bot. 52 (1964) 118—172; EHRENDORFER in Beck (ed.), Origin and
early evolution of the Angiosperms, Columbia Univ. Press, New York (1976) 220—240; EHREN-
DORFER ef al., Taxon 17 (1968) 337—353; GADELLA et al/., Acta Bot. Neerl. 18 (1968) 74—83; GoLpD-
BLATT, J. Arn. Arb. 55 (1974) 453—457; Hair & BEUZENBERG, New Zeal. J. Sci. 2 (1959) 148—156;
LORENCE, A systematic and eco-evolutionary study of the Monimiaceae in the Malagasy region,
Thesis, Washington Univ., St. Louis (1980); Ann. Mo. Bot. Gard. 72 (1985) 1-165; MoRAWETzZ,
Pl. Syst. Evol. 138 (1981) 157-172. ‘

Phytochemistry. Chemical characters were summarized sixteen years ago (HEGNAUER,
1969). The benzyltetrahydroisoquinoline family of alkaloids (abbreviated: benzylisoquinolines)
and essential oils consisting mainly of phenylpropanoids and mono- and sesquiterpenoids were
considered to be characteristic secondary metabolites of the family, but the lack of chemical
knowledge for Hortonioideae, Monimioideae (except Peumus boldus) and Siparunoideae was
stressed. Many members of the family are aluminium accumulators; this character, however,
seems to be lacking in Atherospermatoideae. Phenolics were scarcely known, but predominance
of the flavonols kaempferol, quercetin and isorhamnetin in leaves, and absence of flavones, flavo-
nols with trihydroxylated B-ring and of galli- and ellagitannins had been reported. As a whole
phytochemistry of Monimiaceae agreed perfectly with their inclusion in a group loosely termed
woody polycarps. In the meantime more became known about the chemistry and distribution of
benzylisoquinolines (URzUA & CAssELs, 1978; HEGNAUER, in prep.) and polyphenolic com-

1986 | MonIMIACEAE (Philipson) 261

pounds, especially lignans (including neolignans) (HEGNAUER, in prep.). Siparuna gilgiana and S.
guyanensis synthesize liriodenine and related oxoaporphine alkaloids, and laurotetanine,
N-methyllaurotetanine and leurotitsine were detected in three Palmeria species of New Guinea.
The isolated position of Daphnandra with respect to alkaloid metabolism was stressed (URZUA
& CASSELS, 1978); only bisbenzyltetrahydroisoquinoline alkaloids, including a number of com-
pounds apparently restricted to the genus, have been isolated hitherto from six species. Daphnan-
dra aromatica was transferred by ScHopDE to Doryphota; Doryphora aromatica yielded the apor-
phine isocorydine besides bisbenzylisoquinolines. Dryadodaphne novoguineense which is con-
fined to New Guinea, synthesizes aporphines, oxoaporphines and the bisbenzylisoquinolines dry-
adine and dryadodaphnine. 4-Hydroxyaporphines, alkaloids with a very unusual substitution pat-
tern were encountered in Laureliopsis philippiana (= Laurelia philippiana). It deserves mention-
ing that lignans which were known from Trimeniaceae only (Piptocalyx, Trimenia, HEGNAUER,
1969) have been detected in leaves of Laurelia novae-zelandiae; they yielded pinoresinoldimethyl-
ether and yangambin. Lignans and many different types of neolignans are widespread in Polycar-
picae; they begin to form an outstanding chemical character of the order as a whole. Summar-
izing, old and new chemical evidence conforms with the classification of Monimiaceae with
woody polycarps without contributing much to the question whether the family is nearer to Mag-
noliaceae and Annonaceae or to Lauraceae, i.e., whether inclusion in Laurales is more natural
than inclusion in Magnoliales. Chemical evidence also agrees with the exclusion of Amborella-
ceae, Austrobaileyaceae and Trimeniaceae which all seem to lack benzylisoquinolines.

References: HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 99-107, 431—432, 457; ibid.
8 (in prep.); Urzua & CassELs, Lloydia 41 (1978) 98. — R. HEGNAUER.

Taxonomy. The family as first founded by DE Jussreu (1809) included genera representing
most of the subfamilies at present recognized. The heterogeneous nature of these genera imme-
diately instigated a series of proposals for the division of the family by the recognition of the A the-
rospermataceae or, more recently, into several smaller families. Concurrently other systematists
have retained the original broad view of the family. Early proponents of splitting were R. BROWN
(1814), BaRTLING (1830) and LinDLEy (1853), but the broader view long prevailed among other
systematists, principally ENDLICHER (1837), TULASNE (1855), BENTHAM & HOOKER (1880), Pax
(1891), Perkins & Gri (1901), PERKINS (1911, 1925), and MELCHIOR (1964). The more recent pro-
posals to remove elements as distinct families include Gress (1917), PICHON (1948) and especially
Money, BarLey & Swamy (1950) whose view that Amborella and Trimenia (Piptocalyx) should
form separate families has been accepted ever since. The removal of further elements has con-
tinued, but opinion remains divided on this trend. A most important contribution by SCHODDE
(1969) favoured the recognition of the Atherospermataceae and the same author later proposed
the erection of Siparunaceae (SCHODDE, 1970). This was followed by Hortoniaceae (SmitH, 1972).
Systematists currently favouring the broader view of the family include THORNE (1974), who brief-
ly argues the case for amalgamation concluding ‘The logical alternative treatment would be to ex-
pect five or more separate and obviously closely related families, an exercise in taxonomic infla-
tion that would seem to serve no useful purpose.’ The same view is taken by DAHLGREN (1980)
and Cronguist (1981) and this treatment is adopted here.

Subdivision. The grouping of genera into subfamilies and tribes is still subject to debate. Gener-
ally speaking those authors taking a broad view of the family recognize the same subdivisions as
the splitters but treat them as subfamilies or tribes. PerK1Ns (1925) adopted only two subfamilies,
Monimioideae and Atherospermoideae with four and two tribes respectively. MELCHIOR (1964)
followed Money, BaiLey & Swamy (1950), omitting Amborella and Trimenia, and accepting two
subfamilies, Hortonioideae and Siparunoideae. THoRNE (1974) added a further subfamily by
restricting the Monimioideae to Monimia and Peumus, and forming the Mollinedioideae for the
several remaining genera. His reduction of ScHoppe’s subfamily Peumoideae (ScHoppE, 1970)
into his restricted Monimioideae is accepted by PHILIPSON (Nordic J. Bot., 1986, in press), who
added Palmeria to this subfamily. Most Malesian genera fall into subfamily Mollinedioideae,

262 FLORA MALESIANA [ser. I, vol. 102

seven into tribe Mollinedieae and one Levieria into tribe Hedycaryeae. Two other subfamilies are
represented by one genus each: Atherospermatoideae by Dryadodaphne and Monimioideae by
Palmeria.

Generic limits. The family comprises several small distinctive genera whose definition is not dif-
ficult. It is perhaps only among the genera of the tribe Mollinedieae that generic limits become
problematic and it is these genera which are abundant in Malesia. In the first place it is possible
to distinguish those genera which receive their pollen on a hyperstigma secreted by prominent
glands within the ostiole of the female receptacles. These are (in Malesia) Kibara, Wilkiea and
Faika. Kibara is distinguished from the other two by the regular arrangement of its stamens. In
Wilkiea and Faika the stamens are inserted irregularly over the inner surface of the male recep-
tacles. These two genera are separated by the dehiscence of the anthers: in Wilkiea this is by a
single horizontal or horseshoe-shaped slit, whereas in Faika it is by two vertical slits. The two ge-
nera are also well separated geographically. Parakibara cannot yet be placed by this system be-
cause its female flowers are not known. Three Malesian genera lack a hyperstigma. Of these Ste-
ganthera is the largest and is closely related to Matthaea, a genus with a more westerly range,
which differs by its anthers opening by two vertical slits. The third genus, Kairoa, is immediately
distinguished by the male receptacles which open widely at anthesis to expose the very large num-
ber of stamens.

References: BARTLING, Ordines Naturales Plantarum, Dietrich, Géttingen (1830); BENTHAM &
Hooker, Genera Plantarum 3 (1880); R. Brown in Flinders, Voyage 3 (1814); CRoNQulisT, An inte-
grated system of classification of flowering plants, New York (1981); DAHLGREN, J. Linn. Soc. Bot.
80 (1980) 91—124; ENDLICHER, Genera Plantarum | (1837); Grsps, Contr. phytogeography & flora
Arfak Mts (1917); pe Jussteu, Ann. Mus. Nat. Hist. Nat. Paris 14 (1809) 116—135; LinDLEy, Vege-
table Kingdom, ed. 3 (1853); MELCHIOR in Engler, Syllabus Pflanzenfamilien, ed. 12 (1964);
Money, BAILEy & Swamy, J. Arn. Arb. 31 (1950) 372—404; PaxinE. & P. Nat. Pfl. Fam. 3, 2 (1891)
94—105; PERKINS, Pfl. R. Heft 49 (1911); Ubersicht iiber die Gattungen der Monimiaceae, Leipzig
(1925); PERKINS & Gita, Pfl. R. Heft 4 (1901) 1—22; PicHon, Bull. Mus. Hist. Nat. Paris 20 (1948)
383—384; ScHopDDE, Monograph of the family A therospermataceae R. Br., Thesis, Univ. Adelaide
(1969); Taxon 19 (1970) 324—328; Smirn, J. Ind. Bot. Soc. 50A (1972) 215—226; THoRNE, Aliso 8
(1974) 147—209; TuLasne, Arch. Mus. Nat. Hist. Nat. Paris 8 (1855) 273—436.

Specific delimitation. The much greater amount of material now available, especially from the
mountains of New Guinea, has revealed that many of the species described by earlier authors are
conspecific, and has also brought to light many undescribed species. Unfortunately, a consid-
erable number of the type specimens of earlier species have not been located, so that several spe-
cific names have been treated as unsufficiently known species. The concept of a species in this
family often appears less well defined than in many others. The larger genera include local species
which are well segregated, but also more widespread species with an unusually wide range of varia-
tion. Undoubtedly some regional and local species will eventually be recognized within these large
species, but this is not possible at present. The largest and most variable species are Kibara co-
riacea, Steganthera hirsuta, Palmeria arfakiana and Levieria squarrosa.

Uses. A variety of minor local uses are reported by collectors. The wood of larger species may
be used as stakes and for house-building, and the stems of Palmeria spp. for binding. The aromat-
ic leaves of several Palmeria species are used for smoking or to provide salt. Meat is wrapped in
leaves of Kibara possibly as a tenderizer. Pewmus (non-Mal.) has many uses in Chile: the hard
wood provides handles for implements and is converted to charcoal, the bark is used for tanning
and dyeing, and the leaves for medicinal purposes. Laurelia produces useful timber.

Notes for collectors. Species will usually be located by the characteristic foliage or by the
conspicuous fruits. As the fruits are not sufficient for generic determination, it is important to
search for flowers. These are so inconspicuous that they are commonly dismissed as buds and not
collected. If flowers cannot be found on the plant a search in the forest for other specimens will
usually prove successful. Always search for examples of flowers of both sexes, and bear in mind
that these may occur on separate plants. Examples of the fruits should be preserved in fluid.

1986 | MoNIMIACEAE (Philipson) 263

KEY TO THE GENERA
(11. Lauterbachia not included)

1. Flowers hermaphrodite; anthers valvate; filaments of stamens with a pair of glands; fruits with persistent
elongated plumose style (Subfam. ATHEROSPERMATOIDEAE)...........--0-0000000+ 1. Dryadodaphne

1. Flowers unisexual; anthers dehiscing by slits; filaments without glands; style not persistent in fruit.

2. Lianes; receptacle splitting irregularly when ripe to reveal drupes dispersed over its inner surface (Subfam.
IEE EAE) 5 GOs GO a oid ors SIG Seid Memeo « ache Susi Ne aie le oi es ete ates plore te alee sie 2. Palmeria

2. Trees or shrubs, or if rarely scandent then fruit not as above (Subfam. MOLLINEDIOIDEAE).

3. Male flower with a small, flat receptacle bearing relatively large rotund tepals; after anthesis female re-
ceptacle splitting irregularly from the ostiole; fruit a head of sessile drupes with the irregular rim of the
wecmiacie Tellexed below them (Tribe’HapycaRYBAS) 920)... 250 S20. Ss eee ee 3. Levieria

3. Male flower with a globose or flask-shaped receptacle, either with a small ostiole or (in Kairoa) splitting
open at anthesis to form a cup with sharply lobed margin; after anthesis upper half of female receptacle
abscissing as a calyptra; fruit a head of sessile or stipitate drupes with the annular scar of the calyptra
below them (Tribe MOLLINEDIEAE).

4. Female receptacle with thickened glands inside the ostiole.
5. Style-stigma subulate; anthers dispersed irregularly within the receptacle.

Haantners deniscing by-a horseshoe-shaped slit,....5. 45.2 ...5.. 23... Ses wa ee oe ke ee 4. Wilkiea
Gerensners ochiscing by. two: verticalislits . 22... ae as oe es OE. eae tee ee ae 5. Faika
5. Stigma sessile obtuse; anthers disposed regularly in whorls of 4 or in decussate pairs.

Pameamens m several whorls Of 4 ...62.5.0 oes 8 es Se me ot oe a he cael ee c ehebate ete 6. Parakibara
Se eamiens i two whorls (up to’4 stamens) '<... <0. eS a. POLS: 7. Kibara

4. Female receptacle without glands inside the ostiole.
8. Male receptacle at anthesis a fleshy cup with rim split into acute lobes; stamens numerous (over 100);
stigma short sessile; stems and branches with prominent irregular longitudinal ridges of cork

8. Kairoa

8. Male receptacle at anthesis with an ostiole surrounded by minute tepals; stamens few; branches +
smooth.

9. Anthers dehiscing by a single horizontal or horseshoe-shaped slit............... 9. Steganthera

Pemnness dehiscing: by two verticalshits 0.4... AE ese I ee ee Oe tee 10. Matthaea

1. DRYADODAPHNE'

S. Moore, J. Bot. 61 (1923) 109; A.C. Smitu, J. Arn. Arb. 23 (1942) 442. —
Isomerocarpa A.C. SmitH, J. Arn. Arb. 22 (1941) 250. — Daphnandra PER-
KINS, Bot. Jahrb. 52 (1915) 217, p.p.; Ubersicht Gattungen Monim. (1925) 47,
p.p. — Levieria Koster. Rec. Trav. Bot. Néerl. 34 (1937) 605, p.p.; HUTCH.
Gen. Fl. Pl. 1 (1964) 114, p.p. — Fig. 3.

Tall trees, young branches + tetragenous usually glabrous. Leaves dentate or
almost entire. Inflorescence axillary, of dichasia or few-flowered pleiochasia,
bracteoles broad enclosing the flowerbud, early caducous. Flowers bisexual,
perianth and androecium on the rim of the hypanthium; tepals 8, in 2 whorls,
oblong-elliptic, obtuse. — Androecium tetramerous, the outer 1—2 whorls of
stamens with divergent ovate-deltate, planate to shallowly cupular, obtuse
staminal glands and shortly apiculate to broadly rostrate, latrorsely to + ex-
trorsely dehiscing anthers, the inner whorls of lanceolate-deltoid to almost sub-
ulate staminodes. — Gynoecium of several free carpels, styles terminal; ovule

(1) The account of this genus is based on ScHoppe, A monograph of the family Atherospermataceae R. Br.,
unpublished thesis, University of Adelaide (1969).

264 FLORA MALESIANA [ser. I, vol. 102

Fig. 3. Dryadodaphne crassa SCHODDE. a. Habit, nat. size, b. fruiting hypanthium, x1 1/2, c. anther of outer
whorl, x14, d. embryo, x20, e. open flower, <7 (a, c, e STREIMANN 8643, b NGF 28519, d SCHODDE 3144).

subbasal. Fruiting hypanthium cylindrical-ellipsoid, dehiscing into 4 (some-
times 2) equal valves. Nutlets narrowly ellipsoid to obclavate, silky hirsute on
one side in deep pits, styles terminal, subulate.

1986 |

MONIMIACEAE (Philipson)

265

Distr. Three species, Queensland (1 sp.), in Malesia: New Guinea (2 spp.).

Ecol. Montane rain-forest, (500—)600—2800 m. In New Guinea D. crassa is associated consistently with
Nothofagus forest, and D. novoguineensis with lower altitude mixed forest.

Note. Dryadodaphne is the only representative of the A therospermatoideae in Malesia. The New Guinea
material was originally confused with Daphnandra, and any reports of that genus from Malesia refer to

Dryadodaphne.

KEY TO THE SPECIES

1. Stamens 8 (very rarely 4); outer stamens (including appendages) broader than long with widely divergent
+ acute glands. Leaves obovate (rarely elliptic) with rounded, obtuse, frequently retuse apices. Tepals in

+ equal whorls, usually + pale cream green ....

1. D. crassa

1. Stamens 4, including appendages longer than broad, with slightly divergent + obtuse glands (but see ssp.
occidentale). Leaves lanceolate to elliptic with attenuate to acuminate (rarely rounded obtuse) apices. Tep-
als in unequal whorls, usually pale maroon-brown, pinkish or purplish......... 2. D. novoguineensis

1. Dryadodaphne crassa SCHODDE, sp. nov. — Iso-
merocarpa novoguineensis (PERKINS) A.C. SMITH, J.
Am. Arb. 22 (1941) 251, p.p. — D. novoguineensis
(PERKINS) A.C. SmiTH, ibid. 23 (1942) 443, p.p. —
Fig. 3.

Arbor magna 20-30 m alta, folia comparate lata,
obtusa saepe retusa, series tepalorum aequiforma-
tae, plurumque virellae, stamina fugentia octo rare
quattuor, cum apiculis brevibus, dehiscentia valva-
rum antherarum revera laterali, et glandulis diver-
gentibus, late aliformibus, planis-cupulatis, ovatis-
lanceatis, acutis vel aliquantum obtusis, 0.5—1.2
(—1.5) mm longis, et hypanthia fructificantia compa-
rate brevia, (15—)18—23(—27) mm longa x (5-)
6—10(—11) mm lata, crasse ligneaque. — Type:
SCHODDE & CRAVEN 4816 (CANB), Morobe Prov.,
Menyamya Distr., Angebena ridge, c. 3 miles east of
Aseki.

Tree to 20—30 m; branchlets usually tetragonous,
glabrous, or young parts sometimes unevenly sordid
buff tomentulose, or with scaly incrustation. Leaves
narrowly to broadly ovate; sometimes rounded cu-
neate, infrequently elliptic (0.7—)2.5—6(—10) by
(0.5—)1—3(—4) cm, base + narrowly cuneate, apex
broadly, or less often narrowly, obtuse, frequently
retuse, margin faintly undulate, or obscurely or ob-
tusely crenulate, rarely markedly crenulate or entire,
+ markedly recurved, almost revolute towards the
petiole, coriaceous or rarely thinly coriaceous, prin-
cipal nerves conspicuous below, anastomosing to-
wards the margin, glandular pits sparse to frequent;
petioles up to 9 mm, or sometimes longer, deeply
channelled above, glabrous. /nflorescences axillary,
up to 3 cm long, or rarely longer, glabrous or tomen-
tose distally, bearing few (rarely more than 5 but up
to 11) flowers in a dichasium or a pleiochasium;
bracts lanceolate deltate to ovate or oblong, (1.5—)
2—3(—4) mm long, caducous; pedicels slightly or +
conspicuously expanded under the hypanthium;
bracteoles enclosing the hypanthium in bud. Flowers

with perianth pale cream or yellow-green, rarely
whitish or reddish; hypanthium 1—2(—2.5) mm long,
glabrescent or sparsely tomentulose; tepals 8 in 2 +
equal whorls, broadly obtuse oblong-linguiform to
elliptic, (1.5—)3—4.5(—S) by (1.5—)2—2.7(—3) mm.
— Androecium in several series, the 2 outer whorls
each with 4 stamens, outer stamens broader than
long (1—)1.3—2 by (1.25—)1.5—2.5(—3) mm, with
widely divergent, wing-like ovate to lanceolate-
deltate, + acute staminal glands, and latrorsely
dehiscing valves, inner stamens as broad as long,
with shorter + obtuse glands and longer apiculate
apices; staminodes narrowly to broadly lanceolate or
deltoid to subulate, the outer often with vestigial
glands. — Gynoecium of up to 15 carpels; styles ex-
serted beyond the hypanthium as a slender column,
stigmatic column narrowly conical, protruding be-
yond apices of staminodes to near apices of outer sta-
mens. Fruiting hypanthium thickly lignified, de-
hiscing into 4 (rarely 2) equal valves, sparsely hairy
within. Nutlets (3—)5—6 mm long, styles (5—)7—10
(—14) mm long, exserted beyond the hypanthium.

Distr. Malesia: New Guinea. Through the cen-
tral mountains from Oranje Range to the Owen Stan-
ley Range, and also in the Saruwaged Range, Huon
Peninsula.

Ecol. Primary, or rarely old secondary, montane
rain-forest, between 2000 and 2800 m in the north-
western and central parts of its range, and between
(1350—)1600—2400 m in the Eastern Highlands,
Saruwaged and Owen Stanley Ranges. In fairly well-
drained sites, usually on slopes and towards the
crowns of mountains and ridges. Generally associat-
ed with forest dominated by Nothofagus. Towards
the upper limit of its altitudinal range it extends into
conifer forests dominated by species of Libocedrus,
Phyllocladus and Podocarpus.

Uses. The timber is used in the Mt Hagen and
Chimbu areas.

Vern. Dafa, Telefomin, enman, Banz, katan,

266

FLORA MALESIANA

[ser. I, vol. 102

mongo a kuli, nopohn or pohn, piljim, Mt Hagen,
korgi, Chimbu, mowku, mugu, muk, Enga, munk,
Mendi, onda or onde, Aseki.

Note. Though the distinctness of the two New
Guinea species has been recognized only compara-
tively recently, most individuals can be distinguished
readily on leaf characters such as size, texture and
shape. Leaves of saplings and those in deep shade
tend to be larger and less coriaceous with narrower
apices and planate margins. Trees from lower alti-
tudes may show some features of D. novoguineensis
suggesting the possibility of introgression. The most
westerly collection (BRASS & VERSTEEGH 11194) is
somewhat distinct, and further collections from this
region are needed. Flowering and fruiting occur
throughout the year, though an individual tree usual-
ly does not bear flowers and fruits together.

2. Dryadodaphne novoguineensis (PERKINS) A.C.
SmirH, J. Arn. Arb. 23 (1942) 443, p.p. — Daphnan-
dra novoguineensis PERKINS, Bot. Jahrb. 52 (1915)
217; Ubersicht Gattungen Monim. (1925) 50. — D.
celastroides S. Moore, J. Bot. 61 (1923) 109. —
Daphnanadra perkinsiae Gitc & Diets, Notizbl. Berl.-
Dahl. 9 (1925) 466. — /somerocarpa novoguineensis
(PERKINS) A.C. SmitH, J. Arn. Arb. 22 (1941) 251,

D.p.

KEY TO THE SUBSPECIES

1. Staminal glands ovate oblong, obtuse or shallowly
cupular. Leaves (from crown) with acuminate,
rarely rounded obtuse, apices and distinctly undu-
late-crenate margins. Fruiting hypanthium 2—4
times longer than broad. a. ssp. novoguineensis

1. Staminal glands lanceolate to ovate, rather acute.
Leaves (from crown) with narrowly obtuse, hard-
ly acuminate apices, and obscurely undulate-
crenate, almost entire margins. Fruiting hypan-
thium | 3/4—2 times longer than broad

b. ssp. occidentalis

a. ssp. nOvoguineensis var. novoguineensis

Tree to 40 or 50 m, often buttressed; branchlets
usually conspicuously tetragonous, glabrous or
sometimes hispidulous or tomentulose. Leaves nar-
rowly to broadly elliptic, sometimes lanceolate, (2—)
4—10(—12) by (0.7—)1.5 by 3.5(—4.5) cm, base trun-
cate to widely or rarely narrowly cuneate, apex blunt-
ly acuminate, very rarely narrowed, margin conspi-
cuously undulate crenate, slightly to markedly re-
curved, + coriaceous, principal nerves + prominent
below, anastomosing towards the margin, occasion-
ally with sparse glandular pits; petioles up to 15 mm,
sometimes longer, channelled above, glabrous,
sometimes strigillose. Inflorescences axillary, up to

35 mm long, rarely longer, glabrous or glabrescent
with sparse hairs sometimes persistent on bracteoles
and perianth in buds, bearing few (rarely more than
5, but up to 11 flowers) in a dichasium or a pleiocha-
sium; bracts lanceolate-deltate, rarely linear oblong,
(1.5—)2.5—4 mm long, early caducous; pedicels rare-
ly much expanded under the hypanthium, bracteoles
enclosing the hypanthium in bud. Flowers with peri-
anth reddish (rarely entirely cream-green); hypan-
thium 1—2(—2.5) mm long, glabrous to rather dense-
ly tomentulose; tepals 8, in 2 unequal whorls, the
outer obtusely linguiform, sepaloid, (2—)3—4(—6) by
1.5—3(—4) mm, the inner narrowed towards the
apex, more petaloid, (2—)2.7—3.5(—5) by (1.2—)
1.5—2.5(—3.5) mm. — Androecium in 2 or 3 series,
the outer whorl of 4 stamens; stamens longer than
broad, (1.3—)1.5—2(—2.5) by 1—1.7 mm, with slight-
ly divergent, lanceolate or ovate-oblong obtuse, rare-
ly acute staminal glands, and extrorsely dehiscing
valves, the inner series of lanceolate, deltoid to subu-
late staminodes, the outer often with vestigial glands.
— Gynoecium of up to 16 carpels, styles exserted
beyond the hypanthium, stigmatic column narrowly
conical, reaching apices of inner staminodes, rarely
beyond. Fruiting hypanthium thickly lignified, (11—)
18—25(—35) mm long, dehiscing into 4 (rarely 2)
equal valves, hairy within especially towards the rim.
Nutlets (3—)5—6(—7) mm long, styles (5—)7—14(—15)
mm long, exserted beyond the hypanthium.

Distr. Malesia: Papua New Guinea. Throughout
the central mountains from the Hunstein Range to
the Owen Stanley Range, and also in the Torricelli
Mts and the Saruwaged Range.

Ecol. Primary montane rain-forest, and second-
ary forest as a remnant, 500—1950 m. In general at
lower altitude than D. crassa and preferring gullies,
slopes and fairly well-drained ridge crowns. Only
rarely associated specifically with Nothofagus forest.

Uses. The bark is chewed and rubbed on the chest
by Chimbu natives for certain illnesses.

Vern. Adengambi, Chimbu, anona, Akuna,
anonya, nasapu, Aiyura, kamo, Okapa, onda or on-
de, Aseki.

Note. D. crassa may be distinguished from this
species by its relatively broadly obtuse, frequently re-
tuse leaves, generally cream-green flowers, equal
whorls of tepals, two unequal whorls of stamens,
broad wing-like staminal glands attached near to the
base of the filament, shortly apiculate outer anthers,
latrorse anther dehiscence, and shorter fruit.

var. macra SCHODDE, var. nov.

Laminae foliorum coronarum arborum saepe
longiores quam 100 mm, latioresquam 35 mm, et pe-
tioli (1.5—)2—2.5(—3.5) mm crassi, inflorescentiae
relative tomentulosae dense glauco-griseae vel aluta-
ceae, et hypanthia fructificantia 35—45 mm longa. —

1986 |

Type: SCHODDE & CRAVEN 5081 (CANB), Morobe
Prov., Menyamya Distr., Aseki Valley, c. 3 miles SE.
of Aseki.

Differs from var. novoguineensis in the + densely
glaucous-grey or pale brown tomentulose inflores-
cence, and in the larger leaves and fruiting hypan-
thia.

Distr. Malesia: Papua New Guinea, Enga Prov.,
Wabag Distr., Morobe Prov., Menyamya and Wau
Distr.

Ecol. Montane rain-forest, 1000—2400 m.

Note. Known only from within the range of ssp.
novoguineensis and usually in close proximity to it.

2. ssp. occidentalis SCHODDE, ssp. nov.

Cortex relative asper fissuratusque, folia corona-
rum arborum relative anguste obtusa vix acuminata
ad apices, et obscure undulato-crenata paene integra,
glandulae staminum planae, lanceolate ad ovate

MONIMIACEAE (Philipson)

267

1 3/4—2 plo longiora quam latiora. — Type: Brass
& VERSTEEGH 11984 (L), Irian Jaya, 15 km SW. of
Bernhard Camp, Idenburg River.

Differs from ssp. novoguineensis in the longer and
flatter divergent staminal glands, the rounder apex
and more entire margin of the leaf, the thicker fruit-
ing hypanthium and the rougher dark-brown bark.

Distr. Malesia: Irian Jaya, Vogelkop Peninsula,
Mt Nettoti; Wissel Lakes; Idenburg River.

Ecol. Primary montane rain-forest (Nothofagus)
and mossy forest, 900—1900 m.

Vern. Goekaai, Kapauku lang.

Note. The three localities from which this species
is known are far apart, but the specimens are uni-
form, especially as regards their staminal glands. D.
novoguineensis apparently shows no clinal trend to-
wards the characters of this species at the western end
of its range. A considerable geographical gap (of c.
200—250 miles) separates the ranges of the two sub-

0.7—1 mm longae, et hypanthia fructificantia species, as known at present.

2. PALMERIA

F. v. M. Fragm. 4 (1864) 151; A.DC. Prod. 16, 2 (1864) 657; Bru. Fl. Austr.
5 (1870) 291; PERKINS & Gra, Pfl. R. Heft 4 (1901) 64; Perkins, Bot. Jahrb.
§2 (1915) 214; Ubersicht Gattungen Monim. (1925) 42; Hutcu. Gen. FI. Pl. 1
(1964) 119; PHitipson, Blumea 28 (1982) 85, f. 1—3. — Fig. 4—5.

Woody lianes with opposite, entire leaves, usually bearing stellate hairs. Dioe-
cious, with lateral or terminal cymose panicles or pleiochasia. — Male flowers
bowl-shaped or saucer-shaped, with 4—7 tepals either incurved or spreading at
anthesis to reveal the numerous stamens + sessile on the surface of the recep-
tacle; anthers opening by longitudinal slits. — Female flowers globose or flask-
shaped with c. 5 small obtuse tepals surrounding a minute ostiole; carpels sessile
on the inner surface of the receptacle, interspersed with numerous bristles. Re-
ceptacle enlarging to become a + globose fruit, which splits open irregularly at
maturity to reveal the drupes.

Distr. Australia and Malesia: 11 spp. in New Guinea, of which 3 may also occur in Queensland and one
extends to E. Sulawesi. Three further species have been described from Australia.

KEY TO THE SPECIES

Lanes sustace.of receptacle slabrous. 5. state fie e0ls coi oc ip-oce vis vin 'v gis aw 0/a-vi ve wie ben aan 1. P. hooglandii
1. Outer surface of receptacle bearing hairs.
marmcerouriace Of leaf +. GIGBrOUs ¢ ia\c.< Wai ooo sis pes ov saves nove vinden toda sana 3. P. arfakiana

2. Undersurface of leaf with indumentum.
3. Undersurface of leaf bearing numerous simple hairs (and usually also some stellate hairs) 2. P. gracilis
3. Undersurface of leaf bearing stellate hairs (occasionally also with a few simple hairs).
4, Mature fruits c. 7~8 mm @ (before dehiscing). Achenes about 5 mm long when dry (known only from
me Agel 1 akes Gree). ie \isin tee eacamath ola ie dxcp baie -aiile aes Cetin Bt cachet een A 4. P. angica

268 FLORA MALESIANA [ser. I, vol. 102

Fig. 4. Palmeria clemensae Pui.ipson. a. Habit of male twig, «1/2, b. portion of lower surface omleaivixKs
(a PULLEN 281, b VAN BaLcooy 957).

1986 } MONIMIACEAE (Philipson) 269

4. Mature fruits c. 15—20 cm @ (before dehiscing). Achenes about 7 mm long or longer when dry.
5. Stellate hairs of underside of leaf laxly or densely distributed but not matted to form a close felt.
6. Stellate hairs of the underside of leaf large (0.6—0.9 mm), generally distributed over the leaf surface,
usually rather dense (but the leaf-surface clearly visible through them) ............ 5. P. brassii
6. Stellate hairs of the underside of the leaf small (0.2—0.25 mm) occasionally generally distributed and
dense, but often confined to leaf base and petiole (common throughout New Guinea) 3. P. arfakiana
5. Stellate hairs densely distributed over the undersurface of the leaf and matted together to form a close

felt.

7. Felt on the underside of the leaf with a smooth even surface.
8. Leaves large (mostly over 12 cm long) (known from Morobe and Central Provinces) 6. P. incana

8. Leaves smaller.

9. Leaves small (less than 5 cm long) (known from the Murray Pass area) ....... 7. P. montana

9. Leaves larger (Tari District westwards)...

= 0 RS, SR. Ses. SE eee 8. P. hypargyrea

7. Felt on the underside of the leaf with some larger hairs protruding above the general surface.
10. Veins on the underside of the leaf fringed with long hairs .................. 9. P. clemensae
10. Veins on the underside of the leaf bearing tufted stellate hairs above the felt.

11. Leaves large (9 cm or longer) ..........
11. Leaves smaller (6 cm or shorter) .......

1. Palmeria hooglandii PHitipson, Blumea 28 (1982)
88.

Woody liane to 22 m high, branches and foliage
glabrous. Leaves oblong-elliptic, occasionally lan-
ceolate-elliptic, 7.5—15.5 by 1.7—6 cm, chartaceous,
apex with an apiculum (often long and narrow), base
cuneate, midrib slightly channelled above, promi-
nent below, lateral veins c. 6, well-defined, meeting
within the margin; petiole 0.75—1 cm. Inflorescence
axillary and terminal, either simple unbranched
pleiochasia (often grouped at the ends of branches to
resemble panicles) or with long lateral branches
forming a panicle-like inflorescence, the rachis
4.5—10 cm, with lateral branches up to 4.cm; male in-
florescences often larger and more openly branched
at anthesis than females. — Male receptacles bowl-
shaped becoming almost disk-shaped at anthesis, c.
5.5 mm @ (without the tepals), margin produced into
c. 5—7 irregular tepals, outer surface of tepals
minutely pubescent; stamens c. 18, subsessile; an-
thers c. 0.75 mm. — Female receptacle globular, 2.5
mm @, with a terminal ostiole surrounded by 5 mi-
nute obtuse tepals, a few microscropic hairs present
on the tepals, the inner surface of the receptacle bear-
ing long simple hairs between the carpels; carpels c.
8, tapering to filiform styles which project through
the ostiole. Immature fruit globose or irregularly
bulged, with an asymmetric beak, mature fruit not
seen.

Distr. Malesia: Papua New Guinea (Southern
Highlands, Enga, Western Highlands, Chimbu,
Eastern Highlands, Morobe Prov.).

Ecol. A liane reaching 22 m in primary and sec-
ondary lower montane rain-forest, 1950-2800 m.

Vern. Kende, kent, korinda, all Enga, boma,
Chimbu, abiya, Okapa.

"Note. One of the most distinctive species, with
both the vegetative parts and inflorescence glabrous

seis de RR ee eee 10. P. womersleyi
SER OO See eee 11. P. schoddei

(only the tepals are minutely pubescent). Its distribu-
tion covers most of the Central Highlands from La-
gaip to Okapa, with a single disjunct occurrence in
the Cromwell Mts (Huon Peninsula). The flowers are
described as creamy and scented. The drupes are
purple-black on a bright red torus.

2. Palmeria gracilis PERKINS, Bot. Jahrb. 31 (1902)
745; Pfl. R. Heft 49 (1911) 39; Puttrpson, Blumea 28
(1982) 89, f. 1b. — P. fengeriana Perkins, Pfl. R.
Heft 49 (1911) 39. — P. paniculata Riv. Trans.
Linn. Soc. Bot. II, 9 (1916) 144.

Woody liane, reaching 20 m, young parts covered
with a lax indumentum of simple hairs, together with
small stellate hairs especially on the flowers. Leaves
usually ovate with broad rounded, cordate, or cu-
neate base, and a long narrow apiculum, more rarely
elliptic with a short apiculum, 4—14.5 by 1.8—11.5
cm, membranaceous, midrib and lateral veins well-
defined, upper surface with the remnants of simple
hairs and small stellate hairs, or becoming glabrous
except for simple hairs along the midrib and some-
times the principal veins and the margin, undersur-
face with a dense or sparser covering of curved
simple hairs mixed with a varying number of small
stellate hairs; petiole 2.5—-10 mm, hairy. J/nflo-
rescence axillary and terminal, either simple pleio-
chasia, few-flowered and with a delicate rachis, or
branching to the second degree with rather stouter
rachis, 5—20 cm long, often produced profusely on
lateral branches of limited growth; rachis, branches
and pedicels with lax hairs and a varying amount of
short dense stellate hairs; bracts linear, caducous. —
Male receptacle saucer-shaped, 6—8 mm @, with 5 re-
curved deltoid tepals, outer surface with long simple
hairs and dense stellate indumentum; stamens c.
20—24, sessile, c. 1.25 mm long. — Female receptacle
globose, c. 2.25 mm @, outer surface with indumen-

270

FLORA MALESIANA

[ser. I, vol. 102

tum as in male, inner surface with long bristles be-
tween the carpels; carpels c. 8—10, tapering to a long
filiform reflexed style. Immature fruit subspherical
with an asymmetric beak; mature fruit splitting into
irregular lobes c. 2—2.5 cm long, densely bristly on
both surfaces; drupes spherical, sessile, c. 7 mm long
when dry.

Distr. Malesia: Irian Jaya (Vogelkop Peninsula,
Japen I., Lake Habbema, Idenburg R.); Papua New
Guinea (W. & E. Sepik, Southern, Western & East-
ern Highlands, Morobe & Central Provinces).

Ecol. Woody liane reaching 20 m or more, in
rain-forest, second growth or open scrubby areas
with a wide altitudinal range (SO0—3000 m), but most
frequent in lower montane rain-forest (Castanopsis,
Lithocarpus, Nothofagus, Podocarpus or Elaeocar-
pus dominated).

Uses. For wrapping tobacco before being
smoked in pipes or as cigarettes. When burnt to a
fine ash, used as salt.

Vern. Gapunga, Koroba, hompanofi, Okapa,
kepundom, Maring, obirambiram, Mendi, fown-
dun, Maring.

Note. One of the most widespread and frequent
species, recognized by the simple curved bristles on
the undersurface of the leaf. The size of the leaf is
varied, as is the density of the indumentum, but the
variability has no geographical or apparent ecologi-
cal basis. The flowers are cream, in males with whit-
ish stamens. The red or dark drupes are borne on a
light red receptacle.

3. Palmeria arfakiana Becc. Malesia 1 (1877) 186;
PERKINS, Pfl. R. Heft 4 (1904) 65; Bot. Jahrb. 52
(1915) 215; A.C. Smitu, J. Arn. Arb. 22 (1941) 245;
KANEH. & Hatus. Bot. Mag. Tokyo 56 (1942) 254;
PuiLipson, Blumea 28 (1982) 90. — P. warburgii
PERKINS, Pfl. R. Heft 49 (1911) 37, f. 13D —H. — P.
pulchra Perkins, /.c. 38. — P. myriantha PERKINS,
Bot. Jahrb. 52 (1915) 214. — P. myrtifolia PERKINS,
Ubersicht Gattungen Monim. (1925) 43, nomen. —
P. puberula A.C. Smitn, J. Arn. Arb. 22 (1941) 249.
— P. acuminata KANEH. & Hatus. Bot. Mag. Tokyo
5 (1942) 251, f. 3F—G. — P. parvifolia KANEH. &
Harus. 163,255; f. 3A—E:

Woody liane, young branches hirsute or puberu-
lous. Leaves usually oblong-elliptic or obovate,
sometimes narrower or subrotund, (2.5—)12—20 by
1—9.5 cm, chartaceous to coriaceous, apex attenuat-
ed to a short or long apiculum, base rounded, trun-
cate, or cuneate, upper surface of mature leaves with
remnants of minute stellate hairs, especially at the
base of the midrib, or glabrous, undersurface with
minute stellate hairs forming a dense or open indu-
mentum or virtually absent from the mature blade,
persisting, if at all, on the midrib or near the base of
the leaf, longer simple hairs may be present along the

midrib; petiole 3-15 mm long, either densely cov-
ered with minute stellate hairs or these may be sparse
(almost absent) at maturity, longer simple hairs
sometimes also present and abundant. /nflorescence
axillary and terminal, paniculate, 7-20 cm long
(rarely shorter), lower branches 1.5—3(—5) cm with
few flowers or with tertiary branches, the rachis and
branches usually with a greyish or cream tomentum
or with sparse stellate hairs, rarely more or less gla-
brous at anthesis. — Male receptacle bowl|-shaped
becoming a flat disk at anthesis c. 6-8 mm @, with
c. 5 irregular tepals, outer surface with a short dense
indumentum or with fewer minute stellate hairs, in-
ner surface with short simple hairs; stamens c. 25,
sessile c. 0.75—1.25 mm long. — Female receptacle
urceolate, c. 2—2.5 mm high, outer surface with in-
dumentum as in male, inner surface with long simple
hairs; carpels c. 5—8(—16), with filiform styles. Im-
mature fruit subspherical, beaked, splitting irregu-
larly at maturity to reveal the red inner surface and
black drupes.

Distr. Malesia: Central Celebes (E. Peninsula)
and New Guinea: in uplands throughout the island
from Vogelkop Peninsula to Milne Bay; also on Ma-
nus I. and New Britain.

Ecol. Liane, often strongly growing in primary
rain-forest, mostly in the lower and upper montane
zones, ascending to 3200 m, but occasionally as low
as 100 m. Also in second growth and climbing over
shrubs near the forest margin.

Uses. The leaves are burnt to make salt.

Vern. Papua New Guinea: angore, laso, Mendi,
bengop, pengop, Hattam, gawa kepundom, Maring,
ibiwudediji, puquabou, Tari, kainagent, repalip,
tsik, Enga, kangom, laro R., linore, mulkrima, Tele-
fomin, njeng, Hagen, Wankl, tomenguntong, Sai-
der, xatapi, Guale, Movoi.

Notes. The most frequent and widespread spe-
cies of the genus. It is the only species known to occur
on islands to the north of the New Guinea mainland
and in Sulawesi.

The variety of forms included within the limits of
this species as treated here is greater than is generally
acceptable in one species. It is possible that future
studies will recognize entities within this complex,
and perhaps reinstate species here regarded as con-
specific. However, it has proved impossible to subdi-
vide on the basis of indumentum and leaf-size.
Broadly speaking four groups may be recognized,
though frequent intermediates link all of these: (i) a
large-leaved form with adult leaves more or less gla-
brous except for small stellate hairs on the petiole
and base of the leaf (P. arfakiana sensu stricto), (ii)
a large-leaved form with the lower surface more or
less densely and evenly covered with small stellate
hairs, (iii) a smaller-leaved form (often with nar-
rower more oblong blades) usually with the indu-

1986]

mentum persisting only near the leaf base (P. pul-
chra), (iv) a small-leaved form with few stellate hairs
(P. acuminata, P. parvifolia). The first three forms
are all frequent and widespread. The fourth appears
to be confined to mountains to the west of the island
(e.g. Arfak Mts, Wissel Lakes). The species de-
scribed by KANEHIRA & HaTusmMa are here regarded
as reduced subalpine states and reduced to synon-
ymy; they may prove to be distinctive enough for spe-
cific status when better material becomes available,
but the evidence available is that they are the end of
a reduction series. The type of P. myriantha PERKINS
represents the most glabrous state, with even the
branches of the inflorescence more or less glabrous
and the receptacles bearing a rather sparse coating of
stellate hairs. However, similar collections occur
throughout the range and must be regarded as one
extreme of a continuous range of variation. I have
not seen type material of P. warburgii PERKINS from
Celebes (the only occurrence of the genus west of
New Guinea), but plants collected in Celebes by
Eyma fit the original description. The younger leaves
are rather heavily coated below with small stellate
hairs, but older leaves can approach the glabrous
condition.

Although undoubtedly a liane, like all other spe-
cies of Palmeria, it must vary in habit because a num-
ber of collectors describe it as a tree or sprawling
shrub. The young foliage is tinged with red. The
flowers are cream or yellowish. The black drupes are
borne on a red or pink receptacle.

4. Palmeria angica KANEH. & Hatus. Bot. Mag. To-
kyo 56 (1942) 252; Puitipson, Blumea 28 (1982) 92.

Woody liane to 3 m high, young branches with a
greyish indumentum. Leaves oblong-elliptic, 4—8 by
2—4 cm, chartaceous or coriaceous, base rounded,
apex shortly acuminate, the upper surface + densely
or sparsely stellate hairy, becoming glabrous, the
undersurface densely stellate-pilose; petiole 7—8 mm
long, closely pubescent. /nflorescence axillary, to 10
cm long, the rachis and branches with greyish stellate
indumentum. — Male flowers not seen. — Female
receptacle urceolate, 2.5 mm long (after flowering),
with short dense indumentum on the outer surface,
tepals 5, minute; carpels 7—8. Fruits subspherical, c.
8 mm 9, splitting irregularly; drupes c. 5 mm long
when dry.

Distr. Malesia: West New Guinea (Mt Arfak,
Angi Lakes).

Ecol. Scandent in low spinneys on the burnt and
open summit of Mt Koebre at 2300 m.

Note. The small fruits appear to enclose a single
drupe, which is considerably smaller than those of
the other species.

5. Palmeri brassii Puiipson, Blumea 28 (1982) 92, f.

MOoONIMIACEAE (Philipson)

yy i

ld. — ‘Palmeria fengeriana PERKINS’ A.C. SmiTH, J.
Arn. Arb. 22 (1941) 248, p.p.

Woody liane, to 20 high, young branches with a
thick indumentum. Leaves usually broadly elliptic or
slightly obovate or rotund, with a small blunt apicu-
lum or occasionally with an attenuated apex, 8—22
by 4.8—12 cm, chartaceous, midrib prominent be-
low, lateral veins c. 6, arched and meeting within the
margin, upper surface of mature leaves covered with
the scattered remnants of stellate hairs, which may
form a dense pile above the midrib and principal
veins and on the margins, lower surface covered with
large stellate hairs with lax, shining, bristle-like
arms, sometimes rather densely disposed but not ob-
scuring the surface of the blade; a few simple hairs
may occur among the stellate indumentum; petiole
10—17 mm, densely clothed in brown indumentum.
Inflorescences axillary and terminal, covered with a
short dense creamy or fulvous tomentum, elongate
(to c. 40 cm) with relatively short opposite or subop-
posite lateral branches (S—8 cm long) or the inflo-
rescence more paniculate with lateral branches
15—20 cm long (the male inflorescences frequently
more lax than the female at anthesis); lateral
branches bearing several opposite tertiary branches
and caducous subulate bracts. — Male receptacle
bowl-shaped, becoming almost a flat disk at anthe-
sis, with c. 5 irregular tepals, c. 5mm @, outer surface
with a dense covering of small stellate hairs, inner
surface with short simple hairs between the stamens;
stamens c. 20, sessile on the inner surface of the re-
ceptacle, c. 0.75—1 mm long, apex of the connective
with a tuft of minute hairs. — Female receptacle ur-
ceolate, often asymmetric, 2—3 mm high at anthesis,
outer surface with indumentum as in the male, inner
surface with long simple hairs between the carpels;
carpels c. 15 distributed over the inner surface of the
receptacle, tapering to filiform styles which project
through the ostiole, becoming reflexed. Immature
fruit subspherical, or irregularly bulged, usually
markedly asymmetric with a prominent beak to one
side; at maturity the enlarged receptacle splits to
form c. 5 coriaceous, very irregular lobes c. 2 cm long
to which the ripe achenes are attached; drupes spheri-
cal, sessile, with a shining black surface, mesocarp
succulent, endocarp stony, c. 7 mm long when dry.

Distr. Malesia; Papua New Guinea (Eastern
Highlands, Morobe and Central Provinces).

Ecol. Woody liane on shrubs and trees in pri-
mary and secondary lower montane rain-forest
(dominated by Castanopsis, Lithocarpus, Nothofa-
gus or Podocarpus), occurring in ridge forest and
also in swampy places with dark brown loam,
1200-2450 m.

Uses. Employed as lashing material.

Vern. Nani, Kassam, arawe, Finisterre Mts, bo-
ma, Gumini, arokot, Ueli, Movoi.

212

FLORA MALESIANA

[ser. I, vol. 102

Note. Characterized by its large leaves bearing a
loose indumentum of large stellate hairs with shining,
bristle-like, spreading arms. The flowers are de-
scribed as white. The immature green, white-spotted
fruits later split open to reveal black drupes on a red
torus. P. brassiimay be distinguished from P. gracilis
by the predominance of stellate hairs, by the shape of
the blade and by the longer inflorescence. Some spe-
cimens with more numerous simple hairs may also
approach P. gracilis in leaf shape. They are possibly
of hybrid origin. Of all the New Guinea species, P.
brassii probably is most closely related to the Austra-
lian P. scandens from which it differs principally in
the length of the inflorescence and the leaf shape.

6. Palmeria incana A.C. SmirH, J. Arn. Arb. 22
(1944) 245; Pumipson, Blumea 28 (1982) 94.

Woody liane, with young branches covered in
greyish tomentum. Leaves elliptic-oblong, 9—18 by
3.5—8 cm, chartaceous, base rounded or broadly cu-
neate, apex shortly acuminate, upper surface becom-
ing glabrous (except for puberulence above the mid-
rib), lower surface evenly covered with a close, fine,
greyish-white indumentum, midrib and arched later-
al veins prominent below, impressed above; petiole
6—10 mm, puberulous. — Inflorescence densely
greyish pubescent, axillary and terminal, narrowly
paniculate, up to 12 cm, with branches c. 2—3 cm. —
Male flowers globose, c. 2 mm @ (in bud), apex de-
pressed with 5 tepals, outer surface densely covered
with a close indumentum of stellate hairs; stamens c.
20—24, broadly deltoid, c. 0.75 mm long, sessile. —
Female flowers not known. Immature fruits sub-
spherical with an asymmetric beak; at maturity the
enlarged receptacle ruptures irregularly to form co-
riaceous lobes c. 2 cm long; drupes subspherical, c.
7 by 5 mn, sessile.

Distr. Malesia: Papua New Guinea (Morobe
Prov.: Wau Distr.; Central Prov.: Goilala Distr.).

Ecol. Liane scrambling over trees in primary for-
est at 1200 m.

Notes. Evidently a rare and local species, only
twice collected. Lower surface of leaves covered with
a smooth, even, greyish felt of close-set stellate hairs.
This indumentum is similar to that of P. hypargyrea,
but that western species has smaller leaves.

The original collection lacked flowers, but imma-
ture male flowers were provided by another. The
flowers are described as cream, with a + unpleasant
scent, and the drupes as black on a red torus.

7. Palmeria montana A.C. SmitH, J. Arn. Arb. 22
(1941) 247; Puirpson, Blumea 28 (1982) 95.
Woody liane to 3 m high, with slender branches
covered with a greyish indumentum of minute stel-
late hairs. Leaves ovate to oblong-ovate or broadly
ovate, 1.8—4.7 by 0.6—2.5 cm, chartaceous, base cu-

neate or rounded, apex curving into an acute apicu-
lum, midrib slightly channeled above, prominent be-
low, lateral veins rather obscure, upper surface of
mature leaves glabrous or with sparse remnants of
small stellate hairs, lower surface covered with a
dense fine greyish white felt of stellate hairs; petiole
to 5 mm long, puberulous. /nflorescence: axillary,
few-flowered pleiochasia, c. 2 cm long, covered with
a short dense indumentum, and bearing subulate
bracts; pedicels 3-7 mm. — Male receptacle bowl-
shaped, 4—5 mm @g at anthesis (without the tepals)
with 5 or 6 deltoid, acute tepals, outer surface densely
covered with minute stellate hairs, inner surface with
short simple hairs; stamens 15—20, filaments c. 0.4
mm, anthers c. 0.8 mm long. — Female receptacle
subglobose, 2—2.5 mm g, with a terminal ostiole sur-
rounded by 5 minute obtuse tepals, indumentum of
outer surface similar to that of male, inner surface
covered with long simple hairs between the carpels;
carpels 10—15, tapering to filiform styles which pro-
ject through the ostiole. Immature fruit subglobose,
splitting at maturity to reveal 1—5S drupes.

Distr. Malesia: Papua New Guinea (Central
Provy.).

Ecol. Sprawling over undergrowth in montane
forest or climbing to 3 m, also in disturbed forest and
tree-fern savanna, 2700—2850 m.

Notes. Known only from the vicinity of Murray
Pass. Features which distinguish this small-leaved
montane species from P. schoddei are given under
that species.

The flowers are described as cream, and the drupes
brown on a red receptacle.

8. Palmeria hypargyrea PERKINS, Bot. Jahrb. 52
(1915) 215; Puttipson, Blumea 28 (1982) 95, f. la. —
P. pulleana PERKINS, Ubersicht Gattungen Monim.
(1925) 43, nomen. — P. habbemensis A.C. SmitH, J.
Arn. Arb. 22 (1941) 246. — P. dallmannensis KAN.
& Har. Bot. Mag. Tokyo 56 (1942) 254, f. 4.
Woody liane; young branches with minute stellate
hairs, often 4-angled. Leaves elliptic or narrowly el-
liptic, 5—11.5 by 2.5—4.5 cm, thinly coriaceous, base
cuneate or rounded, apex shortly acuminate, obtuse
or acute, upper surface becoming glabrous or retain-
ing widely spaced remnants of stellate hairs, lower
surface evenly covered with a close, fine indumen-
tum (which may become thinner on the midrib and
principal veins or rarely over the surface of the bla-
de); petiole S—9 mm long, puberulous. /nflorescence
axillary and terminal, pleiochasial, 4—8(—13) cm
long, lateral branches few-flowered, densely covered
in greyish pubescence, except that the peduncle and
rachis may become glabrous or bear only sparse mi-
nute stellate hairs; small caducous bracts below the
branches and some pedicels. — Male receptacle
saucer-shaped, c. 5 mm @, with 5—6 deltoid tepals,

1986]

outer surface with a short dense indumentum; sta-
mens c. 40, sessile, c. 0.75 mm long. — Female recep-
tacle ovoid, c. 1.75 mm long, outer surface with in-
dumentum as in male, inner surface with long simple
bristles; carpels usually 3; style subulate. Fruit not
seen.

Distr. Malesia: Irian Jaya (Nabire; Lake Habbe-
ma; Hellwig Mts); Papua New Guinea (West & East
Sepik and Southern Highlands Prov.).

Ecol. Scrambling in primary rain-forest or over
shrubs in young regrowth, 400—2800 m.

Vern. Oberonk, Mendi.

Note. Although there is considerable variation in
the size of the leaves, correlated with the unusual alti-
tudinal range, the shape of the leaves is rather uni-
form and the smooth, close, but fine indumentum of
the lower leaf surface and of the inflorescence is
characteristic. In the original material collected by
LEDERMANN the fine felt of stellate hairs has partially
or completely disappeared from the lower surface of
some leaves, only a few scattered, slightly larger, stel-
late hairs remaining. However, the typical covering
of hairs has persisted in some areas. The leaves are
described as greyish or whitish beneath, though some
specimens from higher altitudes appear light fawn, at
least when dried. The flowers are cream.

9. Palmeria clemensae PHiLipson, Blumea 28 (1982)
96, f. 2. — Fig. 4.

Woody liane; young branches covered with a ful-
yous or creamy indumentum. Leaves elliptic to
broadly elliptic, with an apiculum (either long and at-
tenuate or short), 7—15 by 3.5—7.3 cm, chartaceous
or slightly coriaceous base truncate or rounded, mid-
rib and principal veins prominent below, sometimes
deeply impressed above, upper surface of mature
leaves with scattered remnants of stellate hairs or gla-
brous, lower surface densely covered with a close felt
of small stellate hairs and with longer soft hairs along
the veins (usually forming a prominent fringe, rarely
almost absent); petiole 5-10 mm, hairy. /n/flores-
cence axillary and terminal, c. 7—20 cm long, covered
with a dense fulvous or creamy indumentum, nar-
rowly paniculate, side-branches few-flowered. —
Male receptacle bow|-shaped becoming almost a flat
disk at anthesis, with 5—7 irregular tepals, c. 8-10
mm @, outer surface with a dense covering of short
stellate hairs, inner surface with short simple hairs
between the stamens; stamens c. 30—45, with broad
filaments up to 0.5 mm long, anthers 0.75—1.5 mm
long; filaments and connectives hairy. — Female re-
ceptacle urceolate, c. 2 mm high at anthesis, outer
surface with indumentum as in the male, inner sur-
face with long simple hairs between the carpels; car-
pels c. 8-12, tapering to filiform style. Immature
Sruit globose with a beak (often asymmetrical); at
maturity the enlarged receptacle splits open to form

MONIMIACEAE (Philipson)

273

c. 5 coriaceous very irregular lobes c. 2 cm long;
drupes subspherical, sessile; mesocarp succulent,
endocarp stony, c. 7 mm long when dry.

Distr. Malesia: Papua New Guinea (Southern
Highlands, Chimbu, Eastern Highlands, Morobe
and Central Provinces).

Ecol. Lower montane and mossy forest (Casta-
nopsis, Lithocarpus, Nothofagus, Podocarpus,
Libocedrus dominated), 1200—2750 m.

Vern. Kari, Chimbu.

Note. A widespread but rather rarely collected
species, distinguished by the close felt of minute stel-
late hairs on the undersurface of the leaves, com-
bined with longer soft bristly hairs along the veins.
The species occurs in two forms. One has rather thin
leaves with a close buff felt on the lower leaf surface,
and with rather few and short bristles, whereas in the
other form the leaf blade is more coriaceous with the
veins deeply impressed above, the felt is paler
(creamy white or fawn), and the bristles form con-
spicuous fringes along the veins. The second form
may also have larger flowers, though the specimens
available are inadequate to establish this conclusive-
ly. The first form is more frequent in the east (Mo-
robe and Central Prov.) and the second in the west
(Eastern and Southern Highlands Proy.), but in both
regions specimens of both forms have been collected.
The black drupes are sessile on a red torus.

10. Palmeria womersleyi PHiLIpsoN, Blumea 28
(1982) 98, f. 3. — Fig. 5.

Woody liane, repeatedly branched, often reaching
the top of medium-sized trees; young branches cov-
ered with a fulvous indumentum. Leaves elliptic to
broadly elliptic, 9-18 by 3.5—10 cm, chartaceous to
coriaceous, base cuneate or rounded, apex broad
with a short obtuse apiculum or more gradually nar-
rowed to an acute apex, midrib slightly channelled
above, prominent below, lateral veins c. 5, arched
and meeting within the margin; upper surface of ma-
ture leaves with widely spaced remnants of stellate
hairs, lower surface with a loose or close felt of stel-
late hairs together with a variable number of larger,
more tufted, stellate hairs especially on the veins;
petiole 8—17 mm long, densely covered with stellate
hairs. Inflorescences axillary and terminal, pleio-
chasial, often produced profusely on short leafy lat-
eral branches, coming to resemble panicles when the
foliage abscisses, covered with a short dense indu-
mentum; pleiochasia 7—15 cm long, bearing opposite
or verticillate short branches (c. 1.5—2 cm) each with
a small number of flowers and minute subulate
bracts mostly caducous before anthesis. — Male re-
ceptacle bow|-shaped, 7-9 mm @ at anthesis (without
the tepals), with 5-7 irregular tepals eventually
opening to disclose the numerous (30—40) stamens,
outer surface with a dense covering of small stellate

274 FLORA MALESIANA [ser. I, vol. 102

Fig. 5. Palmeria womersleyi PHiLieson. a. Habit of male twig, 1/2, b. bud of male flower, c. same at anthe-

sis, both x6, d. anther, x15, e. female flower, f. same in LS, both x6, g. receptacle, bearing achenes, nat.

size, h. portion of lower surface of leaf, x5 (a—c HOOGLAND & SCHODDE 6787, d NGF 14005, e—f PHILIPSON
3721, g HOOGLAND & PULLEN 5429, h PHILIPSON 3690).

1986]

hairs, inner surface with very short simple hairs; an-
thers sessile, c. 0.75—1.25 mm long, the connective
sometimes with short hairs at the base and apex. —
Female receptacle cup-shaped, 2—3 mm @ at an-
thesis, upper surface concave with a small central os-
tiole, outer surface with indumentum as in male, in-
ner surface covered with long simple hairs between
the carpels; carpels c. 10—12, distributed over the in-
ner surface of the receptacle, tapering to filiform
styles (c. 2 mm long) which project through the os-
tiole, becoming reflexed. Immature fruit subspheri-
cal with an unusually asymmetric beak developed
from the tepals; at maturity the enlarged receptacle
ruptures irregularly to form c. 4—6 coriaceous arms
2—2.5 cm long to which the ripe achenes are at-
tached; drupes spherical, sessile, with a shining black
surface, mesocarp succulent, endocarp stony, 7 mm
when dry.

Distr. Malesia: Papua New Guinea (Enga,
Southern Highlands, Western Highlands and East-
ern Highlands Provinces).

Ecol. A liane copiously branched over low
shrubs or climbing high on forest trees in primary or
secondary forest or open scrub, 1800—2600 m.

Uses. Leaves used for smoking (Wabag).

Vern. Kombendegambeka, Hagen, Towopa,
noldunkan, Whaji, Minj, hakappa, Mairi, Wata-
bung, kibekelakkan, Chimbu, Masul.

Note. A moderately large-leaved species which
may be recognized by the close covering of stellate
hairs on the lower leaf surface which are fulvous or
rufous at least in dried specimens. The hairs are of
two sizes: the smaller more numerous and forming a
general felted tomentum; the larger more widely
spaced and giving the leaf surface and especially the
principal veins a tufted appearance. In fresh material
the pubescence on the lower leaf surface appears
either olive green, fulvous or rufous. Flowers fra-
grant, stamens white. Fruit with mature torus green
outside and pink to bright red within; drupes black.

11. Palmeria schoddei Puiipson, Blumea 28 (1982)
100, f. Ic.
Woody liane to 25 m high, with slender branches

MONIMIACEAE (Philipson)

275

covered with a rough fulvous indumentum of stellate
hairs. Leaves ovate, 3.3—6 by 1.5—2.5 cm, charta-
ceous, base rounded, narrowed to an acute apex,
midrib slightly channelled above, prominent below,
lateral veins conspicuous, upper surface of mature
leaves with widely spaced remnants of small stellate
hairs, lower surface covered with dense indumentum
of small fulvous stellate hairs, with scattered slightly
larger stellate hairs along the principal veins; petiole
7-10 mm long, covered in stellate hairs. Jnflo-
rescence: axillary, few-flowered pleiochasia, up to
5.5 cm long, covered with a dense fulvous tomentum
and bearing subulate caducous bracts; pedicels oppo-
site or subopposite, c. 5—~7 mm. — Male receptacle
bowl-shaped, c. 7 mm @ at anthesis (without the tep-
als) with 5 or 6 deltoid tepals, outer surface densely
covered with an uneven stellate indumentum, inner
surface with short simple hairs; stamens 20—25, with
short filaments, anthers c. 0.8— mm long. — Female
flowers not seen. Mature fruits with irregular coria-
ceous receptacular lobes 2 cm long, bearing sessile
drupes 0.8 cm @ when dry.

Distr. Malesia: Papua New Guinea (Southern
Highlands and Western Highlands Provinces).

Ecol. In forest climbing to 8 m or scrambling
over low scrub at the margin of grassland,
2700—2900 m.

Vern. Obe, yaso, Mendi, kena’ugl, Enga, deka-
ruek, Melpa.

Note. Similar to P. montana A.C. SmiTH, a mon-
tane species from the Wharton Range, but distin-
guished by leaf shape and size (larger and lacking an
apiculum); coarser indumentum on the midrib; and
by the larger flowers.

Insufficiently known

Palmeria hypochrysea PERKINS, Bot. Jahrb. 52
(1915) 215.

The species is known only from the type specimen
(LEDERMANN 12404) which was destroyed during
world war II. The description does not appear to fit
any known species.

3. LEVIERIA

Becc. Malesia | (1877) 192; Perkins & Gita, Pfl. R. Heft 4 (1901) 20; PERKINS,
Bot. Jahrb. 52 (1915) 192; Ubersicht Gattungen Monim. (1925) 20; PHILIpson,
Blumea 26 (1980) 373, f. 1-16. — Fig. 6—8.

Trees or shrubs, rarely climbing. Leaves exstipulate, glabrous when mature
or more or less pubescent below, entire or dentate. Dioecious, with terminal or
lateral cymose inflorescences. — Male flowers with a small receptacle bearing
8 rounded tepals enclosing numerous almost sessile stamens; connective project-

276 FLORA MALESIANA [ser. I, vol. 102

Fig. 6. Levieria squarrosa PERKINS. a. Twig of male plant (western form), X1/2, b. male flower, x5, and
stamen, c. twig of female plant (eastern form), 1/2, d. female flower, x4, and in LS, x5, e. fruit, x3/4
(a PULLEN 5227, b Brass 29090, c~d WOMERSLEY & FLOYD 6908).

1986 | MONIMIACEAE (Philipson) 277

ing; anthers opening up by longitudinal slits. — Female flowers + globose with
a small ostiole bounded by 4 irregular tepals. The margin of the receptacle soon
becoming reflexed to expose the ovoid sessile drupes.

Distr. Queensland and Malesia: Celebes, Moluccas (Ceram, Ambon), New Guinea (incl. the Bismarck
Archipelago).

Ecol. Mainly in lower montane rain-forest, between 1200 and 3000 m but descending to sea-level. Frequent
in the shrub layer and lower canopy of rain-forest; persisting in regrowth areas, more rarely on shrubby hill-
sides.

Note. Male plants of the genus may be recognized by the separate rounded tepals borne on a very small
receptacle. Female plants in flower and fruit are distinguished by the developing drupes becoming exposed
by a curling outwards of the receptacle together with the tepals on its margin. This type of fruit development
distinguishes the Hedycaryeae from other tribes of the family. In the Mollinedieae the tepals and the upper
part of the receptacle fall as a calyptra after flowering, leaving a distinct circular abscission scar. In the Moni-
mioideae the receptacle encloses the developing carpels until they are mature.

KEY TO THE SPECIES

1. Undersurface of mature leaf glabrous.

2. Leaves obovate or narrowly elliptic ............

2. Leaves broadly elliptic to elliptic.

Saieafabout:S cmilong, or shorten is: id<nm. a3%i5--
Sel caf about) 7 cm long;:or longer’... 212.506 4%. -

. + Reade cotehe tele eclereeene Ce: 1. L. squarrosa

« fonds bree ee a a 2. L. orientalis
. oe 2h tena an bad eee hee tot 3. L. nitens

1. Undersurface of mature leaf with some indumentum (at least a few hairs at the base of the midrib).
4. Indumentum of lower leaf surface confined to the midrib.

5. Liane with small leaves (3 cm long, or less)....

5. Trees or shrubs, leaves longer than 3 cm.

3. te Aone ce A dc ee ee eee 4. L. scandens

6. Lateral veins clearly defined, blade usually rhombic or obovate ..............--. 1. L. squarrosa

6. Lateral veins indistinct, blade narrowly elliptic

. casita aman PAR eae 5. L. acuminata

4. Indumentum of lower leaf surface extending to the lateral veins (and sometimes to the whole lamina).

7. Leaf with a long narrow apiculum; blade small (usually 7 by 2.5 cm)............. 5. L. acuminata
7. Apiculum, if present, shorter; blade larger (usually more than 8 cm long).
8. Leaf oblong, hairs mostly straight and often sparse............. cee eee eee e eee 6. L. montana
8. Leaf. obovate, hairs crisped and usually copious .... 2.0... 0esecreeeedeeee vee one 7. L. beceariana

1. Levieria squarrosa PERKINS, Bot. Jahrb. 52 (1915)
196; A.C. Smit, J. Arn. Arb. 22 (1941) 232; PHILIP-
son, Blumea 26 (1980) 377, f. 1, 8, 9, 14. — L. parvi-
folia A.C. Smiru, J. Arn. Arb, 22 (1941) 233. — L.
montana (non Brecc.) KANEH. & Hatus. Bot. Mag.
Tokyo 56 (1942) 249, f. 2. — Fig. 6.

Sparsely branched shrub or small tree, occasional-
ly to 18 m; stems more or less densely covered with
minute hairs at least when young, often becoming
glabrous. Leaves chartaceous, obovate, or rhombic,
3-12 by 1—4 cm, in two principal forms with inter-
mediate states frequent: 1) apex rounded or sub-
acute, margin entire, glabrous; 2) with an acute apic-
ulum, upper part of margin dentate, midrib below
with obscure minute appressed hairs (rarely with
dense but very short crisped hairs); midrib promi-
nent, lateral veins arched and meeting within the
margin; petiole 4—8 mm long, glabrous or minutely
hairy. Dioecious. — Male inflorescence either small
lateral pleiochasia (up to c. 4 cm long) or terminal

pleiochasia or panicles (up to c. 8 cm long); rachis
and branches of panicles with a few pairs of pedicels
(c. 8 mm long) and a terminal flower, minutely
hairy, and with pairs of small lanceolate bracts be-
low the pedicels and sometimes along the peduncle
and rachis. — Male flowers more or less globose, 7
mm @; receptacle small, bearing 8 tepals; 4 outer tep-
als rotund, subcoriaceous, with sparse hairs on the
outer surface; 4 inner smaller, narrower, membrana-
ceous, glabrous. Stamens numerous (c, 25 or more);
connective acuminate; anthers subsessile, loculi lat-
eral, separate. — Female flowers lateral or terminal,
solitary or in few-flowered racemes (c. 4 cm long);
receptacle coriaceous, broadly cupuliform (5-8 mm
9) with an irregularly cleft mouth (4 tepals may be
more or less clearly defined); carpels numerous (ec, 25
or more), c. 2 mm long, densely packed over the re-
ceptacle, obovoid, truncated distally and bearing a
short style. Drupes numerous, ovoid, apex acute,
glabrous, c. 9 by 5 mm when dry, sessile on a re-

278

FLORA MALESIANA

[ser. I, vol. 102

curved leathery pilose receptacle.

Distr. Malesia: New Guinea (Vogelkop Penin-
sula to Owen Stanley Range).

Ecol. A laxly branched shrub in the undergrowth
of rain-forest, sometimes semi-scandent, or a lower-
storey tree up to 18 m high with rather dense leafy
branches. Above 1800 m, ascending to 3000 m (rarely
descending to 250 m). Montane and mossy forests of
varying composition (Podocarpus, Libocedrus, No-
thofagus, Castanopsis) and also in regrowth and
marginal forest.

Vern. Ang, Hindenburg Ra., eberak, mungo-
munk, tantan, Mendi, lekiem, Kapauku, kalipka,
main-gobugont, Minj, kammakam, kamokam, En-
ga, kip, kunguma, Togoba, kolkola, Eipomek, kom-
bo, Onim, timonksagu, Wabag.

Notes. The bark is grey-brown, smooth with
very shallow fissures; inner bark pinkish brown. The
wood is heavy, light brown with wide, pale rays.
Flowers yellowish green, drupes orange to reddish
black on yellow receptacle.

The leaf shape is extremely variable, at first sug-
gesting that two species are involved. Obtuse, entire,
glabrous leaves occur in the eastern portion of New
Guinea, whereas apiculate, dentate and obscurely to-
mentulose leaves occur towards the west. There is,
however, a considerable area of overlap of the two
forms (mainly between 144° E and 146° E) and in
this region leaves with intermediate characters are
abundant. Thus, although all specimens from the
two ends of New Guinea are quite distinct in several
characters, these two forms are connected by a broad
zone in which it is impossible to separate them.

2. Levieria orientalis PHILIPSON, Blumea 26 (1980)
SRD

Tree to 10 m, shoots glabrous except for the
youngest parts. Leaves glabrous, chartaceous,
broadly elliptic or elliptic, c. 5 by 2.5 cm, broadly cu-
neate at the base, rounded apex produced in a short
obtuse apiculum; margin with few widely spaced
teeth; midrib prominent below; lateral veins rather
obscure, arched and meeting within the margin; pet-
iole 5—8 mm. Probably dioecious. Male flowers not
seen. Inflorescence of terminal and lateral few-
flowered cymes or flowers solitary. — Female
flowers with a glabrous, coriaceous, globose recep-
tacle, c. 3 mm g, the ostiole small with obscure tepal-
lobes; carpels c. 20, densely packed over the re-
ceptacle, ovoid with a short style. Drupes ovoid, c. 8
by 6 mm, sessile on a reflexed sparsely hairy recep-
tacle.

Distr. Malesia: Papua New Guinea (Milne Bay
Prov.: Goodenough I.).

Ecol. A much branched tree, 8—10 m, occurring
at the edge of forest at 1500 m.

Note. Drupes black on a yellow receptacle. The

glabrous, broad elliptic leaves suggest L. nitens, but
are much smaller, and their short apiculum is obtuse.

3. Levieria nitens PERKINS in Perkins & Gilg, Pfl. R.
Heft 4 (1901) 21, f. 3r—s; Pfl. R. Heft 49 (1911) 7, f.
3R-—S; Bot. Jahrb. 52 (1915) 197, f. 3, 15; A.C.
SmitH, J. Arn. Arb. 22 (1941) 232; Puitirson, Blu-
mea 26 (1980) 379. — Steganthera schlechteri PER-
KINS, Pfl. R. Heft 49 (1911) 2 (non L. schlechteri
PeRKINS). — L. rudolfii PERKINS, Bot. Jahrb. 52
(1915) 196. — Steganthera elliptica A.C. Smiru, J.
Arn. Arb. 22 (1941) 236.

Moderate to large tree, reaching 45 m; bole 25 m,
buttressed, dbh 1 m, ora shrub sometimes with semi-
scandent branches; young shoots with appressed,
often golden hairs. Leaves glabrous, chartaceous, el-
liptic, broadly elliptic or oblong, 7—17 by 2.5—6 cm;
base cuneate or attenuate; apex produced as an apic-
ulum; margin entire or occasionally with a few small
teeth; midrib prominent; lateral veins usually indis-
tinct, arched and meeting within the margin; petiole
1—1.5 cm. Dioecious. /nflorescence of lateral or ter-
minal pleiochasia or usually finely branched pan-
icles, minutely hairy, c. 10—15 cm long; rachis bear-
ing pairs or whorls of branches subtended by small
bracts. — Male flowers globose, 3—4 mm g; recep-
tacle small; tepals 8, rounded; stamens numerous,
connective projecting, anthers sessile. — Female
flowers with a coriaceous cupuliform receptacle, c. 4
mm @, with an irregularly cleft ostiole; carpels nu-
merous, densely packed over the receptacle, ovoid
with short style. Drupes numerous, ovoid, 8—10 by
4—5 mm (when dry), sessile on a leathery, pilose re-
ceptacle with a reflexed margin.

Distr. Malesia: New Guinea (Vogelkop Penin-
sula to Huon Peninsula and Tufi).

Ecol. A tree to 45 m, though normally smaller
and even shrubby; the branches sometimes sprawling
or semi-scandent. Montane and mossy forest, asso-
ciated with Nothofagus, Castanopsis, Lithocarpus,
etc., also in regrowth and on scrubby slopes. Usually
500—1800 m, but occasionally descending almost to
sea-level.

Notes. Bark smooth or becoming fissured, olive-
or grey-brown; under bark pinkish. Wood soft,
straw coloured, with large rays. Flowers greenish
cream, drupes orange to black on yellow or orange
receptacles.

The mature leaves are entirely glabrous, even at
the base of the midrib, a character found in other Le-
vieria species only in L. orientalis (which has smaller
leaves and is confined to Goodenough I.) and in
some specimens of L. squarrosa (which has leaves of
a different shape). The lamina of L. nitens is broadly
elliptic and apiculate, whereas that of L. squarrosa is
narrowly obovate or elliptic and usually obtuse and
not apiculate (in glabrous forms). Confusion is more

1986 |

likely to occur between L. nitens and L. montana. In
the latter species the leaf shape is often very similar
to that of L. nitens and the inconspicuous hairs of the
leaf may be confined to the underside of the midrib
and may be largely lost by abrasion.

4. Levieria scandens PHiLtipson, Blumea 26 (1980)
S815 f2 4, 10.

A woody liane, lateral shoots c. 1.5 mm 9, when
young densely covered with minute appressed hairs.
Leaves closely set along the branches; chartaceous,
lanceolate to lanceolate-ovate, 2.5—3 by 0.8—1 cm,
rounded or cuneate at the base, narrowed to a mu-
cronulate apex; margin subrevolute with few irregu-
lar teeth; midrib prominent below, with a few ob-
scure hairs on its lower part; lateral veins obscure;
petiole c. 2 mm long, bearing minute hairs. Probably
dioecious, female flowers not seen. /nflorescence
usually of simple dichasia, minutely hairy, terminat-
ing the lateral branches and in their upper axils. —
Male flowers globose, 2 mm @ (in bud); tepals 8,
rounded, borne on a small receptacle; stamens c. 20,
connective projecting, obtuse, anthers sessile.

Distr. Malesia: Papua New Guinea (West Sepik
Prov., Telefomin Subprov.).

Ecol. Liane in montane forest, at 2100 m.

Note. The small lanceolate-ovate leaves are un-
matched in any other species of the genus. Other spe-
cies occasionally show straggling growth, but this
species appears to be a true liane.

5. Levieria acuminata (F. v. M.) PERKINS, Bot.
Jahrb. 25 (1898) 570; Puitipson, Blumea 26 (1980)
382, f. 5, 11. — Mollinedia ? acuminata F. v. M.
Fragm. 5 (1866) 155.

Small tree to 15 m high; young shoots minutely pu-
bescent. Leaves elliptic, c. 7-10 by 2—4 cm, mem-
branous; base cuneate or attenuate; apex prolonged
aS a narrow acute apiculum; margin entire or with
small teeth; midrib prominent; lateral veins indis-
tinct, arched and meeting within the margin; midrib
bearing inconspicuous hairs, or the whole lower sur-
face finely tomentose; petiole 10-15 mm, usually
glabrous. Dioecious. Inflorescence of terminal and
axillary pleiochasia or panicles, the male being more
finely branched and usually larger than the female,
pubescent, rachis bearing pairs of small bracts. —
Male flowers globose, 3—4 mm @; tepals c. 8, the
outer broad and rounded; stamens numerous (c.
20—40), crowded on a small receptacle, connective
projecting, reflexed, obtuse, anthers sessile. — Fe-
male flowers with a coriaceous, globose receptacle,
c. 3mm @, the ostiole with 4 irregularly cleft tepals;
carpels numerous, densely packed over the recep-
tacle, ovoid with a short style; drupes ovoid, sessile

“on a reflexed receptacle.
Distr. Australia (N. Queensland) and Malesia:

MONIMIACEAE (Philipson)

m9

Papua New Guinea (Central and Morobe Prov.).

Ecol. Small tree to 15 m, in primary rain-forest
or second growth, 1200—3000 m, but descending to
near sea-level in Queensland. The cream or greenish
flowers are fragrant.

Note. This extension of the range of a Queens-
land species is based on five collections made by
Carr in 1935—1936 and two collections by SCHODDE
& CRAVEN in 1966. While some collections match
Queensland specimens closely, others differ in the
greater extent of the pubescence on the lower leaf
surface.

6. Levieria montana Becc. Malesia | (1877) 193;
PERKINS & Gia, Pfl. R. Heft 4 (1901) 21; PERKINs,
ibid. Heft 49 (1911) 9; Bot. Jahrb. 52 (1915) 197;
PHILIPSON, Blumea 26 (1980) 382, f. 6, 12. — L.
schlechteri Perkins, Pfl. R. Heft 49 (1911) 7. — L.
urophylla PERKINS, Bot. Jahrb. 52 (1915) 93, f. 1. —
L. laxiflora PERKINS, /.c. 195.

Shrub, sometimes semi-scandent, or small tree to
15 m high; young shoots densely covered with minute
appressed hairs. Leaves membranaceous, broadly to
narrowly oblong or elliptic, 4.5—13 by 1.5—6 cm;
base acute or obtuse; apex narrowed more or less
abruptly into an acute or obtuse apiculum; margin
entire or occasionally with a few small teeth; midrib
prominent; lateral veins arched and meeting within
the margin; the midrib below bearing minute crisped
or straight hairs, which usually extend on to the later-
al veins and occasionally on to the entire lower sur-
face of the lamina; petiole 6—14 mm. Dioecious. —
Male inflorescences axillary and terminal, panicu-
late, 1O—15 cm long, densely and minutely hairy; the
rachis bearing pairs of pleiochasial branches
subtended by small bracts. — Male flowers globose,
4—7 mm a, receptacle small; tepals 7—8, outer
rounded, subcoriaceous, hairy, inner smaller, mem-
branaceous; stamens 25—50, connective projecting,
anthers subsessile. — Female inflorescence similar to
male or simple axillary and terminal racemes. — Fe-
male flowers with a coriaceous, globular, cupuliform
receptacle, 5—8 mm 9, aperture irregularly cleft; car-
pels numerous, densely packed over the receptacle,
ovoid with a short style. Drupes numerous, ovoid, 8
by 5mm when dry, sessile on a leathery, pilose recep-
tacle with a reflexed margin.

Distr. Malesia: Celebes, Moluccas (Ceram Am-
bon), New Guinea (Biak, Vogelkop, NW. Irian, Se-
pik Prov., Chimbu Prov., Finisterre Range, Huon
Peninsula, Wagau-Garaina region of Morobe
Prov.).

Ecol. A slender shrub, sometimes semi-scandent
or epiphytic, or a small tree with drooping branches.
In forest or scrub, often in lower montane rain-forest,
1200-1900 m, but in the western part of its range
(NW. Irian, Biak, Moluccas) also near sea-level.

280 FLORA MALESIANA [ser. I, vol. 102

Fig. 7. Levieria beccariana PERKINS. a. Habit of male plant, x 1/2, b. male flower, from side, x3, and from
top, x4, c. twig of female plant, x1/2, d. female flowers after anthesis, x1 1/4, e. fruits, from behind and
side, nat. size (a FisHER 83, b PULLEN 7910, c NGF 23781, d—e Ftoyp & HooGLanp 3997).

1986 |

MONIMIACEAE (Philipson) 281

Uses. In the southern foothills of the Finisterre
Range shrubs are preserved by villagers because the
conspicuous pendulous fruits attract birds which are
then shot from cover.

Vern. Dandanggu, Naho lang., hoppetu, Wagu,
mardichber, mardieber, mardiewer, Biak, woisingai,
Wasuk.

Notes. The bark is smooth grey-brown or fawn.
The flowers are light yellow and the mature drupes
are bright orange or deep purple, borne on firm
fleshy yellow or orange-brown receptacles. The leaf
is characteristically oblong-elliptic and apiculate,
with short crisped or appressed hairs along the lower
surface of the midrib, and sometimes also on the lat-
eral veins, or even on the lower surface of the lamina.
The degree of hairiness is variable, being most
strongly developed on the type from the Arfak Mts.
Size of the lamina and the degree of dentation of its
margin are also variable. Leaves of juvenile shrubs
are larger, broader, thinner in texture, and more den-
tate than those of adults.

PERKINS (1915) did not have the type of L. mon-
tana Becc. available when describing L. urophylla
and was herself doubtful of the distinctness of this
species. Unaccountably, she regarded L. montana as

glabrous, although the original description clearly re-
fers to the indumentum of the lower surface of the
leaf. There has been some confusion also concerning
the use of the name L. schlechteri PERKINS. The type
specimen is given as SCHLECHTER 17176, and is de-
scribed as having glabrous leaves when mature.
However, the labels of this number are named L.
laxiflora PERKINS and the midribs bear minute hairs
(in some specimens rather severely abraided). Levie-
ria schlechteri, therefore, is here regarded as synony-
mous with L. montana.
7. Levieria beccariana PERKINS in Perkins & Gilg,
Pfl. R. Heft 4 (1901) 21; ibid. Heft 49 (1911) 8, f.
3P—Q; Bot. Jahrb. 52 (1915) 193; PHirpson, Blu-
mea 26 (1980) 383, f. 7, 13, 16. — L. forbesii PER-
KINS, Pfl. R. Heft 49 (1911) 7, f. 3L—O. — Fig. 7, 8.
Tree to 20 m, or shrub, sometimes semi-scandent;
young shoots densely covered with minute hairs.
Leaves chartaceous, broadly to narrowly obovate or
elliptic, 6—15 by 2.6—7 cm; base cuneate; apex usual-
ly acute apiculate, sometimes obtuse; margin entire
or occasionally dentate in its upper part; midrib
prominent; lateral veins approaching the margin, the
lower veins often ascending well beyond the middle

Fig. 8. Levieria beccariana Perkins. In fruit. Papua New Guinea (Photogr. PHILIpson).

282

FLORA MALESIANA

[ser. I, vol. 102

of the blade; whole undersurface often russet or
fawn, more or less densely covered in short curled or
rarely lax hairs, or these confined to the principal
veins; petiole 8—15 mm long, pubescent or glabres-
cent. Dioecious. /nflorescence of terminal and axil-
lary panicles, those in the upper axils often combin-
ing to form a massive compound leafy panicle, the
male rather more finely branched than the female;
rachis bearing pairs of pleiochasial branches sub-
tended by small bracts, tomentose. — Male flowers
globose, 4—6 mm g; receptacle small; tepals round-
ed; stamens numerous, connective projecting, an-
thers sessile. — Female flowers with a coriaceous cu-
puliform receptacle 5-8 mm g, with 4 irregularly
cleft tepals; carpels numerous (c. 20—100), densely
packed over the receptacle, ovoid with a short style.
Drupes numerous, ovoid, 8 by 5 mm when dry, ses-
sile on a leathery pilose receptacle with a reflexed
margin.

Distr. Malesia: Papua New Guinea (on the cen-
tral ranges from the Telefomin area to Milne Bay).
This species does not overlap the range of L. monta-
na except in the Wagau-Garaina area.

Ecol. Tree to 20 m, with a clear bole 30 cm @ and
pendulous branches, or a slender, sometimes semi-
scandent shrub, occurring in montane and mossy
forest of varying composition (Nothofagus, Casta-

nopsis, Araucaria); also in regrowth, 1200—2800 m.

Vern. Dangrumon, Chimbu, gokey, Wagau, ho-
ma, Hademari, kiangap, komali, Enga, kombo kom-
bo, Upper Kangel, /upulupu, Sirunki, matam-
matam, Managalese, tangitang, Tomba, tsuk, Por-
get, umgude, Oksapmin.

Notes. Bark smooth with very small fissures,
greyish or reddish brown; inner bark pinkish, aro-
matic. Wood brittle, cream-brown, with prominent
rays. Flowers greenish yellow, drupes orange to
black on a yellow receptacle. The leaves are charac-
teristically obovate with the lower veins ascending
for much of the length of the lamina, but elliptical
leaves and more spreading lateral veins are not infre-
quent. The midrib and lateral veins are usually dense-
ly clothed with minute crisped, often russet or fawn
tomentum, and this frequently covers (if only sparse-
ly) the whole lamina. These hairs may be short and
crisp as in the type, or longer and more lax (as in the
type of L. forbesii), but these differences are not con-
sidered of specific importance. One or two collec-
tions in which the tomentum is practically confined
to the midrib approach rather closely to L. montana
Becc. Since these specimens are from the region
where the ranges of the two species meet (the Wagau-
Garaina region of Morobe Prov.) it is probable that
some admixture of the two species occurs there.

4. WILKIEA

F.v.M. Trans. Phil. Inst. Vict. 2 (1858) 64; PERKINS, Bot. Jahrb. 25 (1898) 569,
t. 6D—E; Perkins & GILG, Pfl. R. Heft 4 (1901) 57, f. 14—15; Perkins, ibid.
Heft 49 (1911) 26; Ubersicht Gattungen Monim. (1925) 37; Puiipson, Blumea
26 (1980) 365, f. 1-5. — Fig. 9.

Shrubs or trees with opposite or verticillate, entire or serrate /eaves. — Male
receptacle + globose with small tepals surrounding an ostiole; stamens up to c.
30 irregularly spaced over the receptacle; anthers with a single horizontal or
horseshoe-shaped slit. — Female receptacle + globose with tepals surrounding
a small ostiole, thickened glands within the ostiole; carpels numerous (up to c.
100), sessile on the lower half of the receptacle, style subulate; upper half of the
receptacle becoming detached by a circular scar. Drupes sessile or shortly stipi-
tate.

Distr. Eastern Australia and Malesia: E. Papua New Guinea.

Ecol. In Malesia in montane rain-forest.

Note. One species occurs in Papua New Guinea, the genus otherwise being confined to Queensland and
New South Wales. For separation of Wilkiea from Kibara see that genus.

1. Wilkiea foremanii PHitipson, Blumea 26 (1980)
365, f. 1-5. — Fig. 9.

Tree, densely hairy in all its parts. Leaves elliptic
to elliptic-oblong, base rounded, apex rounded with

an apiculum to attenuated, margin entire, midrib
prominent, main lateral veins c. 4—6 pairs, arched-
ascending; petiole 7—10 mm. Flowers axillary, singly
or in few-flowered cymes; peduncles (or pedicels)

1986]

MONIMIACEAE (Philipson)

283

Fig. 9. Wilkiea foremanii PuiLieson. a. Twig with female flowers, at time of abscission of calyptra, 1/2,
b. calyptra, seen from above, 2, c. carpel with filiform style, x5, d. male flower, in LS, after dehiscence
of anthers, x5, e. stamen, x12 1/2 (a—c NGF 48404, d—e FropIn 672).

stout, up to 4 cm long, often with the scars of two
bracts below the flower. — Male flowers globose
with an ostiole surrounded by 6 tepals (4+ 2), c. 7
mm @ at anthesis; stamens c. 18—30, inserted over the
inner surface of the receptacle, filaments c. 1 mm
long, anther triangular, c. 1 mm long. — Female
flowers globose, c. 10-15 mm @ at anthesis, leathery,
hairy within, tepals 4, the upper half of the receptacle
becoming detached after anthesis by a circular scar;

carpels numerous (c. 75-100), sessile on the lower
half of the receptacle, 1.5 mm long, pubescent; style
slender, 5 mm. Ripe fruit unknown.

Distr. Malesia: Papua New Guinea (Central
Prov., Goilala Distr.).

Ecol. A small tree with light green, very hairy
leaves (fulvous when dry), in mid-montane forest or
regrowth, 2400—2600 m.

284 FLORA MALESIANA [ser. I, vol. 102

Fig. 10. Range of distribution of six monotypic genera of Mollinedieae. |. Tetrasynandra; 2. Austromatthaea;
3. Kairoa; 4. Lauterbachia; 5. Faika; 6. Parakibara.

5. FAIKA

PHILIPSON, Blumea 30 (1985) 417, f. 1—2. — Fig. 11, 12.

Small tree with villose young branches and leaves. Flowers solitary in the leaf
axils. ?7Dioecious. — Receptacle of male flower ovoid, with a large ostiole sur-
rounded by 3 decussate pairs of tepals. Stamens c. 24, inserted over the inner
surface of the receptacle, anthers opening by two vertical slits; filament short.
— Receptacle of female flower turbinate, with c. 5 decussate pairs of tepals (the
inner swollen and glandular) and with 2—3 pairs of bracts at its base or on the
lower outer surface, the upper part abscissing as a calyptra after anthesis, inner
surface setulose. Carpe/s numerous, sessile, with a long subulate style stigma.

Distr. Malesia: W. New Guinea (Vogelkop Peninsula to the Cyclops Mts). Monotypic. Fig. 10.
Ecol. In rain-forest.

1. Faika villosa (KANEH. & HaAtTus.) PHmipson, Blu- Leaves oblong-elliptic, 24—44 by 6.5—13 cm, charta-
mea 30 (1985) 420. — Steganthera villosa KANEH. & — ceous, base cordate, apex apiculate, acute, bristly
Hatus. Bot. Mag. Tokyo 50 (1942) 259, f. 7. — Fig. | along the veins and on both surfaces, but the upper
LiF 2: surface becoming + glabrous, except for the midrib;

Shrub or small tree, to 3 m; shoots densely villose. | midrib, lateral veins and reticulation prominently

1986 | MONIMIACEAE (Philipson)

Fig. 11. Faika villosa (KANeH. & HATUS.) PHILIPsON. a. Habit,

~0.6, b. young infructescence,
in LS, *1.6 (KANEHIRA & HATUSIMA 13975),

x0.8, c. seed

286 FLORA

MALESIANA

[ser. I, vol. 102

Fig. 12. Faika villosa (KANEH. & HATUS.) PHILIPSON.

a. Female flower, b. the same in LS, c. male flower,

d. the same in LS, all x3 1/4, e. stamen, x10 (a—b

VAN ROYEN & SLEUMER 6082, c—d VAN ROYEN &
SLEUMER 5729).

raised on the lower surface, impressed on the upper
surface; petiole c. S—6 mm, villose. Inflorescence
axillary, flowers solitary, + sessile. — Male flowers
obovoid, c. 55 mm long, glabrous, with 1—2 small
triangular bracts usually near the base; tepals 6, tri-
angular; stamens numerous (c.24) distributed irregu-
larly over the inner surface of the receptacle, fila-
ment terete, connective separating the two anthers;
anthers c. 1 mm long, each locule opening by a ver-
tical slit. — Female flowers turbinate, c. 7 by 8 mm,
glabrous, with 2—3 pairs of small deltoid bracts at or
above the base; tepals c. 5 decussate pairs, the inner
3 pairs swollen and glandular, the inner surface setu-
lose; carpels numerous, c. 3 mm long, sessile, with a
long subulate style and stigma. /nfructescence to c. 3
cm @, woody, setulose, margin revolute; drupes
ovoid, c. 15 by 10 mm, sessile, hirsute.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula, Wandammen Peninsula, Hollandia, Cy-
clops Mts). Fig. 10.

Ecol. Undergrowth
100—1250 m.

Note. The villose stems and leaves are conspicu-
ous. The older branches are smooth with pale grey-
brown bark. The male flowers are yellowish, the fe-
male flowers orange, and the ripe drupes black.

in primary rain-forest,

6. PARAKIBARA

PuiLipson, Blumea 30 (1985) 421, f. 3. — Fig. 13.

Small tree with opposite dentate leaves. Inflorescence an axillary fascicle.
?Dioecious. — Receptacle of male flowers obovoid, tepals 4; stamens c. 18, in
decussate pairs inserted near the base of the receptacle; anthers opening by a
single slit. — Female flowers and fruits not known.

Distr. Malesia: Moluccas (Halmahera). Monotypic. Fig. 10.

Note. The precise relationships of this genus cannot be determined in the absence of female flowers, but
it cannot be doubted that it is a member of the Mollinedieae. The large and numerous stamens arranged in
four regular and closely packed files are very distinctive.

1. Parakibara clavigera PHILIpson, Blumea 30 (1985)
421, f. 3. — Fig. 13.

Small tree, shoots glabrous when mature. Leaves
oblong-elliptic, c. 180 by 80 mm, stiffly chartaceous,
base broadly cuneate, apex obtuse or shortly apicu-
late, margin irregularly dentate in the upper part,
midrib and lateral veins prominent; petiole 10—12
mm. ?Dioecious. /nflorescence axillary, a condensed
cyme forming a fascicle of a few flowers arising from
a short peduncle; peduncle 2—3 mm long, with close-
ly imbricated bracts, sparsely pubescent; pedicels c.

20 mm long, slender below but thickening towards the
flower, sparingly pubescent. — Male flower ovoid, c.
9.5 by 6mm; tepals 4, obtuse; stamens 3 mm long, c.
18 in decussate pairs forming 4 rows inserted on the
lower part of the receptacle, filaments broad and
fleshy, c. 2 mm long, sparingly pubescent, anthers
erect, c. 1 mm long, triangular, opening by a single A-
shaped slit. Female flower and fruit not seen.

Distr. Malesia: Moluccas (Halmahera: Pasir Pu-
tih). Fig. 10.

Vern. O morihuhaka.

1986] MONIMIACEAE (Philipson) 287

Fig. 13. Parakibara clavigera Puitieson. a. Twig with male inflorescence, 1/2, b. male flower, x4, c. the
same in LS, x5, d. stamen, *8 (TAYLOR 2835 A).

7. KIBARA

ENnpL. Gen. PI. (1837) 314; Perkins, Bot. Jahrb. 25 (1898) 570; Pfl. R. Heft 4
(1901) 58; ibid. Heft 49 (1911) 28; Bot. Jahrb. 52 (1915) 207; Puitipson, Blumea
30 (1985) 389, f. 1. — Brongniartia BL. Bijdr. 9 (1825) 423, non KUNTH. — Scia-
docarpus Hassk. Flora 25, ii (1842) Beibl. 1: 20. — Sarcodiscus Grirr. Notul.
4 (1854) 380. — Fig. 14—16.

_ Trees or shrubs, resting buds with cataphylls. Leaves usually pubescent at
first, often becoming glabrous, entire or dentate, principal secondary veins

288 FLORA MALESIANA [ser. I, vol. 102

Fig. 14. Inflorescence types in Kibara. a. Fasciculate in K. oblongata Puiipson, b. condensed cyme in K. mo-
/uccana PERKINS, c. open paniculate cyme in K. macrantha PHILIPSON (a LAE 68853, b Eyma 2900, c HARTLEY
13691).

arched and meeting within the margin. Monoecious. /nflorescence lateral or ter-
minal, cymose (racemose in K. streimannil), often pleiochasial, paniculate or
fasciculate; pedicels usually thickening distally into the receptacle. — Male
flowers usually smaller than the females, with a minute ostiole surrounded by
2—4 decussate pairs of tepals; androecium usually with 4 large outer stamens
and up to 4 inner smaller stamens or staminodes which may be rudimentary and
+ connate, occasionally 5 stamens in the outer whorl or as few as 2 stamens
present; anthers opening by a single slit, with a filament or subsessile. — Female
flowers with the ostiole surrounded by about 5 decussate pairs of tepals; the in-
ner pairs thickened and glandular; the upper half of the receptacle abscissing as
a calyptra after anthesis; carpels numerous, stigma obtuse sessile on the apex of
the ovary. Drupes sessile or stipitate.

Distr. Peninsular Thailand and Nicobar Is. to Queensland; in Malesia: throughout the area. In all 43 spp.,
of which 39 in Malesia.

Ecol. Mostly understorey shrubs and small trees in rain-forest from sea-level to c. 2800 m; occasionally
on sandy or coral beaches.

Notes. Kibara is readily separated from Steganthera and Matthaea by the swollen glandular tissue which
surrounds the inner rim of the ostiole (ENDREss, Pl. Syst. Evol. 134, 1980, 79-120). Wilkiea has female re-
ceptacles very similar to those of Kibara, but these two genera can be distinguished by their male flowers. In
most species the androecium of the two genera is distinctive: in Kibara there is a symmetrical group of four
stamens (2 decussate pairs), within which there is unusually a group of up to four smaller stamens or often
apparently infertile staminodes. In most Wi/kiea spp. the numerous stamens (c. 8 or more) are inserted irregu-
larly over the inner surface of the receptacle. Some species of Kibara occasionally may have 5 stamens in the
outer group, or these may be reduced to 3 or even 2, in which case the inner reduced stamens are often absent.
On the other hand, some individual flowers of Wi/kiea have relatively few stamens (as few as 6 in W. huegelia-
na and as few as 4 in W. macrophylla (see ENDREsS, /.c.). In this event the genera are distinguished by the
irregular insertion of the stamens in Wilkiea and the decussate arrangement in Kibara. The only species in
which some doubt may occur is K. rigidifolia in which individual flowers with 3 or 5 stamens may not have
them clearly arranged in a regular manner. When 6 stamens are present, the decussate arrangement is more
definite. The nature of the stigma also serves to separate these two genera. In all species of Kibara the stigma
forms an obtuse cushion or knob, whereas in Wi/kiea it is more elongated and acute, being often subulate.

1986 | MOoNIMIACEAE (Philipson) 289

KEY TO THE SPECIES

In view of the large number of species, many incompletely known, and the paucity of well defined charac-
ters, it is recommended that more reliance than usual is placed on distribution when identifying specimens.
Most species are local or regional, and while their ranges may not be fully known, they are unlikely to occur
far from their known localities. It must also be borne in mind that several species are not included in this key
because they are inadequately known. In addition to those listed at the end of the genus, 39. K. syimplocoides
is omitted from the key because the nature of its inflorescences is uncertain.

aemmecemriear (eal Index 6) .4 33 fet lek Re Fe ee A et. Sees 1. K. roemeri
1. Leaves broader (leaf index less than 4).
2. Leaves shorter than 5 cm.
3. Leaves coriaceous, apex obtuse or retuse (Vogelkop Peninsula) ................ 2. K. oligocarpella
3. Leaves membranaceous, tapering to a long apiculum (Sepik region) .............. 3. K. myrtoides
2. Leaves longer than 5 cm.
4. Inflorescence shorter than, or little exceeding the petiole (c. 2 cm or less).
5. Stems greatly expanded at the nodes (usually inhabited by ants).
6. Leaves sessile, amplexicaul, margin strongly and sharply dentate (E. Papua New Guinea) 4. K. ferox
6. Leaves petiolate, base cuneate or truncate, margin entire.
Pereaves proadly ovate (Moluccas)! 22 5 Bae 5. Ore, SO en eee 5. K. latifolia
7. Leaves elliptic-oblong (S. Papua New Guinea) ......2...c0c82.c0s cdceeks 6. K. archboldiana
5. Stems not markedly expanded at the nodes.
8. Inflorescence pubescent (young shoots also pubescent and hairs + persisting on mature foliage).
9. Leaves stiffly chartaceous, teeth sharply spinulose, often + bullate (Western Highlands Prov.)
7. K. karengana
9. Leaves softly chartaceous, dentations not spinulose.
eiaweaves bullate (Morobe,Prov:)) . 2 .ccece t OP bi eere State ee eee eae eee 8. K. bullata
10. Leaves flat.
11. Female receptacle and pedicel bearing several pairs of acute imbricating bracts (Moluccas)
9. K. kostermansii
11. Female receptacle and pedicel without bracts (or at most small bracts widely spaced) (New

Guinea).
12. Pubescence of fine, pale, appressed hairs (SE. Papua New Guinea).......... 10. K. leachii
12. Pubescence of loose, curled, brownish hairs (Vogelkop Peninsula) ....... 11. K. versteeghii

8. Inflorescence and mature foliage glabrous (buds and young foliage sometimes pubescent).
13. Leaf blade larger than 10 cm.

beseat margin entire: (Normanby'I.): 20S hii alee sates aoe leas centre erates 12. K. oblongata
14. Leaf margin dentate.
15. Leaf margin spinulose-dentate (Vogelkop Peninsula).................... 13. K. warenensis
15. Leaf margin serrate.
16. Leaf base tapering to the petiole, blade narrowly obovate (Rossel I.) ..... 14. K. rosselensis
16. Leaf base broadly cuneate, blade oblong-elliptic (Vogelkop Peninsula)....... 15. K. royenii
13. Leaf blade shorter than 8 cm.
17. Leaf apex with a slender apiculum (Eastern Highlands Prov.)................ 16. K. hartleyi

17. Leaf apex acute or obtuse, without an apiculum (Mt Shungol, Morobe Prov.) 17. K. shungolensis
4. Inflorescence considerably longer than the petiole (3 cm or usually much longer).
18. Inflorescence with the lateral branches crowded on a short peduncle .......... 18. K. moluccana
18. Inflorescence a simple or branched cyme with evident internodes separating the branches.
19. Leaf blade small, membranaceous (usually under 10 by 3 cm). Inflorescence branches very slender

CUR UICC IL WA AP Ge Gee eat MeUITe eta ea ahs Kb BUG eg hats cle bee cleats 19. K. microphylla
19. Leaf blade larger (usually over 10 cm long, if not, then coriaceous).
20. Male flowers racemosely arranged along the inflorescence branches......... 20. K. streimannii

20. Male flowers cymosely arranged or solitary.
21. Flowers 6 mm long at anthesis or longer, densely pubescent (Eastern Highlands & Morobe Prov.)
21. K. macrantha
. 21. Flowers less than 5 mm long (or if 6 mm, then glabrous).
22. Leaf margin dentate.

290 FLORA MALESIANA [ser. I, vol. 102

23. Stems greatly expanded at the nodes (usually inhabited by ants) (Central Prov.) 22. K. carrii
23. Stems not markedly expanded at the nodes.
24. Dentations) of, leafmarein: sharply/spinuloses. 2. . so aaeee fe terelensieeiee 23. K. elongata
24. Dentations of leaf margin not spinulose.
25. Pedicels strong.
26. Underside of mature leaf pubescent. Inflorescence a few-flowered cyme (Southern Highlands
SafilnevBaveProvayi wet: ios ee eee ee eer Boe en eee 24. K. katikii
26. Underside of mature leaf glabrous or, if pubescent, inflorescence + paniculate.
27. Leaf blade usually broadly elliptic-ovate, more than 10 cm long, rounded to broadly cuneate

bases eedicelsimpid) (throughout Malesia) es ee eee eee 25. K. coriacea
27. Leaf blade usually elliptic, usually less than 10 cm long, base cuneate. Pedicels less rigid (SE.
PapuarNew Ginea) i: oleae nes Since ote. ee eke ats ee 26. K. papuana

25. Pedicels delicate.
28. Inflorescence of simple pleiochasia. Outer stamens inserted at widest part of the receptacle

(Star Miss Nelefominwnesion) aeraaeaccds carlaac cays oe eee eee 27. K. nitens
28. Inflorescence of paniculate cymes. Stamens inserted near base of the receptacle (NW. Irian
AV AN Taare. (citar, ecu daleeara ats Cea eke eens nL ELT. Se eT ois oer eee Ie 28. K. fragrans

22. Leaf margin entire (occasionally some leaves with few teeth near apex).
29. Undersurface of mature leaf pubescent.
30. Pubescence on inflorescence soft, not closely appressed (Southern Highlands & Milne Bay

EOWA) ie ao rate ee ate, oe) kl cic, ote Senet foley > Se RA at es RAT oe ae ee 24. K. katikii
30. Pubescence on inflorescence stiffly appressed.
31. Leaves acuminate, usually broadly elliptic (throughout Malesia) ......... 25. K. coriacea

31. Leaves obtuse, usually narrowly elliptic (Sabah, Celebes, Philippines, Biak I.) 29. K. obtusa
29. Undersurface of mature leaf glabrous.
32. Leaves stiffly coriaceous, broadly elliptic to subrotund (New Guinea highlands)
30. K. laurifolia

32. Leaves not as above.

33. Inflorescence pubescent.

34. Leaves acuminate, usually broadly elliptic (throughout Malesia) ........ 25. K. coriacea

34. Leaves obtuse, usually narrowly elliptic (Sabah, Celebes, Philippines, Biak I.) 29. K. obtusa
33. Inflorescence glabrous (young inflorescence + setulose in K. slewmeri).

35. Pedicels of male flowers elongated (20—45 mm).

36. Male flowers in separate inflorescences (Irian Jaya).............. 31. K. flagelliformis
36. Male flowers on lower branches of inflorescences.
S/ealeatenarnowlyelliptic (Chimbi Prove) - 4-05 ssc ae aeete see 32. K. chimbuensis
37. Leaf broadly elliptic or elliptic (Morobe & Central Prov.).............. 33. K. fugax

35. Pedicels of male flowers shorter than 20 mm.
38. Pedicels of female flowers noticeable thickened for c. 10 mm below the receptacle (New
Britain N eWalnelan Gin eaciecet vac cect bee li cieeh eo NC 34. K. novobritanica
38. Pedicels of female flowers not strikingly thickened.
39. Male receptacle 4—6 mm long; anthers kidney-shaped. Leaf rigidly coriaceous.
40. Leaf oblong, apex rounded or retuse with a minute mucro, often 3-whorled (Western

Prove. alsouni@ueensland) eam Sressat eee eee eke. be ae 35. K. rigidifolia
40. Leaf narrowly elliptic, apiculate, opposite (Sudest I.) ............ 36. K. sudestensis
39. Male receptacle 1.5—2.5 mm long; anthers triangular. Leaf not rigidly coriaceous.
41. Leaves oblong, apex and base rounded (Vogelkop Peninsula) ....... 37. K. sleumeri
41. Leaves elliptic.
AD mibeavesinemibranaceousi(Scpikanhe sion) ia cee eer an iaeieie eit pee 38. K. monticola

42. Leaves firmly herbaceous or chartaceous (SE. Papua New Guinea) . 26. K. papuana

1. Kibara roemeri (PERKINS) PERKINS, Bot. Jahrb. 52

(1915) 212; Pxiipson, Blumea 30 (1985) 394. — _ gin minutely dentate, midrib evident, lateral veins
Matthaea roemeri Perkins, Pfl. R. Heft 49 (1911) numerous and connected by a well-defined reticula-
7 tion of minor veins, glabrous; petiole c. 5—10 mm,

A small tree, glabrous in all its parts. Leaves lance- 2.5 mm wide, channelled above. Inflorescence and
olate or obovate-lanceolate, up to 25 by 4.2 cm, char- _ flowers unknown. Drupes narrowly ovoid, c. 16 by
taceous, base broadly cuneate, apex apiculate, mar- 7 mm, shortly stipitate.

1986 |

Distr. Malesia: SW. Irian Jaya, ?Misool.

Ecol. Forest, 750 m.

Note. The stiff, lanceolate leaves, with sharply
toothed margin are characteristic. A second collec-
tion consisting of a leafy shoot only, may belong to
this species. It was made by TEUSMANN in Mysool I.

2. Kibara oligocarpella (KANEH. & Hatus.) PHILIP-
son, Blumea 30 (1985) 394. — Steganthera oligocar-
pella KANEH. & Hatus. Bot. Mag. Tokyo 56 (1942)
Fey Ey Gia

Shrub to 1.8 m. Leaves oblong-elliptic to narrowly
obovate, up to 3.3 by 1.5 cm, firmly coriaceous, base
cuneate, apex obtuse or retuse, entire, midrib promi-
nent, reticulation of veins evident below, glabrous;
petiole to 4mm. Monoecious. /nflorescence axillary
or supra-axillary; flowers in simple cymes or solitary.
— Male flowers ovoid, c. 2.5 mm long, pedicel 6—7
mm long; tepals 4, stamens 3—4. — Female flowers
+ globose, c. 3 mm long; 4 tepals around the minute
ostiole, several irregularly swollen pairs within; car-
pels 4—10, stigma obtuse. Drupes ovoid, c. 15 by 11
mm.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula, Arfak Mts, Angi Lakes).

Ecol. Open scrub (Tristaniopsis, Dacrydium), at
c. 2400 m.

Note. The small, close-set, thick-leathery leaves
(which dry a dark brown) are quite unlike any other
species. The flowers are yellowish brown, and the
ripe achenes black on a reddish brown receptacle.
KANEHIRA & HaATUuSIMA attributed their material to
Kibara on their herbarium labels, but published the
species as a Sfeganthera. The material collected by
SLEUMER & VINK, with female flowers, removes any
doubt about this species being a Kibara.

3. Kibara myrtoides Perkins, Bot. Jahrb. 52 (1915)
208; PHiLipson, Blumea 30 (1985) 395.

Shrub to 2 m; young branches slender, with ap-
pressed tomentum. Leaves elliptic, 4—5 by 1.3—2.2
cm, membranaceous, base cuneate, apex long-
acuminate, entire, midrib prominent, veins obscure,
becoming glabrous except for the midrib below; pet-
iole 3—5 mm long, pubescent, channelled above. /n-
florescence axillary, in dichasia or solitary, pu-
bescent. — Male flowers cupuliform, 2.5 mm @, ped-
icel 3~5 mm, pubescent outside, glabrous within;
tepals 6, ovate; stamens 2, subsessile. — Female
flowers solitary, arising above the axils, pedicel
10-15 mm, pubescent. Receptacle woody with very
short stipes. Drupes ovoid, 20 by 12.5 mm.

Distr. Malesia: Papua New Guinea (Sepik re-
gion). Only known from type.

Ecol. Montane forest, in scrub with few large
trees, 1400-1500 m. FI. fr. August.

Note. The specimen available to me lacks flowers

MONIMIACEAE (Philipson)

291

so that the parts of the above account are derived
from the original description. Since the androecium
is so reduced and details of the female flower are not
given, the assignment to Kibara must remain tenta-
tive. The foliage is quite unlike any other species.

4. Kibara ferox PHILIPSON, Blumea 30 (1985) 395. —
Fig. 15.

Shrub or treelet to 3 m, glabrous, the nodes dilated
with pores inhabited by ants. Leaves sessile, broadly
cordate to oblong, up to 30 by 23 cm, coriaceous, the
base amplexicaul, apex narrowing to a short or long
apiculum, margin with small or coarse sharp denta-
tions (the upper leaves and those of juvenile plants
narrower and more dentate), veins and reticulations
very prominent on the underside. 7Monoecious. Jn-
florescence of axillary fascicles (or rarely supra-
axillary) borne on a very short bracteate peduncle;
pedicels up to c. 10 mm, slender in male gradually
widening into the flower, female thicker, becoming
woody in fruit. — Male flower ovoid, 2 by 1.5 mm,
3 pairs of tepals; stamens c. 5, subsessile; anther
broadly triangular with a single horseshoe-shaped
opening. — Female flower globose, 4.5 by 4.5 mm,
ostiole surrounded by 2 pairs of obscure tepals (addi-
tional pairs probably within the ostiole); upper part
of the receptacular chamber with very thick irregular
glands; carpels numerous, glabrous; stigma short,
obtuse. Drupes sessile ovoid, c. 15 by 10 mm, verru-
culose when dry.

Distr. Malesia: Papua New Guinea (Madang
Prov.; Ramu Distr.; Eastern Highlands Prov.: Kai-
nantu Distr.; Morobe Prov.: Mumeng Distr., Wau
Distr.; Central Prov.: Port Moresby Distr.).

Ecol. Lowland and lower montane rain-forest,
$00—1550 m.

Vern. Daraboro (Madang Distr., Domainde).

Note. The sessile, cordate, coriaceous leaves are
unlike those of any other species. The leaves on lower
thicker branches are very broad and cordate, while
those on the more slender branches are oblong. Ju-
venile plants have smaller, narrower and more den-
tate leaves. Swollen nodes with pores used by small
black ants are also found in K. carrii, K. latifolia and
K. archboldiana, and resemble those of some species
of Steganthera. The flowers are yellowish green or
pinkish. The drupes are black on an orange torus.
The outer bark is light grey-green to brown, rough
with vertical fissures.

5. Kibara latifolia Puitipson, Blumea 30 (1985) 396.

Shrub 1.5 m high, glabrous, with the branches
prominently dilated at the nodes. Leaves broadly
ovate to subrotund, 20-30 by 13—24 cm, charta-
ceous to coriaceous, base broadly cuneate to trunc-
ate, apex obtuse, entire, principal veins widely
spaced, channelled above, prominent below, arched

292 FLORA MALESIANA [ser. I, vol. 102

Fig. 15. Kibara ferox Puiipson. a. Leaf from upper branch, b. ibid. from middle branch, c. ibid. from lower

branch, all «1/2, d. axillary fascicles of female flowers after anthesis, e.male inflorescence, both nat. size,

f. male flower, g. the same in LS, both x7 1/2, A. stamen, X15, i. female flower, j. the same in LS, both
«4, k. carpel, x7 1/2, /. fruit, x1/2 (a—d, i-/ PHmipson & Katro 3640, e—k LAE 61573).

1986 |

and meeting inside the margin, glabrous; petiole
12—18 mm long, deeply channelled above. Flowers
not seen. Jnfructescence lateral, a subsessile umbel or
compact cyme, peduncle stout, c. 2 mm long,
branches (pedicels) c. 15 mm long; receptacle leath-
ery, c. 1O—13 mma, with c. 12—16 short thick stipes.
Drupes ovoid, c. 10—12 by 6—8 mm.

Distr. Malesia: Moluccas (Halmahera & Obi
Is.). Two collections.

Ecol. Indense, low forest with little undergrowth
at 15 m.

Note. Known from only two gatherings, neither
with flowers. However, the broad leaves, swollen
nodes, and the small infructescences are distinctive.

6. Kibara archboldiana A.C. SmitH, J. Arn. Arb. 22
(1941) 231; PHitrpson, Blumea 30 (1985) 396.

Shrub to 4 m, with glabrous branches conspicu-
ously swollen at the nodes. Leaves elliptic or elliptic-
oblong, 15—36 by 6—14 cm, chartaceous, base
broadly cuneate or rounded, apex with a slender
apiculum 1—2 cm long, entire, midrib and principal
veins conspicuous below, arched ascending and unit-
ing within the margin, glabrous or + ephemeral min-
ute pubescence on young foliage; petiole 5—25 mm,
stout channelled above. Monoecious. — Male inflo-
rescence axillary or supra-axillary, a few-flowered
compact cyme or subfasciculate, c. 8 mm long, min-
utely puberulous, with small bracts; pedicel slender,
c. 3—6 mm long, bracteolate. — Male flowers sub-
globose, 2—4 mm g; tepals 6, obtuse; stamens 6, the
innermost 2 smaller. — Female flowers not seen. Old
infructescence thickened and woody below the fruits;
receptacle leathery 8—12 mm ag, stipes short (1—3
mm). Drupes ovoid or narrowly ovoid-oblong,
16—23 by 8—10 mm.

Distr. Malesia: Papua New Guinea (Western,
Gulf & Central Prov.).

Ecol. Inrain-forest from near sea-level to 500 m.

Vern. Hooanu, Uraru lang.

Note. The two original collections agree in all re-
spects. The only other collection (CONN ef al. LAE
66299) has very similar foliage, but has broadly
ovoid achenes instead of the very characteristic elon-
gate fruit of the type, and apparently lacks the swol-
len nodes.

7. Kibara karengana PHiLipson, Blumea 30 (1985)
396.

Scandent shrub or small tree to 6 m high, glabrous
or with small brown strigose hairs on young parts,
sometimes persisting on the petiole and underside of
the midrib. Leaves ovate to oblong, 9-22 by 4—10
cm, stiffly chartaceous, often more or less bullate,
base broadly cuneate, rounded or subcordate, apex
shortly apiculate, margin with small or coarse indu-
rated, sharp dentations, midrib, lateral veins and re-

MONIMIACEAE (Philipson)

293

ticulation of minor veins prominent below; petiole
channelled above, to c. 8 mm long. Monoecious. Jn-
florescences axillary or supra-axillary, male and fe-
male flowers in separate inflorescences; simple triads
orasmall cymose panicle, 10—20 mm long, peduncle
c. 2 mm long with minute bracts; female a few-
flowered fascicle, pedicels becoming woody in fruit
and elongating to 20-30 mm. — Male receptacle
ovoid, 3 by 2.5 mm, tepals in 4 pairs; stamens 4—6,
subsessile, anthers broadly triangular with a single
horseshoe-shaped opening. — Female receptacle
ovoid, 5 by 4 mm, sometimes pubescent, ostiole sur-
rounded by minute tepals and bearing within large
pendulous glands; carpels c. 12—18, pubescent with
short obtuse stigma. Fruiting receptacle leathery, c.
10mm @. Drupes + sessile, ovoid, c. 14 by 10 mm,
surface (when dry) verruculose.

Distr. Malesia: Papua New Guinea (Western,
Eastern & Southern Highland Provinces).

Ecol. Lower montane and mossy forest (Notho-
fagus, Podocarpus, Pandanus) and at forest/burnt
grassland margin, between 2500—3000 m.

Uses. In the Minj District the plant is considered
to be a male sex stimulant.

Vern. Gegnikl, kong-ambugont, Minj, ogum-
bwarombigl, Hagen.

Note. Theripe drupes are shining black borne on
an orange receptacle.

8. Kibara bullata PuHitipson, Blumea 30 (1985)
347.

Shrub to 2.5 m; young branches finely and densely
pubescent. Leaves elliptic to elliptic-oblong, up to 17
by 8 cm, chartaceous, base broadly cuneate, apex
long acuminate, margin dentate, midrib, lateral veins
and reticulation prominent below, the upper surface
raised between the veins (bullate), the pubescence
persisting on the lower surface especially on the mid-
rib and veins but disappearing from the upper sur-
face; petiole c. 1O—12 mm, channelled above, pubes-
cent. Monoecious. /nflorescences axillary or supra-
axillary, of short few-flowered fascicles; pedicel 2—4
mm long, densely pubescent. — Male flowers ovoid,
1.5 mm long, pubescent on the outer surface; 4 pairs
of tepals; stamens 4 in the outer whorl 0.75 mm long,
with 2 smaller central stamens; filament as broad as
the anther, short and pubescent. — Female flowers
similar to male but stouter, c. 2 mm long; tepals sur-
rounding the ostiole minute, with large pendulous
glands within; inner surface of the receptacle with
hairs between the carpels; carpels c. 10, ovary pubes-
cent, 0.8 mm long, stigma obtuse. Fruiting receptacle
+ sessile (peduncle undeveloped) with long radiating
stipes (receptacle c. 6 mm @, stipes c. 4-8 mm long).
Drupes ovoid, c. 18 by 10 mm.

Distr. Malesia: Papua New Guinea (Morobe
Prov.).

294

FLORA MALESIANA

[ser. I, vol. 102

Ecol. Rain-forest (Anisoptera and Castanopsis),
500—1000 m.

Note. The slightly bullate, pubescent leaves with
a serrate margin, combined with a fasciculate inflo-
rescence and sessile fruiting receptacles are distinc-
tive. The black drupes are borne on an orange recep-
tacle. Collected only twice. The specimen from Ga-
raina has less dentate leaf margins and shorter thick-
er stipes below the achenes but agrees with the type
in other respects.

9. Kibara kostermansii PHILIPSON, Blumea 30 (1985)
398.

Shrub to 4 m; young branches covered with stri-
gose tomentum. Leaves elliptic to obovate (occasion-
ally lanceolate), 14—22 by (3—)5—8(—10) cm, charta-
ceous, base cuneate or rounded, apex with an apicu-
lum, margin dentate (at least in most leaves) with
short, crisp hairs on the midrib, principal veins, and
often + sparsely over the whole lower surface; peti-
ole c. 1O—14 mm, deeply channelled above. 7Monoe-
cious. Inflorescences axillary, supra-axillary or ter-
minal; male and female flowers in separate inflores-
cences. — Male inflorescence a few-flowered cyme,
c. 10 mm long, hispid; peduncle 2 mm long bearing
minute bracts and lateral branches (pedicels) sub-
tended by bracts. — Female flowers solitary, borne
on a short (2 mm long) bracteate peduncle (pedicel).
— Male flowers obovoid, c. 2 mm long, receptacle
hispid on the outer surface and bearing a pair of
rounded bracts (? tepals), ostiole minute, surrounded
by c. 3 pairs of tepals; stamens 4, anthers broadly tri-
angular, subsessile. — Female receptacle hispid on
the outer surface and bearing c. 2 pairs of acute ovate
bracts, ostiole surrounded by 3—4 pairs of decussate
rounded tepals, with large pendulous glands within
the ostiole, inner surface of receptacle hispid, ovules
with a blunt peg-like stigma. Pedicels becoming
woody and longer in fruit (up to 15 mm); receptacle
becoming woody, to 20 mm g, including the stout
stipes. Drupes ovoid or ellipsoid, c. 15 by 10 mm.

Distr. Malesia: Moluccas (Halmahera, Batjan,
Morotai).

Ecol. Primary forest, often at sea-level on lime-
stone but reaching up to 500 m.

Vern. Wajoka gogumini, Halmahera, Tobelo
dial.

Note. The pubescent dentate leaves and the small
inflorescences are characteristic. Acute bracts like
those on the pedicel and receptacle of the female
flowers have not been seen on any other species. The
ripe fruits are blue-black, borne on an orange or red
receptacle.

10. Kibara leachii PHiLipson, Blumea 30 (1985) 399.
Shrub or small tree, to 6.5 m; young branches,
buds and young foliage covered in strigose hairs.

Leaves elliptic or narrowly elliptic, 14 by 5.5 cm,
thinly chartaceous, base narrowly or broadly cu-
neate, apex with a short obtuse apiculum, or tapered
to a long acute apiculum, margin finely or more
coarsely dentate, midrib prominent below, lateral
veins arched and uniting within the margin, traces of
the tomentum persisting on the mature leaves, espe-
cially on the midrib below, or the blade more or less
glabrous; petiole c. 5—10 mm, channelled above,
strigose. Monoecious. /nflorescences terminal, axil-
lary or supra-axillary, of compacted dichasia (fasci-
cles) or with the peduncle evident, c. 10—20 mm long,
strigose and bracteate. — Male flowers ovoid, 3 pairs
of rounded tepals; 4 stamens in the outer whorl, 0—3
in the inner whorl. — Female flowers not seen. Ped-
uncle below fruit remaining short or elongating
slightly (to c. 10 mm), pedicel may also elongate
under fruiting receptacle; receptacle becoming
woody, c. 5 mm @ (without stipes), stipes short (up
to c. 2mm long), strigose or glabrous. Drupes ovoid,
18 by 13 mm.

Distr. Malesia: Papua New Guinea (Central,
Northern & Milne Bay Provinces).

Ecol. Lowland rain-forest and lower montane
forest, 390—1370 m.

Note. The thinly chartaceous dentate leaves with
compact inflorescences are distinctive. Most gather-
ings have narrower leaves than the type (chosen be-
cause in flower) with long narrow apicula.

11. Kibara versteeghii PHitipson, Blumea 30 (1985)
399.

Shrub to 5 m, young branches with dense crisp
brown tomentum. Leaves elliptic, 11 by 3.8 cm, char-
taceous, base cuneate, apex narrowed into an ob-
scure apiculum, margin dentate, midrib and veins
prominent below, lateral veins few, strongly arched
and meeting within the margin, upper surface be-
coming + glabrous, lower surface retaining the crisp
tomentum; petiole c. 10 mm, densely tomentose.
Monoecious. J/nflorescence axillary or terminal, of
solitary flowers, fascicles, or short pleiochasia, pu-
bescent in all parts. — Male flowers ovoid, 1.5 mm
long, strigose on the outer surface; 3 pairs of round-
ed tepals; stamens 2 with 1 central staminode (only 2
flowers dissected); anther with a single horizontal
opening; filament short, strigose. — Female flowers
ovoid, 2.7 mm long, densely strigose on the outer
surface and with hairs on the inner surface between
the carpels, c. 4 pairs of tepals and prominent pendu-
lous glandular swellings within the ostiole; carpels c.
12—15, ovary pubescent, stigma blunt. Fruits not
seen.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula).

Ecol. Rather common shrub in Nothofagus for-
est and in old secondary forest, 1640—1950 m.

1986 |

MOoONIMIACEAE (Philipson)

295

Note. The small, dentate, pubescent leaves are
distinctive. The young leaves are described as brown-
ish purple above and light red below. The dried
leaves are a dull dark brown. The flowers are yellow.

12. Kibara oblongata PHILIPsoN, Blumea 30 (1985)
400, f. la. — Fig. 14a.

Small tree, c. 5 m high, glabrous in all its parts.
Leaves oblong, up to 31 by 12 cm, coriaceous or
chartaceous, base broadly cuneate, apex obtuse to
slightly retuse, margin entire, midrib prominent, lat-
eral veins connected within the margin; petiole
15—20 mm, 4 mm wide, channelled above. ? Monoe-
cious. /nflorescences axillary, fasciculate, pedicels c.
10 mm, wider towards the top; male flowers not seen.
— Female flowers globose, c. 4 mm @, ostiole sur-
rounded by c. 3 pairs of rounded tepals and with
swollen glands within; carpels c. 20, with a short ob-
tuse stigma. Fruiting receptacle enlarged and woody,
c. 15—20 mm g, with short stout stipes 2—3 mm long.
Drupes ovoid, 14 by 10 mm (? fully developed).

Distr. Malesia: Papua New Guinea (Morobe
Provy.; Lae Distr.; Milne Bay Prov.; Esa’ala Distr.,
incl. Normanby I.).

Ecol. Lowland rain-forest, from sea-level to c. 65
m.
Note. The large glabrous oblong leaves resemble
those of K. moluccana but the inflorescence is dis-
tinct. Two collections have been made on Normanby
I., a third collection from north of Lae appears to be
identical.

13. Kibara warenensis KANEH. & Hartus. Bot. Mag.
Tokyo 56 (1942) 249, f. 1; PHiLtipson, Blumea 30
(1985) 401.

Shrub to 2 m, glabrous. Leaves oblong or oblan-
ceolate-oblong, 18—27 by 5.2—8.2 cm, coriaceous,
base rounded to broadly cuneate, apex shortly acu-
minate or acute, margin spinulose dentate, midrib,
lateral nerves and reticulation prominent below; pet-
iole c. 10 mm. Monoecious. /nflorescences axillary,
of compact fasciculate cymes, glabrous, c. 10 mm
long. — Male flowers with pedicels c. 6—10 mm long,
thicker distally; receptacle c. 1.8 mm long; tepals 6,
minute; stamens 6, c. | mm long. — Female flowers
not seen. Drupes ovoid, sessile on a woody recep-
tacle, c. 20 mm @.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula).

Ecol. Lowland rain-forest at c. 100 m.

Note. The glabrous fasciculate inflorescence
combined with the spinulose dentate leaves are dis-
tinctive.

14. Kibara rosselensis PHitipson, Blumea 30 (1985)
401.
Small tree, 4 m; young branches finely pubescent.

Leaves narrowly obovate, 10—15 by 3—4.5 cm, sub-
chartaceous, base narrowly cuneate, apex apiculate,
margin dentate, midrib prominent below, lateral
veins c. 7—9 pairs, arched ascending, glabrous at ma-
turity; petioles to 15 mm long, channelled above.
Monoecious. /nflorescences fasciculate (compacted
cymes) c. 10—14 mm long, with small bracts around
the peduncle and the bases of the pedicels; pedicels
and usually the receptacle also bearing minute
bracts. — Male receptacle subglobose, narrowed into
the pedicel, c. 3 mm g; tepals in 3 pairs; stamens 4 in
outer whorl, c. 3 in inner whorl. — Female receptacle
similar, c. 4 mm 9, ostiole surrounded by c. 3 pairs
of tepals and bearing within large pendulous glands;
carpels c. 18. Drupes not seen, but described by the
collector as ovoid, 20 by 16 mm.

Distr. Malesia: Papua New Guinea (Milne Bay
Prov.).

Ecol. Low forest on ridge crest, 700 m.

Note. Male and female flowers in fascicles in the
axils of the dentate leaves. The flowers are yellow
and the ripe drupes black.

15. Kibara royenii PHILIPSON, Blumea 30 (1985) 401.

Shrub c. 3 m high, glabrous. Leaves narrowly
oblong-elliptic, 21 by 6.5 cm, chartaceous, base cu-
neate, apex apiculate, apiculum obtuse, margin
sharply serrate, midrib and lateral veins and their
junctions prominent, main veins numerous (8—10
pairs) with subsidiary lateral veins between them and
a prominent reticulation; petiole 1S—20 mm, chan-
nelled above. Monoecious. /nflorescences axillary,
of short compacted dichasia (fascicles), c. 10 mm
long, one or more in an axil. — Male flower ovoid,
c. 2mm @g; 4 pairs of tepals, the outer triangular, the
inner rounded; outer stamens 4, inner c. 2; filament
short, glabrous. Female flowers and fruit not seen.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula; Steenkool Distr.).

Ecol. Primary forest, at 50 m.

Note. The narrowly oblong elliptic, dentate
leaves, combined with the small fasciculate inflores-
cence are distinctive.

16. Kibara hartleyi PHiLipson, Blumea 30 (1985) 402.

Tall shrub, glabrous except for unopened buds.
Leaves up to 8 by 3.8 cm, rigidly coriaceous, base cu-
neate, apex with a prominent apiculum, margin
sharply dentate, reticulation of veins very evident on
both surfaces, glabrous; petiole 4—6 mm, channelled
above. Monoecious. /nflorescence axillary; flowers
solitary, pedicels c. 15-20 mm long at anthesis, elon-
gating slightly in fruit. — Male flower obovoid, c. 3
mm 9; tepals in c. 3 pairs; stamens in an outer whorl
of 5 and an inner whorl of 2—3. — Female flower
similar but slightly larger, carpels c. 12-16, ovary
densely covered with appressed hairs. Ripe drupes

296

FLORA MALESIANA

[ser. I, vol. 102

not seen, but developing fruits verruculose and pu-
bescent.

Distr. Malesia: Papua New Guinea (Eastern
Highlands Prov.; Goroka Distr.).

Ecol. Montane forest, c. 2700 m.

Note. This species shows a close but superficial
resemblance to Steganthera ilicifolia. It approaches
Kibara shungolensis which differs in having leaves
without a prominent apiculum, and with a broader
base, the female inflorescence usually bearing a later-
al flower, and the ovary being glabrous.

17. Kibara shungolensis PHiLipson, Blumea 30 (1985)
402.

Small tree c. 4 m high, glabrous in all its parts.
Leaves elliptic, up to 7 by 3.5 cm, rigidly coriaceous,
base broadly cuneate, apex acute or obtuse, margin
shallowly dentate, reticulation of veins very evident
on both surfaces, petiole c. 5—8 mm, channelled
above. ?7Monoecious. /nflorescence of compacted
dichasia (fascicles); peduncles bracteate at the base,
short at anthesis but later elongating (10—12 mm),
pedicels c. 8—10 mm long, those of the male flowers
more delicate than those of the female. — Male flow-
ers obovoid, c. 2mm @, ostiole surrounded by 3 pairs
of rounded tepals; androecium with 4 outer stamens
and fewer smaller inner stamens; filaments glabrous.
— Female flowers globose, c. 3 mm @, ostiole sur-
rounded by c. 3 pairs of minute tepals and with swol-
len glands within; carpels c. 20, ovary and stigma gla-
brous. Fruits not seen.

Distr. Malesia: Papua New Guinea (Morobe
Prov.; Mumeng Distr., Mt Shungol).

Ecol. Lower mossy forest, at about 2300 m.

Note. For features distinguishing K. shungolen-
sis from K. hartleyi see that species.

18. Kibara moluccana PERKINS, Bot. Jahrb. 45
(1911) 425; Pfl. R. Heft 49 (1911) 36; Puitipson, Blu-
mea 30 (1985) 403, f. 1b. — K. vrieseana PERKINS,
Bot. Jahrb. 45 (1911) 424. — K. teijsmanniana PER-
KINS, /.c. 425. — K. ledermannii PERKINS, Bot.
Jahrb. 52 (1915) 213. — Fig. 14b.

A shrub or small tree to 10 m high, glabrous.
Leaves oblong-elliptic, (10—)15—36 by (4.5—)7—12.5
cm, chartaceous or subcoriaceous, base broadly cu-
neate to rounded, apex obtuse to subacute or with a
short obtuse apiculum, entire, principal veins evident
on lower surface, arched and meeting inside the mar-
gin, glabrous; petiole 15—22 mm, deeply channelled
above. Monoecious. Jnflorescence axillary, gla-
brous, with a short peduncle (3—5 mm long) and sev-
eral short branches bearing slender pedicels (3—4 cm)
crowded near their apices, the upper pedicels thicker
and bearing female flowers or all the flowers male. —
Male receptacle obovoid, c. 2 mm long, with brac-
teoles; tepals 4, minute; stamens 6 (2 inner smaller)

or 4. — Female receptacle subglobose, c. 4 mm long,
with bracteoles and 4 rounded outer tepals, enclosing
irregularly swollen glands; c. 10 carpels, stigma
short. In fruit the pedicel becoming stout and woody,
the receptacle with several short, very stout stipes.
Drupes ovoid, 18 by 10 mm.

Distr. Malesia: Moluccas (Buru, Ceram, Am-
bon, Halmahera, Morotai, Obi) and Papua New
Guinea (East Sepik Prov.).

Ecol. Rain-forest, to 1000 m.

Uses. The skin of the fruit is rubbed on the hair
to dye it black (Halmahera).

Vern. Ogoroutu masauru mnauru, Halmahera,
Tobaro lang.

Note. The large, glabrous, oblong leaves with the
almost fasciculate inflorescences are characteristic.
The flowers are yellow. The material I have seen of
K. ledermannii is fragmentary but the leaves and in-
florescence match those of K. moluccana.

19. Kibara microphylla PERKINS in K. Sch. & Laut.,
Nachtr. Deut. Schutzgeb. Stidsee (1905) 268; Pfl. R.
Heft 49 (1911) 33; Bot. Jahrb. 52 (1915) 210; PHitir-
SON, Blumea 30 (1985) 403.

A shrub (?) with glabrous branchlets. Leaves ellip-
tic to narrowly elliptic, 7.5—11 by 2—3.5 cm, mem-
branous, base cuneate, apex acuminate, margin en-
tire or irregularly dentate, midrib evident, principal
veins arched and uniting within the margin, gla-
brous; petiole to 8 mm, slender, channelled above.
Monoecious. /nflorescences axillary or terminal, gla-
brous, c. 2.5—5 cm long, of small simple cymes or
panicles with delicate peduncle and branches; pedi-
cels of the male flowers more delicate than of the fe-
male. — Male flowers c. 1 mm @, ovoid; tepals 4,
minute; stamens 4, subsessile. — Female flowers +
globular with minute tepals with swollen glands with-
in; carpels c. 8, glabrous. Fruits not seen.

Distr. Malesia: Papua New Guinea (West Sepik
Prov.: Torricelli Mts).

Ecol. Forest at 600 m.

Note. I have not seen the type collection, but a
specimen collected at the same locality agrees with
the original description in all respects, except that the
leaf margins are not entire but irregularly dentate.
The small leaves and very delicate inflorescences of
this specimen leave no doubt that it represents K. mi-
crophylla.

20. Kibara streimannii PHiLIpsoNn, Blumea 30 (1985)
403.

Small tree to 7 m, young shoots softly and densely
pubescent. Leaves narrowly oblong or oblong, stiffly
chartaceous, 8—16 by 2.5—4.7 cm, base truncate to
shallowly cordate, apex narrowed into indistinct
acute apiculum, margin entire, becoming glabrous
above, soft pubescence persisting below, midrib

1986 |

channelled above, prominent below, lateral veins
arched and meeting within the margin, prominent,
reticulation well defined; petiole 4-7 mm, densely
short pubescent. Monoecious. /nflorescences axil-
lary; male and female flowers in_ separate
inflorescences. — Male inflorescence consisting of
elongated rachises, c. 8 cm long, setulose, bearing
many pairs of subulate bracts. — Female inflores-
cences shorter (c. 4.5 cm long), peduncle c. 3.5 cm
long, ending in a dichasium. — Male flowers in axils
of bracts, apparently arising in acropetal sequence
and soon caducous, pedicel c. 4 mm long, setulose;
receptacle obovoid, c. 3.5 mm long, hairy outside;
tepals 6, rounded or mucronate; stamens 4 outer and
c. 2 inner, c. 0.75 mm long, anthers triangular with
very short narrow filaments. — Female flowers glo-
bose, c. 4 mm ag, densely tomentose. Fruiting re-
ceptacle c. 8 mm Qg, setulose, without stipes. Drupes
not seen.

Distr. Malesia: Papua New Guinea (Morobe
Prov., Wau & Menyamya Distr.).

Ecol. Nothofagus and Castanopsis forest at 1700
m.
Note. The softly pubescent foliage is distinctive
and the form of the male inflorescence is unique in
the genus. The flowers are yellowish white and the
ripe fruit deep blue-black.

21. Kibara macrantha PHiLipson, Blumea 30 (1985)
404, f. lc. — Fig. 14c.

Tree up to 20 m, with pubescent branches. Leaves
ovate or elliptic, 120—240 by 40—80 mm, charta-
ceous, base rounded to broadly cuneate, apex obtuse
or subacute, margin entire or occasionally with few
coarse dentations, midrib prominent and reticulation
of veins evident, pubescence persisting on the midrib
below and to a less extent on the subsidiary veins;
petiole 10—15 mm, channelled above, pubescent.
Monoecious. Jnflorescence axillary or supra-
axillary, on leafy twigs or clustered on older
branches, simple dichasia or open cymose panicles,
rachis up to 10 cm with several pairs of lateral
branches, or much shorter, pubescent; bracts ovate,
c. 4mm long, at the base the peduncle and lateral
branches and usually at the base of each receptacle.
— Male receptacle obovoid, pubescent outside, c, 5
mm long at anthesis; tepals 6, rounded; stamens 6—8,
subsessile, anther broadly triangular with a single
horseshoe-shaped opening. — Female receptacle
similar, 6—8 mm long; tepals in several pairs with
swollen glands within, inner surface pubescent; car-
pels numerous, pubescent, stigma short, obtuse.
Drupes + sessile, ovoid, c. 12 by 9 mm, pubescent;
fruiting receptacle c. 10 mm @, woody.

Distr. Malesia; Papua New Guinea (Morobe
Prov.: Finschhafen & Wau Distr.; Eastern High-
lands Prov.: Kainantu Distr.).

MONIMIACEAE (Philipson)

297.

Ecol. Lower montane forest (Nothofagus, Cas-
tanopsis, Lithocarpus), 1400—2000 m.

Note. The open inflorescences of large flowers
distinguish this species.

22. Kibara carrii PHILIPSON, Blumea 30 (1985) 405.

A shrub, 1.75 m high, glabrous, in all its parts, the
nodes dilated. Leaves broadly elliptic, up to 27 by 16
cm, coriaceous, base broadly cuneate, apex abruptly
apiculate, margin coarsely dentate, midrib and lateral
veins prominent below, channelled above; petiole c.
10mm long, 4mm wide. ?Monoecious. /nflorescence
axillary or supra-axillary, cymose panicles, rachis c. 5
cm long, lateral branches c. 2 cm long, sometimes
again branched, pedicels 10—12 mm, slender and
slightly thickened below the receptacle. — Male flow-
er ovoid, c.2mmg@, ostiole with 3 pairs of tepals, outer
whorl of stamens 3—4, inner whorl c. 3, anther with 1
horseshoe-shaped opening; filament pubescent. —
Female flower ovoid, c. 3 mm Q, ostiole surrounded
by small obtuse tepals in pairs, the innermost forming
thickened pendulous glands; carpels c. 15—20, witha
short blunt stigma. Fruiting receptacle c. 12 mm 9.
Drupes subsessile, ovoid (not fully mature).

Distr. Malesia: Papua New Guinea (Central
Prov.: Goilala Distr.).

Ecol. Secondary forest at 1000 m.

Note. The large broadly elliptical and coarsely
serrate leaves are distinctive. Similar dilated nodes
are found in a few other species and also in some spe-
cies of Steganthera. The flowers are described as yel-
low.

23. Kibara elongata A.C. SmirH, J. Arn. Arb. 22
(1941) 244; Puitipson, Blumea 30 (1985) 404.

A shrub or small tree, glabrous. Leaves elliptic to
oblong-elliptic, rarely lanceolate (? juvenile); 17—30
by (3—)5.5—11 cm; coriaceous, sometimes rugose,
base broadly cuneate to rounded or cordate, apex
with a short blunt or long slender apiculum, margin
+ distantly spinose-dentate, veins prominent, gla-
brous; petiole 8-20 mm, stout, channelled above.
Monoecious. /nflorescence axillary or above the fo-
liage leaves, paniculate with slender peduncle, rachis
and branches, to 15 cm long and much branched, but
often simpler and much shorter, sometimes cauliflor-
ous. Purely female inflorescences sometimes simple
pleiochasia. — Male receptacle ovoid, c. 3 mm long;
tepals 6; stamens 4—6, c. 1 mm long, filamen short.
— Female flowers globose, c. 2 mm @; minute tepals
surround the ostiole, within which are glandular
swellings; carpels c. 30, glabrous, stigma obtuse.
Peduncle and pedicel thickened and woody in fruit;
receptacle c. 7-12 mm @ (without the stipes), stipes
stout, c. 3-4 mm. Drupes ovoid or narrowly ovoid,
18—27 by 8—12 mm.

Distr. Malesia: West New Guinea (Vogelkop

298

FLORA MALESIANA

[ser. I, vol. 102

Peninsula to Idenburg R.), Papua New Guinea (Jimi
Valley).

Ecol. In primary rain-forest from near sea-level
to 2000 m.

Vern. Kinjoem, Hattam lang., fa/walye, Eipo-
mek Valley.

Note. The coriaceous leaves with spiny teeth on
the margin are distinctive. The dentations on the
leaves from Jimi Valley are less spinose than the col-
lections from Irian Jaya. The inflorescence may be a
large diffuse panicle, or may be reduced in size but
still with open branching. The flowers are yellow or
orange and the ripe drupes are black, borne on an
orange receptacle.

24. Kibara katikii PHiLIPsoN, Blumea 30 (1985) 406.

Small tree to 16 m, young growth densely covered
with buff or fulvous tomentum. Leaves variable in
size and shape, ovate oblong-elliptic or obovate,
7.5—21 by 3.2—8.3 cm, becoming coriaceous, base
broadly cuneate or rounded, apex shortly apiculate,
margin entire or (more usually) irregularly dentate in
the upper part, dentations either small and obscure
or prominent, midrib prominent below, lateral veins
evident and meeting within the margin, upper sur-
face becoming + glabrous above, lower surface soft-
ly pubescent; petiole c. 10—15 mm, densely pubes-
cent. Monoecious. /nflorescence axillary or supra-
axillary, usually of simple, 3-flowered cymes, c. 30
mm long, densely pubescent, pedicels c. 8 mm long,
male and female flowers in separate inflorescences.
— Male flowers scarcely wider than the pedicel (c. 1
mm), pubescent on the outer surface and on the
lower part inside the receptacle; tepals 4, rounded;
stamens 2, attached to the base of the receptacle, c.
0.75 mm long, anther kidney-shaped, filament nar-
row, very short. — Female flowers wider than the
stout pedicel (to 3 mm), pubescent on both surfaces;
tepals 4 (2 tepals or bracts often present on the outer
surface), inner rim of ostiole thickened and glandu-
lar; carpels c. 10—15, c.1.25 mm long, pubescent,
stigma obtuse glabrous. Fruiting receptacle usually
solitary (developing from the terminal flower, side
branches occasionally also with fruits), peduncle and
pedicel becoming thickened and woody, receptacle
enlarging slightly (c. 8 mm @), pubescent, with thick
stipes or achenes + sessile. Drupes ovoid to sub-
spherical, 13—18 by 9—11 mm, sparsely pubescent.

Distr. Malesia: Papua New Guinea (Southern
Highlands, Morobe, Northern, Central & Milne Bay
Provinces).

Ecol. Lowland and montane rainforest, 30—2200
m.
Note. The foliage may have very different as-
pects, sometimes being broadly obovate with an en-
tire margin, at others being ovate and tapering to-
wards the apex. The copious and persistent tomen-

tum on the underside of the leaf is characteristic.
This species has a wider distribution than most in the
genus and a very considerable altitudinal range. The
bark is thickly corky, pale brown and deeply and
closely fissured.

25. Kibara coriacea (BL.) TULASNE, Arch. Mus. Hist.
Nat. Paris 7 (1855) 404; BL. Mus. Bot. Lugd.-Bat. 2
(1856) 89; PERKINS, Bot. Jahrb. 25 (1898) 576; PErR-
KINS & Gia, Pfl. R. Heft 4 (1904) 62, f. 16B; PEr-
KINS, ibid. Heft 49 (1911) 35, f. 11; Rup. Fl. Mal.
Pen. 3 (1924) 75; Back. & BaAKH. f. Fl. Java 1 (1963)
117; Putttpson, Blumea 30 (1985) 406. — Brongniar-
tia coriacea Bu. Bijdr. (1825) 436. — K. blumei
StEuD. Nomencl. Bot. (1840) 846; BL. Mus. Bot.
Lugd.-Bat. 2 (1856) 89. — Sciadocarpus brongniartii
Hassk. Flora 25, ii (1842) Beibl. 1: 20. — Sarcodiscus
chloranthiformis Grirr. Not. Pl. As. 4 (1854) 380. —
K. chartacea Bu. Mus. Bot. Lugd.-Bat. 2 (1856) 89.
— K. cuspidata Bt. I.c. 89. — Mollinedia coriacea
(BL.) Batt. Hist. Pl. 1 (1869) 304. — K. tomentosa
PERKINS, Bot. Jahrb. 25 (1898) 571. — K. trichantha
PERKINS, /.c. 572. — K. macrophylla Perkins, l.c.
571. — K. serrulata PERKINS, /.c. 575; Pfl. R. Heft
4 (1901) 60, f. 16A. — K. angustifolia PERKINS, l.c.
577. — K. grandifolia MERR. Publ. Gov. Lab. Phi-
lip. n. 29 (1905) 15; En. Philip. 2 (1923) 185. — K.
ellipsoidea MERR. Philip. J. Sc. 1 (1906) Suppl. 56.
— K. mollis Mer. ibid. 3 (1908) Bot. 225. — K. cle-
mensiae PERKINS, Bot. Jahrb. 45 (1911) 423; MerrR.
En. Philip. 2 (1923) 184. — K. vidalii PERKins, Bot.
Jahrb. 45 (1911) 423; Merr. En. Philip. 2 (1923) 185.
— K. motleyi Perkins, Bot. Jahrb. 45 (1911) 424. —
K. merrilliana PERKINS, /.c. 424; MERR. En. Philip.
2 (1923) 185. — K. stapfiana PERKINS, Bot. Jahrb. 45
(1911) 424; Merr. En. Philip. 2 (1923) 185. — K.
warburgii PERKINS, Bot. Jahrb. 45 (1911) 424. — K.
macrocarpa PERKINS, /.c. 425. — K. schlechteri PER-
KINS, Pfl. R. Heft 49 (1911) 31. — K. longipes PErR-
KINS, /.c. 31. — K. inamoena Perkins, lI.c. 34. — K.
dichasialis SUESSENG. & HEINE, Mitt. Bot. Staats-
samml. Miinchen 2 (1960) 60.

Tree to 22 m or rarely a scandent shrub, young
branches glabrous or pubescent. Leaves opposite
(rarely subopposite or whorled); broadly ovate to el-
liptic oblong, 9—35 by 5—24 cm, coriaceous or char-
taceous, base cuneate rounded or subcordate, apex
shortly to long acuminate, margin entire or minutely
to coarsely dentate, glabrous or sparingly to rather
densely pubescent beneath, midrib and principal lat-
eral veins prominent beneath, lateral veins arched-
ascending and meeting within the margin; petiole
5—25 mm, channelled above, glabrous or pubescent.
Monoecious. /nflorescences axillary, supra-axillary,
terminal, or cauliflorous, pubescent, cymes solitary
or several arising at the same node, varying from
simple 3-flowered cymes to complex pleiochasia with

1986 |

lateral branches branching to the third or fourth de-
gree, male flowers on the lower branches; rachis and
lateral branches of female part of the inflorescence
stout and thickening and becoming woody after an-
thesis, those of the male parts finer and caducous, up
to 20 cm long but often much shorter, pedicels 15—30
mm, gradually widening below the flowers, of the
male more delicate than those of the female. — Male
flowers globose, 1.5—2 mm ag, only slightly wider
than the pedicel, pubescent; tepals 6—8, apex round-
ed, usually with 4 outer stamens and up to 4 smaller
inner stamens, filaments strap-shaped. — Female
flowers globose, c. 3—5 mm @g; tepals c. 6, with swol-
len pendulous glands within the ostiole; carpels
usually c. 20, stigma short obtuse. Infructescence en-
larged, woody, receptacle c. 20 mm 9. Drupes ovoid,
15 mm long, shortly stipitate or sessile.

Distr. Throughout Malesia.

Ecol. Lowland rain-forest, including swamp for-
est and coral limestone, to lower montane forest,
from sea-level to 1600 m.

Uses. The fruit is said to be edible, and the leaves
are used to flavour meat dishes.

Vern. Malaya: pako kubang tando, p. pakan
Jjantan, Malacca, pako srean puteh, Negri Sembilan,
susu ajam, Pahang; Sumatra: alimau dotan, kaju
mata ole, k. ruang-ruang, k. singumban paya,
sangka-sangka batu, s.-s. sito, sito mérah; Java: bu-
lusan, ki bara, ki éndog, ki kuya, ki sauheum, pata-
tulang, ramat daging, rimik dasing, S; Banka: tam-
pui, M; Sabah: ambibliw, Dusun Banggi, /abak, Du-
sun; Philippines: anonias, Tag., pototan, Manila;
New Guinea: keawon, Sepik, mardiber, Biak lang.,
pu-ala, Managalese, sakunuo, Muswaar I., tor, Bu-
Jolo, Biaro dial.

Note. Although this species varies in respect to
the size and shape of its foliage and inflorescences,
the number of its parts and the degree of its pubes-
cence, it retains a character over its extensive range
which ensures its recognition. This is best expressed
by the broad, pliant, fresh green leaves and the open-
ly branched inflorescence with strong pedicels to the
female flowers which terminate the more distal
branches. Specimens with the largest inflorescences
oecur in Sumatra. The range of variation appears
greatest in the Philippines, and, when more complete
material becomes available, several species which
have been described from there may yet prove to be
valid, though here reduced to synonymy. The flow-
ers are yellow, the male being somewhat greener; the
drupes are black, borne on a yellow to orange recep-
tacle.

26. Kibara papuana A.C. Smirn, J. Arn. Arb. 22
(1941) 242; Pumipson, Blumea 30 (1985) 407.

» Shrub or medium-sized tree to 22 m; branchlets
glabrous or slightly pubescent; buds glabrous or cov-

MONIMIACEAE (Philipson)

299

ered in appressed pubescence. Leaves opposite (or
subopposite), narrowly to broadly elliptic, or ovate,
thinly to firmly chartaceous, (6.5—)10—19 by (2—)
5—6(—9) cm, base broadly to narrowly cuneate or
rounded, apex cuspidate obtuse, or narrowly and
long-apiculate acute, margin entire or with few to
many teeth, midrib prominent below, lateral veins
arched-ascending, the reticulation of minor veins evi-
dent; petiole narrow, to 15 mm, channelled above.
Inflorescence axillary, supra-axillary, or terminal,
often of 3-flowered cymes, or solitary flowers or
pleiochasia, sometimes with secondary or tertiary
branching to form panicles; male flowers on lower
more slender branches or in separate often panicu-
late inflorescences. — Male flowers ovoid, 1.5—2.5
mm long, glabrous; tepals 6; stamens 4—8, the outer
4 large, the inner smaller or absent, anthers broadly
triangular, obtuse, filaments short, as wide as the an-
thers. — Female flowers globose, c. 3 mm long, gla-
brous and sometimes minutely puberulous on both
surfaces; tepals minute with thickened glands within
the ostiole; carpels numerous, glabrous or pubes-
cent, stigma short, obtuse. Fruiting receptacle up to
10 mm @, with stout prominent stipes c. 4-6 mm
long. Drupes ovoid, c. 18 by 12 mm, rugulose.

Distr. Malesia: Papua New Guinea (Central,
Northern, Milne Bay and Morobe Prov.).

Ecol. Primary lowland rain-forest, montane for-
est and mossy forest; understorey with Castanopsis
and Araucaria. Also in secondary growth, 100—2100
m.
Vern. Boakeava, Wagau, popoia, Northern
Prov., Koiari lang., siganapa, Northern Prov., Oro-
kawa lang., saha, northern Managalese.

Note. Similar to K. coriacea but with smaller fo-
liage, and with smaller, less indurated flowers. The
ripe drupes are black on an orange receptacle. In
some specimens the achenes are muricate, but this
may be due to insect infection. The species, as here
treated, includes a considerable range of leaf size,
and most specimens do not show the compact inflo-
rescence which SmirH noted on the type. The extreme
forms come from the islands to the east of New
Guinea (especially Goodenough and Normanby Is.),
but a continuous series of intermediate states unite
all the forms.

27. Kibara nitens PHiLipson, Blumea 30 (1985) 408.

Small sparsely branched shrub usually under 3 m
high, glabrous. Leaves elliptic, elliptic-oblong or
narrowly obovate, 17 by 6.5 cm, coriaceous, base cu-
neate, apex apiculate, apiculum obtuse or acute,
margin obscurely and irregularly dentate, midrib
prominent, lateral veins numerous and close-set, at
first straight but curved and uniting near the margin,
glabrous; petiole 15 by 2.5 mm, channelled above.
Monoecious. /nflorescence an axillary or terminal

300

FLORA MALESIANA

[ser. I, vol. 102

pleiochasium, rachis c. 4cm, peduncle c. 18 mm long,
bracteate at the base, the opposite pairs of pedicels
also subtended by bracts; pedicels c. 5 mm, becoming
thickened and elongating in female flowers after an-
thesis. — Male flowers ovoid, c. 1 mm 9g, ostiole sur-
rounded by 3 pairs of obtuse tepals; stamens 4 insert-
ed halfway up the receptacle, with 2 small central
staminodes. — Female flowers not seen. Fruiting re-
ceptacle woody, c. 8 mm @. Drupes c. 12—15 shortly
stipitate, ellipsoid, c. 16 by 11 mm (when dry).

Distr. Malesia: West New Guinea (Star Mts), Pa-
pua New Guinea (West Sepik Prov.: Telefomin Dis-
trict; Southern Highlands Prov.: Tari Distr.).

Ecol. Undergrowth in primary forest, 750—2150
m.

Vern. Soinok, Telefomin.

28. Kibara fragrans PHiLIpsON, Blumea 30 (1985)
408.

Small tree, c. 3 m, young shoots glabrous. Leaves
elliptic-oblong, to 22 by 8.5 cm, chartaceous, base
broadly cuneate, apex shortly apiculate, apiculum
obtuse, margin irregularly and indistinctly dentate,
midrib prominent, principal lateral veins evident,
arched, ascending, glabrous; petiole to 3 cm. Jn/flo-
rescences axillary, c. 50 mm long, glabrous, small
few-flowered cymes or rather more complex small
paniculate cymes, solitary or clustered at nodes, up-
per flowers female on stouter pedicels; pedicels c. 20
mm, wider distally. — Male receptacle c. 2 mm long,
cup-shaped, widely open; tepals 6, rounded; stamens
4 in outer whorl, c. 0.75 mm long, c. 2 in centre,
shorter, filament strap-shaped, as wide as the anther.
— Female receptacle ovoid, c. 2.75 mm long; tepals
6, rounded, ostiole small, with thickened glands
within. Fruié not seen.

Distr. Malesia: NW. New Guinea.

Ecol. Lowland forest and on hilltop at 150 m.

Note. The foliage resembles that of Steganthera
hirsuta, while the inflorescences are similar to those
of Kibara fugax. The green flowers are fragrant.

29. Kibara obtusa BL. Mus. Bot. Lugd.-Bat. 2 (1856)
89; PERKINS & Gita, Pfl. R. Heft 4 (1901) 61; PEr-
KINS, Bot. Jahrb. 45 (1911) 424; Pfl. R. Heft 49
(1911) 34; Merr. En. Philip. 2 (1923) 185; Puiip-
son, Blumea 30 (1985) 409. — K. depauperata MERR.
Publ. Gov. Lab. Philip. 1. 13 (1906) 13.

Shrub or tree to 20 m, bole 36 cm @ dbh, young
branches with appressed stiff hairs. Leaves narrowly
to broadly elliptic, 8—11.5(—15) by 3.5—5(—6.5) cm,
base narrowly or broadly cuneate, apex obtuse, mar-
gin entire, becoming glabrous or lower surface with
sparse stiff pubescence; petiole 10—18 mm, becom-
ing glabrous or hairs + persistent. Inflorescences
axillary or terminal, pubescent, simple or paniculate
cymes, up to c. 70 mm long, the rachis, lateral

branches and pedicels slender or stronger with a
more compacted inflorescence. — Male receptacle
obovoid, c. 2 mm long, with a pair of small brac-
teoles, outside minutely pubescent; tepals 4, minute;
stamens 4, sometimes with 2 central staminodes, and
with staminodes inside the outer 2 tepals. — Female
receptacle + globose, c. 2.5 mm @, with bracteoles
on the outer surface or on the pedicel, minutely pu-
bescent on outer and inner surfaces; 4 outer tepals,
obtuse, with 4 swollen glands within which project
among the carpels; carpels c. 13, pubescent, with a
blunt stigma. Mature torus swollen and leathery,
with | to c. 13 short stout stipes, slightly pubescent.
Drupes ovoid, 17—24 by 10—12 mm.

Distr. Malesia: N. Borneo (Sabah: Lahad Datu
Distr.), Philippines (Luzon: Benguet Prov.), NE. Ce-
lebes (Minahasa), West New Guinea (Biak I.).

Ecol. Primary rain-forest, sea-level to 700 m.

Vern. Mardieber, Biak lang.

Note. The flowers are yellow or orange; the ripe
drupes are black on an orange receptacle.

30. Kibara laurifolia A.C. Smirn, J. Arn. Arb. 22
(1941) 240; Puitipson, Blumea 30 (1985) 409.

Sprawling shrub or small tree to 7 m, glabrous.
Leaves elliptic, elliptic-oblong or suborbicular, up to
17 by 8.5 cm, coriaceous, base rounded to broadly
cuneate, apex obtuse to slightly apiculate, margin en-
tire or occasionally with obscure dentations, midrib,
lateral veins and reticulation evident, glabrous; pet-
iole 5-10 mm, channelled above. Monoecious. /n-
florescences axillary, supra-axillary, or terminal,
flowers solitary or usually in dichasia, or pleiochasia,
c. 30 mm long, peduncle c. 10 mm, bracteate at its
base, pedicels c. 10—12 mm, subtended by bracts. —
Male flowers ovoid, c. 3 mm g, 3 pairs of tepals, 4
stamens with horseshoe-shaped openings, filaments
very short. Female flowers + globose, c. 4mm Q, os-
tiole with 4 pairs of tepals and prominent swollen
glands within, carpels numerous, ovary glabrous,
stigma obtuse. Fruiting peduncle and pedicels elon-
gated up to 8 cm, receptacle woody, c. 10 mm @.
Drupes ovoid, c. 16 by 12 mm, or subspherical, +
sessile or shortly stipitate.

Distr. Malesia: throughout New Guinea in the
highlands between 139°—148° E.

Ecol. Primary and disturbed lower montane
rain-forest, 1800—2800 m.

Vern. Kamokam, Enga lang.

Note. The broad, often suborbicular, leathery
leaves are distinctive. The ripe drupes are purplish
black on an orange to yellow receptacle. A specimen
from near Wabag approaches K. macrantha in the
large pubescent receptacles, but has foliage similar to
that of K. laurifolia. It may be a hybrid, though it lies
west of the known range of K. macrantha.

1986 |

MONIMIACEAE (Philipson)

31. Kibara flagelliformis PHiipson, Blumea 30
(1985) 409.

Shrub or small tree; young parts minutely puberu-
lous and soon becoming glabrous. Leaves opposite
or subopposite, elliptical, chartaceous, c. 19 by 9 cm
(all damaged), base broadly cuneate, apex not pres-
ent, margin entire, midrib prominent below, lateral
veins strongly arched-ascending; petiole stout, c.
10—12 mm, channelled above. Jnflorescences axil-
lary, central rachis c. 8 cm, lateral branches in oppo-
site pairs or 3—4 at the same level, usually again
branched about the middle; pedicels slender, elon-
gated (up to 45 mm). — Male flowers obovoid, 3 mm
long, with 4 rounded tepals surrounding a widely
open ostiole (sometimes another pair of tepals
(bracts) about the middle of the receptacle), 4 large
stamens and c. 3 smaller stamens in the centre; fila-
ments wide, strap-shaped, c. 2 mm long, wider than
the small triangular anthers. — Female flowers and
fruit not seen.

Distr. Malesia: West New Guinea (Napan
Distr.). Known only from the type.

Note. The specimen bears only male flowers in
strangely branched inflorescences. The stamens are
unusual, with long filaments much wider than the
small anthers.

32. Kibara chimbuensis PHiLipson, Blumea 30 (1985)
410.

Small tree to 7 m, young branches glabrous.
Leaves glabrous, narrowly elliptic to lanceolate,
membranaceous, 9—13 by 2.8—4.8 cm, base nar-
rowed into the petiole, apex narrowed into an indis-
tinct and obtuse apiculum, margin entire or slightly
undulate, lateral veins rather indistinct, steeply
ascending; petiole to 10 mm. Monoecious. /nflores-
cences axillary, supra-axillary or terminal, glabrous,
singly or in groups, either few-flowered cymes or
more branched open, cymose, panicles up to 8 cm
long; pedicels slender, 20—45 mm; male flowers on
lower branches. — Male flowers cup-shaped, c. 1.5
mm long, 2 mm g; rounded tepals widely open; sta-
mens 4 in outer whorl and 4 smaller stamens at
centre, filaments strap-shaped, glabrous. — Female
flowers subglobose, c. 3 mm long, 2.5 mm 9; tepals
6, rounded, inner rim of ostiole with pendulous
glands; carpels c. 10, glabrous, stigma obtuse. /n-
Sructescence with somewhat thickened pedicels and a
woody receptacle c. 15 mm @ (including short thick
stipes). Drupes ovoid, c. 15 by 10 mm when dry,
shortly stipitate.

Distr. Malesia: Papua New Guinea (Chimbu
Prov.: Kerowagi Distr.).

Ecol. A small tree in forest remnant at 1350 m.

Note. The narrowly elliptic to lanceolate leaves
have a thin texture and dry to a dull dark green. The
inflorescences are similar to those of K. fugax.

301

33. Kibara fugax PHiipson, Blumea 30 (1985) 411.
— Fig. 16.

Small tree, up to 10 m, all parts glabrous. Leaves
elliptic to broadly elliptic or obovate, 10—16(—18) by
3.8—7(—8) cm, membranaceous to thinly charta-
ceous, base cuneate, apex slightly apiculate, apicu-
lum obtuse, entire or occasionally with a few obscure
dentations, midrib evident, lateral veins few, strong-
ly arched; petiole 5—8(—10) mm long, channelled
above. Monoecious. /nflorescence terminal or axil-
lary, simple or paniculate cymes, either all flowers of
one sex or with male flowers on the lower branches,
to c. 50 mm long; pedicel of male flowers c. 36—60
mm long, very slender, or female flowers stronger
and shorter (c. 15—20 mm). — Male receptacle sub-
globose, c. 1.5 mm 9; 6 rounded tepals around the
ostiole; stamens c. 4 (5) outer, 2—5 inner, filament
short, broad, anther broad with a single horizontal
opening. — Female receptacle globose, c. 2—2.5 mm
@; tepals minute, 4 around the ostiole, another pair
inside and with thickened glands within; carpels c.
16—20, glabrous, stigma short, blunt. Drupes ovoid,
c. 13 by 9 mm, shortly stipitate.

Distr. Malesia: Papua New Guinea (Morobe &
Central Prov.).

Ecol. Lower montane forest or secondary forest,
between 750 and 1400 m.

Note. A common tree in the Wau District distin-
guished by its membranous leaves (which usually dry
to a blackish colour) and the long delicate pedicels of
male flowers. The male receptacles and pedicels
wither immediately after anthesis, a feature reflected
in the specific epithet. The flowers are yellowish and
the ripe drupes purple-black.

34. Kibara novobritanica PHriLipson, Blumea 30
(1985) 411.

Small tree up to 16 m high, glabrous. Leaves ob-
long to broadly elliptic, 16—20 by 7.5—10 cm, charta-
ceous, base broadly cuneate to rounded, apex with a
short obtuse (or occasionally acute) apiculum, entire,
midrib prominent, principal veins arched ascending
and meeting near the margin; petiole c. 10 mm.
Monoecious. /nflorescences terminal, axillary or
supra-axillary, short pleiochasia often densely aggre-
gated together at the nodes, rachis short (up to 30
mm long) with a few pairs of lateral branches which
may branch again, pedicels thickened for about 10
mm below the receptacle. — Male receptacle subglo-
bose, c. 2 mm 9; tepals 6, rounded; stamens usually
6 (in 3 pairs, the inner small, the outer inserted well
above the base of the cavity); anthers small subses-
sile. — Female receptacle similar but larger, c. 4 pairs
of tepals with swollen glands within the ostiole; car-
pels numerous, glabrous with an obtuse stigma.
Drupes + sessile, ovoid, c. 13 by 18 mm, glabrous;
fruiting receptacle c. 8 mm @, woody.

302

FLORA MALESIANA

[ser. I, vol. 102

Fig. 16. Kibara fugax Puitipson. a. Leafy shoot bearing male, female and mixed inflorescences, 1/2, b.
male flower, c. the same in LS, both x7, d. stamens, x10, e. female flower, f. the same in LS, both x7,
g. carpel, x10 (NGF 14583).

Distr. Malesia: Papua New Guinea (W. & E.
New Britain; New Ireland).

Ecol. Lowland rain-forest to 100 m.

Vern. Gangnan, Puhi River, Kandrian Distr.,
napeewa, Talasea.

Note. Confined to New Britain and New Ireland,
and the only species of Kibara known from that re-
gion. The thickened pedicels are characteristic. The
ripe drupes are black on an orange torus.

35. Kibara rigidifolia A.C. SmirH, J. Arn. Arb. 22
(1941) 243; PHmrpson, Blumea 30 (1985) 412.
Small tree to 8 m, sparsely branched; young
branches glabrous, flattened or triangular. Leaves
opposite or in whorls of 3, elliptic-oblong or lanceo-
late-elliptic, up to 30 by 11.5 cm, base broadly cu-
neate, apex rounded or obtuse with a glandular mu-
cro, coriaceous, light yellow-green when dry, entire
or with acute dentations, midrib prominent beneath,

principal veins numerous, at first straight but arched
and uniting within the margin, glabrous; petiole
13—25 mm long, stout, slightly channelled above.
Dioecious (according to Brass). Inflorescence axil-
lary, the male flowers in panicles c. 40 mm long, the
female pleiochasia up to 60 mm. — Male receptacle
globose, 4—5 by 3—4 mm, tepals 8, obtuse (the inner-
most pair + resemble staminodes); stamens 3-6,
1.5—1.75 mm long, with broad vertical anthers and
a short filament, glabrous. — Female receptacle glo-
bose, 5 by 3.5 mm; tepals, minute, obtuse, with swol-
len glands within; carpels c. 30, glabrous, with a
cushion-shaped stigma. Peduncle and pedicels be-
coming thicker and woody in fruit; receptacle leath-
ery, c. 10 mm 9, bearing short stipes. Drupes ovoid,
12—16 by 8—10 mm.

Distr. Queensland and Malesia: Papua New
Guinea (Western Prov.: Morehead & Balimo Distr.).

Ecol. In coastal scrub or undergrowth of forest

1986 |

near sea-level, on sandy soil and on shaded cliff face.

Note. A sparsely branched shrub or small tree
with pale, shining, coriaceous leaves which are fre-
quently arranged in whorls of three on triangular
stems. Older branches fawn, with soft fissured corky
bark. The drupes are black on an orange receptacle.
The number of stamens is variable, even on the same
plant. They are not arranged irregularly over the re-
ceptacle wall (as in Wi/kiea) but form a central
group. When the number of stamens is reduced, the
inner tepals may resemble staminodes. The form of
the stamens differs from that of most species of Ki-
bara as they are held vertically with the two lips of the
horseshoe-shaped slit + equal (in most species the
anther inclines towards the centre with the lower lip
smaller than the upper).

36. Kibara sudestensis PHiLIpson, Blumea 30 (1985)
412.

Shrub or small tree, c. 4m, glabrous in all its parts,
except that the buds and young foliage are covered in
appressed silky hairs. Leaves narrowly elliptic, up to
17.5 by 5.8 cm, stiffly coriaceous, narrowed to a
truncate base, apex narrowed or slightly apiculate,
margin entire or one or a few teeth near the apex,
midrib prominent, lateral veins numerous, running
straight from the midrib at a slight angle, uniting
near the margin; petiole c. 10 mm, channelled above.
Inflorescences axillary or supra-axillary, one- or few-
flowered cymes, c. 25 mm long; pedicels widening in-
to the base of the flower. — Male receptacle obo-
void, up to c. 6 mm long, glabrous; tepals 6, cavity
small, enclosing 4 large and 2 smaller central sta-
mens, anthers kidney-shaped; filaments short, nar-
rower than the anthers. — Female flowers and fruits
unknown.

Distr. Malesia: Papua New Guinea (Louisiades
& Sudest [.).

Ecol. A small undergrowth tree on ridge crest in
rain-forest, at 150 m.

Note. The narrowly elliptic, rigid leaves, and the
large male flowers with small kidney-shaped anthers
are distinctive.

37. Kibara sleumeri PHiipson, Blumea 30 (1985)
413.

Small tree, 3 m, glabrous. Leaves oblong, 14—20
by 6—11 cm, chartaceous, base rounded or broadly
cuneate, apex rounded with a short apiculum, entire,
principal veins prominent below and meeting inside
the margin, glabrous; petiole to 22 mm, channelled
above. Inflorescences axillary or arising above the
foliage leaves, at first setulose becoming glabrous at
maturity, one or more few-flowered cymes arising
together, peduncles slender, 15-20 mm, with basal
"bracts and a pair of bracts towards the middle; pedi-
cels c. 5 mm, slightly swollen below the female flow-

MONIMIACEAE (Philipson)

303

ers. — Male receptacle obovoid, c. 2.5 mm long, with
6 stamens, the central pair reduced. — Female recep-
tacle subglobose, c. 3 mm long; tepals 4, rounded,
enclosing swollen glands; c. 10 carpels, very faintly
pubescent, stigma short. In fruit the peduncles and
pedicels become slightly woody, c. 45 mm long; the
receptacle in fruit rather small (c. 10 mm @) with
prominent stipes c. 3 mm long. Drupes ovoid, 16 by
10 mm.

Distr. Malesia: West New Guinea (Vogelkop
Peninsula).

Ecol. Dense
450—600 m.

Note. The foliage is similar to that of K. moluc-
cana, but the compact subfasciculate inflorescences
of that species are distinctive. The flowers are pale
yellow and the drupes black on a swollen orange re-
ceptacle.

shade in Castanopsis forest,

38. Kibara monticola PERKINS, Pfl. R. Heft 49 (1911)
32; Puitipson, Blumea 30 (1985) 414.

Small tree, with glabrous branchlets. Leaves nar-
rowly elliptic, 13—20 by 4—6.5 cm, membranaceous,
base cuneate, apex apiculate, entire, midrib promi-
nent, principal veins few, arched and meeting inside
the margin, glabrous; petiole 8-10 mm, slightly
channelled above. Monoecious. /nflorescences axil-
lary or supra-axillary, dichasial or small pleiochasia,
glabrous. — Male flowers ovoid, 2.5 mm long; tepals
4, rounded; stamens 6, the innermost 2 smaller. —
Female flowers ovoid, 2.5—3.5 mm g, pedicels 10—15
mm long; tepals with 4 swollen glands within; carpels
10—15, slightly pubescent with a short stigma. Fruits
not seen.

Distr. Malesia: Papua New Guinea (Sepik re-
gion: Ibo Mts). Only known from the type.

Ecol. In forests at 110 m.

39. Kibara symplocoides Perkins, Bot. Jahrb. 52
(1915) 210; PHitipson, Blumea 30 (1985) 414.

Small tree to 8 m, glabrous. Leaves elliptic to nar-
rowly elliptic, 7-12 by 2.5—4.5 cm, chartaceous,
base cuneate, apex narrowly acuminate, entire or
with a few small remote dentations in the upper half,
the midrib and principal veins deeply immersed
above, prominently raised below, the lateral veins
arched and uniting within the margin, glabrous; peti-
ole 4—6 mm. ?Monoecious. — Male inflorescences
axillary or supra-axillary, fasciculate or solitary, cy-
mose, to 10 mm long, with small ovate bracts. —
Male receptacle ovoid, 1.5—2 mm long; tepals 6; sta-
mens 2, subsessile. — Female inflorescences race-
mose or flowers solitary, axillary or supra-axillary,
to 35 mm long, with lanceolate bracts. — Female re-
ceptacle globose, 4 mm, ostiole with minute tepals
and swollen glands within; carpels numerous, pilose.
Fruit not seen,

FLORA MALESIANA [ser: I, vol

10?

Fig. 17. Kairoa suberosa PHILIPSON
male flowers, nat. size, e. male flow

7 <2/2e
x1 1/2 (a HartTLeEy 12697, b, dNGF
ON 3681, f PHILIpson 3684, g LAE 56322).

1986 |

Distr. Malesia: Papua New Guinea (Sepik re-
gion). Only known from the two original collections.

Ecol. In open mountain forest, about 20 m high,
at 1000 m altitude.

Insufficiently known

The types of the following species have not been
seen, and it is not possible to determine from the de-
scriptions whether they correspond to any known
species or are distinct:

K. olivaeformis Becc. Malesia 1 (1877) 187.

K. aruensis Becc. lI.c. 188.

K. formicarum Becc. l/.c. 188.

K. perkinsiae K. Scu. & Laut. Fl. Deut. Schutz-

MONIMIACEAE (Philipson)

305

Inadequately represented

The collections listed below appear to represent
undescribed species of Kibara, but the material is in-
adequate because no flowers are present.

BARKER LAE 67616, Telefomin Distr.; EymMa
4555, Wissel Lake region; FOREMAN LAE 52220, Ra-
mu Distr.; KOSTERMANS & SOEGENG 853, Baliem Val-
ley; VAN ROYEN & SLEUMER 7773, Vogelkop Penin-
sula.

Excluded species

Kibara borneensis BoERL. = Pycnarrhena (Meni-

geb. Stidsee (1900) 330. spermaceae).
K. elmeri Perxtns, Bot. Jahrb. 45 (1911) 424. Kibara_ timorensis Boert. = Pycnarrhena
K. buergersiana PERKINS, Bot. Jahrb. 52 (1915) (Menispermaceae).
209.
K. neriifolia PERKINS, /.c. 212.
8. KAIROA

PuHiLipson, Blumea 26 (1980) 368, f. 1-10. — Fig. 17, 18.

Small tree or sparsely branched shrub with toothed leaves and flowers in axil-
lary fascicles. Flowers monoecious. — Receptacle of male flower at first globose
with a small ostiole bounded by 8 tepals (in two double pairs), at anthesis split-
ting to form an open bowl-shaped flower with 4—6 radiating lobes, fleshy. Sta-
mens very numerous, inserted over the surface of the receptacle; anthers open-
ing by two longitudinal slits; filament very short. — Receptacle of female flower
oblate, with an ostiole bounded by 4 tepals, splitting into broad segments; the
upper part of the receptacle abscissing by a circular split after anthesis. Carpels
numerous, sessile, with a very short style. Fruits numerous, ovoid, sessile on an
enlarged fleshy receptacle.

Distr. Malesia: Papua New Guinea (Morobe and Northern Provinces, from Finschhafen to Tufi Distr.).

Monotypic. Fig. 10.
Ecol. Understorey of rain-forest.

1. Kairoa suberosa Puitipson, Blumea 26 (1980) 368,
f. 1-10. — Fig. 17, 18.

Shrub or small tree with stout terete branches,
bark becoming corky. Leaves coriaceous, up to 45 by
17 cm, oblong, elliptic or lanceolate-oblong; base
cordate, truncate or cuneate, apex narrowed to an
acute apex; lower surface with dense or sparse pubes-
cence or glabrous; margin with few to many sharp
teeth; midrib prominent; secondary and tertiary
veins forming a distinct reticulation; petiole stout,
§—15 mm. Inflorescence of axillary fascicles which
continue to appear from successive buds at older
nodes; fascicles with a short peduncle bearing crowd-
ed minute bracts, the lower sterile, the upper with

flowers in their axils. — Pedicels of male flowers
fleshy (when fresh), c. 15-20 mm, sometimes with
solitary or paired bracteoles, expanded above; recep-
tacle at first globose with a terminal depression and
an ostiole with 8 tepals in two double pairs, expand-
ing to c. 18 mm @ before opening by 4—6 radial splits
to form a star-shaped flower with the stamens fully
exposed. Stamens numerous (over 100), inserted over
the lower half of the receptacle, filament short; an-
thers to 1 mm long, opening by two longitudinal slits.
— Pedicels of female flowers shorter (c. 5 mm at an-
thesis); receptacle globose, smaller (c. 8 mm @), tep-
als 4, the upper half of the receptacle becoming de-
tached after anthesis by a circular scar. Carpels nu-

306 FLORA MALESIANA [ser. I, vol. 102

Fig. 18. Kairoa suberosa PHiLipson. A group of ripe infructescences (the fruits in this cluster were borne on
three closely associated receptacles), 3/4 (PHILIPSON 3626).

merous (c. 50 or more), sessile on the lower half of
the receptacle, with a short, blunt style. Receptacle
becoming enlarged and fleshy in fruit. Drupes ses-
sile, numerous, ovoid, 20—24 by 11-14 mm.

Distr. Malesia: Papua New Guinea (Morobe &
Northern Prov.). Fig. 10.

Ecol. Locally frequent in the understorey of low-
land rain-forest and occasionally ascending into lo-
wer montane forest to 1200 m, with Castanopsis and

Note. The specific epithet refers to the thick
deeply fissured bark of the older stems, by which fea-
ture this species can most readily be detected in the
field. The stiff leaves, which are usually sharply
toothed, are also distinctive. The male flowers are
soft, fleshy, cream-coloured often flecked with vio-
let. They are rather large for the family. The succu-
lent, fleshy fruiting receptacle is orange and the ripe
drupes are black and shining.

Lithocarpus dominant.

9. STEGANTHERA

PERKINS, Bot. Jahrb. 25 (1898) 564; Pfl. R. Heft 4 (1901) 52; ibid. Heft 49
(1911) 20; Bot. Jahrb. 52 (1915) 197; Ubersicht Gattungen Monim. (1925) 33;
PHILIPSON, Blumea 29 (1984) 486, f. 1. — Anthobembix Perkins, Bot. Jahrb.
25 (1898) 567. — Fig. 19-23.

Trees or shrubs, resting buds with cataphylls. Leaves pubescent at first, some-
times becoming glabrous, entire or dentate, principal secondary veins arched
and meeting within the margin. Monoecious or dioecious. /nflorescences later-
al, cymose or fasciculate. — Male flowers globose, turbinate or patelliform,
with 4 small tepals, stamens (3—)4(—5), or rarely with a second (inner) whorl
of 1—4 smaller stamens; anthers opening by a single horizontal slit. — Female
flowers similar to male but larger and with a smaller ostiole surrounded by 4 tep-

1986 | MONIMIACEAE (Philipson) 307

Fig. 19. Steganthera fasciculata Puipson. a. Habit, with subumbellate male flowers, x 1/2, b. male flowers,
-one in LS, «5, c. female inflorescence, d. infructescence with 4 stipitate achenes, both 1/2 (@ PHILiPsON
& Kaino 3642, b ditto 3643, c ditto 3641, d Pratt 1082).

308 FLORA MALESIANA [ser. I, vol. 102

als, the upper half abscissing as a calyptra after anthesis to reveal numerous car-
pels; ovary pubescent; style glabrous awl-shaped. Drupes sessile or long-
stipitate.

Distr. Queensland, Solomons and Malesia: Celebes, Moluccas, New Guinea (incl. Bismarck Archipel-
ago).

Notes. Steganthera is frequently confused with Kibara, another common and widely distributed genus,
because the foliage, fruits and inflorescences are similar. The technical differences between these two genera
are not readily observable. They are, firstly, the greater number of tepals in Kibara, certain of which are thick-
ened and glandular in the female receptacle, and, secondly, the greater number of stamens in the androecium
(but see S. salomonensis). However, other features usually allow a generic identification to be made. The most
important of these is the appearance of the pedicel: in Steganthera the pedicel is relatively slender and is clearly
distinct from the receptacle, whereas in Kibara the pedicel is thickened distally and has a less clearly defined
junction with the receptacle or none at all. Another useful indication is that the leaves usually dry greenish
(often yellowish green) in Kibara, whereas in Steganthera they are mostly brown when dry.

The species of Steganthera can be placed in three groups on morphological grounds. One is characterized
by the expansion of the female, and to a lesser extent also the male flower, into a turbinate or patelliform
receptacle. This group formerly was segregated as the genus Anthobembix, but KANEHIRA & HATUSIMA (Bot.
Mag. Tokyo 56, 1942, 256) set out valid reasons for uniting the two genera. A second group of species is char-
acterized by their fasciculate inflorescences and by being dioecious. The third group comprises those species
lacking both these characters; that is, the species have subglobose receptacles in cymose inflorescences. It in-
cludes a number of species which are difficult to separate, including S. hirsuta which is very variable in all
its characters. The species of the first two groups are well defined morphologically and geographically, where-
as those of the third group form a complex which has not yet resolved itself into stable species. These three
groups of species are not given formal taxonomic recognition because their limits are not clearly defined. Sée-
ganthera dentata and S. cyclopensis combine fasciculate inflorescences with discoid receptacles and therefore
fall into both of the first two groups. In some species the expansion of the receptacle is a variable character;
for example, in S. hospitans and S. oligantha the male flower may be globose, but this may be due to hybridity
with S. hirsuta.

The association of some species with ants is very striking. These species have enlarged nodes with well-
defined pores leading into the hollow stems. Scale insects line the pith cavities and small black ants abound
over the piant surface and in the hollow stems. The association is most conspicuous in the large-leaved and
abundant species S. hospitans, but also occurs in S. moszkowskii, S. ledermannii and S. royenii.

Pollination biology has not been studied and the structure of the receptacles poses several problems. The
male receptacles have open, if small, ostioles at the time the anthers dehisce, but it is not known what insects
visit them. The carpels in the female receptacles are covered by the ‘calyptra’ at anthesis. The ostiole giving
access to the female receptacle is even smaller than that of the male and it is not known how pollen reaches
the stigmas. These are awl-shaped and converge towards the ostiole. In Kibara and some other genera pollen
is received on a hypostigma outside the receptacle (ENDREss, Experientia 35, 1979, 45) but this is not so in
Steganthera.

KEY TO THE SPECIES
lpaRemalemmibonescenGe; SUDSESSIIG.. anys aic-cis cis bra Ole See MEM o soainoe ev coves cel ee 16. S. insculpta

1. Female inflorescence pedunculate.
2. Male inflorescence umbellate.

3. Leaves elliptic to broadly elliptic (mostly under 10 cm long: W. Sepik Prov.).......... 1. S. hentyi
3. Leaves oblanceolate to narrowly elliptic (mostly longer than 10 cm).
4. Male receptacle globose (Papua New Guinea, east of 146°30’ E)................ 2. S. fasciculata
4. Male receptacle discoid.
Sscavesananrowenmthamid Suiits (Wr gla) eee helene apse) oterale/eteie. ig) eleva acral oe auetedlyaucrts 3. S. cyclopensis
Seeley CSRWA GOTH UN Allis a Clit (VV en LITA) er ove ravers. sits teorapelsusioeedcwsucacwen ty crari chalne Sei cick ares eee 4. S. dentata

2. Male inflorescence of dichasia, pleiochasia or panicles.
6. Pedicels of male flowers mostly in irregular clusters.
7. Nodes dilated, with a pore inhabited by ants (W. Irian).

1986] MONIMIACEAE (Philipson) 309
Saale weceptacleclohoseesntee siete « . ochter oemleiase Sade ioe et aed 5. S. royenii
8. Male receptacle turbinate with a 4-lobed apical disc ........................ 6. S. moszkowskii

7. Nodes not harbouring ants.
9. Leaves large’. Inflorescence large with stout pedicels (Solomon Is. and Papua New Guinea east of
PESO SE) ee SE eee EERIE. . 5, See, Leaet est) sony eae, 4 7. S. salomonensis
9. Leaves smaller (to 10 cm long). Inflorescence small, with slender pedicels (Wau, Goilala & Moresby
ENTS OOS Serene SRR SE BA gtd Bt. os w Semi enoeies) » Saye ack eras 15. S. australiana
6. Pedicels of male flowers mostly solitary.
10. Receptacles discoid or turbinate.
i Leaves 20 cmlong or longer (widely distributed) 2c.) to. J as 8. S. hospitans
11. Leaves considerably shorter.
12. Male inflorescence 1—2-flowered. Leaf 6 cm long or shorter (W. Irian, Idenburg R.) 9. S. myrtifolia
12. Male inflorescence with more flowers. Leaf longer than 6 cm.
Se Maleireceptacle turbinate: (W !\Iniany W..sSepik) ise ae ee, Le ee
Peamale xeceptacie’patelliform) (Central! Prov.) 2.203). ee 8a OG ee eee
10. Receptacles globose (or only slightly depressed).
14. Leaves usually moderate to large (10—32 cm long) and entire (widely distributed, mainly at lower alti-
ides but reaching Upto 2450 mM), <0 e ae os skeet ee eee 12. S. hirsuta
14. Leaves variable in size (usually under 15 cm long, but up to 21 cm), usually coriaceous, often dentate,
but also entire when narrow (widely distributed, usually above 1200 m)......... 13. S. ilicifolia
14. Leaves S—10 cm long, coriaceous, + rugose, entire (Chimbu & Eastern Highlands Prov., above c.
UTD) isk Sa CLASES AG dats Sets oe ARETE Rein wee, Se 14. S. chimbuensis
14. Leaves 5—10 cm long, chartaceous, usually narrowly elliptic, acuminate, entire or dentate (Wau, Goi-

10. S. parvifolia
11. S. oligantha

See eADAOLESDY: PEOV2) Gras sce oe ee. «

1. Steganthera hentyi PHiLipson, Blumea 29 (1984)
487.

Small tree, to 6 m; young branches densely strig-
ose. Leaves elliptic, c. 8—9 by 3—4 cm, base cuneate,
apex with a short obtuse apiculum, margin entire or
with few obscure teeth, becoming glabrous on the
leaf surface but with appressed hairs persisting at
least on the principal veins; petiole to 8 mm. ?Dioe-
cious. Inflorescence axillary or above the axils, of
sessile or shortly pedunculate subumbellate fascicles;
peduncle to 10 mm, pedicels 20—30 mm, slender, stri-
gose, subtended by minute subulate bracts. — Male
flowers unknown. — Female receptacle subglobose,
c.3—4mm @ at anthesis; tepals 4, rounded, outer sur-
face densely strigose except for the upper part, inner
surface with long bristles between the carpels; carpels
c. 15, ovary covered in dense shaggy hairs, style awl-
shaped, glabrous. Drupes ovoid, 12 by 7 mm, with
appressed hairs, short stipitate.

Distr. Malesia: Papua New Guinea (West Sepik
Prov.).

Ecol. Primary lower montane valley forest, at
500 m.

Note. The small leaves with appressed hairs on
the veins of the underside of the leaf, and the fascicu-
late inflorescences are distinctive. The flowers are de-
scribed as pink.

15. S. australiana

2. Steganthera fasciculata PHiLIpson, Blumea 29
(1984) 487, f. 3. — Fig. 19.

Small tree, to 12 m; young branches pubescent.
Leaves oblanceolate to narrowly elliptic, 13—20(—30)
by 3.4—7(—9) cm, base rounded to broadly cuneate,
apex narrowed toa short obtuse, or long acute apicu-
lum, margin entire or dentate especially in the upper
half (leaves of juvenile plants or of suckers more
prominently dentate), pubescence may persist on ma-
ture leaves or may be lost except along the veins; peti-
ole to 10 mm, pubescent. /nflorescences axillary or
supra-axillary; male of sessile or shortly pedunculate
fascicles with 10—20 flowers, often appearing as ses-
sile umbels, but sometimes seen to consist of dense
fascicles of pleiochasia, the principal branches
10-30 mm, tenuous, hairy; female solitary or in
pairs, axillary or supra-axillary, pedicel 20—30 mm
with subulate bracts at the base and sometimes about
the middle, pubescent. — Male receptacle globose,
outside pubescent, c. 4-5 mm @; tepals 4, minute;
stamens 4, c. 1.5 mm long, filament short, hairy, the
anthers 1.5 mm broad. — Female receptacle bowl-
shaped, c. 12 mm @ at anthesis; tepals 4, carpels nu-
merous, ovary hairy, style awl-shaped glabrous.
Drupes not seen. The mature torus with many protu-
berant pubescent carpophores c. 4—7 mm long.

Distr. Malesia: Papua New Guinea (Morobe,

(1) Rarely (in the Solomon Is.) small leaves occur on stunted individuals.

FLORA MALESIANA

[ser. I, vol. 102

310
Northern & Milne Bay Prov.), also in Goodenough I.
Ecol. Primary lower mountain rain-forest,

365—1200 m.

Vern. Mamkananeh, Daga lang.

Note. The young flowers are described greenish
cream and the mature torus red.

3. Steganthera cyclopensis PHiLIPSON, Blumea 29
(1984) 489.

Shrub to 3 m; young branches with strigose ap-
pressed hairs. Leaves oblanceolate to narrowly ellip-
tic, up to 17 by 4.2 cm, base truncate or rounded,
apex shortly apiculate, margin dentate, when dry
grey above, brown below, principal veins prominent,
pubescence persisting on the lower midrib and main
veins; petiole to 5 mm, hairy. ?Dioecious. /nflores-
cences axillary or above the axils, sessile fascicles of
c. 12 flowers, pedicels slender, densely strigose, c. 15
mm. — Male receptacle discoid or bowl-shaped, the
stamens in a small central cavity, c. 8 mm g, pubes-
cent on the outer surface of the disc; tepals 4; sta-
mens 4, c. 1 mm broad, with a very short broad fila-
ment, connective expanded as two wings bordering
the small narrow anthers. — Female flowers not
seen, but fewer per inflorescence. Fruiting receptacle
slightly enlarged, not succulent, on stout pedicel c. 15
mm, upper surface with long hairs between the bases
of the apparently sessile achenes. Drupes c. 13 by 9
mm with a few persistent appressed hairs.

Distr. Malesia: Northern District of West Irian.

Ecol. In primary Nothofagus lower montane for-
est, 1200—1340 m.

Note. Similar to S. dentata in many respects, but
with smaller usually lanceolate leaves. The flowers
are described as carmine and pale orange.

4. Steganthera dentata (VALETON) KANEH. & HATUS.
Bot. Mag. Tokyo 56 (1942) 257; Puitipson, Blumea
29 (1984) 489. — Anthobembix dentatus VALETON,
Bull. Dép. Agric. Ind. Néerl. 10 (1907) 13.

Shrub to 4 m, with spreading branches; young
branches fulvous pubescent, older branches with
swollen nodes. Leaves c. 5 mm petioled or leaves sub-
sessile, pubescent, oblanceolate or elliptic, 16—30 by
7—11 cm, chartaceous, base + auriculate or trun-
cate, apex apiculate, margin with well-spaced, small
dentations especially on the distal half; young leaves
with pubescence which may persist on the mature
leaves or may be lost except along the veins and
around the base. Probably dioecious. Inflorescences
usually inserted well above the nodes (occasionally at
the nodes); male sessile or pedunculate fascicles of
10—25 pedicellate flowers, peduncle (if present) up to
15 mm; pedicels 10—20 mm, slender, pubescent; fe-
male fascicles few-flowered. — Male receptacle
bowl-shaped or discoid, outside hirsute, c. 6-10 mm
@; tepals 4, minute; stamens 4, 1.5 mm long, filament

short, hairy, connective broader than the anthers
(before dehiscence). — Female receptacle similar to
male, carpels numerous, hairy, style awl-shaped, gla-
brous. Fruit unknown.

Distr. Malesia: West New Guinea (Northern &
Southern Distr.).

Ecol. Primary and secondary forest, at low alti-
tudes (to 160 m).

Note. The short-petioled or subsessile leaves and
fasciculate male inflorescences are distinctive. The
flowers are described as red, orange or yellow. The
anthers are considerably narrower than the connec-
tive except in one collection where the more widely
gaping anthers may be at a stage after anthesis with
the lateral lobes of the connective no longer turgid.
It has not been possible to determine whether male
and female flowers occur on the same or on separate
plants.

5. Steganthera royenii PHiLIPSON, Blumea 29 (1984)
489.

Shrub c. 4 m; young branches pubescent; nodes
becoming dilated and developing pores. Leaves
oblong-elliptic, up to 29 by 11 cm, chartaceous, base
cuneate, apex acuminate, margin entire or with min-
ute remote dentations, principal veins channelled
above, prominent below, arched ascending, becom-
ing glabrous or with a minute puberulence persisting
mainly on the veins; petiole 5-10 mm. ?Monoecious.
Inflorescences supra-axillary or axillary, broadly
paniculate, 4—5 cm long, single or grouped, puberu-
lent, rachis and principal branches somewhat thick,
ending in irregularly clustered slender pedicels. —
Male receptacle globose, 2—2.5 mm, sparsely puber-
ulent on the outside; tepals 4, minute, rounded; sta-
mens 4, | mm long, filament broad, pubescent. —
Female flowers not seen. Fruiting receptacle 2—3 mm
9, leathery. Drupes c. 12—20, ovoid, c. 12 by 10 mm,
verruculose, stipitate; stipes 2-3 mm long, 2.5mm@.

Distr. Malesia: West New Guinea (a restricted
area which includes portions of the Vogelkop,
Northern & Southern Distr.).

Ecol. Low-lying primary forest, sometimes peri-
odically flooded, from sea-level to 1200 m.

Vern. Sirochomenwhah, Manikiong dialect.

Note. Similar to S. hospitans in the appearance
of its vegetative parts, but the arrangement of the ul-
timate branches of the inflorescence is distinctive.
The male flowers lack any disk-like extension of the
receptacle but female receptacles have not been seen.
This species is described as possibly monoecious be-
cause one collection bears male inflorescences and
also old fruits. However, as these are not on the same
branches, the monoecious condition is not certain.
All the inflorescences seen consist solely of male
flowers, which suggests the possibility of the sexes
being on separate plants. The flowers are described

1986 |

as yellow, the ripe fruit black. The swollen nodes are
inhabited by ants.

6. Steganthera moszkowskii (PERKINS) KANEH. &
Hatus. Bot. Mag. Tokyo 56 (1942) 255; PHitipson,
Blumea 29 (1984) 490. — Anthobembix moszkowskii
Perkins, Pfl. R. Heft 49 (1911) 26.

Shrub or small tree; branches becoming glabrous;
nodes swollen. Leaves elliptic-ovate to 35 by 12 cm,
chartaceous, base broadly cuneate, apex apiculate,
margin entire or remotely dentate, glabrous above,
minute pubescence persisting below, especially on
the veins, the midrib and principal veins prominent
below; petiole 5—8 mm long, minutely pubescent.
?Dioecious. Inflorescences at or well above the
nodes, diffuse paniculate, to 10 cm long; rachis and
branches minutely pubescent. — Male receptacle tur-
binate, pubescent, 3 mm high, apex forming a
4-lobed disk; tepals 4, minute; stamens 4, filament
broad, hairy, the united anthers small. — Female
flower and fruit not seen.

Distr. Malesia: West New Guinea (Northern
Distr.).

Ecol. Understorey in rain-forest at low altitude.

Note. The male receptacle, with 4 rounded lobes
radiating from the apex, is distinctive. The swollen
nodes are apparently inhabited by ants which gain
entrance through pores.

7. Steganthera salomonensis (HEMSL.) PHILIPSON,
Blumea 29 (1984) 490, f. 2. — Hedycarya salomo-
nensis HEMSL. Kew Bull. (1895) 137. — S. suberoso-
alata KosterM. Gard. Bull. Sing. 22 (1968) 445. —
Fig. 20.

Shrub or tree, 2—20 m; young branches with a
crisp, brownish tomentum or a fine, appressed pu-
bescence which may be quickly lost. Leaves ovate or
elliptic to narrowly elliptic, rarely suborbicular (Fer-
gusson I.), usually rather large (up to 32 by 11.5 cm),
occasionally smaller to much smaller (5 by 2 cm on
stunted trees, crater rim New Georgia Group), base
cuneate to rounded, apex slightly apiculate; margin
entire or rarely a few obscure dentations (even large
serrations on saplings, and on flowering branches in
Sudest I.), with either a tomentum of crisp brownish
hairs which may persist on the undersurface of the
leaves especially on the midrib and principal veins, or
with a fine pubescence which is soon lost, the mature
leaves then becoming glabrous; petiole stout, usually
c. 5—18 mm, but occasionally shorter, pubescent or
becoming glabrous. Monoecious. /nflorescences on
leafy shoots or cauliflorous, variable in size and com-
plexity, frequently with several main branches aris-
ing from a short peduncle, and with the lateral
branches clustered at the nodes, the principal

“branches often flattened; all parts pubescent when
young but the branches may become more or less gla-

MONIMIACEAE (Philipson)

‘

311

brous; the panicles may contain only male flowers
when the female flowers are in smaller inflorescences
(even solitary) or the female flowers may occur in
panicles of mainly male flowers (apparently usually
on terminal branches). — Male flowers ovoid or sub-
globose, c. 3 mm @, outside pubescent; tepals 4,
rounded; stamen number very variable, usually in
two whorls, the outer of larger stamens (3—)4(—5),
2—2.5 mm long, filament hairy, the inner of smaller
stamens 1—4 or absent (male receptacles 4 mm @ with
broad stamens occur in Rennell Is.). — Female flow-
ers subglobose or bowl-shaped, pubescent outside,
pilose within, c. 3—6 mm @ at anthesis but swelling to
c. 7-10 mm before the calyptra is shed; ostiole very
small with 4 minute tepals; carpels numerous, c. 2
mm long, ovary hairy. Peduncle and receptacle
usually becoming enlarged and woody in fruit with
numerous drupes borne on long woody stipes, but
frequently only few (even only 1) fruits ripen, when
the receptacle and peduncle are little enlarged; recep-
tacle up to 3 cm 9; stipes up to 14mm. Drupes ovoid,
12—18 by 10—15 mm, verruculose.

Distr. Throughout the Solomon Is. (incl. Bou-
gainville) to San Cristobal and Rennell I., in Malesia:
Papua New Guinea (Central Prov.; east of Cape
Rodney; Milne Bay Prov.).

Ecol. Primary and secondary lowland rain-
forest; in scrub on coral debris and in stunted forest
on ridges, from sea-level to 750 m.

Note. Considerable variation occurs, one ex-
treme being a handsome tree with large leaves, +
brown-pubescent below, diffuse panicles on leafy
twigs and also on the older stems, and with clusters
of many stipitate achenes (=S. suberosa-alata). On
the other hand many specimens, including the type,
have smaller, greener leaves which become more or
less glabrous, and have fruits with few or even a
single stipitate achene. The latter form is characteris-
tic of higher altitudes. Specimens from a dry crater
rim in the New Georgia Group are so stunted that
their inclusion in this species is open to doubt,
though intermediate forms occur. A number of local
forms are found on the islands off the eastern end of
New Guinea (Fergusson I. to Misima I.). The most
distinctive of these, with coarsely serrate leaves, oc-
curs on Sudest I. This may represent a distinct spe-
cies, but serrate leaf-margins occur elsewhere, espe-
cially on saplings or suckers, and the flowers and in-
florescences are quite typical of S. salomonensis.

8. Steganthera hospitans (Becc.) Kangen. & Hatus.
Bot. Mag. Tokyo 56 (1942) 255, f. 5; Puttipson, Blu-
mea 29 (1984) 491. — Kibara hospitans Becc. Male-
sia 1 (1877) 189. — Anthobembix hospitans (Becc.)
Perkins, Bot. Jahrb. 25 (1898) 567, t. 6B; K. Scu. &
Laur. Fl. Deut. Schutzgeb. Stidsee (1900) 330; Per-
KINS & Gia, Pfl. R. Heft 4 (1901) 55, f. 12; PERKINS,

312 FLORA MALESIANA [ser. I, vol. 102

Fig. 20. Steganthera salomonensis (HEMSL.) PHILIPSON. a. Cauliflorous panicle, with several principal ped-
uncles, central rachis ending with two female flowers, b—c. two male inflorescences, d. purely female inflores-
cence. All x1/2 (a BSIP 10575, b NGF 28733, c—d ScHODDE 5388).

Pfl. R. Heft 49 (1911) 25, f. 10; Bot. Jahrb. 52(1915) — the nodes soon becoming dilated, and pores develop-
205, f. 4. — S. insignis Perkins, Pfl. R. Heft 49 ing. Leaves oblong-elliptic, or broadly elliptic to ob-
(1911) 24, f. 9. — Fig. 21. ovate, coriaceous, 20—41 by 7—18 cm; base cuneate

Shrub or small tree to 8 m; young branches more _ or rounded, apex apiculate to long acuminate, mar-
or less glabrous, or pruinose or minutely puberulous; _—_ gin entire or occasionally with well-spaced small den-

1986]

MONIMIACEAE (Philipson)

Fig. 21. Steganthera hospitans (BECC.) KANEH. & Hatus. Panicle of female flowers. Note ants on blade and
thickened petiole; one flower attacked by mould (after PHitIpson & KATIK 3737).

tations, glabrous or minutely pubescent on the mid-
ribs and principal veins, veins impressed above,
prominent below. Foliage of juvenile plants lanceo-
late, c. 27 by 4 cm, base narrowly cuneate, apex at-
tenuate, irregularly dentate; petiole to 1 cm. Monoe-
cious. Inflorescence axillary or above the nodes,
arising successively at the same position, becoming
cauliflorous, polychasia or paniculate cymes,
minutely pubescent, up to 9 cm long, rachis rather
stout, often flattened, with small caducous bracts;
pedicels slender, 10-15 mm long. — Male recep-
tacles variable in shape, usually turbinate with an ex-
panded flat rim, or this more or less obsolete, 2.5—5
(or more) mm g; tepals 4, minute, obtuse; stamens 4,
1—1.5 mm high, filament broad, puberulous. — Fe-
male receptacle turbinate with a broadly expanded
flat rim, to c. 12 mm 9; tepals 4, minute, carpels nu-
merous, ovary pilose, style awl-shaped, glabrous
above. Fruit with large succulent receptacle (c. 3—4
cm @) bearing many drupes on short thick stipes.
Drupes ovoid to globose, c. 12 by 10 mm, verrucu-
lose.

Distr. Solomon Is. and Malesia: throughout
New Guinea (from Vogelkop to the east).

Ecol. Primary rain-forest and second growth
from low altitudes up to 1200 m.

Vern. Eppareppar, Adelbert Range, Utu lang.,
soreng, ibid., Rawa lang., oloni, wolonyik, Ambun-
ti, Waskuk lang., wasa, Kutubu lang.

Note. The female receptacles are striking, being
broadly expanded into button-like disks of a soft
butter-yellow colour. The male receptacles are green-
er and smaller, with a variable extension of the rim:
in some species the male receptacles have a broadly
expanded rim, but in others the rim may be obsolete,
as in the specimens identified as S. insignis by PER-
KINS (originally placed by her in Anthobembix hospi-
tans). It is possible that these are hybrids with S. hir-
suta. Sterile juvenile plants with lanceolate leaves are
frequent, and this type of leaf may be found on the
lower parts of more adult shrubs, which have begun
to produce flowers. The hollow stems and swollen
nodes are inhabited by scale insects (Adeyrodidae)
and by many small black ants. The receptacle be-
comes succulent and bright orange in fruit, and the
ripe drupes are purple-black.

9. Steganthera myrtifolia (A.C. SmirH) PHILIPSON,
Blumea 29 (1984) 491. — Anthobembix myrtifolia
A.C. SmitH, J. Arn. Arb. 22 (1941) 238.

Small tree to 4 m; young branches with appressed
strigose hairs. Leaves chartaceous, narrowly elliptic
to elliptic, 3-6 by 1—3 cm, base attenuated, apex
acute or shortly cuspidate, margin entire, glabrous,
midrib well-defined below and with numerous fine
lateral veins and reticulations; petiole 4—6 mm, gla-
brescent. ? Dioecious. — Male inflorescences axil-
lary, 10—30 mm long, 2- or often 1-flowered, with

314 FLORA MALESIANA [ser. I, vol. 102

a —aa B
Fig. 22. Steganthera hirsuta (WaARB.) PERKINS. Branch with axillary pleiochasia; the large flowers are female,
the smaller male; terminal bud not yet developed (after PHILIPSON & KarRo 3657).

1986]

MONIMIACEAE (Philipson)

315

dense appressed hairs, slender; receptacle somewhat
disk-shaped, 2 mm high, 5 mm g; tepals 4, minute;
stamens 4, the longer c. 1 mm longer, filaments
densely strigose. — Female inflorescences similar to
male but apparently always 1-flowered; receptacle
turbinate, slightly expanded at the circumference, c.
2—2.5 mm high, c. 3 mm g, densely strigose within,
tepals obsolete; carpels c. 6, 1.3—1.5 mm long, ovary
densely strigose, style subulate, glabrous. Fruit with
a stout pedicel c. 18 mm; receptacle leathery, only
slightly enlarged. Drupes subsessile, ovoid, c. 13 by
9 mm, verruculose.

Distr. Malesia: West New Guinea (Idenburg R.).

Ecol. Mossy forest, 2150 m.

Note. Known only from the original two speci-
mens collected by Brass. The small, chartaceous, en-
tire leaves are unmistakable among species with a
disk-like male receptacle.

10. Steganthera parvifolia (PERKINS) KANEH. & Ha-
Tus. Bot. Mag. Tokyo 56 (1942) 257; Puiipson, Blu-
mea 29 (1984) 491. — Anthobembix parvifolia PER-
Kins, Bot. Jahrb. 52 (1915) 205. — Anthobembix
brassii A.C. SmitH, J. Arn. Arb. 32 (1941) 239. — S.
brassii (A.C. SmMiTH) KANEH. & Hatus. Bot. Mag.
Tokyo 56 (1942) 261, f. 9.

Shrub, sometimes scandent, young branches pu-
bescent. Leaves oblong-elliptic to narrowly elliptic,
to 14 by 6.5 cm, chartaceous, base rounded or broad-
ly cuneate, apex shortly cuspidate to acuminate,
margin entire or with obscure or more prominent re-
mote teeth in the upper part, becoming glabrous or
with slight pubescence persisting below, the veins
and minor reticulations prominent below; petiole
8—22 mm, pubescent or becoming glabrous. Monoe-
cious. Inflorescences axillary or at the ends of leafy
shoots, of few-flowered cymes or larger paniculate
cymes, to 5 cm long, the rachis, branches and pedi-
cels pubescent; female flowers appear to be confined
to the ends of the lower branches. — Male recep-
tacles obconic with a flat glabrous and black apex (in
dried material) to 8 mm 9g, sides of the receptacle his-
pid; 4 minute tepals forming a raised ostiole; stamens
4, sessile; anthers c. 1 mm wide, connective hairy. —
Female receptacle similar but larger (c. 10 mm @ at
anthesis), inside with long hispid hairs between the
numerous (10—20) carpels; carpels c. 2 mm long,
ovary pilose, style awl-shaped, glabrous. Receptacle
not greatly enlarged in fruit, leathery. Mature drupes
subsessile or stipitate, ovoid, to 12 by 10 mm, apex
mucronate, glabrous, rugose.

Distr. Malesia: West New Guinea (Vogelkop,
Northern & Southern Distr.), Papua New Guinea
(West Sepik Prov.).

Ecol. Understorey of primary lower montane
rain-forest, 1900-2800 m.

Vern. Bobinok, Telefomin Distr., Bulindup.

Note. The glabrous disk-like upper surface of
both male and female receptacles contrasts with the
pilose outer surface, especially in dried material in
which it takes on a black coloration. The material
from the West Sepik Prov. has narrower leaves than
the type, but otherwise conforms. The flowers are
described as yellow.

11. Steganthera oligantha (PERKINS) KANEH. & Ha-
Tus. Bot. Mag. Tokyo 56 (1942) 257; Puiipson, Blu-
mea 29 (1984) 493. — Anthobembix oligantha PER-
KINS, Bot. Jahrb. 25 (1895) 568.

Small to large tree, young branches slender, finely
puberulous. Leaves papyraceous, elliptic-oblong to
obovate, 6—11 by 2.5—5 cm, base cuneate, apex acu-
minate, margin entire, midrib and principal veins
evident, reticulations rather obscure, when young
covered in appressed minute hairs which persist on
the midrib and sometimes on the principal veins; pet-
iole 5—10 mm. ? Monoecious. /nflorescences axillary
or terminal, few flowered pleiochasia, rachis,
branches and bracts pubescent, rachis slender, with
a long peduncle before the first branches, at flower-
ing 4—6 cm long. — Male receptacle turbinate with
an expanded rim, 3 m high, 5—6 mm 4g, the upper sur-
face glabrous, the sides puberulous; tepals minute.
— Female receptacle similar to male, carpels numer-
ous, ovary densely pilose, style awl-shaped, gla-
brous. Fruiting receptacle leathery, c. 10 mm @.
Drupes ovoid, 15 by 10 mm, verruculose, c. 4—5 mm
stipitate.

Distr. Malesia: Papua New Guinea (Central
Prov., Moresby Distr., Sogeri region).

Ecol. Forest, 900—1600 m.

Note. A large tree with slender twigs, known with
certainty only from the three collections made by
Forses in 1855—56 and one recent collection from
Efogi. Other collections from the same region closely
match the foliage and fruit, but the male receptacles
are globose. These may be hybrids with S. hirsuta.
The female flowers are described as bright yellow
and the fruit as black on orange stipes.

12. Steganthera hirsuta (WARB.) PERKINS, Bot.
Jahrb. 25 (1898) 567; K. Scu. & Laut. Fl. Deut.
Schutzgeb. Siidsee (1900) 329; Perkins & GILG, Pfl.
R. Heft 4 (1901) 54; Perkins, Bot. Jahrb. 52 (1915)
202; Puitipson, Blumea 29 (1984) 493, f. 4. — Kibara
(?) hirsuta Wars. Bot. Jahrb. 13 (1891) 316. — S.
warburgii Perkins, Bot. Jahrb. 25 (1898) 564. — S.
schumanniana Perkins, /.c. 565. — S. thyrsiflora
Perkins, /.c. 565, t. 6A; Perkins & Gita, Pfl. R.
Heft 4 (1901) 53, f. 11; Perkins, ibid. Heft 49 (1911)
22, f. 8G. — S. oblongiflora Perkins, Bot. Jahrb. 25
(1898) 566; Perkins & Gita, Pfl. R. Heft 4 (1901) 54
(‘oblongifolia’); Perkins, ibid. Heft 49 (1911) 22, f.
8A—E. — S. fengeriana Perkins, Bot, Jahrb. 25

316

FLORA MALESIANA

[ser. I, vol. 102

(1898) 566; Pfl. R. Heft 49 (1911) 22, f. 8F. — S. cris-
pula Perkins, Pfl. R. Heft 49 (1911) 21. — S. torri-
cellensis PERKINS, /.c. 21. — S. forbesii PERKINS, l.c.
23. — S. buergersiana Perkins, Bot. Jahrb. 52
(1915) 199, f. 2. — S. riparia KANEH. & Hatus. Bot.
Mag. Tokyo 56 (1942) 259, f. 8. — Fig. 22.

Tree up to 20 m, occasionally a straggling shrub or
liane; young branches with a dense tomentum or a
fine pubescence which may persist on the leafy
branches or these may quickly become glabrous.
Leaves broadly ovate or elliptic to narrowly elliptic,
rarely lanceolate or more or less orbicular, usually of
moderate size (10—13 cm long) but frequently up to
32 by 13 cm and occasionally much smaller (7.5 by 2
cm); base rounded or cuneate, apex apiculate, mar-
gin entire or occasionally with a few obscure denta-
tions or rarely serrate, either quickly losing all or
most of their original pubescence, or the tomentum
may persist on both sides of the blade, but more espe-
cially below, sometimes as a thick woolly coating
over the entire undersurface; principal veins conspic-
uous below, arching strongly or more gradually to-
wards the apex; petiole 3—14 mm, pubescent or gla-
brous. Monoecious. /nflorescences axillary, supra-
axillary or terminal, usually pleiochasia with the low-
er lateral branches again branched or reduced to
simple dichasia, densely pubescent throughout or
glabrous. — Male flowers ovoid or globose, usually
pubescent on the outside, 2.5—4 mm g; tepals 4,
rounded, the wall thick, leathery, enclosing a cavity
either confined to the upper half of the receptacle or
extending to near the base; stamens 4, 1—1.5 mm
long, filaments pilose. — Female flowers globose or
somewhat depressed, 3.5—4.5 mm g, usually pubes-
cent outside, pilose within, sometimes longitudinally
furrowed when dry; ostiole small, surrounded by 4
minute or obsolescent tepals; the cavity large; carpels
numerous, 1.5—2 mm long, ovary slender, pilose,
stigma subulate. Fruiting receptacle enlarged to c. 2
cm 9. Drupes stipitate or more or less sessile; stipes
woody, to 10 mm long; achenes ovoid to c. 13 by 10
mm, verruculose.

Distr. Australia (Queensland, Iron Range); in
Malesia: SW. Celebes (Mt Bonthain), Moluccas (Bu-
ru, Ceram, Ambon, Halmahera, Ternate, Aru Is.),
New Guinea (throughout the island from Vogelkop
Peninsula to Milne Bay Prov.: Mt Suckling), also on
Manus I., New Britain and New Ireland.

Ecol. An understorey tree in rain-forest, mostly
at low altitudes, but also in montane forest up to
2450 m. It is recorded from seasonally inundated
swamps and from ridge forest. Usually a moderate
sized tree with a spreading crown, it may also be a
straggling shrub and occasionally is described as a
liane.

Uses. The wood is used for clubs (Isago village,
Balimo district, Western Prov.). Pipe tobacco rolled

in the leaves (Butemu village, Saidor Distr., Madang
Prov.).

Vern. Gwa, Wagu, imaitru, Jal, kamakama, ka-
mokam, Enga, kubal, sap, Madang, kuraili, Koro-
pa, kwaffel, Bembi, mankanane, Mt Suckling, met-
jem, mey, Waskuk, namelawe, Kainantu, napiwa,
Talasea, New Britain, ongupinpae, S. New Britain,
sakoboe, Moeswaar I., sore, Naho, tangitang, Tom-
ba, fonona, Kutubu, tsohren, Rawa, tugwambi, Wa-
gu, tunngacook, Wein, vingi-vingi, Bayer R.,
waluwali-i, Gogodala.

Notes. The number of different forms included
within this species cannot be considered satisfactory,
yet they appear to be connected by intermediates so
that any segregation of these forms as species would
seem, at this stage, to be even less satisfactory. This
aggregate has a geographical range extending from
Celebes almost to the eastern limit of New Guinea,
and reaches from sea-level to 2450 m. It is frequent
throughout its range, and the number of separate
collections of it approaches that of all the other spe-
cies combined. Several specific names have been ap-
plied to different forms within the complex and some
of these certainly appear distinctive until interme-
diate forms are compared. Variability effects most
features of the plant: leaf shape and leaf size and the
pattern of the principal veins; the presence or ab-
sence of marginal teeth and their prominence; the de-
gree of development and persistence of the indumen-
tum; the degree of branching of the inflorescence;
the size and shape of the male and female receptacles;
the ratio between the cavity and wall thickness in the
male flower; and the presence and length of the stipes
bearing the achenes. Some combinations of charac-
ter states may occur frequently in one region, encour-
aging the belief that local segregate species may even-
tually become recognizable. An example is the form
named S. oblongiflora (with elliptical male recep-
tacles with small cavities and narrowly elliptical +
glabrous leaves with few, sharply ascending veins)
that occurs in the upland parts of the Northern and
Central Provinces and adjacent parts of the Man-
yamya District of Morobe Province. Similar to it and
occurring sympatrically is the form named S. thyrsi-
flora which is most frequent in the neighbouring
Wau, Mumeng and Lae Districts of Morobe Prov-
ince. This form has broader glabrous leaves with
more spreading principal veins, and with globose re-
ceptacles with proportionally larger cavities. How-
ever, the character states intergrade and occur in dif-
ferent combinations, so that specific distinctions
cannot be maintained. A practical difficulty with
herbarium material is that specimens rarely display
all the significant characters, so that a definitive
treatment of this complex must finally rely on exten-
sive fieldwork. Another distinctive form with densely
hairy leaves (and often with + sessile achenes) has

1986 |

MOoONIMIACEAE (Philipson)

317

Fig. 23. Steganthera ilicifolia A.C. SmitH. a. Habit, 2/3, b. male flowers, one in LS, c. anther from this
flower, all x7 (a PHILIPSON & KARINGA 3720, b—c CARR 15203).

been collected most frequently east of Wau, but is
also found from the Owen Stanley Range to the
Western Highlands Province.

The young foliage is pinkish; the flowers are cream
coloured; and the black drupes are borne on red, yel-
low or orange receptacles.

13. Steganthera ilicifolia A.C. Smiru, J. Arn. Arb.
22 (1941) 235; PHitipson, Blumea 29 (1984) 495, —
Fig. 23.

Shrub or small tree to 12 m; young branches with
strigose or silky appressed hairs. Leaves chartaceous,
elliptic, lanceolate to broadly elliptic, 5.5—21 by
2—8.5 cm, base obtuse rounded or truncate, apex
with a long or short acute apiculum, margin with

prominent, remote denticulations, or with few min-
ute teeth or entire, at first often covered with silky
appressed hairs, becoming glabrous or retaining
some indumentum, especially on the midrib below,
nerves and reticulations prominent on the lower sur-
face; petiole 5—15 mm, strigose or becoming gla-
brous. Monoecious. /nflorescences axillary, supra-
axillary or terminal, 4—6.5 mm long, either solitary
few-flowered pleiochasia, or groups of a few pleio-
chasia, rachis and branches with strigose hairs, min-
ute linear bracts below or on the branches (and also
sometimes on the receptacles), rachis slender with a
long peduncle before the first branching, branches
opposite or subopposite, singly or in clusters and
themselves branching. — Male receptacle spherical c.

318

FLORA MALESIANA

[ser. I, vol. 102

2-3 mm g, slightly strigose outside; tepals 4, round-
ed; stamens 4, c. 1.25—2 mm long, with short, gla-
brous or minutely pubescent filaments. — Female re-
ceptacile larger and slightly flatter than the male, c.
4—5 mm @ at anthesis, ostiole minute; tepals 4, min-
ute or obsolete, inner surface pilose; carpels c. 10—
20, 1.5—2.5 mm long, ovary pilose, style awl-shaped,
glabrous. Fruiting receptacle only slightly enlarged,
with hairs between the subsessile or more usually
stipitate drupes (stipe occasionally to 5 mm long).
Drupes ovoid, c. 12 by 9 mm when dry, verruculose
and slightly pilose.

Distr. Malesia: New Guinea, from Milne Bay
Prov. (incl. Fergusson I. at 900 m) westwards as far
as the Star and Carstensz Mts in West New Guinea;
also in the mountains north of the Huon Gulf in Mo-
robe and Madang Prov.

Ecol. A straggling understorey shrub or small
tree in lower mountain forest (often with Nothofagus
and Castanopsis) or in thickets at the forest margin.
Frequent along the central mountains usually above
1200 m to as high as 3250 m (but rather lower in Cen-
tral Prov.).

Uses. Provides stakes for general purposes, e.g.
for house-building, for digging sticks and firewood.

Vern. Agubporombigl, Hagen, genzphora,
iganaphore, Tairoa lang., kamagam, kamokam, ka-
mokum, Enga lang., kamokamp, Mendi lang., ka-
makama, \alibu, kKombugump, toin bekl, Togoba,
munne yambo, Maring, pundpunda, Melpa lang.,
Mt Hagen, soreng, Nako lang., yuri, Minj.

Note. Acommon and widespread species, usual-
ly a straggling shrub, but sometimes attaining tree
stature. In its usual form the harsh dentate leaves
vary in size and shape, but the prominent venation
below is characteristic as are the delicate, few-
flowered inflorescences with small, globose male
flowers. Female flowers are less frequent and are
either terminal or end the distal branches. In less typ-
ical forms the dentations become fewer and less
prominent, culminating in forms with quite entire,
often lanceolate leaves. A few of the specimens with
entire leaf margin cannot be distinguished from
forms of S. hirsuta with certainty. The young foliage
is described as red or pink, the flowers cream, and
the ripe receptacle orange bearing purple-black
drupes. The flowers are frequently deformed by in-
sect galls.

14. Steganthera chimbuensis PHiiipson, Blumea 29
(1984) 495.

Shrub or tree to 20 m; new growth glabrous or
sometimes very finely pubescent. Leaves glabrous,
chartaceous, elliptic, 5-10 by 2.2—4.2 cm, base
broadly cuneate, apex apiculate, margin entire (rare-
ly some leaves on a plant may have one or a few den-
tations), midrib and principal veins usually im-

pressed above (blade more or less rugose) and elevat-
ed below; petiole 4-10 mm. Monoecious. /nflores-
cences axillary or supra-axillary, dichasia or with one
terminal flower, c. 2—3 cm long, with small linear
bracts subtending the branches or inserted on them.
— Male receptacles globose, 2—3 mm 9g; tepals 4,
rounded, ostiole tending to open widely; stamens 4,
c. 1.75 mm long, filament hairy. — Female re-
ceptacle similar to male, slightly larger and often
solitary, tepals and ostiole + obsolete; carpels c.
12—20, 2 mm long, ovary pilose, stigma glabrous,
awl-shaped. Fruiting receptacle slightly enlarged,
leathery. Drupes ovoid, c. 11 by 8 mm, verruculose,
shortly stipitate.

Distr. Malesia: Papua New Guinea (Chimbu &
Eastern Highlands Prov.).

Ecol. Primary and secondary lower montane for-
est, 2400—2850 m.

Vern. Abangle, Chimbu, Masul, ivananottoi,
Mairi, Watabung, pemble, Wahgi, Minj, pogambeg,
Hagen, Togoba.

Note. Similar to the more widely spread S. ilici-
folia, but the leaves are entire (a dentation very rarely
occurs on a leaf of a plant with otherwise entire
leaves) and the undersurface dries a warm cinnamon
brown in contrast to the buff colour of S. ilicifolia.
The flowers are described as pale yellow, the drupes
purple-black on an orange receptacle.

15. Steganthera australiana C.T. WuirTeE, Proc. R.
Soc. Queens]. 55 (1944) 78; PHimipson, Blumea 29
(1984) 496.

Small tree to 10 m; young branches finely pubes-
cent. Leaves chartaceous, lanceolate or narrowly to
broadly elliptic, c. S—10 by 1.5—5 cm, base atten-
uate, apex long-acuminate, margin entire or re-
motely and finely dentate, becoming glabrous or pu-
bescence persisting on midrib and main veins below
or occasionally also on the lower surface; petiole
5-10 mm, glabrous or pubescence persistent.
Monoecious. Jnflorescences axillary or supra-
axillary, dichasia or few-flowered pleiochasia, ra-
chis, branches, bracts and receptacles pubescent, the
rachis slender with a long peduncle below the first
branches, branches opposite or subopposite, singly
or in clusters. — Male receptacles globose, narrowed
into the pedicel, 2—2.5 mm g; tepals 4, rounded,
minute; stamens 4, 1.5 mm long, filament pilose. —
Female receptacles similar but larger, c. 4 mm @ at
anthesis, at and near the ends of inflorescences (when
present); tepals 4, minute, inner surface long-pilose,
carpels numerous, c. 2.2 mm long, ovary pilose, style
awl-shaped, glabrous. Fruiting receptacle slightly en-
larged, leathery. Drupes ovoid, 11—14 by 9-11 mm,
verrucose, sessile or shortly stipitate.

Distr. Queensland; in Malesia: Papua New

1986 |

Guinea (Morobe Prov.: Wau Distr.; Central Prov.:
Goilala & Moresby Distr.).

Ecol. Lower montane forest (Castanopsis and
Nothofagus dominated) and _ second growth,
1000—2300 m.

Note. A plant of restricted distribution in New
Guinea, characterized by the small, usually more or
less lanceolate and apiculate leaves and delicate in-
florescences, is identified with the species described
from Queensland. The few New Guinea specimens
show more variation in size, shape, dentation, and
indumentum of their leaves than may occur in Aus-
tralia. A specimen from near Wau has more richly
branched inflorescences, the side-branches being
clustered, whereas in the other specimens they occur
in opposite pairs. CARR collected this species on five
occasions, but it has been collected only twice subse-
quently. A small tree with greenish or cream flowers.

16. Steganthera insculpta Perkins, Bot. Jahrb. 52
(1915) 202; PHripson, Blumea 29 (1984) 496.

Shrub to 1.5 m high; young branches densely
brown pilose. Leaves chartaceous, elliptic oblong or
ovate, 12—21 by 5—10 cm, base rounded, apex apicu-
late, margin with regular, wide-spaced, small denta-
tions; young leaves uniformly densely brown pilose,
in mature leaves the upper surface with remnants of
the indumentum especially on the veins, the under-
surface remaining densely pilose on the veins, with
scattered hairs elsewhere, reticulation impressed
above and prominently raised below; petiole 4—6
mm, densely pilose. — Male flowers unknown. —
Female flowers in subsessile axillary or supra-axillary
groups. Fruiting receptacle densely pilose. Drupes
numerous, sessile, ovoid or subglobose, 9 by 6 mm,
densely pilose.

MOoONIMIACEAE (Philipson)

319

Distr. Malesia: Papua New Guinea (Sepik region
& Telefomin). Two collections.

Ecol. In forest at 850—1600 m.

Note. This species is distinguished from densely
hairy forms of S. Airsuta by the virtually sessile fe-
male inflorescences and the regular small dentations
on the leaf margins. It may prove to be best regarded
as a form of that species because a sterile specimen
from much further to the east has leaves identical to
those of the type. However, fertile specimens from
the same locality with similar (but not so regularly
dentate) leaves have long-pedunculate cymose inflo-
rescences and are identified as S. hirsuta.

Insufficiently known

1. Steganthera atepala PERKINS in K. Sch. & Laut.,
Fl. Deut. Schutzgeb. Siidsee (1900) 329. — Ropatz
& Kink 237, Bismarck Range.

2. Steganthera odontophylla Perkins, Pfl. R.
Heft 49 (1911) 23. — SCHLECHTER 17847, Kaui Mts.

3. Steganthera pycnoneura PERKINS, /.c. 23. —
SCHLECHTER 19517, Maboro.

4. Steganthera symplocoides PERKINS, /.c. 23. —
SCHLECHTER 19825, Goridjoa.

5. Steganthera psychotrioides PERKINS, Bot.
Jahrb. 52 (1915) 198. — LEDERMANN 9078, Sepik.

6. Steganthera alpina PERKINS, /.c. 201. — LEDER-
MANN 11968, Sepik.

7. Steganthera ledermannii (PERKINS) KANEH. &
Hatus. Bot. Mag. Tokyo 56 (1942) 255. — Antho-
bembix ledermannii PERKINS, Bot. Jahrb. 52 (1915)
203, f. 3. — LEDERMANN 11412. Very like S. hospi-
tans but leaves with long white hairs, a feature not
seen in S. hospitans.

10. MATTHAEA

Biume, Mus. Bot. Lugd.-Bat. 2 (1856) 89; PERKINS, Bot. Jahrb. 25 (1898) 562;
Pfl. R. Heft 4(1901) 51, f. 10; ibid. Heft 49 (1911) 15; Ubersicht Gattungen Mo-
nim. (1925) 62; PHiLipson, Blumea 28 (1982) 77, f. 1—3. — Fig. 24—26.

Shrubs or small trees. Leaves pubescent or glabrous at maturity, entire or
dentate. Monoecious, with lateral (rarely also terminal) cymose inflorescences,
much shorter than the leaves. — Male receptacle subglobose, the ostiole sur-
rounded by 4 small rounded tepals. Stamens 4, free; filaments short; anthers
opening by 2 longitudinal slits. — Female receptacle more flattened; tepals 4,
upper half abscissing as a calyptra at anthesis to reveal very numerous carpels.
Drupes \ong-stipitate, verruculose.

Distr. Malesia; Malay Peninsula, Sumatra, Anambas Is., Borneo, Celebes, Philippines, N. Moluccas (Ta-
laud Is.).
Ecol. Rain-forest, ascending to 1700 m.

320 FLORA MALESIANA [ser. I, vol. 102

\y ‘\
A :
q Mai Dp
y) WW Wy

Fig. 24. Matthaea sancta BL. a. Habit, developing fruits, b. male inflorescence, both 1/2, c. male flower
n Ls, x7, d. female flowers, on left the calyptra removed, x5, e, carpel with prominent stigma, x12 (a
AHMED s.n., b VAN BALGOoy 2029, c Toxopeus 547, d—e VAN STEENIS EADY.

1986 | MONIMIACEAE (Philipson) 321

Fig. 25. Range of the species of Matthaea Bi. 1. M. pubescens, 2. M. chartacea, 3. M. sancta, 4. M. hetero-
phylla, 5. M. vidalii, 6. M. intermedia.

Note. The distinction between Matthaea and Steganthera is very slight. The two genera are so alike vegeta-
tively that sterile specimens cannot be allocated to a genus with confidence, and their flowers and fruits are
also very similar except for their anthers. In both genera there are four stamens; in Matthaea these bear an-
thers opening by two more or less vertical slits, whereas in Steganthera the anthers open by a single horizontal
slit. It is useful to maintain the two genera because they are geographically separated (though both may occur
in Celebes). Matthaea is restricted to western Malesia while Steganthera, a considerably larger genus, is cen-
tered on New Guinea with outliers in the Moluccas, Celebes and Queensland. Some uncertainties of range
remain as male flowers and fruits are not available from several areas.

KEY TO THE SPECIES

Saucersurface of leaves. pubescent. iiaiv és: ee ws vitisitts OE: Sh Wade Mes ap kN Rana ait 1. M. pubescens
1. Undersurface of leaves glabrous.
2. Leaves more than 15 cm long.
3. Principal lateral veins of undersurface less distinct, scarcely raised above the surface. Leaves usually el-
lintic.to broad-ellintic \ 55:5 ssw aire Save es +e Dineen Paras Obs ae Ae 2. M. chartacea
3. Principal veins on undersurface prominent, raised above the surface, Leaves usually oblong-lanceolate
to lanceolate.
42 Leaves oblong-elliptic to narrowly oblong sic. dele incle ca Viad veel awa oats 3. M. sancta
4 eaves lanceolate (Ni LUzon)) a5 Ooi FA ocd DER ee ode 4. M. heterophylla
2. Leaves less than 15 cm long.

322 FLORA MALESIANA [ser. I, vol. 102

Fig. 26. Leaves of Matthaea species, all x1/3. a. M. chartacea, b. M. pubescens, c. M. heterophylla, d. M.
sancta, e. M. vidalii, f. M. intermedia (a BS 40705, b ELMER 10699, c BS 78395, d RipLey 10770, e BS 30718,
ff PNH 4480).

1986 |

St Leaves lanceolate-oblong . 2.2 5.2.0) 2.022228. .s

5. Leaves elliptic to elliptic-oblong.

MONIMIACEAE (Philipson)

323

Be Se Tea Bes Soin eee 4. M. heterophylla

6. Venation on lower surface prominent, raised above the surface. Lamina, petiole, and young branches

meTRe te) LNT. MOR een SLES Sea «

x) AUPE raat Pees See neee ene at eect nae 5. M. vidalii

6. Venation on lower surface less distinct, scarcely raised. Lamina, petiole, and young branches olivaceous

1. Matthaea pubescens MERR. ex PERKINS, Bot.
Jahrb. 45 (1911) 422; Pfl. R. Heft 49 (1911) 16;
Merk. En. Philip. 3 (1923) 186; PHmipson, Blumea
28 (1982) 80, f. 1 & 2. — Fig. 26b.

Shrub or small tree, to 10 m; young branches
densely brown pubescent. Leaves broadly elliptic to
elliptic, 12—27.5 by 5.S—13.7 cm; apex rounded with
a short acute apiculum, base broadly cuneate, trun-
cate or rounded, densely brown tomentose (but the
upper surface eventually becoming almost glabrous),
margin entire or often with dentations on its upper
part, lateral veins arched, obscure above; petiole
5-15 mm, densely tomentose. /nflorescences axil-
lary, solitary or few in the same axis; male rachis
densely brown tomentose, c. 20 mm, bearing a ter-
minal flower, with usually 2 lateral branches sub-
tended by minute bracts; receptacle c. 6 mm @ (when
dry), tomentose; tepals 4, minute; stamens 4, free. —
Female inflorescences similar, but apparently only
the terminal flower develops fruit (lateral branches
frequently bear male flowers) and usually only one
inflorescence develops fruit in each axil; rachis be-
coming stout, elongating to c. 30 mm; receptacle tur-
binate, 10O—15 mm gat anthesis, larger in fruit, inner
surface with pilose hairs between the carpels; carpels
very numerous, ovary densely strigose; stigmas prom-
inent. Drupes numerous, 30 by 15 mm (when dry).

Distr. Malesia: Philippines (Mindanao: Agusan,
Surigao, Bukidnon & Davao Prov.). Fig. 25.

Ecol. In dense, often mossy forest, 300—1700 m.

Vern. Kalagau, mangilas, Bukidnon, baringoras,
bayung-bayung, Bag.

Note. The wood is described as white and moder-
ately hard; the bark as yellowish grey, fissured. Ripe
fruits are blue-black.

2. Matthaea chartacea Merr. Publ. Gov. Lab. Phi-
lip. nm. 35 (1905) 14; Perkins, Pfl. R. Heft 49 (1911)
18; Merr. En. Philip. 3 (1923) 186, f. 1—2; PHitp-
son, Blumea 28 (1982) 80. — M. williamsii PERKINS,
Bot. Jahrb. 45 (1911) 422. — M. pinchotiana PrEr-
KIns, /.c. 423. — Fig. 26a.

Shrub or small tree to 10 m; young branches be-
coming glabrous. Leaves oblong-ovate, (13—)15—20
(—25) by 6—9.5(—12) cm, apiculate with an obtuse or
acute apex, chartaceous, glabrous when mature
(rarely a few hairs retained at the base of the blade);
margin with small dentations on the upper part or en-
tire; lateral veins arched, obscure above, veins of a
lower order obscure; petiole 15-20 mm, glabrous

6. M. intermedia

(rarely with some persistent hairs). /nflorescences
axillary, solitary or few in the same axil. — Male ra-
chis strigose, to 20 mm, with minute scales at the
base, bearing a terminal flower, side branches 1—2 or
absent, c. 10 mm long, subtended by minute strigose
bracts; receptacle c. 5mm @ (when dry), slightly strig-
ose; tepals 4, minute; stamens 4, free; filaments
broad, short; anthers broadly triangular. — Female
inflorescence similar, but apparently only the termin-
al flower develops to fruit (lateral branches frequent-
ly bear male flowers); rachis becoming stout and di-
lated distally, elongating to c. 35 mm; receptacle tur-
binate, c. 10 mm gat anthesis, slightly larger in fruit;
carpels very numerous; ovary densely strigose; stig-
ma prominent. Hairs persist on the young fruit, but
the numerous ripe drupes are glabrous, shining
black, wrinkled, c. 20 by 15 mm (dry), long-stipitate
(c. 18 mm long).

Distr. Malesia: Philippines (Luzon: Laguna &
Tayabas Prov.; Mindoro; Leyte; Mindanao: Zam-
boanga Prov.) and N. Moluccas (Talaud Is.: Karake-
long). Fig. 25.

Ecol. Dense forest and second growth, to 700 m.

Uses. The stem is scraped and applied for head-
aches. The plant is used in hunting rituals.

Vern. Alukba, barau-barau, Mang., molikot-
lang, Mindoro, matagusa, selimbwang, Zamboanga;
laba, Moluccas, Talaud.

Note. This species is similar to M. pubescens ex-
cept for the lack of indumentum. Although both
these species occur on Mindanao, their ranges do not
overlap, M. chartacea being confined to the west and
M. pubescens to the east.

3. Matthaea sancta Blume, Mus. Bot. Lugd.-Bat. 2
(1856) 90, t. 10; Perkins, Bot. Jahrb. 25 (1898) 563,
t. 5d; Pfl. R. Heft 4 (1901) 52, f. 10; ibid. Heft 49
(1911) 18, incl. var. mindanaoensis PERKINS et var.
venulosa PERKINS, /.c.; MERR. En. Philip. 3 (1923)
186; Rip. Fl. Mal. Pen. 3 (1924) 73; Pxtipson, Blu-
mea 28 (1982) 82, f. 1-3. — Ficus pulchra WALL.
Cat. (1831) n. 4518, nomen. — Mollinedia sancta
(Bt.) Bat. Hist. Pl. 1 (1869) 306. — M. latifolia
Perkins, Bot. Jahrb. 25 (1898) 563, t. Sd; Pfl. R.
Heft 4 (1901) 52, f. 10; ibid. Heft 49 (1911) 17, f.
7B—C, — M. calophylia Perxins, Bot. Jahrb. 25
(1898) 563; Pfl. R. Heft 4 (1901) 52; ibid. Heft 49
(1911) 17, f. 7A. — Fig. 24, 26d.

Shrub or small tree, rarely to 15 m; young
branches becoming glabrous. Leaves lanceolate-

324

FLORA MALESIANA

[ser. I, vol. 102

oblong to oblong, 15.5—31 by 3.5—9.5 cm, acumi-
nate, base broadly cuneate, truncate or rounded,
chartaceous, often somewhat bullate, margin entire
or dentate distally, glabrous; lateral veins arched and
uniting far from the margins, impressed on upper
surface, prominently raised above the lower surface;
petiole 2—3 cm, glabrous. /nflorescences axillary,
solitary or fascicled. — Male rachis 2—3 cm, pubes-
cent, bearing a terminal flower and usually 1—2 pairs
of lateral flowers on short branches (which occasion-
ally also bear side-branches); receptacle subglobose,
c. 3—5 mm Q, slightly strigose; tepals 4, minute; sta-
mens 4, free; filaments broad, short; anthers triangu-
lar. — Female inflorescences similar, some lateral as
well as the terminal flower female and developing
fruit, or (more usually) the lateral branches bear
male flowers which are soon shed; rachis becoming
stout and dilated distally; receptacle turbinate, c. 12
mm @ at anthesis (dry), inner surface pilose between
the carpels; carpels very numerous, ovary densely pi-
lose, stigma prominent. Drupes numerous, up to 25
mm stipitate, ovoid, 18 by 11 mm.

Distr. Malesia: Sumatra, Malay Peninsula (Pe-
rak & Pahang to Singapore), Anambas Is., Borneo
(Sarawak, Sabah, Kalimantan), Philippines (Luzon,
Negros, Cebu, Mindanao) and Celebes. Fig. 25.

Ecol. Small tree or shrub, sometimes scandent in
forest, from low altitudes to 1200 m.

Uses. The wood is heavy and branches are used
in building houses. The leaves are smoked with to-
bacco to relieve headaches.

Vern. Malacca: poko churow, p. lumsoo; Anam-
bas Is.: kayu sama; Philippines: babang di putukan,
If., colog-colog, Negros, bago-bago, C. Bis.

Note. The only species with a wide geographical
range. Leaf size and breadth are variable, as is the de-
gree of toothing of the margin. Typical leaves are
narrowly-oblong, apiculate, and with distant, promi-
nent, strongly arched lateral veins. Broader and
more coarsely toothed leaves appear to occur more
frequently to the east of the range, but no basis for
subspecific taxa can be discerned. The wood is said
to be reddish and moderately hard. The ripe fruit is
blue-black.

4. Matthaea heterophylla Quis. & MErR. Philip. J.
Sc. 37 (1928) 145; Puitipson, Blumea 28 (1982) 82, f.
1—2. — Fig. 26c.

A shrub with appressed whitish hairs on the devel-
oping parts but becoming glabrous. Leaves oblong-
lanceolate to lanceolate, 10—30 by 2.5—6.5 cm, acu-
minate, base broadly cuneate to rounded, charta-
ceous, margin entire, glabrous, lateral veins widely
spaced, prominent on the lower surface, arched; pet-
iole 13—20 mm, glabrous at maturity. Flowers un-
known. Jnfructescences terminal or axillary, ped-
uncle becoming woody, c. 6 cm long with a flattened

receptacle c. 20 mm @ at maturity, inner surface pi-
lose. Drupes c. 15 mm stipitate, ovoid, c. 25 by 18
mm, verruculose (when dry).

Distr. Malesia: Philippines (Luzon: Isabella &
Cagayan prov.). Fig. 25.

Ecol. In forests, about 1200—1350 m.

Note. A local species from northern Luzon ap-
parently outside the range of M. sancta, character-
ized by the narrow elongated leaves, with major lat-
eral veins widely spaced and running out straight
from the midrib very nearly at right angles.

5. Matthaea vidalii PErKins, Bot. Jahrb. 45 (1911)
422; Pfl. R. Heft 49 (1911) 17; Merr. En. Philip. 3
(1923) 186; PuHmipson, Blumea 28 (1982) 82, f. 1—2.
— Fig. 26e.

Shrub, about 2 m; young branches glabrous.
Leaves elliptic to elliptic-oblong, 11—15 by 3.5—6 cm,
acuminate, base broadly cuneate, somewhat coria-
ceous, margin entire or rarely with a few teeth in the
distal part, principal veins prominent on the lower
surface and strongly arched; petiole 15S—20 mm, gla-
brous. Inflorescences axillary, solitary or fascicled.
— Male rachis c. 35 mm, sparsely pilose, bearing a ter-
minal flower and a pair of lateral flowers on short
branches; receptacle turbinate, c. 5mm, sparsely pi-
lose outside; tepals 4; stamens 4, free. — Female in-
florescences similar, apparently only a single terminal
flower producing fruit; receptacle at anthesis not
seen; rachis becoming stout and woody at fruiting.
Drupes numerous, c. 15—18 mm stipitate, ovoid, c. 30
by 20 mm, glabrous, wrinkled, purple-black.

Distr. Malesia: Philippines (Luzon: Camarines
Sur & Sorsogon Prov.; Panay: Capiz Prov.). Fig. 25.

Ecol. Forest, from 600 to 1600 m.

Vern. Salapula, Bik.

Note. Occurs within the range of M. sancta,
which it approaches. The smaller, coriaceous and
more elliptic leaves appear consistently distinct.

6. Matthaea intermedia Merr. Philip. J. Sc. 13
(1928) Bot. 11; PHitipson, Blumea 28 (1982) 82, f.
1-2. — Fig. 26f.

Shrub, about 4 m; young branches glabrous.
Leaves elliptic to oblong-elliptic, c. 8—13 by 3—6 cm,
acuminate, base broadly cuneate, margin entire, gla-
brous, lateral veins arched, slightly raised below, not
impressed above; petiole c. 15 mm, glabrous. Jnflo-
rescences axillary; rachis c. 15—20 mm, with scat-
tered appressed hairs, bearing a terminal flower and
usually one pair of lateral flowers on short branches.
— Male receptacle turbinate, sparsely pubescent;
tepals 4; stamens 4, subsessile. — Female receptacle
similar, inner surface pilose between the carpels; car-
pels numerous, ovary densely pilose, stigma promi-
nent. Drupes 8—10 mm stipitate, ovoid, c. 20 by 15
mm, verruculate.

1986] MONIMIACEAE (Philipson) 325

Fig. 27. Lauterbachia novoguineensis Perkins. a—b. Habit, with female flowers, nat. size, c. open flower,
d. opening flower, both «3, e. flower in LS, x4 (after PERKINS).

326

FLORA MALESIANA

[ser. I, vol. 102

Distr. Malesia: Philippines (Luzon: Tayabas
Prov.). Fig. 25.

Ecol. Forests, about 1000 m.

Note. A local species, from the central Philip-
pines, resembling the neighbouring M. vidalii in size
and shape of leaf, but differing in the less prominent
venation on the lower surface, and particularly in the
brown-olivaceous colour of the leaves, petioles and
branches.

Insufficiently known

Matthaea philippinensis PERKINS, Bot. Jahrb. 45
(1911) 423; Pfl. R. Heft 49 (1911) 18. Of this species,
described from Leyte, no specimens have been seen.

11. LAUTERBACHIA

PERKINS in K. Sch. & Laut., Fl. Deut. Schutzgeb. Siidsee (1900) 330, t. 6; PER-
KINS & Gita, Pfl. R. Heft 4 (1901) 63; PERKINS, Ubersicht Gattungen Monim.
(1925) 40; Hutcu. Gen. Fl. Pl. 1 (1964) 119. — Fig. 27.

Trees or shrubs with entire leaves. Inflorescence axillary or terminal. — Male
flowers not known. — Female flowers with 4 minute tepals and a velum sur-
rounding the ostiole; upper part of the receptacle abscissing as a calyptra. Car-
pels numerous, with a long subulate style.

Distr. Malesia: Papua New Guinea (Madang Prov.: Ramu Distr.). Fig. 10.

Ecol. Lower montane rain-forest.

Note. The single species was known only from the type which was destroyed in World War II. No duplicate
has been located so the above description and that of the species is based on information published by PER-
KINS. The presence of a velum in what appears to be a member of the Mollinedieae is so exceptional that the
interpretation of PERKINS must be considered doubtful until further material can be examined.

1. Lauterbachia novoguineensis PERKINS in K. Sch.
& Laut., Fl. Deut. Schutzgeb. Siidsee (1900) 331;
PERKINS & GILG, Pfl. R. Heft 4 (1901) 63. — Fig. 27.

A shrub or tree, the young branches with brown
tomentum. Leaves elliptic-oblong or oblong, 13—21
by 5—8 cm, base cuneate or rounded, apex broadly
acuminate or acute, glabrous above except for sparse
hairs on the nerves, below clothed with long greyish
hairs chiefly on the nerves, becoming glabrous, en-
tire, margin revolute (when dry); nervation promi-

nent below, nerves arched and meeting within the
margin; petiole 10—12 mm, tomentose. /nflorescence
axillary or terminal, 4—8-flowered pleiochasia, c. 30
mm long, clothed with brown tomentum; pedicels c.
5 mm. — Female receptacle turbinate, 8—9 mm 9;
tepals 4, minute, a velamen surrounding the wide
ostiole; carpels c. 38, narrowed above into a long
subulate style.

Distr. Malesia: Papua New Guinea (Madang
Prov.: Ramu Distr.; Bismarck Range). Fig. 10.

Excluded

Idenburgia Gipss = Sphenostemon Batu. (Sphenostemonaceae).

Scyphostegia STAPF = Scyphostegiaceae.

Tambourissa ficus (TuL.) A.Dc. (Ambora ficus TUL.) was mentioned by MiquEL, FI. Ind. Bat. 1, 2 (1859)
75 and DC. Prod. 16, 2 (1868) 659 to have been collected in Java by LESCHENAULT. PERKINS, Pfl. R. Heft
4 (1901) 70 raised already doubt about this record, which certainly rests on an erroneously localized specimen

from Madagascar.

Trimenia SEEM. (Piptocalyx OLiv. ex BTH.) = Trimeniaceae.

TRIMENIACEAE (W. R. Philipson, Christchurch)

Trimenia was first described by SEEMANN as a genus related to the Ternstroemiaceae. BENTHAM
& Hooker f. (1880) regarded it as more closely related to the Monimiaceae without definitely plac-
ing it there. This was done by PERKINS & Gitc (1901) who formed the tribe Trimenieae of that
family. Gress (1917) created the family Trimeniaceae without stating grounds for the separation.
Gitc & SCHLECHTER (1923) disagreed, thinking the differences between Trimenia and other Moni-
miaceae too slight. However, a more complete study by Money, BatLey & Swamy (1950) firmly
established the family which is now generally accepted. The work of ENDREss & SAMPSON (1983)
strengthened this conclusion and demonstrated the isolated position of the family by drawing at-
tention to a number of features deviating from those generally found in the Laurales. These in-
clude absence of a floral cup; spiral floral phyllotaxis; caducous tepals; utriculate carpels; poly-
forate pollen; tectate-columellate exine; capitate stigma with multicellular papillae; vascularized
outer integument. Chromosome number n=8.

References: BENTHAM & HOOKER f., Gen. PI. 3 (1880) 143; ENpREss & Sampson, J. Arn. Arb.
64 (1983) 447—473; Gipss, Fl. Phyt. Arfak Mts (1917) 136; Gitc & SCHLECHTER, Bot. Jahrb. 58
(1923) 245; Money, BAILEY & Swamy, J. Arn. Arb. 31 (1950) 372—404; PERKINS & GILG, Pfl. R.
Heft 4 (1901) 21.

1. TRIMENIA

SEEMANN, FI. Vit. (1871) 425, t. 99; B. & H. Gen. Pl. 3 (1880) 143; Pax in E.
& P., Nat. Pfl. Fam. 3, 2 (1889) 98; Perkins & Gia, Pfl. R. Heft 4 (1901) 21,
f. 4a—c; Rww_. Trans. Linn. Soc. Bot. II, 9 (1916) 144; Gress, Arfak (1917) 135;
Gitc & ScuHiTr, Bot. Jahrb. 55 (1919) 195, f. 1—2; ibid. 58 (1923) 245, f. 1; PER-
Kins, Ubersicht Gattungen Monim. (1925) 22; A.C. Smiru, J. Arn. Arb. 23
(1942) 442; RopDENBURG, Blumea 19 (1971) 3. — Piptocalyx OLIv. ex Bru. FI.
Austr. 5 (1870) 292; PERKINS & Gita, Pfl. R. Heft 4 (1901) 22, f. 4d—f. — Fig.
1-3.

Small trees, shrubs or lianes, up to 20 m or more, young parts tomentose or
glabrous. Leaves opposite, petiolate, exstipulate, ovate to ovate-lanceolate
(obovate, extra-Mal.), base cuneate, apex acute to long acuminate, entire or ser-
rate, with translucent dots, nerves connected near the margin. /nflorescence
axillary or terminal, cymose, pleiochasial (racemose) or paniculate. Flowers
unisexual or bisexual; receptacle continuous with the pedicel, slightly convex,
glabrous; tepals caducous before or at anthesis, spirally arranged (outermost
sometimes decussate), imbricate, 10—38, the lower ovate to + orbicular or reni-
form, up to 3 mm long, base swollen and sometimes peltate, apex rounded or
obtuse, grading upwards into longer, narrower and more membranous tepals,
the uppermost spathulate, up to 5 mm long. Stamens 7—25, spirally arranged,
filament shorter or as long as the anther, connective produced at apex, anthers
tetrasporangiate, extrorse or latrorse, opening by two longitudinal slits. Carpel
solitary (rarely 2), rudimentary or absent in male flowers, superior, barrel-
shaped, glabrous or sparsely strigose; stigma sessile tufted-papillose, 1-celled;
ovule 1, pendulous, anatropous. Fruit a small spherical, succulent berry. Seed
hard, smooth or ridged; embryo small, apical; endosperm abundant.

Distr. Eastern Pacific (Marquesas), Polynesia (Samoa, Fiji), New Caledonia, E. Australia (New South

(327)

328 FLORA MALESIANA [ser. I, vol. 102

Fig. 1. Trimenia papuana Rit. A. Habit, nat. size, B. bud, x6, C—E. outer, middle and inner tepals, x8,
F. flower, after removal of tepals, x7, G. stamen, from inside, x10, H. ovary, x7, J. ditto in LS, x10,
K. ditto in CS, x12, L. fruit, x3 (after Gitc & SCHLECHTER, 1919).

Wales), Solomons (Bougainville) and Malesia: New Guinea (incl. New Britain & New Ireland), Moluccas (Ce-
ram, Batjan) and Central Celebes. In all 5 spp. Fig. 2. ;

Fossils. MULLER (1981) considered that Cretacaeiporites scabrabut from West Africa and Brazil, which
first appears in the Cretaceous (Albian-Cenomanian) (JARDINE & MAGLOIRE, 1963; HERNGREEN, 1973), may
represent 7Trimenia pollen.

References: HERNGREEN, Pollen et Spores 15 (1973) 515—555; JARDINE ef al., Coll. Int. Micropal. Dakar
(1963); MULLER, Bot. Rev. 47 (1981) 9.

Ecol. Small trees or climbers in forests, often on ridge crests or other exposed places and usually at moder-
ate to high altitudes.

Floral biology. The flowers of T. papuana are scentless, produce no nectar and the pollen is dry. No insect
visitors were observed (ENDRESS & SAMPSON, 1983). Presumably wind plays a major role in pollination. The
outer tepals fall before flowering and at anthesis all the tepals have been shed. The other species probably
have a similar biology. In all species some flowers are male, but the degree of separation of the sexes varies.
In T. papuana most flowers are hermaphrodite with some male flowers present. In 7. neocaledonica and T.
moorei the proportion of male flowers is greater. The sexes are more completely separated in 7. weinmannii-
folia in which species all flowers are functionally either male or female. Trimenia weinmanniifolia ssp. wein-
manniifolia is monoecious but the other two subspecies may be dioecious (RODENBURG, 1971).

TRIMENIACEAE (Philipson) 329

SL Bs p
ne) ar pis
: Us ; >
.t
eae i :
a a oa ear
eee apa Nae
asi -* F ‘ ans >. s RNY : : Ey. ..
i oS, hr ~ = 3 — E i
wn ager fa es NA eons nds
kab oe yi ia va
= : ce y “
AS xX. . x ;

= Rn ‘af a
6
w4 pay
= eo f e

Fig. 2. Range of Trimenia SEEMANN: T. papuana (dots), T. neocaledonica (square), T. moorei (oval), T. wein-
manniifolia (triangles), T. macrura (in New Guinea, not indicated).

Dispersal. The one-seeded succulent fruit has dark red to purple juice, and no doubt is distributed by birds.

References: ENpress & SAMpPson, J. Arn. Arb. 64 (1983) 447; RoDENBURG, Blumea 19 (1971) 3-15.

Morph. The leaves are in decussate pairs and exstipulate. The lamina is tapered to an acute apex which
in T. macrura is elongated into a delicate drip-tip. Inflorescences are lateral, or in some species also terminal.
The inflorescence axes bear a terminal flower and either pairs of opposite flowers or branches which in turn
bear a series of flower pairs, or even branches of a third order, when the inflorescence becomes a diffuse pan-
icle.

Anat. Anatomical features are described by Money, BAILEY & Swamy (1950), METCALFE & CHALK (1950),
RopeNBuRG (1971) and Cartouist (1984). Hairs are unicellular or tricellular uniseriate and non-glandular.
Stomata are paracytic. Oil cells and mucilage cells occur in the mesophyll of the leaf and in the axis. The leaf
blade has no hypodermis and a palisade layer is not clearly defined from the spongy mesophyll. The leaf trace
which departs from the single nodal gap comprises 2 or 4 bundles. Young stems develop no hippocrepiform
sclereids (or only vestiges of them) in the pericycle opposite the interfascicular sectors. Elongated sclereids
develop precociously in the secondary phloem. In the secondary xylem the vessel elements are long with high
incidence of scalariform intercellular pitting. There are numerous uniseriate rays in addition to multiseriate
rays (5 or 6 cells wide). Gelatinous fibres occur in the tension wood (KUCERA & PHILIPSON, 1977). Phloem
plastids are of the S-type (BEHNKE, pers. comm.).

References: Car.ouist, Pl. Syst. Evol. 144 (1984) 103-118; Kucera & Puitipson, New Zeal. J. Bot. 15
(1977) 649-654; Metca.re & CHALK, Anatomy of the Dicotyledons 2 (1950); Money, BaiLey & Swamy, J.
Arn. Arb. 31 (1950) 372—404; RopeNBURG, Blumea 19 (1971) 3—15.

Floral anatomy. The structure and development of the flower have been carefully described and discussed
by Enpress & Sampson (1983). The tepals, stamens and carpel(s) are initiated in a spiral phyllotaxis. The peri-
anth members show a gradation of form from below upwards without differentiation into sepals and petals.
The anthers have neither lateral glands nor valvular dehiscence. The middle layer of the anther wall is only
1 or 2 cells thick, and the tapetum is glandular. Cytokinesis is successive and pollen is shed in the 2-celled con-
dition. The mature carpel is markedly utriculate although early developmental stages pass through a chair-like
form. The pendent, anatropous ovule is crassinucellar and bitegmic with the micropyle directed upwards, The

* archesporium is multicellular, but only one megagametophyte reaches maturity forming a long tube growing
towards the micropyle. The fruit is a berry, the very juicy carpel wall enclosing a single seed with a very stony

330 FLORA MALESIANA [ser. I, vol. 102

outer coat. The small embryo is embedded in the apical part of the abundant endosperm. The cotyledons are
rudimentary in the ripe seed and diverge slightly.

Reference: ENpRrEss & SAMPSON, J. Arn. Arb. 64 (1983) 447—473.

Palyn. Earlier accounts of the pollen by MonNEy, BAILEY & Swamy (1950), ERDTMAN (1952) and WALKER
(1976) are extended and fully discussed by SAMPSON & ENDRESS (1984). They report that 7. macrura, T. moorei
and 7. neocaledonica have disulcate, globose-elliptic or globose-spherical to globose-elliptic grains, with fine-
ly reticulate or rugulose (7. neocaledonica) structure. Trimenia papuana has dimorphic pollen, with only one
type on an individual plant, consisting of either globose-elliptic inaperturate grains, or globose-spherical poly-
forate grains. Both types have weakly rugulose structure, 7. weinmanniifolia has globose-spherical polyforate
pollen, with similar structure. Exine is tectate-columellate. In 7. papuana and T. weinmanniifolia the inner-
most tectum, columellae and foot layer have a partly granular form. All taxa have a lamellate endexine in
non-apertural regions. SAaMpsoN & ENpREss compare the pollen of 7rimenia with that of other families and
conclude that pollen morphology confirms the comparatively isolated position of the family within the Laura-
les.

References; ERDTMAN, Pollen morphology and plant taxonomy (1952) 272—273, f. 157A; Monery, BAILEY
& Swamy, J. Arn. Arb. 31 (1950) 372—404; Sampson & ENpRESS, Grana 23 (1984) 129—137; WALKER in Beck,
Origin and early evolution of Angiosperms (1976) 241—299.

Chromosomes. Trimenia papuana (GoLDBLATT, 1974) and 7. moorei (GOLDBLATT & BricGs, 1979) both
give counts of n=8.

References: Gotpsiatt, J. Arn. Arb. 55 (1974) 453—457; Go_pBLatt & Briccs, Ann. Mo. Bot. Gard. 66
(1979) 898-899.

Phytochem. The lignans (or neolignans) veraguensin and calopiptin have been isolated from Trimenia
papuana and Piptocalyx (= Trimenia) moorei (‘bitter vine’) respectively. The nature of the bitter principles
of bitter vine is still unknown, but a slightly sweet glucoside was isolated from its leaves and called piptoside;
its aglucone was shown to be structurally related to the proteaceous metabolite leucodrin. CHENERY reported
aluminium accumulation for one species of Trimenia (two investigated), but not for Piptocalyx (one species
tested). The lack of benzylisoquinoline alkaloids in Trimeniaceae conforms with the exclusion of the taxon
from Monimiaceae.

References: HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 99—107, 457, and ibid. 8 (in prep.); family
treated together with Monimiaceae. — R. HEGNAUER.

Taxon. Generic limits. Hitherto a second genus of the Trimeniaceae, Piptocalyx, has been recognized.
First described from Australia (P. moore?) a second species (P. macrurus) is known from New Guinea. The
climbing habit of these two species contrasts with the arboreal or shrubby habit of Trimenia, but the floral
characters are closely alike. Careful comparisons of Trimenia and Piptocalyx by ENDRESS & SAMPSON (1983)
revealed that some characters thought to separate the genera are indecisive. In particular they showed that
the tepals of Piptocalyx are spiral, as in Trimenia, and not decussate. As T. neocaledonica, previously inade-
quately known, was found to be closer in some respects (e.g. pollen) to Piptocalyx than to other species of
Trimenia they concluded that the only characters which separate the two genera are habit and the number
of tepals (more than or fewer than 11). They preferred to leave the genera intact, to avoid name changes, but
it is concluded here that the recognition of two genera is unjustifiable and the necessary new combination is
made. I should add that also in Monimiaceae habit (erect or climbing) is variable within genera and sometimes
even within a single species.

Trimenia moorei (OLIV. in Bru.) Puitipson, comb. nov. — Piptocalyx moorei OLtv. in Bru. Fl. Austr. 5
(1870) 292.

Specific delimitation. The treatment of the species adopted here follows that of RoDENBURG (1971). A.C.
SMITH (1978) was critical of RODENBURG’s broad specific concept as regards both the Malesian and Polynesian
species. RODENBURG’s sinking of T. arfakensis and T. myricoides into T. papuana appears entirely justified.
This conclusion has been reached after examination of very ample material. The treatment of RODENBURG’S
subspecies of JT. weinmanniifolia does not concern this account except for ssp. bougainvilleensis which SMITH
elevated to specific rank. In view of the wide geographical range of this complex and the indecisive nature
of the characters involved, it appears wise to follow the more conservative treatment of RODENBURG.

References: BENTHAM & Hooker f., Gen. Pl. 3 (1880) 143; ENpREss & Sampson, J. Arn. Arb. 64 (1983)
447—473; Gite & SCHLECHTER, Bot. Jahrb. 58 (1923) 244-248; Grpps, Fl. Phyt. Arfak Mts (1917) 317;
Money, BaliLey & Swamy, J. Arn. Arb. 31 (1950) 372—404; Sampson & ENprREss, Grana 23 (1984) 129-137;
PERKINS & GitG, Pfl. R. Heft 4 (1901) 1—122; RopENBURG, Blumea 19 (1971) 3—15; A.C. Situ, Allertonia
1 (1978) 311-314.

TRIMENIACEAE (Philipson)

331

Fig. 3. Trimenia papuana RipL. In Papua New Guinea (PHILIPSON 3692).

KEY TO THE SPECIES

1. Woody climber. Tepals 10 or fewer.............

1. Trees or shrubs. Tepals 13 or more.

«abe dap bk wid a de 6 OCR ee 3. T. macrura

2. Filaments relatively long and thin; mature anthers once or twice as long as the filaments. Seed smooth

1. T. papuana

2. Filaments relatively short and broad; mature anthers at least 4 times as long as the filaments. Seed with

longitudinal or reticulate ridges...............

1. Trimenia papuana Ript. Trans. Linn. Soc. Bot.
Il, 9 (1916) 144; Guo & Scuitr, Bot. Jahrb. 55
(1919) 199, f. 1; A.C. Smiru, J. Arn. Arb. 23 (1942)
442: RopensurG, Blumea 19 (1971) 9. — T. arfaken-
sis Gripes, Arfak (1917) 136; Kangen. & Hatus. Bot.
Mag. Tokyo 56 (1942) 262, f. 10. — 7. myricoides
Guec & Scuirr, Bot. Jahrb. 58 (1923) 248. — Fig. 1,
a

Shrub or tree up to 20 m or more high; young

eG cad ave a POSE a Oe 2. T. weinmanniifolia

branches reddish brown villous, becoming + gla-
brous. Leaves narrowly elliptic to lanceolate, 2—1.25
by 0.7—3.5 cm, base cuneate, apex acuminate or
acute, margin serrate or entire, yellowish brown
when young, green at maturity becoming reddish,
nerves numerous (c. 10-20 pairs), villous when
young, becoming + glabrous except for the midrib
and nerves; petiole c, 6-12 mm, villous. /nflores-
cence axillary and terminal, paniculate, shorter than

332

FLORA MALESIANA

[ser. I, vol. 102

the subtending leaf, c. 6.5 by 5.5 cm, peduncle up to
15 mm long, villous at first. Bracts c. 3 mm long,
lower ovate-lanceolate, upper broader, caducous.
Flowers uni- or bisexual, up to 4 by 2 mm, pedicel c.
1 mm long, villous; tepals 11—25(—28), in terminal
flowers c. 6 outer decussate, in lateral flowers the
outer 2 + opposite, the remainder spirally arranged,
the outer broadly ovate c. 1.5 mm long, peltate, grad-
ing into longer narrower tepals, the uppermost
spathulate, c. 3 mm long, ciliate on the upper part,
dark brown before anthesis, caducous. Stamens
9—25, c. 2—3.5 mm long, filament slender, c. 1 mm
long, anthers to 2 mm long, white, pinkish white or
cream at maturity, connective produced. Carpel c. 2
mm long, sparsely strigose, carpellode in male
flowers rudimentary or absent. Berry to 7.5 by 5mm,
crimson to dark purple-black when ripe. Seed ovoid,
c. 4 by 3 mm, smooth.

Distr. Malesia: Central Celebes, Moluccas (Ce-
ram, Batjan), and throughout New Guinea. Fig. 2.

Ecol. Common on ridge crests and exposed
places in primary low to mid-montane forests,
1000—2700 m. Also in regrowth after landslide and
fire, and on infertile stony sand in riverside or stream
bank vegetation.

Uses. The wood is used for fence posts and build-
ing. The leaves provide a treatment against dysentery
(Okapa area).

Vern. Arunan, nerch, tuna, Enga, butulye, Eipo-
mek, daloe, moble, Dani, edigea, Kapauko, gial,
Chimbu, guiamak, Kasanombe, kiluwe, kohbig, Ha-
gen, kora kiyei, Wonatabe, kuje, porlyporl, Mendi,
niebalaa, Kebar, pymbug, Melpa, paribara, Gerebi,
sakolo, Wapi, taingaa, tan-ja, Huli, wonnai, Ma-
ring.

Field notes. The bark is smooth and grey-
brown, the blaze is pinkish straw to red-brown with
few wide rays. The sapwood is pale straw to light red-
dish brown; the heartwood dark pink to red with
conspicuous light brownish to white rays. The bark
and crushed leaves have a peppermint-like odour and
the leaves are bitter when chewed.

2. Trimenia weinmanniifolia SEEMAN ssp. bougain-
villeensis RODENBURG, Blumea 19 (1971) 14. — T.
bougainvilleensis (RODENBURG) A.C. SmiTH, Aller-
tonia 1 (1978) 154.

Small tree or shrub, up to 10 m; young branches
reddish brown villous becoming + glabrous. Leaves
narrowly elliptic to lanceolate, 5—9 by 1.6—3 cm,
base cuneate, tapered to an obtuse apex or acumi-
nate, margin serrate, lateral nerves c. 14—20 pairs,
glabrous at maturity, at first sparsely villous; petiole
c. 10—15 mm, at first villous. Inflorescence axillary,
paniculate, equal to or somewhat shorter than the
subtending leaf, up to c. 85 by 55 mm, peduncle up
to 25 mm, villous; lower bracts ovate, c. 2 mm long,

upper broadly ovate, c. 1 mm long, strigose. ? Dioe-
cious. Flowers up to 3.5 by 2 mm, pedicel 1 mm
long; tepals 12—23, spirally arranged, the outer sub-
orbicular, c. 1.5 mm @, upper narrower and longer,
the uppermost spathulate c. 2.5 mm long, sparsely ci-
liate. Stamens 9-16, c. 3 mm long in male flowers,
filaments short, broad, anthers c. 2 mm long, whitish
at anthesis, connective produced, staminodes in fe-
male flowers c. 2 mm long. Carpel c. 2 mm long,
sparsely strigose, carpellode in male flowers c. 1 mm
long. Berry dark crimson to black-purple. Seed
ovoid, 2.5 by 1.7 mm, with distinct ridges.

Distr. Solomon Is. (Bougainville) and Malesia:
Papua New Guinea (E. New Britain: Pomio Sub-
distr.). The two other subspecies of 7. weinmannii-
folia occur in Fiji and Samoa and the widely distant
Marquesas Is. Fig. 2.

Ecol. In lower montane rain-forest and cloud
forest, especially on exposed ridges, 1500—1700 m.
Vern. Naligugu, naromalalawe, New Britain.

Note. RODENBURG (1971) reserved judgement on
material from New Britain, thinking it might repre-
sent another subspecies. Further material now avail-
able from New Britain indicates that this subspecies
extends outside the island of Bougainville. The iden-
tification of collections from New Ireland remains in
doubt.

3. Trimenia macrura (GILG & SCHLTR) PHILIPSON,
comb. nov. — Piptocalyx macrurus GILG & SCHLTR,
Bot. Jahrb. 55 (1919) 200, f. 2; ibid. 58 (1923) 246,
its Ihe

Woody climber, with terete brown tomentose
branches. Leaves elliptic-lanceolate, to c. 7-11 by
1.7—3 cm, chartaceous, base broadly cuneate to
rounded, apex produced into long delicate acumen
(c. 3 cm long), margin entire, nerves numerous, im-
pressed above, reticulations of smaller veins promi-
nent, becoming + glabrous above, densely rufous-
sericeous below; petiole c. 6-10 mm long, tomen-
tose. Inflorescence axillary and terminal, shorter
than the leaves, pleiochasia on long, brown-
tomentose peduncles. Flowers evidently uni- and bi-
sexual, c. 5 mm long, subsessile; tepals c. 8—10,
ovate to obovate, obtuse, glabrous. Stamens c.
10—12, 3.5—4 mm long, filaments slender, the an-
thers somewhat longer than the filaments, white or
pinkish at maturity, connective produced. Carpel
solitary, c. 2—2.5 mm long, sparsely rigose, carpel-
lode in male flowers rudimentary or absent. Fruit a
succulent berry c. 7 by 5 mm, deep crimson.

Distr. Malesia: Papua New Guinea (West Sepik,
Enga, Madang & Morobe Prov.).

Ecol. Climber in dense montane rain-forest and
mossy forest, between 2000—3000 m.

Field notes. The tepals are brown and cadu-
cous. The stamens are white or pinkish.

1986] TRIMENIACEAE (Philipson) | 333

Excluded

Idenburgia Gress, Fl. Arfak Mts (1917) 136 was reduced by vAN STEENIS, Svensk Bot. Tidskr. 49 (1955)
21 to Sphenostemon Batt. (Sphenostemonaceae).

‘Trimenia’ grandifolia Wars., Index Kewensis, Suppl. 1 (1906) 439, sphalm. for Trimeria grandifolia
(Hocust.) Wars. (Flacourtiaceae).

» ar,

i, aa

; an | tol, 4, 94198

A Rey: . ed Vcr viens CO &
Bar, bahwi i ri (rae DCMT. 44 7

D4
Pata tite. mnctionc some fn parrigas

nies
ere m bran ve enthe Meshy, oaslinulale;p alas fal ne
. | Mary a

Mtermet phous, &= Paes, uvdally Serenity a
eiMdieaiss racemes (up eno ay aa ya tew Br

paragin saath ”

oR gre. Aer

as wily, math, ' om

4 va ee ee

ALSEUOSMIACEAE (C.G.G.J. van Steenis, Leyden)

Until recently this small family was only known to occur in New Zealand and New Caledonia,
but in 1982 I have shown that it occurs in New Guinea and in 1984 that it is also represented in
East Australia.

Its phytographic history is complicated through the former confusion about the systematic af-
finity. Wittsteinia was described by F. VON MUELLER (1861) as probably belonging to Ericaceae
(or Pyrolaceae). Periomphale was described by BAILLON (1888) and has been affiliated to Caprifo-
liaceae or Gesneriaceae. In the ‘Pflanzenfamilien’ it was ranged among incertae sedis (Nachtr.,
1897). GitG & SCHLECHTER (1906) described two other genera from New Caledonia which have
appeared not to be different from Periomphale.

Recently I could show that Wittsteinia, which was by BurtTrT (1949) relegated to Epacridaceae,
is the oldest name for Periomphale and that, in addition to Alseuosmia from New Zealand, there
occurs in Queensland a new, peculiar, third genus, Crispiloba STEEN., of this family. The assem-
blage of the three genera is now a coherent, typically Australasian one.

In my mind the affinity is with Escalloniaceae cq. Grossulariaceae, and pending further system-
atic research in that group, we can maintain it as a family of its own, in agreement with CRONQUIST
(1981).

I am particularly satisfied that recent anatomical work on the three genera by Dr. W.C. Dick-
ISON, Chapel Hill (in /itt.) endorses my opinion.

1. WITTSTEINIA

F. v. M. Fragm. Phytogr. Austr. 2 (1861) 136; ibid. 3 (1863) 166; BurRTT, Kew
Bull. 3 (1949) 493; SrEveNs, Bot. J. Linn. Soc. 64 (1971) 45; J.H. WILIs,
Handb. PI. Vict. 2 (1973) 497; STEEN. Blumea 29 (1984) 391. — Periomphale
BaILL. Bull. Mens. Soc. Linn. Paris n. 92 (1888) 731; Hist. Pl. 10 (1888) 85;
GUILLAUMIN, Fl. Nouv. Caléd. (1948) 318; Airy SHAaw, Kew Bull. 18 (1965) 250;
GARDN. Blumea 24 (1978) 141; STEEN. ibid. 24 (1978) 480; Fl. Mal. I, 9 (1982)
556; v. ROYEN, Alp. Fl. New Guinea 4 (1983) 2659. — Memecylanthus GiLc &
SCHLTR, Bot. Jahrb. 39 (1906) 269. — Pachydiscus GiLc & SCHLTR, /.c. 270. —
Fig. 1.

Small shrubs. Leaves spiral, sometimes some in pairs or in false whorls, entire
or dentate, membranous or rather fleshy, exstipulate; axils puberulous. Flowers
fragrant, actinomorphous, 4—7-merous, usually 5-merous, axillary, solitary or
in pauciflorous racemes (up to 5 flowers), sustained by a few bracts, small, often
in part cleistogamous. Calyx persistent. Disk absent. Corolla sympetalous,
barrel-shaped, lobes + valvate, short, carunculate inside apex. Stamens mostly
5, free, alternipetalous; filaments filiform; anthers introrsely, lengthwise de-
hiscent. Ovary inferior, 2—3-celled, with flat apex, style columnar, as long as
the corolla, the globular, rugose stigma closely enveloped by the corolla seg-
ments. Ovules flattened, attached to the septum, (?1—)4—6 in each cell. Berry
small, globose, crowned by the calyx, few-seeded.

Distr. About 3—4 spp., Australia (Victoria), New Caledonia, in Malesia: Papua New Guinea.

Ecol. In forests, the Papuan species epiphytic.

Notes. A clearly Australasian, East Gondwanic genus, in which the Papuan species is manifestly allied
with the Australian species, not with the New Caledonian one(s).

The occurrence of cleistogamous flowers is highly peculiar and should be more closely studied in the field.

(335)

336

FLORA MALESIANA

[ser. I, vol. 102

BAILLON (1888) mentioned the occurrence of an orange disk but in herbarium specimens I cannot distinguish

this and assume it is the bulging apex of the ovary.

Whether the flowers are always bisexual is also uncertain; there is probably variability and they may appear

to be polygamous. This must be checked in the field.

Fig. 1. Wittsteinia papuana (STEEN.) STEEN. Small
apical part of the very slender habit, nat. size (type,
LAE 67068).

1. Wittsteinia papuana (STEEN.) STEEN. Blumea 29
(1984) 391. — Periomphale papuana STEEN. Blumea
24 (1978) 481; Fl. Mal. I, 9 (1982) 556; v. Royen,
Alp. Fl. New Guinea 4 (1983) 2660, f. 767. — Fig. 1.

Branched, very thin and slender, completely gla-
brous, epiphytic shrublet, c. 75 cm long. Twigs angu-
lar. Leaves scattered and in pseudoverticils, lanceo-
late-oblong, cuneate at base, acute at apex, margin
entire or mostly with 1—2 short gland-tipped teeth;
nerves 2—3 pairs, very erect; venation impressed
above, indistinct beneath; petiole 3—4 mm, much
widened at base. Flowers solitary, axillary; pedicel
emerging between a few minute bracts, 1—2 mm.
Calyx lobes 5, thickish, blunt deltoid, 1.25 by 1.5
mm. Corolla (in mature bud) narrowly barrel-
shaped, 6 mm long; lobes carunculate inside apex, 1
mm. Stamens 5, filaments 4 mm, anthers roundish,
cordate, 0.75 mm. Ovary c. 3 mm, 2-celled; ovules
4(—?5) in each cell.

Distr. Malesia: Papua New Guinea (W. Sepik,
Telefomin Subdistr., 5°S 141°E). One collection.

Ecol. Podocarpus-Phyllocladus woodland with
Gahnia tussocks undergrowth, 3000 m.

Affinity. The species is differing from the New
Caledonian one(s) by having dentate leaves and an
ascending habit producing many roots. These char-
acters are also found in the Australian W. vaccinia-
cea F. v. M. The latter is a + creeping or ascending,
rooting small shrub with larger, more coarsely and
more densely dentate leaves, an attenuate leaf base
with a hardly developed petiole, and hairy twigs and
lower portion of the leaves.

Field notes. Pedicels pink, gynoecium creamy
white, petals pinkish light green, stamens creamy
white.

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