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LIBRARY SERIES I - SPERMATOPHYTA
APR 26 1877 Flowering Plants
Vol. 7, part |
NEW YORK
BOTANICAL GARDEN
"+e
INDEX TO REVISED FAMILIES
Aceraceae . 4: 3, 592
Actinidiaceae s.str. . 4: 37
Aizoaceae 4: 267
Alismataceae . . . §: 317
Amaranthaceae 4: 69, 593
Ancistrocladaceae. 4: 8
Aponogetonaceae 4: 11, 7: 213
Basellaceae 5: 300
Batidaceae 5: 414
Betulaceae 52207
Bixaceae s. str. 4: 239
Burmanniaceae . 4: 13, 592
Burseraceae 5; 209
Butomaceae 5: 118
Byblidaceae oR
Callitrichaceae 4: 251
Campanulaceae 6: 107
Cannabinaceae 4: 223
Capparidaceae 6: 61
Caprifoliaceae . 4: 175, 598
Cardiopteridaceae fet
Celastraceae . . 6: 227, 389
Centrolepidaceae 5: 421
Ceratophyllaceae 4: 41
Chenopodiaceae . 4: 99, 594
Clethraceae pa es (A) Behe
Cochlospermaceae 4: 61
Combretaceae 4: 533
Connaraceae . . . 5: 495
Convolvulaceae . 4: 338, 599
Corynocarpaceae 4: 262
Crassulaceae . 4: 197
Datiscaceae 4: 382
Dichapetalaceae . 5: 305
Dilleniaceae 4: 141
Dioscoreaceae 4: 293
Dipsacaceae 4: 290
Droseraceae 4: 377
Elatinaceae
Epacridaceae .
Ericaceae .
Erythroxylaceae
Ficoidaceae
Flacourtiaceae
Flagellariaceae
Geraniaceae
Gnetaceae .
Gonystylaceae
Goodeniaceae
Haemodoraceae
Haloragaceae
Hamamelidaceae .
Hippocrateaceae .
Hydrocaryaceae
Hydrocharitaceae.
Hydrophyllaceae .
Icacinaceae
Juglandaceae .
Juncaceae .
Juncaginaceae.
Lemnaceae
Loganiaceae
Lophopyxidaceae
Malpighiaceae
Martyniaceae .
Molluginaceae
Moringaceae .
Myoporaceae .
Myricaceae
Najadaceae
Nyctaginaceae
Nyssaceae .
Ochnaceae
Oxalidaceae
Papaveraceae .
Pedaliaceae
: 203
“422
.
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Pentaphragmataceae. 4: 517
Pentaphylacaceae . 52121
Philydraceae 4:
Phytolaccaceae 4: 229
Pittosporaceae 5: 345
Plumbaginaceae 4: 107
Podostemaceae 4: 65
Polemoniaceae 4: 195
Pontederiaceae 4:.255
Portulacaceae Tin
Primulacea 6: 173
Proteaceae 5: 147
Punicaceae 4: 226
Restionaceae . 5: 416
Rhizophoraceae 5: 429
Salicaceae . 5: 107
Salvadoraceae. 4: 225
Sarcospermaceae . 4s. 32
Saururaceae 4: 47
Scyphostegiaceae . 5: 297
Simaroubaceae . . 6: 193
Sonneratiaceae . 4: 280, 513
Sparganiaceae 4: 233
Sphenocleaceae As (27
Stackhousiaceae . . 4: 35
Staphyleaceae 6: 49
Stylidiaceae . . 4: 529
Styracaceae es
Thymelaeaceae 4: 349, 6: 1
Trigoniaceae . .. & azar
Turmeracede . ... \ = eee
Typhaceae’ . 4 —Siaeas
Umbelliferae . . 4: 113, 595
Valerianaceae . 4: 253
Violaceae mp
Xyridaceae . 4: 366, 598
Zygophyllaceae . . 4: 64
ICACINACEAE (H. Sleumer, Leyden)
Trees, or whether or not climbing shrubs, or lianas. Leaves spirally arranged,
rarely opposite, simple, entire or lobed (in Mal. never crenate or serrate), penni-
or palmatinerved, exstipulate. Jnflorescences mostly axillary, sometimes terminal,
rarely extra-axillary, or from old wood, in spikes or spike-like racemes, or often in
cymes, both spikes and cymes not rarely collected to panicles or heads, very rarely
reduced to few-flowered fascicles or to a solitary flower. Flowers bi- or unisexual,
in the latter case at least functionally so, i.e. the plants dioecious, actinomorphic,
(4-)5(-6)-, by reduction rarely in part 3-merous, cyclic (with sepals or calyx lobes
and petals) or rarely spiral (with petals only in Pyrenacantha, or without petals in the
2 flowers of Platea and some spp. of Jodes and Gomphandra). Pedicels, if any,
articulated with the calyx. Sepals 4-6, free or mostly connate below to various
degree to a 4-6-lobed calyx, the lobes imbricate or valvate, generally persistent.
Petals 4-6, free or connate below to various degree, sometimes to a tube, the
lobes valvate, very rarely subimbricate, tip inflexed, mostly caducous, sometimes
persistent. Stamens as many as sepals or petals, episepalous, inserted basally or
sometimes in the upper part of the tube; filaments subulate, fleshy, often flattened,
or filiform, not rarely with clavate subglandular elongate hairs distally; anthers
2-celled, cells often diverging below, basifixed, latrorse or introrse, in Polyporandra
dismissing the pollen from numerous operculate pores. Disk whether or not
present, either annular or cup-like, free or adnate to the ovary, or a unilateral
fleshy scale. Ovary free, 1-celled (in Pseudobotrys, Gonocaryum and Citronella
2-celled with an empty tube-like unilateral cell) (in Mal.); ovules 2 (rarely 1
abortive), apical, pendent, anatropous, apotropous, unitegmic; style 1 or none;
stigma punctiform, subcapitate or peltate, entire or slightly 2—5-lobed or -crenate,
often depressed to one side. Drupe ellipsoid to globose, often laterally compressed
and almond-like; exocarp generally thin-fleshy; endocarp thin-crustaceous to
thick-woody, sometimes spongious or fibrous, often veined or ribbed lengthwise
or reticulate-lacunose outside, smooth or with tubercles or blunt aculei inside,
the seed pitted then. Seed 1, exarillate, generally with abundant endosperm, which
rarely is ruminate; embryo straight; cotyledons whether or not foliaceous.
Distribution. About 56 genera with c. 300 spp., all woody, predominantly in the tropics, rapidly
decreasing in number towards the subtropics; 5 genera with part of their species in the temperate zones
of Africa, Asia, Australia and S. America.
In Malesia a total of c. 100 spp. in 21 genera, of which 3 are strictly endemic, viz Cantleya (W. Malesia),
Hartleya and Pseudobotrys (both in New Guinea); 8 other genera find their main area of distribution and
generally their greatest number of species in Malesia, but occur also in parts of S. and SE. Asia, viz
Gonocaryum, Platea (both also in New Britain), Codiocarpus, Stemonurus (also in the Solomon Is.),
Miquelia, Nothapodytes, Phytocrene, and Sarcostigma; 3 genera, viz Apodytes, lodes and Pyrenacantha,
are found in Africa (also Madagascar), SE. Asia and Malesia. The genus Citronella is amphipacific
(Malesia, E. Australia, Melanesia, New Caledonia, New Hebrides, Fiji, Samoa, Tonga, Costa Rica,
Venezuela, Brazil, Paraguay, N. Argentina, Bolivia, and Central Chile); 6 genera, viz Gomphandra,
Medusanthera, Merrilliodendron, Polyporandra, Rhyticaryum, and Stemonurus, extend from Malesia into
Micronesia and Melanesia or even W. Polynesia; of these only Gomphandra and Rhyticaryum are also
found in NE. Australia. Whitmorea, so far known, is limited to the Solomon Is., but might occur also in
SE. New Guinea.
Icacinaceae of Malesia show a strong affinity with those of tropical South Asia (SW. India, Ceylon)
and Southeast Asia (Sikkim to Assam, Bengal, Burma, Thailand, Indo-China, W., Central and S. China
incl. Hainan and Formosa, S. Japan). Most of the members of the family in Australia point to an
Asiatic-Malesian origin; Irvingbaileya, limited to Queensland, is taxonomically very close to Codio-
(1)
2 FLORA MALESIANA [ser. I, vol. F
carpus and Medusanthera. Pennantia in New Zealand and on Norfolk I. is more distinct from the Asiatic-
Malesian bloc of genera. New Caledonia has, besides Citronella, 2 endemic genera, viz Anisomallon
(allied to Apodytes) and Gastrolepis (related to the group of genera around Medusanthera).
Ecology. Most Malesian spp. occur in the substage of the primary or secondary rain-forest, a few
are canopy trees (Stemonurus, Platea, Cantleya, Citronella p.p.). They are mainly found in the lowlands,
but may ascend to the montane and even the mossy forest (Platea p.p., Rhyticaryum p.p., Gomphandra
p.p.) up to 2400 m (higher altitudes mentioned on collector’s labels being rather doubtful); several genera
reach their highest altitude on Mt Kinabalu or in New Guinea. The distribution is scattered in general
for the tree species, even more so for the climbers, and can cover enormous areas as for instance Platea
latifolia, a species ranging from Sikkim, Himalaya, to New Guinea. Several species (Stemonurus, Gom-
phandra; Platea even with stilt roots) are clearly limited to peat swamp forest. Some Jcacinaceae are found
also in coastal forest, and such species may possibly derive their wide range of distribution from the
floating capacity of the fruit.
Large dimensions are reached only by a few species of Platea, Cantleya, Stemonurus, Citronella, Gom-
phandra, Apodytes, or rarely by such of Nothapodytes and Medusanthera.
Among the rain-forest species none is reported to be dominant in any vegetation type, though oc-
casionally single species may be locally frequent.
Few species occur in the drier, mainly seasonally dry parts of Malesia, all of them being shrubs or
climbers.
Dispersal. Little is known of the dispersal of fruits, part of which are eaten by wild animals (e.g.
of Gonocaryum, Cantleya, Gomphandra, Medusanthera). Buoyancy of fruits is certain for Merrillioden-
dron, less so for Gonocaryum and Stemonurus, and may have contributed to their dispersal.
Galls. No galls specific for a distinct genus or species are hitherto known from Malesian Icacinaceae
(cf. DoCTERS VAN LEEUWEN, Zoocecidia, 1926, 332, f. 597-599: Gomphandra, Phytocrene, Platea; Ned.
Kruidk. Arch. 51, 1941, 175, f. 48: Stemonurus).
Anatomy. Wood. Structure, properties and identification: DEN BERGER, Determinatietabel Malesié,
Veenman, Wageningen (1949) (hand lens), BALAN MENON, Res. Pamphl. For. Dept Malaya 18 (1955)
1-16 (Cantleya), CHATTAWAY, Trop. Woods 102 (1955) 55-74, ibid. 104 (1956) 100-124 (crystals),
Desc, Mal. For. Rec. 15 (1941) 233 (Cantleya, Stemonurus), FEI-TAN & Cuu, Malayan Forester 32
(1969) 287-293 (fibres, Stemonurus), GosH, RAO c.s. Indian Woods 2 (1963) New Delhi (Apodytes,
Gomphandra, Nothapodytes as ‘Mappia’), JANSSONIUS, Key to Javanese woods, Leyden (1952), and MOLL
& JANsSoNIuS, Mikr. 2 (1911) 234-254 (Apodytes, Gomphandra, Platea, Stemonurus), VAN DER WALT C.S.
T. Nat. Wetensk. Suid-Afr. (1970) 173-199 (Pyrenacantha, anomalous growth). For general surveys
also comprehensively covering the older literature see SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899)
227-237 and ibid. (1908) 81-83, and METCALFE & CHALK, Anat. Dicot. Oxford (1950) 367-377.
The vegetative anatomy of the Icacinaceae is extremely diverse and no character from leaf, twig or
wood anatomy is consistent or even typical for the family. Because of the enormous range in structure,
anatomical characters have been extensively used in discussing the relationships and subdivisions
within the family. ENGLER’s wood-anatomical characterizations of the tribes Icacineae, lodeae, Sar-
costigmateae and Phytocreneae (Sitz. Ber. Preuss. Akad. Wiss. Berlin 18, 1893, 1-23) were not based on
enough material and therefore inaccurate. BAILEY & Howarp (J. Arn. Arb. 22, 1941, 125-132, 171-1 87,
432-442, 556-568) distinguished three groups based on a combination of the nodal anatomy and the
type of vessel perforation. Group I with trilacunar nodes and exclusively scalariform vessel perforation
includes Apodytes, Citronella, Pittosporopsis and Platea (all Icacineae), Group II with trilacunar nodes
and a mixture of simple and scalariform vessel perforations includes Cantleya, Gonocaryum, Medusan-
thera, Gomphandra (as ‘Stemonurus’) and Stemonurus (as ‘Urandra’) (all Icacineae). Group Il with
unilacunar nodes and simple vessel perforations includes Iodes, Mappianthus, Merrilliodendron, Miquelia,
Natsiatum, Nothapodytes, Phytocrene, Polyporandra, Pyrenacantha, Rhyticaryum, and Sarcostigma
(from all four tribes sensu ENGLER). Other xylem characters such as vessel distribution, vessel member
length, type of fibres, parenchyma arrangement and ray structure are more or less related to these groups
of increasing xylem specialization. Codiocarpus, Hartleya, Pseudobotrys and Whitmorea were not in-
cluded in the studies of BAILEY & Howarpb. Whitmorea has a trilacunar node and mixed vessel perfo-
rations and therefore belongs to group II. This observation supports SLEUMER’s view that Whitmorea
is related to Stemonurus (Blumea 17, 1969, 264). All data on wood anatomy provided by BAILEY &
Howarb suggest a more or less continuous range of xylem characters within Icacinaeceae rather than the
presence of distinct boundaries.
Hairs. HEINTZELMANN & Howarp (Amer. J. Bot. 35, 1948, 42-52) distinguish ten hair types that
occur in floral parts or on leaves and young twigs of Icacinaceae. Noteworthy are the two-armed or
Malpighiaceous hairs in Nothapodytes and some genera from outside Malesia. This hair type intergrades
with the so-called Icacinaceous hair with one much reduced and one well developed arm occurring in a
majority of the genera. Rhyticaryum has clustered hairs; Platea stellate-peltate hairs (called ‘scales’
in the taxonomic part) (see also UpHoF, HUMMEL & STAESCHE, Handb. Pflanzenanat. IV, 5, 1962).
Stomata are very imperfectly known in Icacinaceae. Both anomocytic and paracytic stomata have
been recorded in literature, but actinocytic (Merrilliodendron) and anisocytic (Medusanthera and Ste-
monurus) types also occur.
1971] ICACINACEAE (Sleumer) 3
The vascularization of the petiole and midrib would also merit further investigation. Species of
Apodytes, Citronella, Medusanthera, Phytocrene, and Stemonurus have central strands with or without
latero-dorsal bundles, but the range in the whole family must be much greater since Pennantia shows a
very complex pattern (cf. Blumea 18, 1970, 217).
Crystals occur as druses, solitary rhomboids and crystal sand in various combinations (cf. HEINTZEL-
MANN & Howard, l.c.).
The fine translucent dots of the leaves of Merrilliodendron are caused by large intercellular spaces in
the spongy tissue of the mesophyll.
Domatia have been recorded for several Icacinaceous genera but are absent from all Malesian species.
— P. BAAS.
Phytochemistry. The few chemical data available about the chemistry of this family were summariz-
ed in my ‘Chemotaxonomie der Pflanzen’ 4 (1966) 275-277, 494. In the meantime, oleanolic acid was
isolated from the bark of Apodytes dimidiata E. MEYER (= A. beddomei Mast.). The scanty chemical
information available about /cacinaceae at this time, prevents a chemotaxonomic discussion. — R.
HEGNAUER.
Uses. In Malesia but a few tree species grow to big dimension with a clear bole (Cantleya, Stemonurus,
Platea). Of these only Cantleya corniculata (BECC.) HoOwARD has a marketing value and is exported from
Sarawak and Brunei. /cacinaceae have a hard or mostly rather soft, often whitish or cream, sometimes
aromatic wood, and are apparently only locally used by the natives, as can be deduced from the many
vernacular names known.
The leaves of Rhyticaryum species are eaten as a vegetable. Medicinal use is recorded for several
species but needs confirmation. The seeds of Cantleya, Phytocrene, Stemonurus and Sarcostigma are
edible, but of a poor quality. The stems of lianas (Miquelia, Phytocrene) hold fresh edible water.
Taxonomy. Four of the genera found in Asia and Malesia were revised by R. A. HOwARD (Cantleya:
J. Arn. Arb. 21, 1940, 479; Codiocarpus: Brittonia 5, 1943, 60; Medusanthera: Lloydia 6, 1943, 133;
Nothapodytes: J. Arn. Arb. 23, 1942, 66); these revisions were based on rather scarce materials as far as
Malesia is concerned. A precursory paper with revisions of most of the genera concerned and based on
practically all Asiatic and Malesian specimens available today was published by myself in Blumea 17
(1969) 181-264, supplementing my previous studies on the family (Notizbl. Berl.—Dahl. 15, 1940, 228-257;
ibid., 1942, 359-365; in E. & P. Natl. Pfl. Fam. ed. 2, 20 b, 1942, 322-396).
KEY TO THE GENERA
1. Trees or (not climbing) shrubs.
2. Sepals essentially free at least in their upper 34 and imbricate.
3. Flowers bisexual.
4. Petals free, valvate, or subimbricate in their upper part. . . wie 1. Citronella
4. Petals connate below into a tube, their upper part free and valvate ii in bud. 2. Pseudobotrys
3. Flowers unisexual. (Petals connate below.)
5. Filaments free, fixed to the very base of the petals. Leaves with a layer of rounded to star-shaped
appressed scales underneath at least in the young state. . . .......+4.4.2.-. 3. Platea
5. Filaments adnate for almost their entire length to the lower tubular part of the petals. Scales
LT ei SRP re Sr re Pier . . 4. Gonocaryum
2. Sepals connate into: a cup- -like calyx, its upper free part (or lobes), if any, short and not imbricate.
6. Flowers unisexual (or functionally so).
7. Drupe ovoid-ellipsoid or oblongoid, without a fleshy lateral appendage. Flowers in rather short
cymes. . a hee. no whee) a Wane ie te ie oP ne oe
7. Drupe laterally compressed (almond- like).
8. Drupe without a fleshy lateral appendage. Flowers in spikes (very rarely in panicles composed
OF scee. OF SUNOS TASCICIEd) (oo cs. aes inp tees Bue ty age F Rhyticaryum
8. Drupe with a thick fleshy, laterally borne, practically entirely adnate appendage. Flowers in
cymes.
9. Disk unilateral, thick-squamular ......... Pred ree Sa ener . 7. Hartleya
9. Disk absent.
10. Filaments glabrous. Fleshy appendage of drupe covering two pronounced ribs of the endocarp.
8. Codiocarpus
10. Filaments with apical, longish, club-shaped hairs at least in the fertile stamens. Prominent
ribs under the appendage of the drupe less pronounced or absent. . . 9. Medusanthera
6. Flowers bisexual.
11. Ovary with a lateral swelling which in the fruit developes into a thick succulent appendage.
Denies wheal aft ass BFR) Attias awe doa Ye thw eas Se er eg 10. Apodytes
11. Ovary and fruit without such an adnate appendage. Disk whether or not present.
12. Connective surpassing the anther cells as a marked glabrous apiculus. Outer part of the endocarp
finally spongious-corky and deeply irregularly lacunose. Disk absent. 11. Merrilliodendron
4 FLORA MALESIANA (ser. I, voip ae
12. Connective, if any, hardly or not surpassing the anther cells. Outer part of the endocarp fibrous,
slightly ribbed or grooved lengthwise, or smooth outside.
13. Peduncle of inflorescence with numerous small knob-like bracts which form alveoles. Stigma
peltate. Disk absent. . . 2 a Sh) 12 @antiaga
13. Peduncle quite smooth. Stigma ‘small, ‘subcapitate or r point- “like. Disk a cup-shaped.
14. Inflorescence usually terminal. Anthers glabrous... . . . . . 13. Nothapodytes
14. Inflorescence axillary. Anthers with an apical tuft of penicillate hairs.
15. Flowers sessile. Petals up to 6 mm, free to almost the base. Stigma point-like at the top of
the +--+ attenuate (sometimes shortly style-like) part of the ovary. . . . . 14. Stemonurus
15. Flowers 1-2 mm pedicelled. Petals (12—)13—15 mm, free in the distal part only. Stigma small
on one side of the inverted, i.e. cup-like distal part of the ovary. . . . 15. Whitmorea
1. Climbing shrubs, or lianas, sometimes with tendrils. Flowers unisexual or functionally so.
16. Leaves opposite.
17. Anthers broadly club-shaped to subglobular, many-celled, with numerous pollen-bearing alveoles.
16. Polyporandra
17: ‘Anthers;\as ‘usual; with 2 ‘cellsy SOE C90 Oa ee WO
16. Leaves spirally arranged.
18. Flowers in elongate spikes or spike-like racemes, these solitary or sometimes composed to panicles.
19. Leaves markedly prominently tessellate on both faces. Sepals persistent. Albumen absent;
cotyledons thick-fleshy. . . . . . 18. Sarcostigma
19. Leaves with rather lax and but slightly raised reticulation: ‘Sepals. absent. Albumen thick, ru-
minate; cotyledons foliaceous. . . ... . 19. Pyrenacantha
18. Flowers in peduncled heads or umbels, these solitary ¢ or composed to racemes or panicles.
20. Style absent, i.e. stigma sessile, thick-peltate. . . . » leh: SS ORE
20. Style (very) shortly thick-columnar, with 2—4 stigmatic lobes. bE MS . ZLB ioeecns
1. CITRONELLA
D. Don, Edinb. New Phil. J. 13 (1832) 243; SLeum. Blumea 17 (1969) 186. —
Villaresia R. & P. Fl. Per. Chil. 3 (1803) 9, t. 231, non R. & P. 1793. — Pleuropeta-
lon BL. Mus. Bot. Lugd.-Bat. 1 (1850) 248, non Hook. f. 1845. — Chariessa
Mia. FI. Ind. Bat. 1 (1856) 794. — Fig. 1.
Trees or shrubs, bisexual (Mal.). Leaves spiral, entire and slightly revolute at
the edge (Mal.), petiolate. Inflorescences terminal and/or (supra-)axillary, panic-
ulate or thyrsoid, cymes of secondary branches scorpioid, whether or not forked,
in Mal. greyish to rusty hirsute. Flowers 5-merous, sessile, subtended by a minute
bract. Calyx imbricately lobed 14 to 14, persistent. Petals free, valvate, or subim-
bricate in the upper part, apex inflexed, midrib inside much prominent, sometimes
wing-like. Stamens 5, free; filaments fleshy, subulate; anthers elliptic, or ovate, or
subcordate, introrse, basifixed. Disk 0. Ovary subgibbous, 1-celled, rarely 2-celled
by the presence of a pseudoloculus. Style 1 (rarely 2), slender, stigma small,
capitate, subbilobed. Drupe moderately fleshy; endocarp woody, rather thin.
Seed 1, longitudinally plicate around the vertical pseudoloculus, hippocrepiform;
embryo in the copious endosperm small.
Distr. About 21 spp. of which c. 12 occur in tropical Central and South America, and 6 spp. in
Australia (New South Wales, Queensland), Solomon Is., New Caledonia, Loyalty Is., New Hebrides,
Fiji, Samoa and Tonga Is.; in Malesia 3 spp. Fig. 2.
Ecol. Understorey of lowland and lower montane rain-forest.
Taxon. The genus was revised by R. A. Howarp (Contr. Gray Herb. 142, 1942, 60-89, t. 4-6). The
inflorescence and floral characters used there to separate the Malesian spp. have been found of little value
after a reinvestigation on the base of much more material than seen by HowarD. This is the reason
why in the key fruit characters have been preferred for the discrimination of species. HOWARD keeps the
New World spp. apart from the Old World ones on the sectional level; the characters given for the dis-
crimination of these sections are rather vague or do not hold.
1971] ICACINACEAE (Sleumer) 5
an
Fig. 1. Citronella suaveolens (BL.) Howarb. a. Inflorescence, x 14, b. flower bud, x 5, c. petal from the
inside, «714, d. ovary, X714, e. stamen, x7'4, f. habit and infructescence, x 4, g. drupe, x1 (a-e
ACHMAD 1181, f/-g ACHMAD 220).
KEY TO THE SPECIES
1. Drupe essentially ellipsoid-oblongoid, (1.6-)1.8-2.2(-2.4) by 0.9-1.2(-1.3) cm. . 1. C. suaveolens
1. Drupe ovoid, or ellipsoid-ovoid, or broadly ellipsoid, 2-3 by 1.8-2 cm.
2. Leaves broadly ovate, base + obtuse to rounded. Solomon Is., Samoa, Tonga, New Hebrides.
C. samoensis (A. Gray) Howard
2. Leaves oblong- to obovate-elliptic, base + cuneate.
3. Leaves + obtuse or shortly and rather abruptly acuminate, tip bluntish. . . 2. C. latifolia
3. Leaves rather gradually and + acutely acuminate at apex... ..... .- 3. C. philippinensis
6 FLORA MALESIANA
1. Citronella suayeolens (BL.) Howarp, J. Arn.
Arb. 21 (1940) 475; Contr. Gray Herb. 142
(1942) 82; BAcK. & BAKH. f. Fl. Java 2 (1965) 59;
SLEUM. Blumea 17 (1969) 187. — Pleuropetalon
suaveolens Bu. Mus. Bot. Lugd.-Bat. 1 (1850) 248.
— Chariessa suaveolens (BL.) Mia. Fl. Ind. Bat. 1
(1856) 794; Becc. Malesia 1 (1877) 118; SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 229; AMsH. in
Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135,
p. 2. — Villaresia suaveolens (BL.) VALET. Crit.
Overz. Olacin. (1886) 199, t. 5, f. 32 a-g; BACK.
Schoolfl. Java (1911) 228. C. brassii HOWARD,
Contr. Gray Herb. 142 (1942) 81. — Fig. 1.
Tree, (8—)15—36 m, trunk up to 14 m 9, rarely
with thin buttresses; bark grey-brown, irregularly
fissured. Branchlets smooth or striate, older parts
lengthwise set with elongate lenticels. Leaves
elliptic to oblong- or ovate-elliptic, variable in
shape and size, apex generally short-acuminate
(tip subacute and often plicate then), or obtuse-
rounded, base broadly attenuate to rounded, often
subequal, coriaceous, shining above, (6—)10-24(30)
by (3-)5-13(-17) cm, nerves (4-)5-6 curved-
ascendent and anastomosing pairs, lower 1(-—2)
rather close to the base of the lamina, slightly
impressed above, much raised beneath, retic-
ulation fine to rather coarse, a little prominent
above, more distinctly so beneath; petiole 6-15 by
2-3 mm. Panicles terminal, greyish-rusty hirsutu-
lous, mostly solitary and spike-like initially, some-
times 2 or 3 forming a -++ pyramidal inflorescence,
lateral branches of the single panicles of + the
same length (0.5-2 cm), mostly 2-(rarely more-)
4 1 ———
(ser. I, vol 7
forked, each branch recurved and bearing several
flowers arranged in a cyme. Flowers bisexual,
fragrant. Calyx c. 1.8 mm, _ lobes ciliate.
Petals elliptic-oblong, white to cream, or some-
times suffused with red, (4-)5(-6)mm at full
anthesis, midrib inside raised considerably, some-
times almost wing-like in the lower 14-34 part.
Filaments thick-subulate, c. 3 mm; anthers ovate-
elliptic or subcordate, 1mm. Ovary ovoid,
glabrous, style 1-2 mm, stigma small, a little
oblique. Drupe ellipsoid to oblongoid (subcylin-
drical), sometimes a little oblique, (1.6—)1.8—2.2
(2.4) by 0.9-1.2(—1.3) cm, smooth or irregularly
lengthwise, very slightly many-ribbed in the lower
part, purple when ripe, blackish when dry.
Distr. Malesia: NW. Sumatra (Simalur I.), W.
& Central Java, N. & E. Borneo, Celebes, S.
Moluccas (Kai Is.), New Guinea.
Ecol. Primary rain-forest on well drained soil,
from the lowland up to 1600m, apparently
scattered.
Uses. Wood light brown, hard and heavy, not
used. Known by numerous vernacular names.
Vern. Simalur: awa iseum-iseum, iseum batu, i.
iseum fatuh, i. pajo, i. sito bulung, sitenheur délok,
tutun surimanu, M; Java: jérukie, S; Celebes:
kekér, Menado (Tt), balula lembo, Tobela,
lenguru tanru, pano, Bugin., sokko, Makassar;
New Guinea: garus, Dumpu, mara, Faita, pomak,
Armat, sakohukwa, Manikiong, sibeer, Hattam,
soromma rinde, Depapre, tambeu, Iko (Hol-
landia), ulumon, Amele, yewel, Bilia.
EE
4 ee ——————l
Fig. 2. Distribution of Citronella.
2. Citronella latifolia (MERR.) Howarp, J. Arn.
Arb. 21 (1940) 472; Contr. Gray Herb. 142 (1942)
80, t. 6, f. 9-12; SLEUM. Blumea 17 (1969) 188. —
Villaresia latifolia MERR. Philip. J. Sc. 14 (1919)
415; En. Philip. 2 (1923) 488. — Chariessa latifolia
({MERR.) SLEUM. Notizbl. Berl.-Dahl. 15 (1940)
229.
Large tree. Branchlets dark brown, with sparse
lenticels. Leaves obovate or oblong-elliptic, apex
obtuse or rounded, the bluntish tip sometimes
abruptly short-acuminate and folded, base cu-
neate, a little decurrent, coriaceous, shining above,
(12-)14-18 by (6—-)8-11 cm, nerves 5—6 anasto-
mosing pairs, reticulation rather dense, minutely
raised on both faces; petiole 1-1.5(-2.5) cm by
2(-3) mm. Jnfructescence terminal, consisting of
1 or 2, sometimes 3 almost pyramidal panicles,
up to 15cm, rusty hirsute; each panicle with
1971]
ICACINACEAE (Sleumer) 7
numerous racemosely arranged branches which
generally are 2(-3)-forked and bear several
flowers crowded at the curved end of the branches,
the latter up to 5cm in the lowest, and c. 1 cm
(and usually not forked) in the uppermost part
of the panicle. Drupe ovoid to broadly ellipsoid-
ovoid, c. 2 by 1.8 cm in submature state, brown
when dry, very slightly and irregularly many-
ribbed lengthwise in the lower 24; endocarp
hard, c. 1 mm.
Distr. Malesia: Philippines (Samar, in the
Catubig R. area), one collection in fruit.
Ecol. Damp forest near river at low altitude.
Vern. Malaampipi, S. L. Bis.
3. Citronella philippinensis (MERR.) Howarp, J.
Arn. Arb. 21 (1940) 474; Contr. Gray Herb. 142
(1942) 85, t. 6, f. 1-8; Dani, J. Arn. Arb. 33
(1952) 260, f. 20 & 20 A (pollen); SLEUM. Blumea
17 (1969) 188. — Villaresia philippinensis MERR.
Philip. J. Sc. 14 (1919) 414; En. Philip. 2 (1923)
488. — Chariessa_ philippinensis (MERR.)
Steum. Notizbl. Berl.-Dahl. 15 (1940) 229;
Hatus. Mem. Fac. Agr. Kagosh. Un. 5,3 (1966) 38.
Tree, 4-5 m. Branchlets smooth. Leaves elliptic
to elliptic-oblong, apex rather gradually acu-
minate for 1—-1.5 cm and subacute, base cuneate
and a little decurrent on the petiole, + inequal,
coriaceous, shining on both faces, 6-12 by (2—)4~—
6.5 cm, nerves 4-5 rather steeply curved-ascending
pairs, reticulation dense, distinctly raised mainly
on the undersurface; petiole 2(-3) cm by 2 mm.
Panicles terminal, solitary or in twos, 5—8(—10) cm,
lateral branches numerous and of equal length
(c. 1 cm), rather crowded, each bearing at apex
5-8 crowded cymosely arranged flowers. Calyx
1.5 mm, lobes ciliate. Petals oblong, white, gla-
brous, 5(-6) mm, the midrib extending as a keel
in the lower half of the petal. Filaments c. 5 mm;
anthers broadly ovate-cordate, 1mm. Ovary
ovoid, glabrous, 5mm, style slender, 3-4 mm,
stigma capitate, minute. Drupe ovoid, rather
oblique, blackish, 2.5—3 by 2 cm; endocarp shallow-
ly and irregularly many-ribbed lengthwise, 1—1.5
mm @.
Distr. Malesia: Philippines (N. Luzon).
Ecol. On forested slopes, c. 1000 m, apparently
rare.
Excluded
Villaresia scandens Hassk. Nat. Tijd. N. I. 10
(1856) 152 = Dichapetalum timoriense (DC.)
BoeERL. (Dichapetalaceae).
2. PSEUDOBOTRYS
MOESER in Fedde, Rep. 10 (1912) 310; SLEUM. in E. & P. Nat. Pfl. Fam. ed. 2,
20 b (1942) 360, f. 101, A & B; Blumea 17 (1969) 248. — Fig. 4.
Trees. Leaves spirally arranged, coriaceous, entire, penninerved. Flowers bi-
sexual, born in short fascicles or panicles on the trunk. Pedicel articulated with the
calyx, subtended by 2 small bracteoles. Sepals 5, connate at base, imbricate.
Petals 5, linear, valvate, tips a little inflexed inside, connate into a tube in the
lower 1-14, caducous. Stamens 5, almost equalling the petals in length; filaments
filiform, dilated towards the base and fixed there to the petals; anthers linear,
subsagittate, introrse, almost basifixed. Ovary ovoid-subglobose; style filiform;
stigma small, depressed-subcapitate. Drupe: exocarp thin, fleshy, early dissoluted;
endocarp woody, verrucose outside, forming a pseudoseptum along the funiculus
inside. Seed 1; embryo small.
Distr. Malesia (New Guinea), 2 spp. Fig. 3.
KEY TO THE SPECIES
1. Ovary densely brownish-pubescent. Petals 2.8-3.2 cm. Anthers 5-6 mm. .
1. Ovary glabrous. Petals c. 2.2 cm. Anthers 3-4mm..............
1. P. dorae
x if tk pew
1, Pseudobotrys dorae Moeser in Fedde, Rep. 10 coriaceous, of a dull olivaceous-green colour and a
(1912) 310; Steum. Notizbl. Berl.-Dahl. 15 (1940)
235; in E. & P. Nat. Pfl. Fam. ed. 2, 20 b (1942)
360, f. 101, A & B; Blumea 17 (1969) 249.
Treelet or shrub, 1.5-6 m, glabrous; bark grey,
squamular. Branchlets terete, striate, slender.
Leaves lanceolate to oblong or elliptic-oblong,
apex gradually more shortly or long acuminate,
tip blunt, base broadly cuneate to rounded,
minutely tubercled undersurface when dry, greenish
yellowish in fresh specimens, entire, 16—32(-44)
by 5-14(-18) cm, midrib slightly raised above,
strongly so beneath, nerves (6—)8—10(—12) pairs,
lower ones curved, upper ones generally more
straight, faintly inarching before the edge, hardly
raised or obscure above, slightly prominent beneath,
veins lax, generally rather inconspicuous; petiole
8 FLORA MALESIANA [ser. I, vol. 71
Fig. 4. Pseudobotrys cauliflora (PULLE) SLEUM. a. Habit, x 14, d. inflorescence, x 1, c. young infructescence,
x1, d. flower, x 114, e. ovary, x1 Y, f. endocarp of mature drupe, x1 (a & c DocTERS VAN LEEUWEN
9127, b, d & e ditto 9307, f ditto 11243).
1971]
ICACINACEAE (Sleumer) 9
rugose, 1—1.5(—2) cm by 2-3 mm. Flowers in short
fascicles or corymbs from swollen parts of the
trunk (occasionally also from the axils of leaves?).
Pedicels thickish, short. Sepals subovate, blunt,
dull purple, c. 3mm. Petals linear, blunt, thin,
connate irregularly in the lower part, i.e. early
splitting from top after the bud stage, spreading or
recurved at full anthesis, white all over, or pur-
plish at tube, 2.8-3.2 cm by 2-2.5 mm. Filaments
2-2.4 cm; anthers 5-6 mm. Ovary 34 mm, densely
brownish-pubescent; style 2.5—2.7 cm. Drupe only
known in submature state, obliquely obovoid,
apiculate, laterally a little compressed, c. 2 by 1.7
by 1.3cm, subtended by the reflexed sepals,
brownish-hairy, finely verrucose outside; endocarp
hard; pseudoseptum protruding for c. 1 mm. Seed
1.
Distr. Malesia: New Guinea (Morobe, Central
and Northern Distr.). Fig. 3.
Fig. 3. Distribution of Pseudobotrys. P. dorae
Moeser (crosses), P. cauliflora (PULLE) SLEUM.
(dots).
apparently very local. F/. May-Sept., fr. Nov.
2. Pseudobotrys cauliflora (PULLE) SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 235; DAHL, J. Arn. Arb. 36
(1955) 160, f. 1 & 1 A (pollen); SLEUM. Blumea 17
(1969) 249. — Chariessa cauliflora PULLE, Nova
Guinea 8 (1912) 657; SCHELLENB. Bot. Jahrb. 58
(1923) 159; BirNiE, Nova Guinea 14 (1926) 275.
— Fig. 4.
Tree, (4-)6—-18 m; bark dark grey-green; wood
white, with prominent rays. Leaves very similar to
those of P. dorae. though sometimes more ovate,
up to 36 by 19cm. Flowers massed on dome-like
swellings of 1-3(-10)cm o@ on lower trunk,
numerous, set in small fascicles. Pedicels 1-3 mm,
minutely bracteolate at base. Sepals ovate, ciliate,
purple, 2(-3) mm. Petals linear, apex acute, in
bud stage coherent in a tube which splits down-
wards at full anthesis to the lower third or less,
white, or white inside and purplish outside, with 3
longitudinal darker lines, c. 2.2 cm by 2 mm.
Filaments c. 1.6 cm; anthers 3-4 mm. Ovary gla-
brous, c. 3 mm; style slender, 16-18 mm. Drupe
ovoid, attenuate-apiculate, 2.5-3 by 2cm, on
thickish pedicel 8-12 mm; exocarp fleshy, dark
blue at full maturity, wrinkled in dry fruit; en-
docarp woody (2mm), outside with knob-like
emergences in numerous longitudinal rows, knobs
smaller towards apex of the fruit; inside with a
large woody pseudoseptum around which the
seed is formed.
Distr. Malesia: New Guinea. Fig. 3.
Ecol. Rain-forest substage, river flood-plains,
or hillsides, 3-175 m (once said to be collected at
1645 m), scattered in general, though sometimes
locally common. Fi. fr. Jan.—Dec.
3. PLATEA
BL. Bijdr. (1826) 646; SLEUM. Blumea 17 (1969) 242. — Platystigma R. BR. in
Wall. Cat. (1832) n. 7523, nom. nud.; ex BENTH. in Benth. & Hook. f. Gen. Pl. 3
(1880) 283, nota; Hook. f. Ic. Pl. 18 (1887) t. 1707. — Fig. 6.
Trees, generally tall and with straight, angular or -+- deeply grooved bole, rarely
treelets or shrubs, dioecious. Leaves spiral, almost distichous, entire, penninerved,
at least initially covered with fascicled or star-shaped, early going hairs, and/or
subpersistent star-shaped, flat, finally silvery scales with a distinct darker centre,
as are branchlets and inflorescences; petioles not rarely transversely wrinkled as in
Gonocaryum. Flowers axillary, the 3 ones in interrupted, glomerulose, mostly to
panicles arranged spikes, sessile, subglobular in bud, each subtended by a small
persistent bract, the 2 ones in few-flowered short cymes, each on a distinct pedicel
and subtended by a subpersistent bract immediately below the calyx lobes. Calyx
lobes (4-)5, slightly imbricate, small, persistent. Petals (4—)5, small, valvate,
glabrous, inflexed at apex, shortly connate at base, absent in the 2 flowers. —
3 Flowers: Stamens (4-)5, inserted with short filaments at the tube of the petals;
anther cells ovoid—2-celled, basifixed, laterally (almost extrorsely) dehiscent. Ru-
diment of an ovary generally absent. — 9 Flowers: Ovary thick-cylindric to ob-
10 FLORA MALESIANA [ser. ie vol. at
conical, blunt, crowned by a large sessile discoid windy Drupe elongate-ovoid,
crowned by the large stigma, on short pedicel; exocarp thin-fleshy, red-orange to
finally purplish blackish; endocarp woody, thin, with 1 or 2 slight longitudinal
grooves, reticulately wrinkled; embryo small, in the apex of the albumen.
Distr. About 5 spp., one endemic in Hainan, another extending from Sikkim into SE. Asia and
Malesia; in Malesia 4 spp., of which 2 endemic in Borneo and the other 2 from the Malay Peninsula and
Sumatra to New Guinea and New Britain (not known yet from the Solomon Is.). Fig. 5.
Ecol. Mainly rain-forest, from the lowland up to c. 2200 m (on Mt Kinabalu up to 2895 m?), often on
wet soil, scattered. F/. fr. Jan.—Dec.
Uses. The wood is whitish cream, available in big dimensions due to the large size of the trees, but
soft and suitable only for inside house constructions.
Note. The distinction of species in this genus is mainly based on the type of indument, clearly rec-
ognizable only with a lens. This can only well be examined on young leaves, where the full set of in-
dument characters is still present, as hairs and/or scales go rather early, and old leaves may become
practically glabrous. The scales give the undersurface of the leaves a metallic silvery lustre similar to that
found in leaves of certain Myristicaceae, Euphorbiaceae, Sapotaceae, and even Lauraceae, with which
they are sometimes confused in sterile sheets. Young leaves may show fine, round, translucent dots
against strong light (ENDERT, Med. Proefst. Boschw. Ned. Ind. 20, 1928, 193).
KEY TO THE SPECIES
1. Indumentum on the undersurface of young leaves giving a woolly appearance to the naked eye, and
under a lens representing two distinct strata, i.e. an upper layer of partly flattish, though mostly oblique-
ly erect star-shaped or fascicled hairs (with a very smallor generally invisible dark centre), and a lower
by ise of scales.
. Leaves shortly cordate at base, almost peltate, lateral nerves, veins and veinlets -- a impressed
~ above, the lamina ++ bullate, hairs persistent. 1. P. bullata
2. Leaves never properly cordate at base, nerves sometimes, veins and veinlets never impressed above,
hairs generally early disappearing except along midrib and nerves. 2. P. latifolia
1. Indumentum on the undersurface of young leaves giving a scaly appearance to the naked eye, seemingly
representing a single layer of scales, but under the lens appearing to consist of two layers of flat star-
shaped scales, i.e. a lower layer of + persistent, small and densely arranged scales, and an upper layer
of similar, though larger and more scattered, rather early caducous ones.
3. Leaves (elliptic or ovate-elliptic, rarely ovate) coriaceous, rigid .
3. Leaves (of various shape) coriaceous to subcoriaceous, + flexible. .
3. P. sclerophylla
4. P. excelsa
1. Platea bullata SLEUM. Blumea 17 (1969) 243.
Tree 13-14m, c. 15cm 9; bark smooth, grey-
brown. Branchlets rufous-brownish tomentose and
slightly many-grooved lengthwise as are petioles
and rachises in dry specimens. Leaves lax, sub-
obovate-elliptic or -oblong, apex subacutely acu-
minate for c. 1.5 cm, base rounded in general, the
very base short-cordate and the lamina almost
peltate, firmly chartaceous, olivaceous-brownish
Fig. 5. Distribution of Platea.
and dull above, more rufescent beneath in dry
specimens, + deeply bullate by nerves, veins
and partly also veinlets above, these well pro-
minent beneath, glabrous above except the short-
tomentose midrib and maybe nerves, brownish-
rufous-tomentose and soft to the touch by obliquely
erect stellate or fascicled longish hairs all over the
undersurface especially on midrib, nerves and veins,
moreover covered all over beneath with a coherent
greyish silvery layer of small star-shaped, flat,
appressed scales each with a minute dark centre
(similar to stellate flattish hairs), 14-32 by
(6-)7-13 cm, nerves 7-10(—12) pairs moderately
curved-ascending and looping, reticulation fine and
dense, minutely raised above, rather obscure
beneath; petiole 2-2.5cm by 2-3mm. ¢ In-
florescences (only known in juvenile state) axillary,
paniculate, with few dense-flowered, much spaced,
subspicate racemes or short-branched panicles,
1.5—3 cm; rachis 6-12 cm by 2-3 mm.
Distr. Malesia: Borneo (Sarawak), once
found.
Ecol. In mixed Dipterocarp forest of steep-
sided valley on basalt beside waterfall, c. 900 m.
1971]
ICACINACEAE (Sleumer) 11
2. Platea latifolia BL. Bijdr. (1826) 647; WaALP.
Rep. 1 (1842) 378; Hassk. Cat. Hort. Bog. (1844)
214: Miers, Ann. Mag. Nat. Hist. II, 10 (1852)
110, repr. Contr. Bot. 1 (1852) 97; Mia. Fl. Ind.
Bat. 1 (1856) 793, incl. var. sumatrana (BL.)
Miq.; Becc. Malesia 1 (1877) 116; VALET. Crit.
Overz. Olacin. (1886) 251, 253, t. 6, f. 40 & 41;
K. & V. Bijdr. 5 (1900) 156; Hocnr. PI. Bog. Exs.
(1904) 45; Bull. Inst. Bot. Btzg. 22 (1905) 50;
Back. Schoolfi. Java (1911) 227; Koorp. Exk. FI.
Java 2 (1912) 531; Koorp.-ScHum. Syst. Verz. 1,
Fam. 162 (1912) 1, p.p.; Koorp. Atlas 1 (1913)
t. 119; Ripi. Fl. Mal. Pen. 1 (1922) 426; Burk.
Dict. (1935) 1768; Merr. J. Arn. Arb. 19 (1938)
44: CHUN, Sunyatsenia 4 (1940) 229; GAGNEP.
Fl. Gén. I.-C. Suppl. (1948) 754; Amsu. in Back.
Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 4;
Heyne, Nutt. Pl. 1 (1950) 986; Back. & BAKH.
f. Fl. Java 2 (1965) 60; SLEuM. Blumea 17 (1969)
243; Fl. Thail. 2 (1970) 76. — P. sumatrana BL.
Mus. Bot. Lugd.-Bat. 1 (1850) 249; WaALp. Ann.
2 (1851) 182; Miers, Ann. Mag. Nat. Hist. II, 10
(1852) 110, repr. Contr. Bot. 1 (1852) 97. —
Sideroxylon gamblei C. B. CLARKE in Hook. f.
Fl. Br. Ind. 3 (1882) 538. — Platystigma myristi-
ceum R. Br. in Wall. Cat. (1832) n. 7523, nom.
nud.; ex HooK. f. Ic. Pl. 18 (1887) t. 1707; Fl. Br.
Ind. 5 (1887) 381. — P. excelsa (non BL.) KING, J.
As. Soc. Beng. 64, ii (1895) 111. — P. fuliginea
Evo. Leafi. Philip. Bot. 8 (1915) 2790; Merr. En.
Philip. 2 (1923) 489. — P. myristicea (R. BR. ex
Hook. f.) HALL. f. Med. Rijksherb. 36 (1918) 5;
SLeuM. Notizbl. Berl.-Dahl. 15 (1940) 360. —
Planchonella gamblei (CLARKE) H. J. LAM, Bull.
Jard. Bot. Btzg III, 7 (1925) 199. — P. ledermannii
SLeuM. Notizbl. Berl.-Dahl. 15 (1941) 360. —
P. hainanensis HOWARD, Lloydia 6 (1943) 149, t. 2;
Dani, J. Arn. Arb. 33 (1952) 261; CHUN &
CHANG, FI. Hainan. 2 (1965) 449, f. 512. — P. par-
viflora (non K. & V.) DAHL, J. Arn. Arb. 33 (1952)
261 (pollen). — ‘*Gomphocarpus’ gamblei (CLARKE)
VAN RoyeEN, Blumea 8 (1957) 425. — Gomphandra
gamblei (CLARKE) VAN ROYEN ex Ind. Kew.
Suppl. 13 (1966) 61.
Large tree, (10—-)20-45 m, bole columnar, an-
gular and with shallow to rather deep grooves, up
to 60 cm @g, buttresses, if any, short, rarely up to
1.5m; crown high inserted, irregular. Bark grey
to brown, rather smooth, with irregular shallow
fissures and numerous lenticels, 12-16 cm thick,
outer part in cross-section red, inner one yellow-
ish white, with a peculiar aromatic agreeable smell
(of cumarine?) and very bitter taste as they are also
found in the leaves. Tip of branchlets, inflorescen-
ces, petioles and young blades covered with a
rather coarse woolly rusty and detersile tomentum
of flattish to obliquely erect star-shaped or fas-
cicled hairs, and with minor scales, which go
early. Leaves elliptic, or ovate-, rarely oblong-
elliptic, apex generally shortly subacutely acumi-
nate, base broadly attenuate to rounded, thin-
coriaceous, at maturity glabrous and shining
above, and dull yellowish-greenish-greyish to
almost bluish (as in Lauraceae and Myristicaceae)
beneath, still covered there with sparse flattish to
obliquely erect star-shaped or fascicled hairs
especially along midrib and nerves, and besides
over the whole undersurface with a skin-like layer
of small, thin, + confluent, finally greyish-silvery
scales (these with a minute or scarcely recognizable
darker centre), which lasts for a while, finally al-
most glabrous even under a lens, (7—)10—20(-25)
by (3—)5—1l cm, midrib and 6—-10(—12) pairs of
rather straight robust nerves -+ sunken above,
distinctly raised beneath, reticulation of veinlets
dense, fine and generally more visibly raised on
both faces of fully mature leaves; petiole (1.2-)
2-4 cm. — ¢ Inflorescences + pyramidal in cir-
cumference, (2—)4—6 cm, consisting of spikes with
spaced glomeruli or short panicles. Petals
light green, at least finally glabrous, c. 1.5 mm. —
2 Inflorescences in 2—5(-8)-flowered cymes. Pedicel
stout, 2-5 mm, slightly elongated in fruit. Ovary
obconical, whether or not covered initially with
fine stellate appressed hairs, base included by the
dorsally pubescent sepals (c. 2mm). Drupe nar-
rowly ovoid-ellipsoid, glabrous, variable in size,
2.5-4.5 by 2-2.5cm; fleshy exocarp first lemon
yellow to reddish orange, finally purplish-blackish
especially when dry, of a bitter taste; stony endo-
carp acuminate, with 2-3 longitudinal grooves and
oblongish to subquadrangular markings between;
albumen white, slimy, becoming bluish when cut.
Distr. Sikkim, East Bengal (Sylhet), Indo-
China (Tonkin, Annam), China (Kwangsi, Hai-
nan), and Thailand; in Malesia: Sumatra, Malay
Peninsula, Banka, Java, Borneo, Philippines
(Biliran, Mindanao), Celebes, and New Guinea;
also in Melanesia (New Britain).
Ecol. Primary lowland and montane rain-
forest, also oak forest, scattered, not rare in Java,
mainly on constantly wet, even swampy places,
found in Borneo in heath forest up to 560 m, in
Java also on volcanic sand or loam, up to 1750 m.
Uses. Timber of large size, whitish, a little
lustrous, with a smell of cumarine, soft, not
durable, locally used for house building in po-
sitions under the roof. For descriptions of the
wood of P. latifolia cf. MoLL & JANSSONIUS, Mikr.
2 (1908) 248, f. 108; the description of the wood of
P. excelsa (l.c. 253) also refers here, as the cited
Koorpers coll. no. actually belongs to P. latifolia.
Galls. Docrers VAN LEEUWEN (Zoocecidia,
1926, 332, f. 599) mentions a gall on the stem and
the leaves caused by a gall-midge.
Vern. Sumatra: katji pako, Batak, pandan, M.
(Indragiri); Banka: kédang tjabé, medang tjabe,
ménjiur, pajit, M; Malay Peninsula: kadondong,
k. hérong (Pahang); Java: huru, h. kapas, ki
kadantja, mandalaksa, 8, wuru, J; Madura: buru;
Borneo: kaju bulu, M, pangak, Kayan; Celebes:
mawi, Ramp., songku, Tado, talang laki, Bugin.;
Philippines: caliso, Bag.; New Guinea: bengok,
Hattam, kabugd, kaburé, Kapauku, siera, Mani-
kiong.
3. Platea sclerophylla SLeuM. Blumea 17 (1969)
245.
Tree, 9-18 m, with spreading branches. Branch-
lets rather slender, tips lepidote. Leaves elliptic
12 FLORA MALESIANA
[ser. i vol. yt
or ovate-elliptic, rarely ovate, apex shortly at-
tenuate or obtuse, sometimes emarginate, base
broadly cuneate to rounded, coriaceous, rigid,
brown in dry specimens, glossy above sometimes,
dull beneath, early glabrous above, subdensely set
with a lower layer of small substellate, almost
contiguous scales, and a so to say upper layer of
scattered, major stellate scales, these early dis-
appearing, whilst the smaller scales go tardily, the
undersurface too finally practically glabrous, 5-9
by (2.5—-)3-6 cm, midrib and nerves slightly to
more deeply impressed above, prominent beneath,
nerves 6-8(—9) pairs, rather straight or more
curved from the midrib, not properly looping,
reticulation of veins and veinlets dense, slightly
or hardly impressed above, well visibly raised
beneath; petiole + distinctly transversely rugulose,
deeply grooved above, 5-10 by 2(-3) mm. —
3 Inflorescences from the upper axils, paniculate,
with spike-like lax-flowered 1—3 cm long branches,
densely scaly on axes, more laxly so on bracts and
calyx lobes; rachis angular, c. 1 mm @. Pedicels
almost none. Calyx lobes ovate-acuminate, co-
riaceous, ciliate, 1.5 mm. Petals deep red, glabrous,
2.5mm. Anther cells elliptic, 1.5mm. — 2 IJn-
florescences raceme-like, few-flowered, (1—)2-3.5
cm. Pedicels thick, 2-5 mm. Ovary thick-cylindric,
densely minutely stellate-lepidote.
Distr. Malesia: Borneo (Mt Kinabalu, twice
found on the eastern shoulder).
Ecol. Forest, 2440-2895 m.
4. Platea excelsa BL. Bijdr. (1826) 697; WALP.
Rep. 1 (1842) 378; Hassk. Cat. Hort. Bog. (1844)
214; Miers, Ann. Mag. Nat. Hist. IJ, 10 (1852)
110, repr. Contr. Bot. 1 (1852) 97; Mia. FI. Ind.
Bat. 1 (1856) 793; BEcc. Malesia 1 (1877) 116;
VALET. Crit. Overz. Olacin. (1886) 250, 253, t. 6,
f. 39; K. & V. Bijdr. 5 (1900) 153; Hocur. Pl.
Bog. Exs. (1904) 45; Bull. Inst. Bot. Btzg 22
(1905) 50; Bacx. Schoolfl. Java (1911) 227;
Koorp. Exk. Fl. Java 2 (1912) 530; Koorp.-
ScHuM. Syst. Verz. 1, Fam. 162 (1912) 1; Koorp.
Atlas 1 (1913) t. 118; BAker f. J. Bot. 62 (1924)
Suppl. 21; Amsu. in Back. Bekn. Fl. Java (em. ed.)
6 (1948) fam. 135, p. 3; Back. & BAKH. f. FI.
Java 2 (1965) 60; SLEUM. Blumea 17 (1969) 245. —
Fig. 6.
See for synonyms under the varieties.
KEY TO THE VARIETIES
1. Leaves oblong to elliptic-lanceolate, rarely sub-
ovate-elliptic or -oblong, generally widest in
the middle, and up to 4.5 (rarely -6) cm wide.
2. Leaves + acutely acuminate at apex, nerves
10-14 pairs, rather straight from the midrib.
Drupes 3.5-4.5 by 1-2 cm. 1. var. excelsa
2. Leaves obtusely acuminate or rounded at
apex, nerves 6-8(-10) pairs + curved-as-
cending from the midrib. Drupes 2.5-3 by
1-1.3 cm.
3. Leaves markedly finely reticulate on both
faces, generally up to 6 cm wide.
2. var. riedeliana
3. Leaves more faintly or hardly reticulate,
2-3.5(-4.5) cm wide. 3. var. microphylla
1. Leaves ovate or ovate-elliptic, or, if elliptic or
oblong-elliptic, generally wider than 6 cm.
4. Leaves shortly subacutely or obtusely acu-
minate at apex, faintly reticulate in general,
nerves 10—12(—14) pairs, these usually rather
straight and/or rather closely subparallel to
each other, (9—)12—17(—26) by (4.5—)6-9(-12)
Cie wah . . 4 var. borneensis
4. Leaves subcaudate- acuminate and subacute at
apex, + distinctly reticulate on both faces,
nerves 6-8 pairs, these curved-ascending and
rather distant from each other, 8—12 by 3.5—5.5
cmp “ea. ee . . 5. var. kinabaluensis
1. var. excelsa.
Tall tree, (12—)25-37 m, trunk + angular, deep-
ly channelled or fissured, 30—60 (rarely -100) cm 9,
with buttresses up to 1.5m. Bark dark grey to
brown, smooth, with fine longitudinal fissures and
small lenticels, thin, with a certain aromatic
smell and bitter taste. Tips of branchlets, young
leaves and inflorescences densely covered with ap-
pressed star-shaped major and minor scales (each
with a + distinct dark, almost sooty centre),
which disappear late. Leaves oblong- or elliptic-
lanceolate, apex subacutely acuminate, base +
broadly cuneate, subcoriaceous or thin-coria-
ceous, at maturity glabrous and shining above,
still rather densely scaly beneath especially on
nerves and petiole, and of a dull pale yellowish-
green colour in fresh state, flush deep pink, 10-17
by 3.5—4.5(—5.5) cm, midrib impressed above, bold
beneath, nerves 10-14 pairs, rather straight from
the midrib, + flat above, not much raised beneath,
reticulation fine, but slightly raised on both faces
or beneath only; petiole 1.5—2 cm by c. 2 mm. —
3 Inflorescences in simple spikes or by them com-
posed (then almost pyramidal), covered with a
rusty short tomentum of stellate hairs and scales,
flowers glomerulose, 3—5 cm. Petals pale greenish
yellowish or pale pink, glabrous, c. 2 mm. Anther
cells elliptic, 1.5mm. — 2 Inflorescences in 2-6-
flowered cymes (these sometimes reduced to a
single flower). Pedicels stoutish, 5-8 mm, elongate
in fruit. Ovary glabrous, base included by the
calyx lobes (c. 1.5 mm). Drupe very similar to that
of P. latifolia, (sub)ovoid-oblongoid, manifestly
narrowed upwards, 3.5—-4.5 by 1—2 cm; fleshy exo-
carp shiny, dark red when ripe, finally blackish,
taste aromatic-bitter; endocarp hard, with the same
markings found in P. J/atifolia.
Distr. Malesia: Sumatra (Tapanuli: Nias L.,
Westcoast, Bencoolen), Java (Bantam to Banju-
mas).
Ecol. Primary forest, often on fertile volcanic
soil, scattered, 240—1800(—2200, only c. 6m tall
there) m.
Uses. Timber whitish, with an aromatic smell,
apparently not in use.
Vern. Béntenu, talas éndriung, Lampong, rasak
bareh, Minangkabau, tjutjuho, Nias, M; gémpél,
meérji, J, durénan, dudurénan, mandalaksa, S.
1971]
2. var. riedeliana (BECC.) SLEUM. Blumea 17 (1969)
245. — P. riedeliana BEcc. Malesia 1 (1877) 116;
VALET. Crit. Overz. Olacin. (1886) 253. — P.
excelsa (non BL. s. str.) ANDERSON, Gard. Bull.
Sing. 20 (1963) 166.
Tree, up to 24 m, up to 45 cma. Leaves elliptic
to subovate-elliptic, apex obtuse or rounded, rarely
subemarginate, 7—15(-21) by 4.5-6 (in sterile
specimens or suckers rarely up to 8.5) cm, nerves
6—8(—10) pairs, curved-ascending and rather spaced
from each other, reticulation fine and dense,
generally markedly raised on both faces. Drupe
c. 3 by 1.3 cm. Flowers as in var. excelsa.
Distr. Malesia: Sumatra (Eastcoast), Malay
Peninsula (Johore), Banka, Billiton, Borneo (Sa-
rawak, Brunei, W. part of North Borneo).
Ecol. Generally in lowland, often (in Borneo
always) in peat swamp forest (then usually with
irregular spreading stilt-roots), once found on
Mt Kinabalu at 1525 m.
Vern. Médang tjabek, Banka, ménpisang, P. Ke-
bal, randjan antu, Bengkalis, sérangop, P. Mendol,
M.
3. var. microphylla (SLEUM.) SLEUM. Blumea 17
(1969) 246. — P. microphylla SLEUM. Notizbl.
Berl.-Dahl. 15 (1941) 360. — P. montana Howarp,
Lloydia 6 (1943) 150, t. 3.
ICACINACEAE (Sleumer) 13
Tree up to 30 m, up to 50 cm g. Leaves oblong
to elliptic-oblong, apex obtusely short-acuminate
or obtuse, base + broadly attenuate, thin-co-
riaceous, (5—)6—-10 by 2—3.5(-4.5) cm, nerves 6-8
pairs, rather straight to moderately curved-as-
cending, subparallel to and rather distant to each
other, often a little depressed above, reticulation
fine, rather faintly raised, sometimes obscure even
beneath. Drupe 2.5 by 1—-1.3 cm. Inflorescences as
in var. excelsa.
Distr. Malesia: New Guinea.
Ecol. Subcanopy tree in primary montane rain-
forest (Conifer-Nothofagus-mossy oak forest), on
slopes, 700-2200 m, scattered or sometimes rather
frequent locally.
Vern. Kaderei, sawera, terori, Ambai, uwar,
Kebar.
4. var. borneensis (HEINE) SLEUM. Blumea 17 (1969)
246. — P. latifolia BL. var. borneensis HEINE in
Fedde, Rep. 54 (1951) 238; Pfl. d. Samml. Cle-
mens Kinabalu (1953) 58. — P. parviflora K. & V.
Bull. Inst. Bot. Btzg 2 (1899) 3; Bijdr. 5 (1900)
155; Nat. Tijd. N. I. 60 (1901) 383; Back. School-
fl. Java (1911) 228; Koorp. Exk. Fl. Java 2
(1912) 530; Koorp.-ScHuM. Syst. Verz. 1, Fam.
162 (1912) 3; Koorp. Atlas 1 (1913) t. 120. —
P. latifolia (non BL. s. str.) MERR. Philip. J. Sc. 1
d =‘ 4J1'68
Fig. 6. Platea excelsa BL. var. borneensis (Heine) SLEUM. a. Habit with ¢ inflorescences, 14, b. 9 inflo-
rescences, X14, c. & c’. § flower buds, «10, d. 2 flower, «5, e. drupe, * 14 (a, c & c’ Brass 11564,
b & d BW 6477, e KosteRMANS 18753).
14 FLORA MALESIANA
[ser. I, voliw*
(1906) ae 58; HALL. f. Med. Rijksherb. 1
(1910) 13; SCHELLENB. Bot. Jahrb. 58 (1923) 159.
P. excelsa (non BL. s. str.) Koorp. Junghuhn
Gedenkb. (1910) 175. — P. philippinensis MERR.
Philip. J. Sc. 9 (1914) Bot. 313; En. Philip. 2
(1923) 489; Howarp, J. Arn. Arb. 21 (1940) 488,
t. 2, f. 26; DAHL, ibid. 33 (1952) 260, f. 23 & 23 A
(pollen). — P. apoensis ELM. Leafl. Philip. Bot. 8
(1915) 2789; Merr. En. Philip. 2 (1923) 489. —
Fig. 6.
Leaves similar in shape and size to those of
P. latifolia, elliptic or ovate-elliptic, rarely ovate,
apex shortly subacutely or obtusely acuminate,
thin-coriaceous, (9—)12—-17(—26) by (4.5—)6—9(-12)
cm, nerves 10—12(—14) pairs, generally rather
straight or only slightly curved-ascending. Flowers
as in var. excelsa. Drupe ovoid-oblongoid, much
attenuate towards the apex, (2.5—)3-3.5(-4) by
1.5(—2) cm.
Distr. Malesia: Sumatra, Java, Lesser Sunda
Is. (Bali, Lombok, Sumbawa, Flores), Borneo,
Philippines, Celebes, Moluccas (Ambon, Batjan,
Buru), New Guinea, and New Britain.
Ecol. Primary (or rarely secondary, also mon-
tane Quercus and mossy Nothofagus) forest or
lowland (Dipterocarp) forest, from sea-level up to
2000(—2400?) m, scattered, often in water con-
taining soil along rivers or on temporarily inun-
dated flats.
Uses. Wood whitish or yellowish; hardly in
use for house-construction anywhere. For ana-
tomy of the wood cf. MoLL & JANssontius, Mikr. 2
(1908) 254 (P. parviflora K. & V.) where is
said that the wood is very similar to that of
P. latifolia.
Vern. Sumatra: balunan, hoting, Batak, ku-
rungan tendi, sibaubakan, Karo-Batak, malenbu,
Nias, médang sékarut, Kubu, sitépu, Toba,
tjémpaka gading utan, M; Java: ki kadantja, ki
tjémpaka, S, njampoh budéng, rékisi, J; madas,
udu, Bali, tarokeli, Endeh; Borneo: balubug, La-
buk, /alak, Sampit, maparé, Balikpapan, njawan,
Bulungan; Celebes: bakalohi, Toradja, kapalore,
Tado, /ari-lari, Bela, marasulo, Torongkong, talang
birang, Bugin., wuamendu maeto, Tobela; Philip-
pines: kalisan, kaliso, Bag., pagpdgo, Bon.;
Moluccas: boba, Ambon, mahu, Buru; New
Guinea: bilou, Mooi, emmok, tegému, Muju,
kidap, Mendi, maar, nokimbuwo, Amberbaken,
mesimeski, Arfak, niebatek, Kebar, sira, Mani-
kiong, subar, Wandammen, tindokopa, Enga.
5. var. kinabaluensis (SLEUM.) SLEUM. Blumea 17
(1969) 247. — P. kinabaluensis SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 234. — P. latifolia (non BL.)
Dau., J. Arn. Arb. 33 (1952) 260 (pollen).
Leaves similar in texture and nervation, espe-
cially reticulation to those of var. riedeliana,
ovate-oblong or -elliptic, apex subcaudate-
acuminate and subacute, 8-12 by 3.5—5.5 cm,
nerves 6-8 pairs curved-ascending and rather
distant from each other. ¢ Flowers only known,
as in var. excelsa.
Distr. Malesia: Borneo (Mt Kinabalu), in
montane forest at 1430-1525m, twice found.
Excluded
Platea oblonga KorTH. ex VALET. Crit. Overz.
Olacin. (1886) 252, in text & t. VI, f. 42 a-d;
SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 234, in
text; Blumea 17 (1969) 248 = Prunus javanica
(T. & B.) Mia. (Rosaceae).
4. GONOCARYUM
Mia. Fl. Ind. Bat. Suppl. 1 (1861) 343; Steum. Blumea 17 (1969) 209. — Phlebo-
calymna GRiFF. ex Miers, Ann. Mag. Nat. Hist. II, 10 (1852) 109, repr. in Contr.
Bot. 1 (1852) 96, in syn. gen. Platea BL., nom. inval.; ex BENTH. in Benth. & Hook.
f. Gen. Pl. 1 (1862) 353, descr. — Fig. 8,
Dioecious, slender trees or shrubs, + horizontally branching. Tete spirally
arranged, entire, penninerved; petiole yellow and always characteristically trans-
versely wrinkled. /nflorescences axillary, often from defoliate parts of the branch-
lets, rarely from trunk. Flowers sessile or very shortly pedicelled, subtended by 1
or 2 bracts, collected to interrupted, solitary or fascicled, sometimes elongate
spikes, or congested to axillary fascicles or glomerules. Calyx small, deeply im-
bricately 5-lobed, persistent. Petals 5, valvate in bud, either united up to the lower
half and cup-like spreading, or connate more highly and forming a tube, tips
inflexed, slightly keeled and sometimes papillose inside. — ¢ Flowers: Stamens 5;
filaments adnate to the petals except at tips; anther cells oblong-elliptic, introrse,
dorsifixed. Rudiment of ovary hairy, on a thin glabrous disk. — 9 Flowers:
Stamens 5, rudimentary. Ovary sessile; style short-conical; stigma rather large or
small, pad-like or capitate, oblique, unilaterally incised or subbilobed. Disk
thin-coriaceous, crenulate, sometimes inconspicuous. Drupe subglobose to ellip-
1971] : ICACINACEAE (Sleumer) As
soid-oblongoid, sometimes obtusely 3 (or sub-5)- ie Sai few to numerous
longitudinal ribs; exocarp either thick and predominantly spongious, showing but
a few fibres, or thinner and with more numerous fibres; endocarp either hard and
with numerous longitudinal ++ sharp crests, and + alveolate besides, or thinner
and but shallowly crested or ridged, whether or not with an inner lateral lon-
gitudinal swelling mainly in the lower part, which houses a small roundish abortive
loculus. Seed 1, with a thin testa; endosperm copious, much corrugate-lobular or
ruminate; embryo thin, with 2 foliaceous lanceolate cytoledons.
Distr. About 9 or 10 spp., all in Malesia, eastwards to the Bismarcks, 2 spp. extending into SE.
Asia (Burma to Kwangsi, Hainan and S. Formosa), a 10th unlocalized and cultivated in Hort. Bog.,
imperfectly known, and possibly 2 undescribed new spp., one in New Britain, the other in Burma and
Malaya. Fig. 7.
Ecol. In the understorey of lowland and montane, primary and occasionally secondary forest, rarely
in swamp forest or mangrove, from sea-level up to 1220 m. Fi. fr. Jan.—Dec.
The drupes seem to be able to float, which may contribute to their dispersal.
Note. The spp. in the group with tubular flowers are very close to each other; with the materials at
hand their discrimination remains rather unsatisfactory. It seems, that mature fruits allow to separate
these species by such characters as form and size of the drupe and the structure of the exocarp and
endocarp. Many fruits, however, are found without seeds, and thus possibly not or not always enough
developed to show the above mentioned characters.
KEY TO THE SPECIES
1. Petals of the 3 flowers almost free or united below up to the middle, cup-like expanded. Anthers as
long as or longer than filaments. Drupe, as far as known, without a distinct abortive loculus*, reddish
to orange at maturity; exocarp thick, almost exclusively spongious.
2. Petals of the 3 flowers almost free, smooth inside, c.2 mm. . . . 1. G. crassifolium
2. Petals of the 3 flowers united below to !4—!/4 their length, with papillose emergences all over the
inside, (4-)5S-6mm ... choke cee Gn tOrale
. Petals of the 3 flowers united below to ‘about yy, forming a 1 tube, not ‘papillose inside. Filaments
distinctly longer than the anther cells. Drupe dark green to olivaceous or turning blackish at maturity,
with a distinct abortive loculus (round in @); exocarp rather thin, spongious or predominantly fibrous.
3. Leaves suddenly narrowed at apex for c. 2 cm by 3-4 mm (at base), tip subacute, nerves, and gen-
erally also veins, much impressed above, sharply raised beneath. Mature drupe acuminate, tip
acute, longitudinal ribs numerous and + sharply outstanding. . . . .. . 3. G. impressinervium
3. Leaves less distinctly or shortly acuminate-attenuate at apex, tip -+ bluntish, nerves whether or not
impressed above, veins but slightly or not so. Mature drupe short-apiculate or obtuse at apex,
longitudinal ribs few to rather numerous and + obtuse, whether or not prominent.
4. 3 spikes solitary in general. Mature drupe obtusely trigonous in 9.
5. Spikes (both 3 and @) generally elongate, up to 30 cm. Leaves (6—)8—18(—20) by 3.5—-6.5(—9.5) cm,
greyish green in dry specimens. Petals 56mm. . . . . 4 G. gracile
5. Spikes (both g and ®) relatively short, rarely up to 6c cm. .. Leaves 6-12 by 3-4(-5) cm, usually
yellowish green especially beneath in dry specimens. Petals c.4mm. ... . . 5. G. minus
4. 3 Spikes generally several in a fascicle, or flowers glomerately arranged. Mature drupe + round in @.
6. Drupe ovoid, or rarely ellipsoid-subovoid, green at we: with several obtuse prominent
longitudinal ribs .. . siay 6. G. cognatum
6. Drupe oblongoid to ellipsoid, or ‘rarely subovoid- ellipsoid or -oblongoid, dark olivaceous to +
blackish at maturity, longitudinal ribs few to numerous, all but little or hardly prominent.
7. Ovary in the 9, or rudiment of ovary in the 3 flower glabrous. Leaves 15—20(—25) by (5—)6-10(—14)
cm, lateral nerves generally slightly impressed above, reticulation rather weakly or not prominent
above. Drupe (1.5—)2-2.5 (rarely —-3) by 1—1.5 cm; exocarp 1-2 mm; endocarp thin.
7. G. macrophyllum
7. Ovary in the 2, or rudiment of ovary in the 3 flower densely to more sparsely hairy. Lateral nerves
of leaves not impressed above. Drupe larger in general.
8. Leaves markedly finely and densely prominent-reticulate on both ries or certainly above.
Exocarp plus endocarp of well developed drupes 1-2mm. .. . . . 8 G. calleryanum
8. Leaves rather obscurely or not prominent-reticulate above, never so beneath. Exocarp plus
endocarp of well developed drupes (2-)3-4mm............ + 9 G.lobbianum
(*) Cf. also 10. G. melanocarpum.
16 FLORA MALESIANA
[ser. I, vol. 72
1. Gonocaryum crassifolium RiDL. J. Str. Br. R. As.
Soc. n. 75 (1917) 18; Fl. Mal. Pen. 1 (1922) 432;
SLeuM. Blumea 17 (1969) 210.
Shrub with flexuous branches, glabrous except
the inflorescence. Leaves reminding of those of
G. litorale, ovate to oblong, apex + abruptly
acuminate for c. 2 cm, tip bluntish, base broadly
cuneate to rounded, coriaceous, 15-17 by 5-9.5
cm, midrib strongly deepened above, sharply
prominent beneath, nerves (5—)6—7 pairs, slightly
impressed above, raised beneath, proper retic-
ulation obscure, both faces of the lamina finely
tubercled; petiole c. 1.5 cm by 2-3 mm. —
3 Spikes solitary or several from tubercles in
defoliated axils of branchlets, lax-flowered, slender,
2.5-5 cm, all over shortly appressedly hairy, not
yet known in fully developed state. Calyx lobes
ovate, obtuse, c. 1.5 mm. Petals ovate-oblong,
subacuminate, thickish, free to almost the base,
glabrous and smooth inside, c. 2 mm. Rudiment
of ovary hairy. 2 Inflorescences and drupe not yet
known.
Distr. Malesia: Malay Peninsula (Selangor:
Sempang Mines), once found.
Fig. 7. Distribution of Gonocaryum. The figure
above the hyphen indicates the number of en-
demic species in that island or area, the figure
below the hyphen the number of non-endemic
species.
2. Gonocaryum litorale (BL.) SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 233; Blumea 17 (1969) 211.
— Stemonurus ? litoralis BL. Mus. Bot. Lugd.-Bat.
1 (1850) 250; Warr. Ann. 2 (1851) 182; Mrers,
Ann. Mag. Nat. Hist. II, 10 (1852) 36, repr.
Contr. Bot. 1 (1852) 87. — Lasianthera ? litoralis
(BL.) Miq. Fl. Ind. Bat. 1 (1856) 792; ScHEFF.
Ann. Jard. Bot. Btzg 1 (1876) 14; Becc. Malesia 1
(1877) 111. — Villaresia macrocarpa SCHEFF. Ann.
Jard. Bot. Btzg 1.(1876) 13; Lc. 102, obs. —
G. pyriforme ScuHeFF. Ann. Jard. Bot. Btzg. 1
(1876) 100; Becc. Malesia 1 (1877) 123; VALET.
Crit. Overz. Olacin. (1886) 246; Hocur. PI.
Bog. Exs. (1904) 43, incl. var. corrugatum HOCHR.
lc. et var. planifolium Hocur. l.c. 44; Bull. Inst.
Bot. Btzg 22 (1905) 39, incl. var. corrugatum
Hocnur. et var. planifolium Hocur.; Ann. Jard.
Bot. Btzg Suppl. III, 2 (1910) 857, incl. var.
genuinum f. planifolium (Hocur.) Hocnr. et f.
corrugatum (HocuHr.) Hocur., var. obovatum
(Hocnure.) Hocure., var. fuscum (HOCHR.) HOcCHR.
Lc. et var. fusiforme (HOCHR.) Hocnre. l.c. 858;
Back. Schoolfl. Java (1911) 227; Koorp. Exk.
Fl. Java 2 (1912) 532 (‘piriforme’); PULLE, Nova
Guinea 8 (1912) 659; Wars. Pflanzenwelt 2
(1916) 349, f. 231 D, 1-6; SCHELLENB. Bot. Jahrb.
58 (1923) 166; WETTSTEIN, Handb. Syst. Bot. ed.
4 (1935) 842, fig. (fr., ‘pyrospermum’); AMSH. in
Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135,
p. 2; Back. & BAKH. f. FI. Java 2 (1965) 59. —
G. selebicum Becc. Malesia 1 (1877) 124; VALET.
Crit. Overz. Olacin. (1886) 247; Koorp. Minah.
(1898) 394. — G. affine Becc. Malesia 1 (1878)
256; VALET. Crit. Overz. Olacin. (1886) 247;
PULLE, Nova Guinea 8 (1912) 659; SCHELLENB.
Bot. Jahrb. 58 (1923) 166. — G. macrocarpum
(SCHEFF.) SCHEFF. ex WARB. Bot. Jahrb. 13 (1891)
299: K. Sco. & Laut. FI. Schutzgeb. (1900) 415.—
G. fuscum Hocnre. Pl. Bog. Exs. (1904) 42; Bull.
Inst. Bot. Btzg 22 (1905) 5, 49; Back. Schoolfl.
Java (1911) 227; DAHL, J. Arn. Arb. 33 (1952) 265
(pollen). — G. fusiforme Hocnre. Pl. Bog. Exs.
(1904) 42; Bull. Inst. Bot. Btzg 22 (1905) 50.
— G. obovatum Hocnr. Pl. Bog. Exs. (1904) 43;
Bull. Inst. Bot. Btzg 22 (1905) 4, 51; VALET. Bull.
Dép. Agr. Ind. Néerl. 10 (1907) 30. — G. sp.,
PuLLE, Nova Guinea 8 (1912) 659.
Slender tree, rarely erect shrub, (2—)6—20(—30) m,
rather sparsely and + horizontally branched;
trunk with several small grooves, 3—15(—30) cm @;
bark rather smooth, mottled, light to dark greyish-
brown. Branchlets slender, practically glabrous as
are the leaves. Leaves ovate or ovate-elliptic,
sometimes elliptic and even oblong, apex shortly
acuminate, top blunt, base cuneate to rounded,
slightly inequilateral, subcoriaceous to coriaceous,
dark green to olivaceous and shining above, paler
and + dull beneath, (10—)12-20(—28) by 6-12(-19)
cm, midrib a little depressed above, much pro-
minent beneath, nerves (4~-)5—7(—8) rather steeply
curved-ascendent pairs, only the upper ones
somewhat looping, raised within a depression in
dry specimens above, prominent beneath, reticu-
lation rather dense, often obscure in dry speci-
mens, or slightly depressed above, or a little
raised on both faces; petiole 4-8(—10) by 1.5—2(—3)
mm. — 3 Spikes 1-4 (rarely —-8) from a foliate or
mostly already defoliate axil, or from tubercles
on branchlets and branches, slender, (3—)5—15(—30)
cm, flowers numerous, laxly arranged, sparsely
appressedly short-hairy or subglabrous. Calyx lobes
ovate, ciliate, greenish-brown, 1.5(—-2) mm. Petals
narrowly ovate-acuminate, united in the lower third
or half, cup-like expanding at full anthesis,
slightly keeled and all over set with papillose
emergences inside, white to cream or greenish,
sometimes pinkish, fragrant, (4-)5—6 by 2-2.5 mm.
Filaments somewhat dilated towards the base,
fleshy, 1.5-2mm; anther cells elongate-oblong,
2-2.5 mm. Rudiment of ovary hairy. — 2 Spikes
generally shorter than the g ones. Calyx less deeply
lobed, (2-)3 mm. Petals 6-7 by 2-2.5 mm. Anther
cells 1 mm, devoid of pollen. Ovary ovoid, densely
fulvous-hairy; stigma rather large, pad-like. Drupe
ellipsoid or obovoid, or obovoid-oblongoid, some-
1971]
ICACINACEAE (Sleumer) 17
times almost globular, often somewhat angular or
laterally compressed, finally quite glabrous and
shining, orange to reddish, (3—)4~-5(-6.5) by
2-3(-4.5) by 2-3.5 cm; exocarp spongious, with
adstringent sap, finally dissolute, c. 3 mm; endo-
carp hard, 1(—2) mm thick, with few to numerous
irregular longitudinal sharp crests 1-5 mm high,
+ marked in the immature fruit by the drying
process, abortive loculus practically absent. Seed
pear-shaped or ellipsoid.
Distr. Malesia: Lesser Sunda Is. (Timor), Ce-
lebes incl. Buton I., Moluccas (Morotai, Halma-
hera, Ambon, Ceram, Kai and Tanimbar Is.),
Aru, Salawati, Miso6l, New Guinea & New Bri-
tain, and Mussau I. in the Bismarcks.
Ecol. In second storey of primary rain-forest,
mostly on hill slopes, also in -+- open secondary
growth or sunny slopes, in coastal and swamp
forest, often near creeks or temporarily inundated
river flats, on clayey soil, scattered, though locally
common, from sea-level up to c. 1200, in New
Guinea up to 1650 m.
Uses. Wood hard, heavy, dark cream or
yellowish or light orange, with broad rays and an
oak-like grain, apparently not used due to the
small dimensions.
Vern. Tanimbar Is.: twajagunmore, Otimmer;
Celebes: balula, Tobela, karo, Buton, kondo,
Topadu; Aruls.: kartomadin; New Guinea: akwaai,
Kawerawedja, aria, rasui, Ambai (Japen), atile,
ulumu, Amele, bisip, sarenki, Dumpu, briengbas,
Karoon, dindo, gadun, gwanawa, Faita, gaun,
mobo, Bilia, kaffadioma, Menawi (Japen), kami,
Maprik, kKawénmatak, Maibrat, kojari, sakowgo
matémuk, wobrega, wobreika, Manikiong, kur-
mardi, Japen, kwydjarak, Kebar, mesobas, Hatam,
owok, New Britain, sabobar, Etnabay, sanganiet,
Tehid, sapiai, Biak, sienbal, Mooi, ubakka, Mata-
paili.
3. Gonocaryum impressinervium SLEUM. Blumea
17 (1969) 212. — Fig. 8.
Shrub or treelet, (2—-)3-8 m, trunk c. 5cm g;
bark smooth. Branchlets obtusangular, slender,
practically glabrous. Leaves oblong-elliptic, apex
subabruptly acuminate for c. 2cm and 34mm
wide at base of acumen, base broadly attenuate,
subinequilateral, thin-coriaceous, glabrous with
age, dilutely green-olivaceous in dry specimens,
(12-)15-23 by 5-10 cm, midrib, nerves and veins
distinctly impressed above in dry specimens, well
raised beneath, nerves 6-8 pairs curved-anasto-
mosing and markedly looping, reticulation of
veinlets dense and prominent on both faces, or
sometimes almost obscure; petiole 8-10 by c.2 mm.
Spikes axillary, solitary or more rarely in twos,
erecto-patent, (2—)3-8 cm, laxly many-flowered,
lower flowers pedicelled up to 2 mm; rachis slen-
der, | mm @, puberulous. Calyx 5-lobed to almost
the base, lobes ciliate, c. 2mm. — ¢ Flowers:
Petals connate to a tube for their lower %4, 5-6
mm, white in fresh, reddish in dry state. Filaments
3mm; anther cells 1.5mm. Rudiment of ovary
hirsute. — 92 Flowers only known in developed
state. Calyx lobes 2.5mm. Petals not known.
Ovary short-pubescent. Drupe (possibly not quite
mature) obliquely elongate-ovoid, subtrigonous,
slightly compressed laterally, more attenuate to-
wards the apex than the base, acute, sparsely hairy,
green with 12-15 subirregular longitudinal sub-
acute and 1—2 mm high ribs, 5—5.5 by (2—)2.5 cm;
exocarp spongious, |—-1.5 mm, with longitudinal
fibres; endocarp hard, thin, longitudinally many-
ribbed.
Distr. Malesia: Borneo.
Ecol. Forest (Dryobalanops), scattered though
obviously locally common, on sandy loam, 50-150
m.
4. Gonocaryum gracile Mia. Fl. Ind. Bat. Suppl. 1
(1860) 137, nom. nud.; ibid. (1861) 343; Becc.
Malesia 1 (1877) 122; VALET. Crit. Overz. Olacin.
(1886) 245, excl. syn.; SLEUM. Blumea 17 (1969)
213. — G. longeracemosum KiNG, J. As. Soc.
Beng. 64, ii (1895) 120; RipL. Fl. Mal. Pen. 1
(1922) 431; BAKER f. J. Bot. 62 (1924) Suppl. 21;
Burk. Dict. (1935) 1099; DAnL, J. Arn. Arb. 33
(1952) 265, f. 44 & 44 A (pollen).
Shrub or small, low-branching tree, up to 12 m;
trunk up to 5cm g; bark smooth, light greyish
brown. Branchlets slender, tips finely pubescent,
lower parts glabrescent, striate, early covered with
greyish cork. Leaves elliptic-oblong, sometimes
subobovate-oblong, apex rather suddenly at-
tenuate, mostly somewhat curved, tip blunt, base
broadly cuneate, slightly inequilateral, subco-
riaceous to coriaceous, practically glabrous with
age, greyish green in dry specimens, shining above,
dull and paler, sometimes tubercled beneath,
(6—)8-18(—20) by 3.5—6.5(—9.5) cm, midrib slightly
sunken above, much prominent beneath, nerves
4-5(-6) pairs, lower 2 or 3 pairs curved-ascending,
upper ones more straight, all slightly or hardly
raised above, more distinctly so beneath, reticu-
lation of veins and veinlets dense, finely raised on
both faces; petiole 8-10 by 1.5—2 mm. Jnflorescen-
ces both from foliate and lower defoliate axils,
erect, (3—)5—15(—30) cm, solitary or rarely in twos
or threes, laxly many-flowered; rachis 1—-1.5 mma,
short-hairy as are the bracts and lower part of the
calyx lobes, the latter ovate-deltoid, 1.5 mm. —
3 Flowers: Petals 5-6 mm, high connate to a tube
of 2mm 9, greenish-white, lobes short, finally
reflexed. Filaments as long as the tube; anther
cells narrow-elliptic, 1mm, finally a little ex-
serted. Rudiment of ovary hirsute, on a _ thick
glabrous disk. — 2 Flowers: Tube of petals more
urceolate, 3—-4(—5) mm, lobes 1—1.5 mm, reflexed.
Anther cells much reduced in size, empty. Ovary
ovoid-conical, densely grey-pubescent, attenuate
to a short -+ unilateral style with a flattish or
subcapitate bilobed stigma. Disk pubescent at
upper margin. Drupe ellipsoid-oblongoid, some-
times subobovoid, green, obtusely trigonous, apex
broadly attenuate or obtuse, rarely apiculate, each
of the 3 faces with 2 or 3 longitudinal low obtuse
ribs, (3—)3.5—4(—4.5) by 1.5-2(-2.8) cm; exocarp
spongious; endocarp hard, thin, slightly longi-
tudinally many-ribbed.
18 FLORA MALESIANA [ser. I, vol. 7
Distr. Malesia: Sumatra, Malay Peninsula (also Vern. Rambai ayam, r. hantu, ruai gajah,
P. Tioman), Banka. tampong bési, toyoh, M.
Ecol. In forest, mainly in lowland, rarely up to
1000 m, in river valleys by streamside, scattered. 5. Gonocaryum minus SLEUM. Blumea 17 (1969)
Uses. Poultices are said to be made from the 213.
fruit for application to the head for head-ache. Treelet up to 12 m, rarely tree up to 30 m, trunk
FEB
EW
at
oo
LES .
EE FRE
~ DORs 3
ere yt
Fig. 8. Gonocaryum impressinervium SLEUM. 4. Habit, x14, b. seed, showing the ruminate endosperm,
x1, c. drupe, x1, d. d flower, from the inside, expanded, x714 (a & d KOSTERMANS 10435 (type),
b-c KOSTERMANS 10480).
1971]
ICACINACEAE (Sleumer) 19
up to 18 cm 9; bark smooth or minutely fissured,
greenish-grey to dark brown. Branchlets obtus-
angular, very slender, glabrescent. Leaves elliptic-
oblong, subcaudate-acuminate for 5-15 mm, tip
blunt, base broadly cuneate or obtuse, subcori-
aceous, glabrous, paler and often yellowish-green
beneath in dry specimens, 6-12 by 3-4(-5) cm,
midrib slightly impressed towards the base of the
lamina above, prominent beneath, nerves 4—-5
irregular pairs, suberect, a little raised on both
faces, reticulation of veins and veinlets dense and
finely raised above, generally less so beneath;
petiole c. lcm by 1-1.5mm. Spikes axillary,
solitary, lax-flowered; rachis slender, sparsely ap-
pressedly hairy, (1—)2-5(-6)cm. Calyx lobes
ovate, ciliate, 1.5 mm. Peta/s greenish, connate to
a tube 4 by 1—1.5 mm, free and erecto-patent for
the distal 1 mm. — ¢ Flowers: Filaments 3 mm;
anther cells 1 mm. Rudiment of ovary hairy. Disk
thickish, hairy. — 2 Flowers: Petals connate to
an urceolate-cylindric tube for their lower 4,
c. 4mm. Ovary ovoid, densely short-hairy; stigma
capitate. Drupe obovoid-oblongoid, obliquely ob-
tusely subtrigonous, apex obtuse or apiculate,
green to brownish, 2.5-3.5(-4) by 1.8-2.5 cm;
exocarp spongious and fibrose, 2(—3) mm; endo-
carp 0.5mm, 8-10-costulate lengthwise, the ribs
not much showing off in the exocarp.
Distr. Malesia: Borneo (northern part).
Ecol. Primary, lowland (Dipterocarp) forest,
also in old secondary forest, up to 245m, on
sandy clay, on tertiary sand or sandstone, on
leached soil.
Uses. Wood yellowish, soft, not used. Fruit
said to be much favoured by barking deer (kijang).
Vern. Kamuning, mélanin, Dusun.
6. Gonocaryum cognatum Evo. Leafl. Philip. Bot.
1 (1908) 315; Merr. En. Philip. 2 (1923) 491;
Pl. Elm. Born. (1929) 172; Exm. Leafl. Philip.
Bot. 10 (1939) 3808; Dani, J. Arn. Arb. 33
(1952) 265, f. 45 & 45 A (pollen); SLEUM. Blumea
17 (1969) 214. — G. grandifolium MeErr. Philip. J.
Sc. 17 (1920) 278; En. Philip. 2 (1923) 491.
Shrub or treelet, (1-)2-6m, practically gla-
brous; stem 4-8 cm o. Branchlets slender, smooth,
green, older parts covered with a thin greyish-
brownish longitudinally fissured cork. Leaves
ovate- or oblong-elliptic, sometimes ovate, apex
subabruptly and for 1-2 cm obtusely subacumi-
nate or apiculate, base + rounded, slightly inequi-
lateral, coriaceous, lucid green above, much paler
beneath, 15—20(-30) by 8-12(-18) cm, midrib
grooved only in the lower part above, strongly
raised beneath, nerves 4~5(—6) pairs, the lower 2
(or 3) pairs usually closer together than the upper
ones and found in about the lower third of the
lamina, curved-ascending and + looping, slightly
raised above, more distinctly so beneath, retic-
ulation of veins and veinlets dense and finely
prominent on both faces; petiole 8—15(-20) by
c. Imm. — 3 Flowers (B.S. 44073) glomerate on
slightly supra-axillary swellings, tubercles or very
short axes, sessile or almost so. Calyx lobes ovate,
ciliate, c.2 mm. Petals united to their lower % into
a tube, white, c. 5mm. Filaments 3 mm; anther
cells elliptic, 1.5 mm. Rudiment of ovary cylindric
with a broadened base, glabrous as is the thick
crenulate disk. — 92 Flowers solitary or few
(rarely up to 8), details not known. Drupe ovoid,
or rarely ellipsoid-subovoid, (2—)2.5—3 by (1.5-)
22.5 cm, round in 9, green, with c. 5 prominent
obtuse ridges from base to the subtruncate or very
shortly apiculate apex, and may be with a few
less distinct ones between; exocarp thin, pre-
dominantly spongious; endocarp hard, 1—-1.5 mm,
with c. 5 obtuse longitudinal crests.
Distr. Malesia: E. and NE. Borneo, Philippines
(Luzon: Sorsogon; Samar, Leyte, Panay, Dinagat
I., Mindanao, Sulu Arch.).
Ecol. Primary, rarely secondary lowland (Dip-
terocarp) forest, up to 300 m, along river, appar-
ently scattered.
7. Gonocaryum macrophyllum (BL.) SLEUM. Blumea
17 (1969) 214. — Stemonurus macrophyllus Bu.
Mus. Bot. Lugd.-Bat. i (1850) 250; WALP. Ann. 2
(1851) 182. — Lasianthera macrophylla (BL.) Mia.
Fl. Ind. Bat. 1 (1856) 792; Suppl. 1 (1860) 137;
Becc. Malesia 1 (1877) 111. — G. melanocarpum
(non Hocur.) WyYATT-SMITH, J. Ecol. 41 (1953)
213, 214.
Treelet or shrub, 2-6(-10)m; bark brown.
Branchlets slender, glabrous. Leaves elliptic to
subobovate-elliptic, apex rather abruptly short-
acuminate (1 cm), tip obtuse, base broadly cuneate,
slightly inequilateral, coriaceous, ++ dull on both
faces in dry specimens, glabrous, 15—20(—25) by
(5—)6—10(—14) cm, midrib slightly impressed above,
strongly raised beneath, nerves 5-6(-7) pairs
curved-ascendent, the upper ones more distinctly
looped, shallowly or not sunken above, pro-
minent beneath, reticulation of veins and veinlets
finely or hardly prominent above, hardly so
beneath, or obscure on both faces; petiole 10-12
by (2—)3 mm. Jnflorescences in foliate or defoliate
axils; ¢ racemes several, sometimes very short and
glomerate (galled then?), 2 ones solitary or few;
rachis 0.5—1.5 (rarely —3.5) cm, slender at anthesis,
stoutish and somewhat accrescent in fruit, covered
with short hairs or almost glabrous; pedicels 1 mm
or less. — 3d Flowers (mainly from L6ORZING 5452,
Sumatra): Calyx lobes ovate, fleshy, ciliate, 1.5 mm.
Petals united in their lower 34 part to a distally
slightly enlarged tube, c. 5mm including the
erect free part, fleshy, white or greenish. Filaments
3 mm; anther cells elongate-elliptic, 1.5 mm. Ru-
diment of ovary cylindrical, base broadened,
practically glabrous, as is the flat disk. — &
Flowers: Calyx \obes and petals as in 3 flowers.
Anther cells reduced in size. Ovary ovoid, glabrous,
style thick, 1 mm; stigma oblique, flattish-peltate.
Disk thin or obscure. Drupe subovoid-ellipsoid or
oblongoid, apex obtuse or apiculate, slightly
longitudinally many-ridged or -keeled (generally
more distinctly so on one side), blue-black,
(1.5—)2-2.5(rarely —3) by 1—1.5 cm; exocarp spon-
gious and fibrous, 1-2 mm; endocarp hard,
thin.
Distr. Malesia: Sumatra (Eastcoast), Mentawei
20 FLORA MALESIANA
Is. (Siberut), Berhala and Jarak islets in Malacca
Straits, Banka (also on Lepar I.), Borneo.
Ecol. Coastal and hillside inland lowland
(also Dipterocarp) forest, sometimes on rather
dry land, sandy loam soil, up to 400 m.
Vern. Kaju napa, Asahan, minyak berok,
Jarak I., M.
8. Gonocaryum calleryanum (BAILL.) BEcc. Ma-
lesia 1 (1877) 123; VALET. Crit. Overz. Olacin.
(1886) 247; Merr. Philip. J. Sc. 3 (1908) Bot.
243, 418; Brown, Min. Prod. Philip. For. 3 (1921)
203; Merr. En. Philip. 2 (1923) 491; Brown,
Useful Pl. Philip. (1950) 356; KRAEMER, Tr. W.
Pacif. Region (1951) 208, f. 74 (wood);
Dau, J. Arn. Arb. 33 (1952) 265, f. 46 & 446A
(pollen); Lr, Pac. Sci. 7 (1953) 184; Liu, SASAKI
& KENG, Quart. J. Taiwan Mus. 8 (1955) 305;
Kao, /.c. 345; Hatrus. Mem. Fac. Agr. Kagosh.
Un. 5, 3 (1966) 38; SLEUM. Blumea 17 (1969) 215.
— Phlebocalymna calleryana BAILL. Adansonia 9
(1869) 147. —G. teysmannianum ScuerFF. Ann. Jard.
Bot. Btzg 1 (1876) 98; Becc. Malesia 1 (1877) 123;
VALET. Crit. Overz. Olacin. (1886) 245; Hocur.
Pl. Bog. Exs. (1904) 44, n. 91; Bull. Inst. Bot. Btzg 22
(1905) 51. — G. tarlacense VIDAL, Sinopsis Atlas
(1883) 20, t. 30, f. C; Rev. Pl. Vasc. Philip. (1886) 86;
CERON, Cat. Pl. Herb. Manila (1892) 46; Merr.
Philip. J. Sc. 1 (1906) Suppl. 86. — G. diospyro-
sifolium HAYATA, Ic. 2 (1912) 106 (‘diospyrifolia’);
KANEH. Formos. Tr. (1917) 119; ibid. (1936) 400,
f. 359; Iro, Ill. Formos. Pl. (1927) t. 12; SAsaAKt1,
Cat. Gov. Herb. Formosa (1930) 324.
Treelet or slender tree, 5—12-(20) m, trunk rarely
up to 25cmg, with greyish bark. Branchlets
slender, glabrous. Leaves elliptic to elliptic-ovate,
apex obtusely apiculate, often plicate, base broadly
cuneate to rounded, coriaceous, glabrous, shining
above, paler and rather dull beneath in dry spe-
cimens, (7—)10-18 by (4-)5-8(-10) cm, midrib
very slightly grooved above, strongly raised
beneath, nerves 4—-5(—6) rather steeply curved-
ascending pairs, the lower 2(—3) ones closer together
than the upper ones and often + bending down-
wards to the petiole in a narrow angle, a little
prominent above, more distinctly so beneath,
reticulation of veins and veinlets rather dense and
markedly raised on both faces especially above;
petiole 12-20 by c. 2 mm. Racemes spiciform,
solitary or several in a fascicle, from foliate
and/or defoliate axils, slender, few- to many-
flowered, laxly appressedly hairy, (1—)2(-4) cm.
Pedicels 0.5—-1.5(—2) mm. Calyx lobes ovate, dorsal-
ly + glabrous, ciliate, 1.5-2(-2.5) mm. Petals
(4-)5 mm, whitish, or light green, united to a
tube for 34 their length, lobes suberect. — ¢
Flowers: Filaments 3mm; anther cells elliptic,
1.2-1.5 mm. Rudiment of ovary cylindric, sparsely
hairy, on a thick broadened glabrous disk. —
© Flowers: Anther cells rudimentary. Ovary ovoid-
subconical, densely short-hairy, on a thick gla-
brous cushion-like disk; style hardly 1mm;
stigma peltate, oblique, attenuate towards one
side. Drupe subovoid-oblongoid, or generally
ellipsoid, apex obtuse or usually attenuate-
[ser. I, vol. 7!
apiculate, 2.5-3.5(-4.5) by 1.5-2(—2.5) cm, with
numerous longitudinal slightly raised lines or
ribs, purplish-blackish when ripe; exocarp spon-
gious-fibrous, 1-1.5 mm; endocarp thin-woody,
with numerous longitudinal low ribs which but
slightly appear as such through the exocarp, and
with may be one shallow deeper lateral groove.
Seed white, hard.
Distr. S. Formosa (incl. also Lan Yu = Botel
Tobago I.); in Malesia: Philippines (Babyuan &
Alabat Is., Luzon, and Samar), Moluccas (Moro-
tai, Halmahera, Batjan), N. Celebes (Gorontalo),
and SW. Borneo (Sampit region).
Ecol. Primary littoral or lowland (Dipterocarp)
forest, also secondary growth, from sea-level up
to c. 750m, on clayey, sometimes volcanic soil.
Uses. The plant is said to be used as a cure for
stomach troubles in the Philippines. Wood durable
for interior work.
Vern. Philippines: ampdleng, gohan, malagohan,
Klg., angkak, malatapdi, Bik., basa-basa, busigan,
gozzang-kalinga, karasoéko, maragaudk, mara-
gauéd, \bn., lunas, malapinggan, malasdmat,
rogrogsd, saling-bato, taingan-bdbui, Tag., ma-
lasitim, Pamp., uratdn, Ilk., yaya, Ibn. & Neg.
9. Gonocaryum lobbianum (MiERS) Kurz, J. As.
Soc. Beng. 395 i (1870) 723 Prelakepweeor
& Veg. Pegu (1875) 37; PIERRE, Fl. For. Cochinch.
(1892) in obs. sub t. 268; Cras, Fl. Siam. En. 1
(1926) 274; SLEUM. Blumea 17 (1969) 215; FI.
Thail. 2 (14970) 80. — Stemonurus macrocarpus BL.
Mus. Bot. Lugd.-Bat. 1 (1850) 250; WALP. Ann. 2
(1851) 182; Miers, Ann. Mag. Nat. Hist. II, 10
(1852) 36, repr. Contr. Bot. 1 (1852) 86, non G.
macrocarpum (SCHEFF.) SCHEFF. ex WARB. 1891. —
Platea lobbiana Miers, Ann. Mag. Nat. Hist. II,
10 (1852) 111, repr. Contr. Bot. 1 (1852) 97, t. 17.
— Platea griffithiana Miers, I.c. 110, repr. /.c. 97.
— Lasianthera macrocarpa (BL.) Mia. FI. Ind.
Bat. 1 (1856) 792. — G. gracile (non MiQ.) KuRz,
J. As. Soc. Beng. 39, ii (1870) 72; ibid. 44, ii
(1875) 155; For. Fl. Burma 1 (1877) 240; Kina,
J. As. Soc. Beng. 64, ii (1895) 120; BRANDIs, Ind.
Trees (1906) 152. — G. griffithianum (MIERS}
Kurz, J. As. Soc. Beng. 44, ii (1875) 155; For. Fl.
Burma 1 (1877) 241; PierRE, Fl. For. Cochinch.
(1892) in obs. sub t. 268. — Phlebocalymna lobbiana
(Miers) Mast. in Hook. f. Fl. Br. Ind. 1 (1875)
590. — Phlebocalymna griffithiana (Miers) MAST.
l.c.; BRANDIS, Ind. Trees (1906) 152. — G. wal-
lichii Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 590.
— G. subrostratum Pierre, Fl. For. Cochinch.
(1892) t. 268 B; Gacnep. FI. Gén I.-C. 1 (1911)
828, f. 97, 5-10; Crats, Fl. Siam. En. 1 (1926)
275; GAGNEP. Fl. Gén. I.-C. Suppl. (1948) 755.
— G. harmandianum Pierre, Fl. For. Cochinch.
(1892) t. 268 A; GAGnep. Fl. Gén. I.-C. 1 (1911)
829; Suppl. (1948) 755. — G. siamense WARB.
in Fedde, Rep. 16 (1919) 254. — G. maclurei
Merk. Philip. J. Sc. 21 (1922) 348; Lingn. J. Sc.
5 (1927) 116; CHUN & How, Act. Phytotax. Sin. 7
(1958) 59, f. 5; CHuN & CHANG, Fl. Hainan. 2
(1965) 448, f. 510. — G. sp., CratB, Fl. Siam. En. 1
(1926) 275. — G. poilanei GAGNEP. Fl. Gén. I.-C.
1971]
ICACINACEAE (Sleumer) 21
Suppl. (1948) 755. — G. pavieanum GAGNEP.
l.c. 755, f. 89, 8-9.
Shrub or tree (2—)3-7(-15)m; bark smooth,
grey to brown. Branches slender, tips puberulous,
glabrescent. Leaves oblong to elliptic, sometimes
subobovately so, apex shortly (c. 1 cm) obtusely
and rather suddenly acuminate, base cuneate,
rarely almost rounded, slightly inequilateral, coria-
ceous, glabrous, often somewhat shining above,
greyish-greenish-olivaceous in dry specimens,
(8—)10-16 by (2.5—)3-8(-11) cm, midrib slightly
impressed above, bold below, nerves 4—6(—7) pairs,
rather straight-ascending, upper ones looping, a
little raised on both faces, veins and veinlets
forming a coarse reticulation which is slightly
raised or obscure above, mostly indistinct be-
neath; petiole 1-1.5 cm by c. 2 mm. Inflorescences
axillary, solitary or 2—3(—4) in a cluster, generally
rather few-flowered, sparsely appressedly hairy;
rachis slender, 0.3-1.5(—3) cm. Calyx lobes sub-
ovate, blunt, ciliate, 1.5mm. Petals greenish
white, connate to a tube in their lower 34 part,
(4-)5—6(—7) by 2 mm in the 3, and by c. 2.5 mm
in the 2 flowers. — ¢ Flowers: Filaments 3 mm;
anther cells oblong-ovate, 1.2-1.5 mm. Rudiment
of ovary thick-cylindric, sparsely to subdensely
hairy, on a broadened almost!glabrous subcrenu-
late disk. — 2 Flowers (KERR 5832): Anther cells
hardly 1mm. Ovary ovoid, rather densely hairy
especially at apex; style a little lateral, short;
stigma small. Drupe in mature state + oblongoid
or ellipsoid, rarely subovoid-oblongoid, + round
in @, 3-4(-5) by 2-2.7(-3) cm, apex and base
roundish or + attenuate, though apex even short-
ly apiculate or subrostrate especially in immature
or empty drupes, green for a long time, finally
becoming bluish-purplish or -blackish, smooth,
i.e. ribs of the endocarp in the dry fruit merely
showing as shallow grooves or low ribs, with one
deeper apical lateral groove or flattish part op-
posite the lateral style rudiment; exocarp spon-
gious-fibrous, 2-3(-4) mm; endocarp hard, 1-1.5
mm. Seed whitish, ovoid-acuminate.
Distr. S. China (Hainan), Burma (from Pegu
southwards), Thailand, Indo-China; in Malesia:
Malay Peninsula (Perlis), S. and SE. Borneo.
Ecol. Primary lowland rain-forest or dry
evergreen forest, in mixed moist woods, woody
ravines, scrub jungle, often near river-banks, rare-
ly up to 1200 m.
Uses. Kernel of drupe bitter, much liked by
wild animals, apparently not eaten by natives.
Imperfectly known
10. Gonocaryum melanocarpum Hocnr. Pl. Bog.
Exs. (1904) 42, n. 87; Bull. Inst. Bot. Btzg 22
(1905) 49; Ann. Jard. Bot. Btzg Suppl. III, 2
(1910) 855; SLEUM. Blumea 17 (1969) 217.
Tree, 6-8 m, with + conical crown and short
trunk c. 15 cm g; bark grey with numerous
horizontal small lenticels. Branchlets brownish,
glabrous. Leaves ovate, shortly subacutely or
more bluntly acuminate, base rounded, thin-
coriaceous, shining above, greenish-olivaceous
when dry, 13-18 by 7-12.5 cm, midrib shallowly
depressed above, much prominent beneath, nerves
5-6(-8) curved-ascending pairs, slightly pro-
minent above, more distinctly so beneath, retic-
ulation of veins and veinlets dense, finely raised
on both faces especially above; petiole c. 1 cm by
2(-3) mm. — ¢ Inflorescences not known. —
2 Inflorescences axillary, solitary, spike-like, nod-
ding, flowers 6-10, spaced; rachis slender, prac-
tically glabrous, 6-11 cm by 1 mm; pedicels hairy,
c. 1 mm. Calyx lobes ovate, subacuminate, dorsal-
ly hairy, ciliate, c. 2mm. Petals 5, fleshy, green,
glabrous outside, papillose inside, 6 mm, connate
about halfway to a subcampanulate tube, lobes
spreading. Staminodes void of pollen, hardly 1 mm.
Ovary ovoid, tomentose, 3mm; stigma large,
peltate. Drupe ellipsoid, a little flattened on one or
two sides, attenuate at both ends, apex subacute,
c. 6 by 3.2 by 2.8 cm, black at full maturity, with
several low longitudinal ribs; abortive loculus
absent; exocarp plus endocarp c. 2 mm.
Distr. Cultivated in Hort. Bogor, but origin not
known; apparently from Malesia.
Note. Certainly related to G. litorale in flower
characters and the absence of an abortive loculus,
though drupes are blackish and the leaves are
more similar to those of G. calleryanum.
Excluded
Gonocaryum monostachyum K. Scu. & Laut.
FI. Schutzgeb. (1900) 415; SCHELLENB. Bot. Jahrb.
58 (1923) 166; SLEUM. Blumea 17 (1969) 217.
To guess from the description, the species must
be excluded from Gonocaryum by its very short
(5-6 mm), almost terete petioles, and the very
short (2mm) petals, which besides are hairy
inside, characters all absent from the genus so far
known. The type specimen, which was collected
in New Guinea, Sattelberg area, is apparently lost.
5. GOMPHANDRA
WALL. ex LINDL. Nat. Syst. ed. 2 (1836) 439; em. Becc. Malesia | (1877) 109;
SLeuM. Blumea 17 (1969) 189. — Stemonurus BL. Bijdr. (1826) 648, p.p. —
Platea (non Bu.) Miers, Ann. Mag. Nat. Hist. II, 10 (1852) 108, p.p. — Lasianthera
(non P. Beauv.) Mia. FI. Ind. Bat. 1 (1856) 790, p.p. — Fig. 10.
Dioecious trees, rarely shrubs. Branchlets usually slender, smooth, without
lenticels. Leaves spirally arranged, entire, chartaceous to coriaceous, generally
De FLORA MALESIANA
dark green above and paler beneath in fresh, often olivaceous or brownish,
rarely blackish in dry specimens, often tubercled on the undersurface, generally
pinnate-, exceptionally tripli-nerved, midrib and nerves often a little sunk above,
reticulation generally absent; petiole often somewhat decurrent on branchlet,
stoutish. Flowers functionally unisexual, articulate with the short, often distally
thickened pedicel, in cymes loose to subcapitate, these 2—3(—4)-chotomously
arranged on top of a peduncle, normally in the axils of leaves, rarely (also) op-
posite a leaf, or from defoliate branchlets, generally many-flowered in the 3,
few-flowered in the 2 specimens; bracts small. Ca/yx cupular, minute, 4—5-toothed
to almost entire, subpersistent. Petals 4 or 5 (rarely —7), valvate in bud, either
remaining united below and forming a cup or distally dilated tube, or free to
almost the base, tips inflexed, midrib on the inside rather obscure, caducous.
Disk 0 or obscure. — 3 Flowers: Stamens 4 or 5 (rarely —7), free, often exserted;
filaments thick, flattened, dilated upwards, apex attenuate, generally bearing shorter
papillose or longer clavate hairs in front below the cells, or (also) on the back or
connective; anther cells ovate-elliptic, basifixed, cells slightly or hardly diver-
gent downwards. Rudiment of ovary conspicuous. — 2 Flowers: Calyx and
petals as in the 3 flowers, the petals sometimes absent. Staminodes present or
absent; filaments linear, hairs shorter and less dense than in the ¢ flowers, cells
reduced in size and void of pollen. Ovary cylindric; stigma large, thick-discoid,
slightly 4- or 5-lobed or -crenate. Drupe + oblongoid or ellipsoid, sometimes
shortly attenuate (beaked) distally and slightly swollen (almost apophysate) and
a little asymmetrical at base, crowned by the large persistent stigma; pericarp
smooth, thin, fleshy or juicy; endocarp crustaceous or hard, smooth or generally
with 8—10(—14) longitudinal ridges outstanding especially in the dry fruit. Seed 1;
embryo minute in the top of the fleshy albumen.
Distr. About 33 spp., of which 6 spp. in SE. Asia (SW. India and Ceylon to Burma, Thailand, Indo-
China, and S. China incl. Hainan); in Malesia 27 spp., one of which extends to the Solomon and St. Cruz
Is. Fig. 9.
Ecol. Primary and secondary forest, often as substage trees, both in lowland and in the montane zone
up to c. 1800 m, occasionally also in mossy forest up to 2200 m (on Mt Kinabalu possibly higher),
generally on clayey soil, very rarely limited to peat-swamp forest. Fl. fr. Jan.—Dec.
Disp. Many species have a wide range. There is no evidence yet, that the fruits are eaten by animals
or of the possibility of buoyancy.
Uses. The wood is whitish-creamy and apparently not durable enough to be generally used; besides,
sizes are usually small.
Note. Specific distinction is in the present state far from satisfactory, mainly due to the still rather
scarce specimens collected and their incompleteness. Too many species have been described either from
$ or fruiting specimens only, and their pertaining 9 flowers or fruits are not yet known. Collectors should
care to obtain as complete as possible materials and make ample field notes.
KEY TO THE SPECIES
1. Leaves persistently velutinous or tomentose all over the undersurface, soft to the touch (cf. also
5. G. papuana).
DPN EVEST=2 (So) DGITS: .-) ts. pine eucchelan sath em 1 ccna bets) iat ichamtel Sp elena taal as 1. G. fuliginea
DPPINELVES: "G— 1S) 0 DAITS). «to met ts) eed chat ter GE peute ha) au cheek RA Yolen ou Ries weet 2. G. velutina
1. Leaves covered with scattered hairs underneath, + glabrescent with age, or glabrous fromthe beginning.
3. Ovary in @ flowers laxly to densely hairy. (Rudiment of ovary in ¢ flowers whether or not hairy).
4. Inflorescence conspicuously head-like, its branches (especially in the ¢ specimens) very short.
5. Leaves elliptic or oblong-, rarely ovate-elliptic, 12-20 by 4-8 cm, nerves 5—6(—7) pairs. Drupe
(U8 )2 = 2) 4F bys (O56 JO gOS. pCINe isis Suk ec pect © ALA Menken sta Acacia ee 3. G. capitulata
5. Leaves oblong, rarely ovate-lanceolate, 5-8(-9) by 1.5-3 cm, nerves 3-4(-5) pairs. Drupe 1-1.3
bye OS=OF Gms vhost Ak CMS Eee ch! Rite i Poe ase 1 OO ee aainst Belaate 4. G. lysipetala
1971] ICACINACEAE (Sleumer) 23
4. Inflorescence (especially in the J specimens) generally more loose, its branches distinct and divergent
6. Petals hairy in their upper part, 5—7(-10) mm. Rudiment of ovary in the ¢ flowers hairy on top.
Drupe 3.54 by c. 1.5 cm... . . . 5. G. papuana
6. Petals glabrous or practically so, rarely up to 5 mm. n. Drupe much smaller.
7. Both 3 and Q inflorescences many-flowered, cymes together on top of a peduncle, 1.5—2 cm
(Rudiment of ovary in the g flowers glabrous). Leaves coriaceous... . . 6. G. simalurensis
7. 3 Inflorescences rather many-, 2 ones few-flowered, cymes together on top of a generally rather
short peduncle. Leaves chartaceous to coriaceous.
8. Leaves + laxly covered underneath for a long time with appressed weak hairs, nerves 5-7 pairs.
9. Rudiment of ovary in the f flowers with a tuft of hairs on top. Very base of drupe not swollen.
7. G. cumingiana
9. Rudiment of ovary in the § flowers glabrous or practically so. Base of drupe swollen to almost
apophysate forc.|1mm.. . . . 8. G. pseudojavanica
8. Leaves practically glabrous from the beginning. (Rudiment of ovary in the ¢ flowers glabrous).
10. Leaves with (5—)6-8 pairs of nerves. Base of drupe not swollen. . . .. 9. G. javanica
10. Leaves with 8-10 pairs of nerves. Base of drupe distinctly swollen . . . 10. G. fusiformis
3. Ovary (and rudiment of ovary in the 3 flowers) glabrous or practically so.
11. Filaments glabrous or practically so.
12. Filaments widened upwards and relatively short, i.e. c. 2 times as long as are the anther cells.
11. G. australiana
12. Filaments elongate-linear, at least 3 times as long as arethe anthercells . .. . 12. G. pallida
11. Filaments ventrally and/or dorsally hairy.
13. Filaments with short-clavate or papillose hairs, these never surpassing the anther cell.
14. Drupe (1.6—)1.8-2(-2.2) by 0.8-l cm. ................ 13. G. mappioides
14. Drupe as far as known 1.3-1.6 by 0.5—0.6 cm.
15. Leaves practically glabrous. Calyx + obconical, entire, glabrous. . . . . . 14. G. apoensis
15. Leaves + densely short-pubescent all over the undersurface, especially at midrib and nerves.
Calyx cup-shaped, dented, hairy . . . . . IS. G. flavicarpa
13. Filaments with elongate-clavate (‘glandular’ ) or r penicillate hairs (those from the connective often
surpassing the anther cell).
16. Inflorescence very lax, peduncle and branches elongate, slender and rather few-flowered both
ing and 2 specimens. . . sucst, aha KGe oligantha
16. Inflorescence lax to dense, peduncle. and ‘branches relatively short at least i in the 3 specimens,
generally many-, rarely few-flowered.
17. Petals + densely appressedly hairy outside. . ............. =%17. G.sawiensis
17. Petals sparsely hairy on top (in bud), or glabrous.
18. Drupe (2—)2.5—3(-4) cm long.
19. Inflorescence rather compact, umbelliform. Drupe fusiform, 3.54 by 0.5-0.6 mm g.
18. G. dolichocarpa
19. Inflorescence rather lax. Drupe obovoid-oblongoid-ellipsoid, at least 0.8 cm @.
20. Leaves narrow-lanceolate, chartaceous to subcoriaceous, nerves 3(—4) pairs.
19. G. lancifolia
20. Leaves (obovate-) elliptoc to oblong or lanceolate-oblong, ++ coriaceous, nerves 4 or more
pairs.
21. Leaves elliptic to obovate-elliptic .............. . 20. G. luzoniensis
21. Leaves elliptic- to lanceolate-oblong . . . . wits. (aw GA, RO eee ONE e
18. Drupe as far as known 1-2 (rarely —-2.2) cm long.
22. Inflorescence (at least in the ¢ specimens) rather compact, + umbelliform, on peduncle up
to 5 mm. Leaves + net eCEReD nerves ceiigbi te Mad pairs. Very base of drupe swollen
and smooth . . . .. 22. G. parviflora
22. Inflorescence (at least i in the 3 ‘specimens) with more laxly arranged and diverging branches,
generally on peduncle longer than 5 mm.
23. Drupe ovoid- to oblong-ellipsoid, base generally swollen. Inflorescences (at least in the ¢
specimens) much-branched, generally many-flowered and distinctly scorpioid (i.e. flowers
arranged on one side), rarely with fewer flowers and rather obscurely scorpioid then.
24. Leaves dark to blackish brown in dry state, and rufous- to rusty-tomentellous on midrib
and nerves beneath, as are the branchlets. . . . . 23. G. subrostrata
24. Leaves green to wornes brownish in dry state, and practically glabrous beneath, as are the
branchlets... . . . . 24. G. quadrifida
23. Drupe as far as known i in mature state, oblongoid-ellipsoid, base broadly attenuate and not
swollen. Inflorescences never scorpioid.
25. Leaves brittle and blackish in dry specimens, nerves and veins -- obscure on both faces.
Inflorescence with red-brown hairs. . . + atnleiBBn Gi schoepfiifolia
25. Leaves greenish-brownish and generally not brittle in dry specimens, with a finely im-
pressed reticulation aboveinfully maturestate.. . .... . . . 26, G. pseudoprasina
24 FLORA MALESIANA
[ser. I, vol. 7?
1. Gomphandra fuliginea (ELM.) Merr. En. Philip .2
(1923) 490; SLEUM. Blumea 17 (1969) 192. —
Urandra fuliginea ELM. Leafl. Philip. Bot. 2 (1908)
491. — Stemonurus fuligineus (ELM.) HOWARD, J.
Arn. Arb. 21 (1940) 468.
Shrub or small tree. Branchlets with brownish-
velutinous tips. Leaves lanceolate- to obovate-
elliptic, apex shortly obliquely acuminate, tip
bluntish, base attenuate and a little decurrent on
the petiole, slightly inequilateral, subcoriaceous,
lucid and glabrous above, evenly short fuligineous-
pubescent and soft to the touch beneath, 7—-12(-14)
by 2.5-5 cm, nerves 2 or 3 steeply ascending pairs,
the lowest pair from a little above the base,
other pairs distant, slightly raised beneath; petiole
pubescent, 6-10 by 1.5-2 mm. Flowers unknown.
Infructescence pubescent, on 1 cm long peduncle,
which bears 2 or 3 branches (3mm). Drupe +
ellipsoid, slightly oblique, with 12-15 low ridges
of the hard endocarp, 1.7—2 by 0.9-1.2 cm.
Distr. Malesia: Philippines (N. Luzon: Ben-
guet).
Ecol. In thickets, 1400-1700 m.
Vern. Danka, Ig.
Fig. 9. Distribution of Gomphandra. The figure above the hyphen indicates the number of endemic
species in the island or area, the figure below the hyphen the number of non-endemic species.
2. Gomphandra velutina SLEUM. Blumea 17 (1969)
192.
Tree c. 10 m. Branchlets fulvous-velutinous.
Leaves lanceolate to oblong, apex shortly acutely
acuminate, base cuneate, chartaceous, glabrous
above, fulvous- or ferrugineous-velutinous all
over beneath, dark brown in dry state, 12-17 by
(3—)4-6 cm, nerves 6—7(—8) curved pairs, slightly
raised beneath; petiole velutinous, 8-13 by 1.5 mm.
— ¢ Flowers inc. 15-flowered peduncled (5 mm)
heads composed of several abbreviate cymes,
fulvous-tomentellous, the lower 24 of the petals
excepted. Calyx cupular, 2.5mm. Petals 5,
forming a campanulate tube which slits irregularly,
4.5-5.5 mm. Stamens 5, hardly exserted; filaments
bearded below the anther cells on the inside and
dorsally at connective. Rudiment of ovary with
erect hairs. — 2 Flowers: Calyx and petals as in 3
flowers. Filaments bearded distally only. Ovary
short-tomentose. Drupe unknown.
Distr. Malesia: Central E. Celebes (Porema
at base of Mt Mengkoka), one collection.
Ecol. Rain-forest, 1200 m.
3. Gomphandra capitulata (JUNGH. & DE VRIESE)
Becc. Malesia 1 (1877) 111; SteuM. Blumea 17
(1969) 193. — Aralia capitulata JUNGH. & DE
VrRIESE, Ned. Kruidk. Arch. 1 (1846) 17. —
Lasianthera capitulata (JUNGH. & DE VRIESE)
Mia. FI. Ind. Bat. 1 (1856) 791; Suppl. 1 (1860)
137; Wars. Pflanzenwelt 2 (1916) 349, f. 231 B,
1-4. — Stemonurus capitulatus (JUNGH. & DE
VRIESE) O. KTZE, Rev. Gen. Pl. 1 (1891) 112. —
G. nyssifolia KING, J. As. Soc. Beng. 64, ii (1895)
114; Ripi. Fl. Mal. Pen. 1 (1922) 427; BAKER
f. J. Bot. 62 (1924) Suppl. 21. — Nyssa sessiliflora
(non Hook.) Koorp. Pl. Jungh. ined. n. 57,
cf. HALu. f. Med. Rijksherb. 1 (1910) 13. —
Stemonurus nyssifolius (KING) HowArb, J. Arn.
Arb. 21 (1940) 468.
Tree, 5—20(—28) m, up to 40cm g. Branchlets
slightly winged below the insertion of the petioles,
tips puberulous. Leaves elliptic or oblong-, rarely
ovate-elliptic, apex shortly subabruptly and acutely
acuminate, base cuneate, the very base folded and
decurrent to the grooved petiole, subcoriaceous
to firmly chartaceous, dark olive-brownish and the
edge slightly revolute in dry specimens, glabrous,
12-20 by 48cm, nerves 5-6(—7) pairs rather
straight-ascending, much paler than the lamina in
dry specimens beneath, transverse veins distinct
beneath; petiole 7-12 by 1.5-2 mm. Inflorescences
all over short-pubescent, on top of a slender pedun-
cle (1-2 cm), consisting of several short cymes
1971]
ICACINACEAE (Sleumer) 25
crowded into a kind of head or umbel, which
bear up to 15 flowers in the 3, up to 7 flowers in
the 2 ones. — ¢ Flower: Calyx cupular, ciliate,
1mm. Petals 5, coherent to a narrow tube,
whitish, laxly hairy distally only, c. 4 mm. Stamens
5, much exserted; filaments long-hairy in the upper
half below the anther cells on the ventral side
only. Rudiment of ovary short-hairy. — 2 Flower:
Calyx and petals as in 3 flowers. Staminodes 5,
hairy. Ovary short-hairy except the glabrous base.
Drupe ellipsoid-oblongoid, apex shortly attenuate,
slightly or not curved, the very base swollen and
smooth, whilst the endocarp shows numerous
outstanding ridges above, (1.8—)2-2.4 by (0.6-)
0.7-0.8 cm; exocarp pale pink or greenish-white.
Distr. Malesia: Sumatra (incl. Simalur I.),
Malay Peninsula (Perak, Pahang).
Ecol. Forest, in Sumatra at 150-915 m, in the
Malay Peninsula at 1065-1370 m.
Vern. Sumatra: simarsimata, Batak; Simalur:
awa suki, pului, p. pajo, samsam dotan, surin
uding, tutun ansan délok, M.
4. Gomphandra lysipetala Starr, Trans. Linn. Soc.
II, Bot. 4 (1894) 139; SLEuM. Blumea 17 (1969)
194. — Stemonurus lysipetalus (STAPF) MERR.
En. Born. (1921) 356.
Tree up to 30m high, trunk up to 45cm 9;
bark dark grey. Leaves oblong, or sometimes more
ovate-lanceolate, apex shortly obtusely attenuate
or subacuminate, base + abruptly narrowed to the
grooved petiole, (sub)coriaceous, glabrous at ma-
turity, dark brown when dry, 5-8(-9) by 1.5-3 cm,
nerves 3—4(—5) curved-ascendent pairs, slightly
raised beneath; petiole 7-10(-12) by c. 1 mm. —
3 Cymes 2 or 3, very shortly peduncled, all con-
densed into a head-like or subumbellate inflores-
cence on top of a slender peduncle (5-10 mm).
Calyx cup-shaped, minutely 5-dented, hardly 1 mm.
Petals 5, becoming free from each other both from
top and base at the same time, whitish-greyish,
c.3 mm. Stamens 5, hardly exserted; filaments with
numerous short-papillose hairs in the middle of
the ventral, glabrous on the dorsal side. Rudiment
of ovary laxly hairy. — 2 Cymes similarly arranged,
though few-flowered. Calyx and petals as in 3
flowers. Staminodes with numerous papillose hairs
in front. Ovary densely set with whitish short
hairs. Drupe ellipsoid-oblongoid, apex (sometimes
a little oblique) and base gradually attenuate,
1-1.3 by 0.5—0.7 cm; exocarp thin-fleshy, cream;
endocarp with several longitudinal shallow ridges.
Distr. Malesia: Borneo (Mt Kinabalu area).
Ecol. Primary and secondary forest, on slopes,
1220-1830 m.
5. Gomphandra papuana (Becc.) SLeum. Notizbl.
Berl.-Dahl. 15 (1940) 238; Steum. Blumea 17
(1969) 194. — Lasianthera litoralis (non Mia.)
F. v. M. Descr. Not. 2 (1877) 53. — Platea pa-
puana Becc. Malesia | (1883) 257; VALer. Crit.
Overz. Olacin. (1886) 254. — Stemonurus pu-
berulus K. Scu. & Laur. Fl. Schutzgeb. (1900)
414. — Stemonurus zygomorphus PuLte, Nova
Guinea 8 (1912) 658; ScHeLLens. Bot. Jahrb. 58
(1923) 162. — Stemonurus papuanus (BECC.)
SCHELLENB. /.c. 161. — G. zygomorpha (PULLE)
SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 239.
Tree, 5—10(—30) m, up to 20 cm 9; bark smooth,
mottled grey-brown. Branchlets short-pubescent
to tomentulose distally, glabrate below, pith
hollow. Leaves elliptic to oblong-, rarely subovate-
elliptic, apex generally shortly, rather abruptly and
often obliquely acuminate, base cuneate, rarely
almost rounded, stiff-chartaceous or subcoria-
ceous, greyish-green to pale olivaceous-brownish
in the dry state, early glabrescent above, + persis-
tently covered with numerous short hairs and
rather soft to the touch all over beneath, finally
partly glabrescent there, 15-25 by (5—)6-11.5 cm,
nerves 5—7(—8) curved-ascending and beneath some-
what prominent pairs, transverse veins subparallel
to and distant from each other, slightly raised
beneath, reticulation obsolete; petiole deeply
grooved above, glabrescent, 10-12 by 2(—3) mm.
— § Inflorescences consisting of 2 or 3 short-
branched 2—5-flowered cymes on top of a common
peduncle (3-6 by 1 mm), all over densely covered
with appressed ferrugineous hairs, the lower part
of the petals excepted. Calyx cup-shaped, green,
4(—-5)-denticulate, 2 by 3 mm. Petals 4 or 5 (rarely
~7), coherent in the lower 24, though partially
free from each other by 1 or 2 slits, thickish,
greenish to white, 5-7 mm. Stamens 5(—7), much
exserted; filaments densely hairy both in front
below the anther cells (1 mm) and dorsally on
connective. Rudiment of ovary with erect hairs on
top. — 2 Inflorescences 2-3-flowered, together on
3-5 mm long peduncle. Calyx and petals (7-10
mm) as in ¢ flowers. Staminodes slightly exserted.
Ovary + laxly appressedly hairy. Drupe obovoid-
oblongoid, strongly beaked in immature, much
less so in mature state, yellow, 3.5—4 by c. 1.5 cm,
somewhat hairy distally only; endocarp rather
thin, with c. 8 obtuse longitudinal ridges.
Distr. Malesia: New Guinea.
Ecol. Primary and secondary forest, also on
flooded plains, 25-450 m, locally common in
undergrowth.
Vern. Assiei, Astrolabe Bay area, behwah,
sinagapa, Orokaiva (Mumuni), numinti, Rawa,
sisinam, Koropa.
6. Gomphandra simalurensis SLEUM. Blumea 17
(1969) 194.
Tree, 13-25 m. Branchlets robust, tips laxly ap-
pressedly hairy. Leaves elliptic, sometimes sub-
obovate-elliptic, apex shortly and bluntly acumi-
nate, base cuneate and a little decurrent on the
petiole, coriaceous, dark brown when dry, gla-
brous at full age, 10-17(-21) by (4-)4.5-9(-11)
cm, nerves 5-6(-7) pairs, lower ones rather
straight, upper ones more curved-ascendent, al-
most flat above, raised beneath, transverse veins
rather obscure beneath; petiole 1-1.8(-2.3) by
c. 2.5mm. ¢ & & Inflorescences consisting of 3-5
many-flowered cymes collected on top of a rather
slender peduncle (1.5-2 cm), pale rusty-tomentel-
lous except the petals. — ¢ Flowers: Calyx cu-
pular, 4-5-dented, sparsely pubescent, ciliate, |
26 FLORA MALESIANA
[ser. I, vol. 74
Fig. 10. Gomphandra javanica (BL.) VALET. a. Habit, with g inflorescence, x 14, b. @ inflorescence,
x 314, c. § flower, <5, d. stamen, ventral side, x 10, d’. ditto, dorsal side, x10, e. drupe, x1 (a, c-d
KOsTERMANS & VAN WOERDEN 128, b WINCKEL 336, e JAcoBS 4797).
1971]
ICACINACEAE (Sleumer) P-4 |
mm. Petals 4 or 5, white or greenish, still connate
at anthesis to a 4- or 5-lobed distally dilated tube,
4.5 mm. Stamens 4 or 5, much exserted; filaments
laxly set with elongate-clavate hairs ventrally
below the anther cells, and penicillate dorsally at
connective. Rudiment of ovary glabrous. — 2
Flowers: Calyx and petals as in 3 flowers. Sta-
minodes present. Ovary glabrous at the very base,
otherwise densely set with forward directed hairs.
Drupe oblongoid, a little oblique, slightly rostrate,
pale yellow, 1.6—1.8(—2) by 0.7-0.8 cm; endocarp
longitudinally 10—12-costulate.
Distr. Malesia: Simalur I. (off the coast of NW.
Sumatra).
Ecol. Primary lowland forest.
Vern. Bajut uding, rubi silai, tutun bahuhu
alafai, t. b. délok, t. b. dotan, t. b. uding, M.
7. Gomphandra cumingiana (Miers) F.-VILL. Nov.
App. (1880) 46; VipAL, Phan. Cuming. Philip.
(1885) 102; Rev. Pl. Vasc. Filip. (1886) 86;
VALET. Crit. Overz. Olacin. (1886) 229; MERR. En.
Philip. 2 (1923) 490; SLEuM. Blumea 17 (1969) 194.
— Stemonurus cumingianus Miers, Ann. Mag. Nat.
Hist. I], 10 (1852) 42, repr. Contr. Bot. 1 (1852) 92;
O. Krze, Rev. Gen. Pl. 1 (1891) 112; DAuL, J.
Arn. Arb. 33 (1952) 268, f. 50 & 50 A (pollen). —
Stemonurus agusanensis Ei. Leafl. Philip. Bot. 7
(1915) 2689. — G. agusanensis (ELM.) MERR. En.
Philip. 2 (1923) 489.
Tree (3—)5-15 m, up to 30cm 9; bark smooth,
yellowish-brown. Branchlets flexuous, tips rusty-
tomentellous. Leaves oblong to ovate- or elliptic-,
rarely subobovate-oblong, apex shortly rather
abruptly and sometimes obliquely acuminate,
tip blunt, base broadly attenuate, slightly inequi-
lateral, very base + abruptly narrowed to the pe-
tiole, membranous to chartaceous, dark brown
above, much paler beneath when dry, dull, gla-
brous above, the midrib and nerves excepted,
undersurface all over set with scattered short + ap-
pressed soft fine hairs, more densely so at midrib
and nerves, (7.5—)9-16(-24) by 5-8(-10) cm,
nerves in 5—7 generally strictly, or rarely more
curvedly ascending pairs, slightly raised beneath,
cross veins faintly prominent beneath; petiole
puberulent, 1-1.5 cm by c. 1.5mm. — ¢ Inflores-
cences composed of several divaricate few-flowered
cymes, together on top of slender peduncle (0.5—1.5
cm), all over rusty-puberulent, the petals excepted.
Calyx cup-shaped, hardly dented, ciliate, 1 mm.
Petals (4—)5, connate to a tube for their lower 24,
glabrous except some hairs on top in bud stage,
yellowish-white, (4-)5 mm. Stamens 5, exserted;
filaments with numerous club-shaped hairs ven-
trally below the anther cells and_penicillate
dorsally at connective. Rudiment of ovary with
erect hairs on top. — 9 Inflorescences more \ax-
flowered. Calyx and petals as in 3 flowers. Sta-
_minodes (4—-)5, penicillate. Ovary ferrugineous-
tomentellous. Drupe subovoid-oblongoid, either
equally attenuate on both ends, or generally
slightly beaked distally, still laxly pubescent for
quite a while, (1.3—)1.5-1.7 (rarely —2) by 0.7-0.8
cm; whitish-yellowish, pinkish or salmon-coloured;
endocarp with 10-12 shallow longitudinal ribs.
Distr. Malesia: Borneo, Philippines.
Ecol. Primary (also mixed Dipterocarp) forest
at low and medium altitude, in the Mt Kinabalu
area up to 1525 m.
Vern. Philippines: Bardbo, Mang., bayanotan,
S. L. Bis., lambuan, Mbo.., lipid, Sul., mavgoi, Bik.
8. Gomphandra pseudojavanica SLEUM. Blumea 17
(1969) 195.
Tree 4-15m. Branchlets puberulous at tips.
Leaves oblong-elliptic, apex -+ abruptly acu-
minate for 1-2cm, tip bluntish, base broadly
attenuate, subcoriaceous, olivaceous-brown above,
paler beneath in dry state, glabrous above, sparsely
set with short hairs initially, more laxly so or at the
lower part of the midrib only at maturity and
minutely tubercled beneath, (8—)10-16 by (4-)4.5-
8(—10) cm, nerves 5—6(—7) pairs, curved-ascendent,
distinctly looping before the edge, slightly raised
beneath; petiole puberulous initially, 1-1.5(-2) cm
by 1.5-2 mm. — ¢ Inflorescences composed of 3
or 4 divergent, rather dense flowered cymes to-
gether on top of a slender peduncle (0.5-1.3 cm),
short-hairy on peduncle and branches. Calyx
cup-shaped, minutely 5-dented, tube glabrous,
hardly 1 mm. Petals connate in a distally widened
and shortly 5-lobed tube, whitish-greenish, gla-
brous, c. 3.5 mm. Stamens 5, exserted for c. 1 mm;
filaments laxly penicillate below the anther cells
ventrally and at connective dorsally. Rudiment
of ovary glabrous or practically so. — 9 Inflores-
cences not known. Immature infructescences 1—3-
flowered, peduncle c. 5 mm, pedicels (branches of
the reduced cyme) 3—5 mm. Developed ovary
densely appressedly hairy. Drupe ellipsoid-oblon.-
goid, a little curved, shortly and rather narrowly
attenuate at apex, more broadly so at base, very
base swollen or subapophysate for c. 1mm,
1.7-1.9 by 0.6-0.7 cm, whitish; exocarp with few
and rather low longitudinal ridges.
Distr. Malesia: Simalur I. (off the coast of
NW. Sumatra).
Ecol. Primary lowland forest.
Vern. Ansan délok, a. d. uding, bajut uding,
bélujan étém, pului silai, sosot manu, surimanu
uding, tutun surimanu délok, M.
9, Gomphandra javanica (BL.) VALET. Crit. Overz.
Olacin. (1886) 217, t. 4, f. 1-9c, t. 5, f. 23a, t. 6, f.
46; K. & V. Bijdr. 5 (1900) 150; Hocur. Bull.
Inst. Bot. Btzg 22 (1905) 42; Mott & JAnss.
Mikr. 2 (1908) 238, f. 106 (wood-anat.); BACK.
Schoolfl. Java (1911) 225; Koorp. Exk. Fl. Java 2
(1912) 531; Koorp.-Scuus. Syst. Verz. 1, Fam. 162
(1912) 3; Koorp. Atlas 1 (1913) t. 116; Docr. v.
LEEUWEN, Zoocecidia (1926) 332, f. 597 (galls);
Amsu. in Back. Bekn. Fl. Java (em. ed.) 6 (1948)
fam. 135, p. 4; Back. & Baku. f. Fl. Java 2
(1965) 60; SLeUM. Blumea 17 (1969) 196. — Ste-
monurus javanicus BL. Bijdr. (1826)) 649, incl. var.;
WALp. Rep. | (1842) 378; Hassk. Cat. Hort. Bog.
(1844) 214; Miers, Ann. Mag. Nat. Hist. I, 10
(1852) 36, repr. Contr. Bot. 1 (1852) 86, incl.
var.; DAHL, J. Arn. Arb. 33 (1952) 268 (pollen).
28 FLORA MALESIANA
— Stemonurus pauciflorus Bu. Bijdr. (1826) 648;
WALP. Rep. 1 (1842) 378; Muers, Ann. Mag.
Nat. Hist. II, 10 (1852) 35, repr. Contr. Bot. 1
(1852) 86. — Lasianthera javanica (BL.) Mia.
Fl. Ind. Bat. 1 (1856) 790, incl. var. pauciflora
(BL.) Mia. lc. 791; Suppl. 1 (1860) 137. — G.
javanica (BL.) VALET. var. lanceolata K. & V.
Bijdr. 5 (1900) 151; Hocnr. Pl. Bog. Exs. (1904)
41; Bull. Inst. Bot. Btzg 22 (1905) 42; MoLL &
JANss. Mikr. 2 (1908) 243 (wood-anat.); Koorpb.-
ScuumM. Syst. Verz. 1, Fam. 162 (1912) 4. —
G. javanica (BL.) VALET. var. dolichocarpa K. &
V. Bijdr. 5 (1900) 151; MoLL & Janss. Mikr. 2
(1908) 243 (wood-anat.); Koorp.-ScHumM. Syst.
Verz. 1, Fam. 162 (1912) 4. — G. comosa (non
KING) BAKER f. J. Bot. 62 (1924) Suppl. 21. —
Fig. 10.
Tree, (3—)5—15 m, suberect, rarely up to 40 cm 9;
bark olive green. Branchlets appressedly puber-
ulous at tips as are the petioles and may be the
young leaves. Leaves variable in form and size,
generally elliptic to oblong, sometimes obovate-
elliptic or lanceolate, apex shortly acuminate to sub-
caudate, base + acutely attenuate, subcoria-
ceous, shining, glabrous or very early so, 5—12(-16)
by 2.5-6.5 cm, nerves (5—)6—-8 pairs, lower ones
more erect, upper ones more spreading and mostly
rather straight from the midrib, + obscurely in-
arching before the edge, moderately raised beneath,
veins lax, generally obsolete; petiole 8-15 by
1-1.5 mm. — 3 Inflorescences consisting of 2-3-
chotomously arranged cymes, (5—)10—15-flowered,
finely pubescent, on common slender peduncle
0.5-1.5 cm. Calyx cupular-patelliform, minutely
4—5-dented, puberulous, 1 mm. Petals connate to
a distally widened tube, free in the upper third,
a little fragrant, light green, glabrous, 4-5 mm.
Stamens 4 or 5, a little exserted; filaments long-
hairy below the anther cells ventrally, and more
laxly so at connective dorsally. Rudiment of ovary
glabrous. — 2 Inflorescences 2—3-chotomous, 3—5-
flowered. Calyx and petals as in the 3 flowers.
Staminodes 5, hairy mainly on the ventral side
below the anther cells. Ovary + densely appressed-
pubescent. Drupe narrowly to broadly ellipsoid,
rather variable, gradually or more abruptly nar-
rowed at both ends, 1.5-2 by 0.6-0.8 cm in dry
state, said to measure 1.7—2 by 1-1.5 cm in fresh
state; exocarp fleshy, milky-white, almost van-
ishing at full maturity; endocarp fibrous, longi-
tudinal ribs numerous, low.
Distr. Malesia: Java and Lesser Sunda Is.
(Bali, Lombok).
Ecol. Primary and secondary forest, scattered -
in the lowland, more frequent in montane forest,
(50—)600—1200(—1700) m.
Vern. Bardbdj, haréndeuj, ki johonn, ki sékél,
ki tangkil, mitongérrét, S, djirak ireng, J, tés
katésan, Mad.
10. Gomphandra fusiformis SLEUM. Blumea 17
(1969) 196.
Treelet, 3m, stem 2-4cm g. Branchlets very
slender, tips shortly fuscous-hairy. Leaves ob-
long, apex per 1-2 cm subabruptly and narrowly
[ser. I, vohe?*
acuminate (tip often curved and bluntish), base
cuneate, chartaceous, lead-coloured greenish, but
slightly brownish, dull, glabrous, 8-13 by 2.5-3.5
cm, nerves 8-10 pairs, lowest pair slightly supra-
basal and high curved-ascendent, upper pairs
from the midrib in a wider angle, more straight
and looping, forming an intramarginal nerve with
the basal pair, hardly raised beneath; petiole 6-10
by c. 1 mm. — Inflorescences consisting of 1 or 2
few-flowered cymes, together on top of a slender
peduncle (1 cm), very shortly pubescent. Calyx
cupular, minutely 4-dented, ciliate, 1 mm. Petals
connate for their lower 34 part into a distally
widened tube, glabrous, white, 3.5 mm. Stamens 4,
shortly exserted; filaments penicillate on upper
ventral side. Rudiment of ovary glabrous. —
2 Inflorescences reduced to a few-flowered, almost
umbelliform and shortly peduncled cyme. Calyx
and petals as in J flowers. Staminodes 4, laxly peni-
cillate. Ovary densely shortly rusty-pubescent, the
glabrous very base excepted. Drupe fusiform, sub-
abruptly rostrate-attenuate at apex, more gra-
dually so at base, very base somewhat cupuliform-
inflated and quite smooth, whilst above the
endocarp shows several longitudinal low ridges,
(2—)2.3-2.5 by 0.6—-0.7 cm.
Distr. Malesia: Central W. Sumatra (Mt Sago
near Pajakumbuh).
Ecol. Forest, 900-1200 m.
11. Gomphandra australiana F. v. M. Fragm. Phyt.
Austr. 6 (1867) 3; VALET. Crit. Overz. Olacin.
(1886) 226, t. 4, f. 20 ah, t. 5, f. 23b; F. M. BAILEY,
Compr. Cat. Q: Pl. (1913) 933°C. Weide,
Proc. R. Soc. Queens]. 53 (1942) 212; SLEUM. Blu-
mea 17 (1969) 197. — G. axillaris (non WALL.)
Becc. Malesia 1 (1877) 110, pr. specim. papuan.:
t. 4, f. 8-13. — Stemonurus australianus (F. v. M.)
O. KtTze, Rev. Gen. PI. 1 (1891) 112. — G. poly-
morpha (non WiGHT) F. M. BatLey, Bot. Bull.
Queensl. Dep. Agr. Brisbane 8 (1893) 71; Compr.
Cat. Q. Pl. (1913) 93. — Stemonurus ramuensis
Laut. in K. Sch. & Laut. Nachtr. (1905) 305;
SCHELLENB. Bot. Jahrb. 58 (1923) 163; BIRNIE,
Nova Guinea 14, 2 (1926) 275..— ? Stemonurus
viridis SCHELLENB. Bot. Jahrb. 58 (1923) 163. —
G. ramuensis (SCHELLENB.) SLEUM. Notizbl. Berl.-
Dahl. 15 (1940) 238. — ? G. viridis (SCHELLENB.)
SLEuM. /.c. 239.
Tree, 6-25 m, sometimes with buttresses up to
3m high. Branchlets generally quite glabrous.
Leaves oblong- -to- ovate-elliptic, apex shortly,
bluntly and gradually, more rarely + abruptly
acuminate, base broadly cuneate toalmost rounded
slightly inequilateral, coriaceous, practically gla-
brous, (6—)10—17(—20) by 5—7(-8) cm, dull greenish-
olivaceous when dry, the undersurface often min-
utely tubercled, nerves (4-)5—6 curved-ascending
pairs, hardly depressed above, slightly prominent
beneath, veins lax, rather obscure; petiole (0.7—)
1-1.5cm by 1.5-2 mm. — ¢ Inflorescences con-
sisting of 2 or 3 divaricate (3—)5—7-flowered cymes,
together on peduncle 0.5-l1 cm, covered with
short substrigose hairs, the petals excepted. Calyx
cup-shaped, hardly dented, 1-1.3 mm. Petals white
1971]
ICACINACEAE (Sleumer) 29
or greenish, coherent in the lower 74 for a long
time into a narrow-campanulate tube 2.5—3 mm.
Stamens hardly or not exserted; filaments widened
upwards, 2—2.3 mm, glabrous; anther cells 1 mm.
Rudiment of ovary glabrous. — 2 Inflorescences
with 2—4 flowers on top of slender peduncle (c. 1
cm). Calyx and petals as in 3 flowers. Staminodes
present. Ovary glabrous. Drupe ellipsoid, obtuse at
both ends, pale reddish, with 12-15 rather marked
longitudinal ridges down to the very base, 1.8-2
(-2.3) by 1-1.2 cm.
Distr. Australia (North Queensland); in Ma-
lesia: Kai Is. and New Guinea.
Ecol. Primary forest, scattered from the low-
land up to 500 m.
12. Gomphandra pallida SLEum. Notizbl. Berl.-Dahl.
15 (1940) 238; Blumea 17 (1969) 197. — Stemo-
nurus gracilis SCHELLENB. Bot. Jahrb. 58 (1923)
164, non G. gracilis KING, 1895.
Shrub or treelet, 5-7 m. Branchlets quite gla-
brous. Leaves oblong-elliptic or elliptic, apex sub-
abruptly acuminate for 1-2cm, base broadly
cuneate, slightly inequilateral, subcoriaceous or
firmly chartaceous, glabrous, dull olivaceous in
dry state, minutely tubercled beneath, 8-18 by
(3.5—)4-6(—7) cm, nerves 4~-S(-6) curved-ascen-
dent pairs, a little raised beneath only, veins and
veinlets faintly prominent-reticulate in fully ma-
ture leaves; petiole 1-1.5cm by 1-1.5mm. —
3 Inflorescences with 2 or 3 few-flowered cymes
together on top of slender peduncle (0.5-1 cm),
very shortly pubescent, the petals excepted. Calyx
cup-shaped, hardly dented, 0.5mm. Petals 5,
thin, white, connate in the lower 7% to a slightly
dilated tube, 3.5 mm. Stamens 5, widely exserted;
filaments elongate-linear, glabrous, 4.5 mm; anther
cells 0.5 mm. Rudiment of ovary glabrous. — 92
Inflorescences few-flowered; flowers unknown.
Juvenile drupe glabrous.
Distr. Malesia: New Guinea (Sepik Distr.),
twice found.
Ecol. Swampy alluvial forest, 20-50 m.
13. Gomphandra mappioides VALET. Crit. Overz.
Olacin. (1886) 227, f. 21, a-f; SLEUM. Blumea 17
(1969) 198. — Stemonurus mappioides (VALET.)
O. Kize, Rev. Gen. Pl. 1 (1891) 112).— G.
australiana F. vy. M. var. celebica VALET. in Koord.
Minah. (1898) 392; Ic. Bog. 1 (1908) t. 89;
Ho.tH. & LAm, Blumea 5 (1942) 205. — G. ?
javanica (non (BL.) VALET.] HoLgu. & LAM Lc. 205.
Tree, up to 25 m, up to 45 cm @; bark olive
green. Branchlets appressedly puberulous in their
younger parts, as are petioles and _inflores-
cences. Leaves lanceolate to oblong-lanceolate or
elliptic, rarely oblong-elliptic, apex attenuate-acu-
minate, tip acutish, base + broadly narrowed, firm-
ly subcoriaceous, dark olivaceous and a little shin-
. ing above, paler, more brownish and dull beneath in
dry specimens, glabrous above, shortly appressedly
hairy all over the undersurface initially, finally
glabrescent there, the midrib excepted, finely
tubercled underneath, (10—)12-16 by (2.5-)3-4.5(-8)
cm, nerves 4-6(-8) spreading to rather steeply
ascendent pairs, obscurely inarching before the
edge, reticulation obsolete; petiole 1-1.5 cm by
c. 1 mm. — 6 Inflorescences with (2—)3 or 4 cymes
together on top of slender peduncle (l—2 cm),
each cyme expanded and + recurved, dense
flowered, very shortly branched distally, the
flowers rather laxly arranged. Calyx widely cup-
shaped, subtruncate or shortly 5-dented, laxly
hairy, 1-1.5mm. Petals 5, connate for c. *%
their length to a subcampanulate tube, white,
glabrous, 3 mm. Stamens 5, slightly exserted; fila-
ments densely set with rather short glandular hairs
below the anther cells ventrally and dorsally at the
connective, the hairs not exceeding the anther cells.
Rudiment of ovary glabrous. — 2 Inflorescences
with 3—Scymes on top of a slender peduncle
(0.5—1 cm), each cyme reduced to a single flower,
the latter pedicelled for 3-5 mm and arranged ina
kind of umbel. Calyx and petals as in ¢ flowers.
Staminodes 5, short-hairy below the void anther
cells. Ovary cylindric, becoming ovoid, glabrous.
Infructescences with 2—3(—5) branches (0.5—1 cm)
together on top of a peduncle (1-1.5 cm). Drupe
subovoid-ellipsoid or ellipsoid, yellowish, (1.6—)
1.8—2(—2.2) by 0.8-1 cm, apex broadly attenuate
and slightly curved, rounded, rarely attenuate;
endocarp with numerous longitudinal low ribs.
Distr. Malesia: Lesser Sunda Is. (Sumba,
Flores, Wetar, Timor), Celebes, Moluccas (Mo-
rotai, Talaud, Halmahera, Ternate, Tidore, and
Tanimbar Is.).
Ecol. Forests, from lowland to the montane
region, occasionally up to 1600 m.
Vern. Flores: saka, Endeh, te-u4, Manggarai;
Celebes: danoan, makapojdan, makuranga (Ts.),
sansalan, Minahasa (T1.), lowa pute, taluha, Mailili
(Tb.), tohaka, Makassar, benu, Buton; Moluccas:
anuwuka, Talaud, mali mali, Tidore, mador,
Morotai, momodja, obu banga, Ternate.
14. Gomphandra apoensis (ELM.) Merr. En.
Philip. 2 (1923) 490; SLeuM. Blumea 17 (1969) 198.
— Stemonurus apoensis Em. Leafl. Philip. Bot. 7
(1915) 2690; Howarp, J. Arn. Arb. 21 (1940)
487, t. 1, f. 11-14; Dani, ibid. 33 (1952) 268
(pollen).
Tree, 15-20 m, trunk up to 45 cm g. Branchlets
puberulent distally. Leaves oblong, apex short-
acuminate, tip often bluntish, base broadly at-
tenuate to roundish, subcoriaceous, rather dark
brown when dry especially above, initially sparsely
set with short + appressed pale hairs at midrib
and nerves beneath, glabrous with age, 6-13 by
2.5—4.5 cm, nerves 5—7 pairs, rather straight, ob-
scurely inarched before the edge, slightly raised
beneath, net of veins coarse and rather obscure;
petiole c. 1 cm by 1-1.5 mm. — 3 Inflorescences
2(—3) cm long, consisting of 2 or 3 shortly branched
cymes on top of common peduncle (0.51 cm),
subdensely 5—12-flowered, short-hairy except calyx
and petals. Calyx +. obconical, entire or almost so,
1.5(-2) mm. Petals 5, connate for the lower 7%
toa narrowly campanulate tube, white or yellowish
green, 3.5-4mm. Stamens 5, a little or hardly
exserted; filaments with short-clavate or papillose
30 FLORA MALESIANA
[ser. I, vol. 7?
hairs both in front below the anther cells and on
connective. Rudiment of ovary glabrous. — @ In-
florescences and drupe unknown.
Distr. Malesia: Philippines (Luzon, Negros,
Mindanao; possibly also Bohol).
Ecol. Primary forest on slopes and along
streams, 900 m.
Vern. Marumai, Bag.
15. Gomphandra flavicarpa (ELM.) MEerR. En.
Philip. 2 (1923) 490; SLEuM. Blumea 17 (1969) 198.
— Stemonurus flavicarpus ELM. Leafl. Philip. Bot.
7 (1915) 2691; DAHL, J. Arn. Arb. 33 (1952) 268
(pollen).
Shrub-like or slender tree, 3-5 m, 2-10 cm g;
bark smooth, brown. Branchlets short-pubescent.
Leaves almost distichous, oblong to obovate-
oblong, apex shortly and + bluntly acuminate,
base broadly attenuate to almost rounded, firmly
chartaceous to subcoriaceous, dark brown above,
much paler beneath when dry, initially sparsely
hairy along lower part of the midrib, with age
practically glabrous above, shortly subappressedly
pubescent all over the undersurface especially
on midrib and nerves (though less obviously than
in G. cumingiana), 8-14 by 2.5-4(—5.5) cm, nerves
5-6 ascendently curved and obscurely inarching
pairs, slightly raised beneath, net of veins coarse,
faintly prominent beneath; petiole puberulent,
0.8-1 cm by 1 mm. Inflorescences from foliate and
defoliate axils, both g and 2 ones composed of
2(-3) few-flowered cymes on common peduncle
(c. 5mm), short-brownish-pubescent, the petals
excepted. —- g¢ Flowers: Calyx cup-shaped, 4(—5)-
denticulate, 1 mm. Petals 4 or 5, united for the
lower 74 to a subcampanulate tube, white or
cream, 3-4 mm. Stamens 4 or 5, exserted for 1.5—2
mm; filaments with rather few papillose hairs in
front below the anther cells and at connective.
Rudiment of ovary glabrous. — 92 Inflorescences
2-3-flowered. Calyx and petals as in ¢ flowers.
Staminodes 4 or 5, papillose. Ovary glabrous.
Drupe oblongoid-fusiform, apex slightly oblique
and a little beaked, base gradually attenuate, pale
yellow when fresh, 1.3—1.6 by 0.5-0.6 cm; endo-
carp thin, with several slightly outstanding longi-
tudinal ribs.
Distr. Malesia: Philippines (Mindanao).
Ecol. Montane, also mossy forest, (700—)1200-—
2200 m.
Vern. Mantangali, Manobo, maramai, mata-
matd, Bag.
16. Gomphandra oligantha SLEuM. Notizbl. Berl.-
Dahl. 15 (1940) 238; Blumea 17 (1969) 199. —
Urandra pauciflora MERR. Philip. J. Sc. 3 (1908)
Bot. 243. — G. pauciflora (MERR.) MERR. En.
Philip. 2 (1923) 491, non Crap, 1914.
Small tree, glabrous except the slightly puber-
ulent tips of branchlets, the young petioles and the
inflorescences. Leaves almost distichous, ovate
to ovate-elliptic, or elliptic, apex -+ abruptly
narrowed to a subcaudate bluntish acumen
(1-2 cm), base broadly cuneate, the very base
often contracted and a little decurrent on the
petiole, membranous to subchartaceous, dark
olive-brownish when dry, 6-10 by 2.5—5 cm,
nerves 5—6 moderately curved-ascendent pairs,
minutely raised beneath, veins + obscure; petiole
5-10 by hardly 1 mm. Both 3 and @ inflorescences
very lax, slender as are the peduncles (2—3 cm)
and the branches. — g Flowers (ELMER 12508)
in a 3—5-flowered very lax cyme; pedicels in fully
developed inflorescences 2—3mm. Calyx cup-
shaped, base truncate, laxly ciliate, hardly 4-
dented, 1 mm. Petals 4, glabrous, coherent in the
lower 34, glabrous, 3 mm. Stamens 4; filaments
glabrous on the ventral, and with a few penicillate
hairs on the dorsal side at the connective. Rudi-
ment of ovary glabrous or maybe with a few
hairs along edges. — 2 Flowers in 2 or 3 very lax
cymes, each cyme reduced to one flower; branches
(simulating pedicels) 1-2 cm. Calyx and petals as
in ¢ flowers. Staminodes 4; filaments laxly long-
hairy on ventral side immediately below the anther
cells and at connective. Ovary sparsely hairy in
upper part, practically glabrous. Submature
drupe ovoid-oblongoid, apex very shortly sub-
abruptly attenuate and maybe very sparsely hairy,
base rounded, seen up to 1.3 by 0.7 cm; endocarp
thin, with numerous low longitudinal ridges.
Distr. Malesia: Philippines (Mindoro, Si-
buyan).
Ecol. Primary forest at low altitudes.
17. Gomphandra sawiensis (BIRNIE) SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 238; Blumea 17 (1969) 199.
— Stemonurus sawiensis BIRNIE, Nova Guinea 14,
2 (1926) 275.
Tree, c. 6m. Branchlets fulvous-tomentellous
at tips. Leaves oblong, apex shortly acuminate or
sometimes obtuse, base cuneate, chartaceous,
initially sparsely appressedly long-hairy all over
especially beneath, at maturity practically gla-
brous, brownish when dry, with numerous pale
tubercles on both faces, 10-15(-18) by 3-—5(-—7)
cm, nerves 7—8(-9) pairs, spreading and rather
straight, raised beneath, transverse veins visible,
no reticulation; petiole descending on the branchlet
as a slight ridge to the next leaf, 1-1.5cm by
1.5-2 mm. — ¢ Inflorescences few-flowered cymes
on very short common peduncle, all over densely
appressedly fulvous-pubescent, 1-1.5 cm long in
all. Calyx widely cup-shaped, base truncate, edge
subentire, 2mm. Petals 4, remaining connate for
their lower 24 at anthesis, -- densely appressedly
hairy outside, 5—5.5 mm. Stamens 4, a little ex-
serted; filaments densely clavate-hairy in the upper
half on the ventral, less so on the dorsal side.
Rudiment of ovary glabrous. — 2 Inflorescences
and drupe unknown.
Distr. Malesia: West New Guinea (Hollandia),
once found.
Ecol. Forest, 100 m.
18. Gomphandra dolichocarpa MERR. Pap. Mich.
Ac. Sc. 23 (1938) 183; SLEUM. Blumea 17 (1969)
199. — Stemonurus dolichocarpus (MERR.) Ho-
WARD, J. Arn. Arb. 21 (1940) 468; DAHL, ibid.
33 (1952) 268 (pollen).
1971]
Tree or treelet. Branchlets densely short-hairy
in youngest parts. Leaves oblong, rarely elliptic-
oblong, apex rather suddenly narrow-acuminate
for 1-2 cm, tip bluntish, base cuneate to the pet-
iole, a little inequilateral, firmly chartaceous to
subcoriaceous, olivaceous to brown above, paler
beneath when dry, (12—)16—-25 by 5-8 cm, glabrous
above, hairy at midrib and nerves, glabrous other-
wise beneath, nerves 10-12 pairs, the lowest pair
generally more steeply ascending than the other
ones, all nerves markedly looping before the edge,
generally a little impressed above, raised beneath,
transverse veins finely prominent beneath, no
reticulation; petiole hairy initially, c. 1cm by
1.5-2 mm. — ¢ Inflorescences composed from
several short few-flowered cymes on top of a pe-
duncle (5-8 mm), densely fuscous short-hairy all
over, the petals excepted. Calyx cup-shaped, base
truncate, obscurely 4—S-dented, 1.5mm. Petals
forming anarrow-campanulate tube 4—5-lobed half-
way or partly further down, glabrous, 3.5 mm.
Stamens 4(—5), slightly exserted; filaments with pe-
nicillate hairs in upper half of the ventral, less
densely so on dorsal side. Rudiment of ovary
glabrous. — 2 Inflorescences bearing 3-4 flowers
clustered on top of peduncle (5 mm). Calyx and
petals as in 3 flowers. Staminodes not seen.
Ovary glabrous. Drupes 2 or 3 per infructescence,
known only in submature state, fusiform, 3.54
by 0.5—0.6 cm, the much attenuate distal rostrate
and the basal part each c. 5 mm.
Distr. Malesia: Sumatra (Eastcoast).
Ecol. Apparently in montane forest, no altitude
given by the collectors.
Vern. Kaju atimang, k. barik, k. rawet, M.
19. Gomphandra lancifolia Merr. Philip. J. Sc. 17
(1920) 277; En. Philip. 2 (1923) 490; SLEUM.
Blumea 17 (1969) 199. — Stemonurus lancifolius
(Merr.) Howarp, J. Arn. Arb. 21 (1940) 468.
Tree, c. 6 m, practically glabrous. Leaves nar-
row-lanceolate, apex subcaudate-acuminate, tip
bluntish, base acuminate-attenuate to the petiole,
chartaceous to subcoriaceous, brown when dry,
7-10.5 by (1.5—)2-2.8 cm, nerves 3(-4) pairs, the
lowest pair from c. 5 mm above the base and steep-
ly ascending halfway the blade, upper ones more
curved and shorter, all slightly raised beneath;
petiole c. lcm by 1.5mm. Jnflorescences not
known. Drupe 1 or 2 per infructescence, each
on 5—10 mm long branches, these on a common
robust almost glabrous peduncle (5 mm), obo-
void-oblongoid, 2.5—3 cm by c. 1 cm; fleshy exo-
carp salmon; endocarp with several coarse longi-
tudinal ridges.
Distr. Malesia: Philippines (Luzon: Mt
Dingalang in Tayabas Prov.), once found.
Ecol. Primary forest, 200 m.
Vern. Paranuyog, Neg.
20. Gomphandra luzoniensis (Merr.) Merr. En.
Philip. 2 (1923) 490; Steum. Blumea 17 (1969) 199.
— Elaeocarpus ? integrifolius BLANCO, FI. Filip. ed.
2 (1845) 306; ibid. ed. 3, 2 (1878) 202, non LamK,
1788. — Urandra luzoniensis Merr. Philip. J. Sc.
ICACINACEAE (Sleumer) 31
3 (1908) Bot. 242; Spec. Blanc. (1918) 237. —
Stemonurus luzoniensis (MERR.) HOWARD, J. Arn.
Arb. 21 (1940) 468; Dani, ibid. 33 (1952) 268
(pollen).
Tree, (5—)12-30 m, up to 40cm go. Branchlets
usually reddish-brown in dry specimens, glabrous.
Leaves elliptic to obovate-elliptic, apex + abruptly
acuminate for c. lcm, tip generally bluntish,
base gradually cuneate-acuminate to the petiole,
coriaceous, dark brown and shining above when
dry, practically glabrous with age, 7-15 by 4—6(-8)
cm, nerves c. 5 (rarely —7) pairs, rather straight
in the lower, curved ascending in the distal part,
slightly raised beneath, reticulation almost ob-
solete; petiole strongly grooved above, 1—1.5 cm
by 1.5-2 mm. — ¢ Inflorescences with 3—S cymes
on a common stoutish peduncle (1-3 cm), the
rather numerous flowers + scorpioidly arranged
along the branches (5-10 mm), subdensely set
with rusty appressed hairs on peduncle and
branches. Calyx widely cup-shaped, base truncate,
edge minutely 5-dented (sometimes more deeply
disrupted), subglabrous, 1-1.5mm. Petals 4,
greenish-whitish, glabrous, connate in their lower
24 part, 4-Smm. Stamens 5, finally somewhat
exserted; filaments with long clavate-penicillate
hairs on the inner side below the anther cells and
on the connective. Rudiment of ovary glabrous. —
Q Inflorescences more lax and with fewer flowers.
Calyx and petals as in 3 flowers. Staminodes 5,
less densely hairy than in g flowers. Ovary gla-
brous. Drupe oblongoid to broadly ellipsoid, very
apex a little oblique, otherwise equally rounded
at both ends, 2-2.5 by (0.8—-)1-1.2 cm; exocarp
thin-coriaceous, becoming blackish when dry;
endocarp with numerous longitudinal low ridges
down to the very base, thin-woody.
Distr. Malesia: Philippines (Luzon, Mindoro,
Marinduque, Panay, Palawan).
Ecol. Primary forest under partial shade or
forest edge, at low altitudes, rarely up to 850 m,
locally common.
Vern. Bibislakin, imus, rogrogso, \\k., bitlag,
mabinot, manankdlau, Tag., maratiba, \bn.
21. Gomphandra montana (SCHELLENB.) SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 238; Blumea 17
(1969) 200. — Stemonurus montanus SCHELLENB.
Bot. Jahrb. 58 (1923) 162. — G. carrii SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 239. — G. sp.,
WALKER, For. FI. Br. Solomon Is. (1948) 127.
Shrubby treelet or tree, 3-20 m, up to 25 cm @;
bark smooth, light grey to creamy brown. Branch-
lets appressedly hairy at tips. Leaves elliptic to
lanceolate-oblong, apex short-acuminate to sub-
caudate (2—3cm), tip acutish, base attenuate,
thin-coriaceous, glabrous above, laxly short-
hairy initially and early glabrescent beneath, dark
to olivaceous-brown, dull and often minutely
tubercled above, paler and usually more greenish-
olivaceous beneath in dry specimens, (8—)10—16
(-18) by (3.5-)4—6(rarely -8) cm, nerves 5—6(—7)
rather straight ascending pairs, raised beneath,
no reticulation; petiole 1-1.5cm by c. 2mm. —
3 Inflorescences consisting of 2 or 3(-4) few-
32 FLORA MALESIANA
to many-flowered cymes, together on a rather
stoutish peduncle (l1—-2cm by 1mm), fulvous-
puberulous. Calyx widely cup-shaped, subentire,
mostly laxly hairy, 1-1.5 mm. Petals 5, remaining
connate to a subcampanulate tube for their lower
%, part, greenish-whitish, glabrous (maybe
with a few hairs at apex), (3-)4-5 mm. Stamens
5, exserted for 1-2 mm; filaments set with few to
numerous longish hairs ventrally below the anther
cells, much less or not so dorsally at connective.
Rudiment of ovary glabrous. — 9 Inflorescences
with 3 or 4 one-flowered branches (3-6 mm)
on top of a common peduncle (c. 1 cm). Calyx
and petals as in 3 flowers. Staminodes 5, laxly or
not hairy. Ovary glabrous. Drupe_ subelliptic-
oblongoid, slightly asymmetrical, -+ obtuse at
both ends, base sometimes slightly (or uni-
laterally) swollen, pale yellowish green, sometimes
with reddish or orange tinge, 2—2.5(rarely —3.2) by
(0.9—)1-1.2(-1.5) cm, ribs of endocarp marked
and down to or almost to the very base.
Distr. Melanesia (New Britain, New Ireland,
Solomon Is. and Santa Cruz Group); in Malesia:
New Guinea.
Ecol. Primary lowland and montane, also
swampy, sometimes secondary or devastated
forest, usually in understorey, locally not rare,
from sea-level up to 1675 m.
Uses. Wood creamy to whitish, rays prominent,
no use known, apparently due to its small di-
mensions.
Vern. Aiyalo, Kwara’ae.
Note. Difficult to separate from G. australiana
F. y. M. in leaf and fruit characters, though cer-
tainly different in flower structure.
22. Gomphandra parviflora (BL.) VALET. Crit.
Overz. Olacin. (1886) 218, t. 4, f. 10 & 11; SLEUM.
Blumea 17 (1969) 200. — Stemonurus parviflorus
BL. Mus. Bot. Lugd.-Bat. 1 (1850) 250; WaALpP.
Ann. 2 (1851) 182; Miers, Ann. Mag. Nat. Hist.
II, 10 (1852) 36, repr. Contr. Bot. 1 (1852) 87;
O. KTZzE, Rev. Gen. Pl. 1 (1891) 112. — Lasian-
thera parviflora (BL.) Mia. Fl. Ind. Bat. 1 (1856)
791; Suppl. (1860) 137.
Treelet. Branchlets with minutely pubescent
tips. Leaves oblong to elliptic-oblong, rarely lan-
ceolate or oblanceolate-oblong, apex shortly
rather abruptly acuminate for 1—2cm, base
broadly cuneate and slightly inequilateral, firmly
membranous to subchartaceous, glabrous, green-
ish-greyish to olivaceous and dull in dry speci-
mens, (10—)15—22(-30) by 4-8cm, nerves (8—)
10-12(-14) rather straight and spreading pairs,
-- parallel to each other and looping before the
edge, the lowest pair in part of the leaves (in the
same specimen!) ascending in a more acute angle
from the midrib, reticulation none; petiole often
yellowish as is the midrib beneath in dry speci-
mens, 1-1.3cm by 1.5mm. — ¢ Inflorescences
with 2(—3) few-flowered cymes, these -+_ umbel-
lately arranged on top of a slender peduncle
(c. 5mm), shortly appressedly rusty-pubescent
on peduncle and branches. Calyx cup-shaped, base
truncate, laxly short-hairy, minutely 5-dented,
[ser. I, volyg#
1mm. Petals 5, united to a cup-shaped tube for
their lower 24, glabrous, c. 3mm. Stamens 5,
slightly exserted; filaments practically glabrous
ventrally, but set with penicillate hairs dorsally
at connective. Rudiment of ovary glabrous. —
2 Inflorescences with 2 or 3 shortly pedicellate
flowers on top of a common peduncle (3—5 mm).
Calyx, petals and staminodes not known. Ovary
glabrous. Drupe subfusiform-oblongoid, apex
broadly attenuate and a little oblique, base more
slenderly narrowed, very base cup-like swollen
for c. 1.5mm and smooth, i.e. the rather low
longitudinal ridges of the endocarp ending there,
1.7—2(—2.2) by 0.6 cm.
Distr. Malesia: Central
Singgalang area), twice found.
W. Sumatra (Mt
23. Gomphandra subrostrata Merr. Pap. Mich. Ac.
Sc. 19 (1934) 164, t. 27; SLEUM. Blumea 17 (1969)
200. — Stemonurus subrostratus (MERR.) HOWARD
J. Arn. Arb. 21 (1940) 469.
Treelet or tree, 2.5--7 m. Branchlets rusty- to
rufous-tomentellous at the younger parts, as are
the petiole and midrib and nerves of mature
leaves beneath, older parts glabrate. Leaves ob-
long to elliptic-oblong, apex gradually or more
abruptly acuminate (often slightly falcate) for
1-3 cm, base broadly attenuate to rounded, a
little inequilateral, firmly chartaceous, dark brown
to blackish brown and somewhat shining above,
paler and dull beneath in dry specimens, 15-25
by 4.5-9 cm, nerves 8-10 pairs, rather straight
below, more curved and + obscurely inarching
before edge, raised beneath, veins obsolete;
petiole 5-8(-10) by 2mm. — ¢ Inflorescences
with 2 or 3 + scorpioid several-flowered cymes
together on top of peduncle (0.5—1 cm), peduncle
and branches rusty-tomentellous except the upper-
most part of branches (or pedicels) immediately
below the calyx which is glabrous and black in
dry specimens. Calyx (L6RZING 6690) cup-
shaped, base truncate, sparsely short-hairy in
upper half, ciliate, 4-5-dented, 1.3 mm. Petals 4
or 5 connate about halfway to a narrow-cam-
panulate tube, glabrous, whitish or greenish, 3.5
mm. Stamens 4 or 5, exserted for c. 1 mm;
filaments penicillate both below the anther ventral-
ly and dorsally. Rudiment of ovary glabrous. —
Q Inflorescences with 1 or 2 few-flowered cymes on
top of a common peduncle (c. 5mm). Calyx and
petals as in g flowers. Staminodes not observed.
Ovary glabrous. Submature drupe oblong-ovoid,
apex shortly rather abruptly attenuate or sub-
rostrate (beak erect), base rounded, 10-15 by
5-6 mm; exocarp glaucous, paler on both ends
of drupe; endocarp with very low iongitudinal
ridges.
Distr. Malesia: Sumatra (Eastcoast).
Ecol. Primary forest, rare, 350-850 m.
24. Gomphandra quadrifida (BL.) SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 238; Blumea 17 (1969) 200;
Fl. Thail. 2 (1970) 78. — Stemonurus quadrifidus
Bi. Mus. Bot. Lugd.-Bat. 1 (1850) 249; WALP.
Ann. 2 (1851) 182; Miers, Ann. Mag. Nat. Hist.
1971]
ICACINACEAE (Sleumer) 33
II, 10 (1852) 36, repr. Contr. Bot. 1 (1852) 86. —
Stemonurus prasinus BL. Mus. Bot. Lugd.-Bat. 1
(1850) 249; Warp. Ann. 2 (1851) 182; Mrers,
Ann. Mag. Nat. Hist. IT, 10 (1852) 36, repr. Contr.
Bot. 1 (1852) 86; O. Krze, Rev. Gen. Pl. 1 (1891)
112. — Stemonurus penangianus Mters, Ann.
Mag. Nat. Hist. II, 10 (1852) 39, repr. Contr.
Bot. 1 (1852) 90; Kurz, J. As. Soc. Beng. 44,
ii (1875) 155; For. Fl. Burma 1 (1877) 239;
O. Krze, Rev. Gen. Pl. 1 (1891) 112. — Lasian-
thera javanica (BL.) MIQ. var. quadrifida (BL.)
Mia. Fl. Ind. Bat. 1 (1856) 790. — Lasianthera
prasina (BL.) Mia. /.c. 791; Suppl. 1 (1860) 137.
— Lasianthera ? ovalifolia Mia. Fl. Ind. Bat.
Suppl. 1 (1860) 137, 342; Becc. Malesia 1 (1877)
111. — G. penangiana WALL. Cat. (1832) n. 7204,
nom. nud.; ex Mast. in Hook. f. Fl. Br. Ind. 1
(1875) 587; VALET. Crit. Overz. Olacin. (1886)
223; Kina, J. As. Soc. Beng. 64, ii (1895) 113;
BRANDIS, Ind. Trees (1906) 151; RipL. Fl. Mal.
Pen. 1 (1922) 427; Burk. Dict. (1935) 1096. —
G. prasina (BL.) Becc. Malesia 1 (1877) 110;
VALET. Crit. Overz. Olacin. (1886) 228, incl. var.
ovalifolia (M1Q.) VALET. l.c. 225, t. 4, f. 17, a—b.
— G. oppositifolia PIERRE ex GAGNEP. Not.
Syst. 1 (1910) 198; Fl. Gén. I.-C. 1 (1911) 825.
— G. puberula Rip. J. Fed. Mal. St. Mus. 6
(1915) 142. — G. affinis sensu Rip. Fl. Mal.
Pen. 1 (1922) 427, incl. var. floribunda RIDL.;
Burk. Dict. (1935) 1096; DAHL, J. Arn. Arb. 33
(1952) 268 (pollen). — G. maingayi KING yar.
pubescens Ripi. Fl. Mal. Pen. 1 (1922) 429. —
G. lanceolata (MAstT.) KING var. tenuifolia CRAIB,
Fl. Siam. En. 1 (1926) 273. — Stemonurus op-
positifolius (P1ERRE ex GAGNEP.) HowArbD, J. Arn.
Arb. 21 (1940) 469. — G. scorpioidea GAGNEP.
Not. Syst. 13 (1947) 133; Fl. Gén. I.-C. Suppl.
(1948) 749, f. 89, 7.
See for further synonyms under the varieties.
KEY TO THE VARIETIES
1. Leaves oblong to elliptic-oblong, rarely elliptic
or subovate-oblong, 10—-15(—25) by 4-6(-11)
cm, nerves 6—-8(-10, rarely more) pairs, +
markedly looping before the edge.
1. var. quadrifida
1. Leaves often smaller, nerves (3—)4—5(-—6) pairs.
2. Leaves with a pair of slightly suprabasal,
high curved-ascending ‘melastomaceous’
nerves, other nerves -+- rectangularly from the
midrib faint or obscure. 2. var. triplinervis
2. Leaves with more numerous, straight or more
curved pairs of nerves subparallel to each
other.
3. Leaves narrowly oblong-lanceolate, some-
times almost linear, 5-12(—18) by (0.5—)0.8—2
ee bieounerd Rive 3. var. angustifolia
- 3. Leaves lanceolate to oblong, rarely sub-
ovate-lanceolate, (4—-)5—10(—14) by (1.5-)2—4
cm.
4. Nerves + spreading except the basal pair,
generally obscurely looping before the edge.
4. var. ovalifolia
4. Nerves all steeply ascending, prominent
and markedly looping before the edge.
5. var. maingayi
1. var. quadrifida.
Shrub or treelet, 0.7-Sm. Branchlets often
yellowish in dry specimens, tips only appressedly
short-hairy. Leaves oblong to elliptic-oblong,
rarely elliptic or subovate-oblong, apex sub-
acutely or obtusely and rather abruptly acu-
minate for 1-2 cm, base broadly attenuate, gen-
erally slightly inequilateral, membranous to
chartaceous, green or greenish brownish and
rather dull in the dry state, glabrous, 10—15(—25)
by (3-)4-6(-11) cm, nerves 6—8(—10, rarely more)
pairs, rather straight and subparallel below, mark-
edly looping before the edge, a little raised be-
neath, veins subinconspicuous, reticulation obscure
or generally none; petiole 10(-15) by 1.5 mm. —
3 Inflorescences consisting of 3-6 (rarely more)
shortly peduncled and repeatedly branched +
scorpioid cymes, these subumbellately collected
on top of a rather slender short-hairy peduncle
(1-2 cm), each cyme rarely few-, generally rather
many-flowered, the flowers in the latter case more
distinctly inserted on one side. Calyx widely cup-
shaped, shortly 4~-5-dented, hardly 1 mm. Petals
4(—-5), connate to a tube for the lower 3% part,
white or cream, glabrous, (3—)4(—5) mm. Stamens
4(—5), slightly to more manifestly exserted; fila-
ments densely covered with longish clavate hairs
ventrally below the anther cells and dorsally at
connective. Rudiment of ovary glabrous. — @
Inflorescences similar to the 3 ones, though gen-
erally less branched and few-flowered. Calyx and
petals as in 3 flowers. Staminodes 4(-5), hairy.
Ovary glabrous. Drupe in immature state +
ovoid, its apex markedly attenuate and the base
suddenly contracted into a kind of (very) short
foot, in mature state ellipsoid to subovoid-ellip-
soid, rarely more oblongoid-ellipsoid, apex grad-
ually attenuate or subapiculate, base broadly
narrowed, the very base generally distinctly
swollen and smooth, whilst the upper part of the
endocarp bears several longitudinal ribs, 0.8-1
(-1.2, -1.4) by 0.6-0.7(-0.8) cm.
Distr. Lower Burma, Thailand, and Indo-
China; in Malesia: Sumatra (incl. also Mentawei
Is.) and Malay Peninsula.
Ecol. Primary, also swampy forest, from low-
land up to 900 (in S. Thailand up to 1200) m,
occasionally on limestone, locally common.
Uses. A decoction of the root is used in Ma-
laya after childbirth (BurKILL & Hanirr, Gard.
Bull. S. S. 6, 1930, 184), as is that of var. angusti-
folia.
Vern. Sumatra (Asahan): kaju barik barik, k.
minak minak, k. topu lisak, k. si topu minak;
Malaya: akar taring pélandok, hémpedu jawa, ulan
hitam, M.
2. var. triplinervis (KiNG) Steum. Blumea 17
(1969) 202. — G. lanceolata (MAst.) KiNG var.
triplinervis Kina, J. As. Soc. Beng. 64, ii (1895)
112; Rip. Fl. Mal. Pen. 1 (1922) 428.
34 FLORA MALESIANA
(ser. I, vane
Leaves oblong to elliptic, rarely subovate-ob-
long, 10-15(-18) by 2-4(-6) cm, with a slightly
suprabasal nerve on either side of the midrib,
steeply ascending to almost the top of the blade,
other lateral nerves straight from the midrib,
numerous and subparallel, obscure above, rather
faint beneath.
Distr. Malesia: Sumatra (W. Indragiri), Ma-
lay Peninsula (Kedah incl. also Langkawi, Pe-
nang, Perak, Trengganu).
Ecol. Primary lowland (Dipterocarp) forest, up
to 150 m.
3. var. angustifolia (KING) SLEUM. Blumea 17
(1969) 202; Fl. Thail. 2 (1970) 79. — G. lanceolata
(MAstT.) KING var. angustifolia KING, J. As. Soc.
Beng. 64, ii (1895) 113; Ripi. Fl. Mal. Pen. 1
(1922) 428. — G. salicifolia RiDL. Fl. Mal. Pen. 1
(1922) 429; Burk. Dict. (1935) 1097. — Ste-
monurus salicifolius (RIDL.) HOWARD, J. Arn. Arb.
21 (1940) 469.
Shrub or small tree (rarely up to 20m tall)
with wiry branches. Leaves narrowly (oblong-)
lanceolate, sometimes almost linear, thinly sub-
coriaceous, 5-12 (rarely —18) by (0.5—)0.8-2 cm,
nerves 4-5 pairs rather inconspicuous beneath.
Flowers and drupe apparently slightly smaller than
in var. quadrifida.
Distr. S.Thailand;in Malesia: Malay Peninsula.
Ecol. Lowland and mainly montane forest,
often along riverside, 100-1220 m.
Vern. Daun ekur bukit dérimba, M.
4. var. ovalifolia (RIDL.) SLEUM. Blumea 17 (1969)
203; Fl. Thail. 2 (1970) 79. — G. lanceolata
(MastT.) KiNG var. ovalifolia RiDL. Fl. Mal. Pen.
1 (1922) 428. — Stemonurus affinis Miers, Ann.
Mag. Nat. Hist. II, 10 (1852) 44, repr. Contr.
Bot. 1 (1852) 94, t. 15. — G. affinis (Miers)
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 586. —
Lasianthera lanceolata Mast. I.c. 585. — Ste-
monurus ? tomentellus VALET. Crit. Overz. Olacin.
(1886) 237. — G. lanceolata (MAsT.) KiNG, J. As.
Soc. Beng. 64, ii (1895) 112; Rip. Fl. Mal. Pen. 1
(1922) 428; Burk. Dict. (1935) 1096. — G. pu-
bescens Ripi. Fl. Mal. Pen. 1 (1922) 429. —
G. ophirensis Ri . l.c. 427, nom. illeg.
Small tree or shrub. Leaves lanceolate to oblong-
or subovate-lanceolate, firmly membranous to
subcoriaceous, sometimes sparingly pubescent
beneath, (5—)8-14 by 24cm, nerves (3—)4—5(-6)
+ spreading and generally obscurely looping
pairs. Inflorescence mostly with less numerous
flowers than and not as manifestly scorpioid as in
var. quadrifida, with which it shares the characters
of flowers and fruits.
Distr. Thailand; in Malesia: Malay Peninsula.
Ecol. Primary rain-forest, and also rather dry
Dryobalanops forest, from lowland up to 1200
(-1525) m, locally not rare.
Uses. A decoction of the roots is said to be
administered for rheumatism, and wood-tar, made
from the stems, to be used to blacken teeth.
Vern. Bédara, bua sa puteh, chémperai ayam,
ch. batu, ch. hitam, kayu gérang jantan, k. késturi
hutan, k. ménghilang api, k. mérésék hitam, k.
séréndah, lada lada, lambas, lémpédu tanah jantan,
pokok lilan hitom, sébasah paya, M.
5. var. maingayi (MAST.) SLEUM. Blumea 17 (1969)
203. — Lasianthera maingayi Mast. in Hook. f.
F]. Br. Ind. 1 (1875) 585. — Stemonurus ? maingayi
(MAsT.) VALET. Crit. Overz. Olacin. (1886) 236.
— Urandra maingayi (Mast.) O. Krtze, Rev.
Gen. PI. 1 (1891) 113. — G. maingayi (MAst.)
Kina, J. As. Soc. Beng. 64, ii (1895) 114; RipL.
Fl. Mal. Pen. 1 (1922) 429.
Leaves lanceolate to subovate-lanceolate, apex
subcaudate, subcoriaceous, nerves c. 6 steeply
ascending pairs, shallowly or hardly impressed
above, raised beneath, manifestly looping before
the edge, the veins more visible than in var.
ovalifolia, (4-)5-8 by (1.5—)2-3(-4) cm. Flowers
and drupes apparently matching those of var.
quadrifida.
Distr. Malesia: Malay Peninsula (Penang).
Ecol. Forest on hillsides, 900-1500 m.
25. Gomphandra schoepfiifolia SLEUM. Notizbl.Berl.
— Dahl. 15 (1940) 241; Blumea 17 (1969) 204.
Tree, 7-20m, trunk 10-30cm g. Branchlets
puberulent at tips, becoming blackish in dry state.
Leaves oblong, apex rather abruptly acuminate
for c. lcm, tip bluntish, base cuneate, firmly
chartaceous to subcoriaceous, blackish above,
brownish-blackish beneath, dull and brittle in
dry specimens, 6-9(-10) by 24cm, nerves 5(-6)
pairs, curved-ascending, a little raised beneath,
no reticulation; petiole 6-8 by 1 mm, glabrescent.
— 6 Inflorescences with 3-4(—5) branches (0.5—1
cm) together on top of slender peduncle (1—1.5
cm), each branch bearing a 3-—5-flowered cyme,
densely covered with minute red brown appressed
hairs. Calyx obconical, minutely 5-dented, laxly
hairy, 1.5mm. Petals 5, firm in texture, whitish,
remaining connate in the lower 34 part, glabrous,
c. 4mm. Stamens 5, hardly or not exserted; fila-
ments with rather few longish hairs ventrally
below the anther cells. Rudiment of ovary gla-
brous. — 2 Inflorescences shorter than g ones and
with less numerous flowers. Calyx and petals as in
3 flowers. Staminodes 5, subglabrous. Ovary gla-
brous. Drupe elongate-ellipsoid, known only in
immature state.
Distr. Malesia: New Guinea (Central & Mo-
tobe Distr.).
Ecol. In montane rain-forest, 1500-1950 m.
26. Gomphandra pseudoprasina SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 240; Blumea 17 (1969) 204.
— G. prasina [non (BL.) Becc.] WaARB. Bot.
Jahrb. 13 (1891) 299. — MStemonurus prasinus
(non BL.) K. Sco. & Laut. Fl. Schutzgeb. (1900)
414; SCHELLENB. Bot. Jahrb. 58 (1923) 163.
Tree, 5-10 m. Branchlets practically glabrous.
Leaves oblong, apex shortly acuminate, subacute,
base + broadly cuneate, coriaceous, olivaceous-
brownish and dull in dry specimens, glabrous
except some sparse hairs on midrib and finely
tubercled beneath, 10-17(-22) by 4-6(-8) cm,
nerves 5-6 curved or more straight pairs, obscurely
or not anastomosing, a little raised beneath, gen-
1971]
ICACINACEAE (Sleumer) 35
erally no reticulation visible, though fully
mature leaves show a very fine impressed reticu-
lation above; petiole 0.7—-1 cm by 1.5-2 mm. —
3 Inflorescences composed of several few-flowered
shortly branched cymes together on a stoutish
peduncle, 1.5-2.5cm long in all, covered with
minute rusty hairs. Calyx cup-shaped, 5-denticu-
late, 1.5mm. Petals 5, free only in their upper
part at full anthesis, fleshy, very laxly short-hairy
to practically glabrous, 4mm. Stamens 5, a
little exserted; filaments hairy ventrally below the
anther cells. Rudiment of ovary glabrous. —
° Inflorescences with 3-4 one-flowered branches
(c. 5mm) together on top of stoutish peduncle
(1-1.5 cm). Calyx and petals as in ¢ flowers.
Staminodes 5. Ovary glabrous. Drupe subovoid-
ellipsoid in submature state, apex + abruptly
attenuate and a little curved, almost beaked, base
attenuate, the endocarp slightly ribbed longitudi-
nally, c. 1.9 by 0.9 cm, probably somewhat longer
and more markedly ribbed in fully mature state.
Distr. Malesia: New Guinea (Morobe & Ma-
dang Distr.: Sattelberg and Kani Mts).
Ecol. Montane rain-forest, 600-1500 m.
Imperfectly known
27. Gomphandra oblongifolia MerRrR. Philip. J. Sc.
17 (1920) 276; En. Philip. 2 (1923) 491; SLEuM.
Blumea 17 (1969) 206. — G. sorsogonensis ELM.
Leafi. Philip. Bot. 10 (1939) 3748. — Stemonurus
oblongifolius (MERR.) Howarpb, J. Arn. Arb. 21
(1940) 469.
Tree or treelet, up to 10m. Branchlets pu-
berulent at tips. Leaves oblong to oblong-elliptic,
apex shortly acute to acuminate, base subequally
narrowed, subacute to obtuse, firmly chartaceous
to subcoriaceous, glabrous, brownish, shining and
with flattish tubercles above, paler and dull be-
neath when dry, (6—-)7-15 by (2.7-)3.5-6 cm,
nerves 4—5(—6) pairs, curved-ascending, the upper
ones obscurely anastomosing, slightly raised
beneath, no reticulation; petiole c.1cm by 1.5 mm.
Inflorescences not known. Infructescences spar-
ingly pubescent, with 2 or 3 cymose branches
(0.5—1 cm, each with 1 or 3, rarely 3 fruits) on a
common stoutish peduncle (1-1.5cm). Drupe
oblongoid-ellipsoid to slightly obovoid-ellipsoid,
i.e. the base more attenuate than apex, very base
swollen, 1.5-1.8 by 0.7-0.9cm; endocarp with
low longitudinal ridges which end before the
smooth swollen base.
Distr. Malesia: Philippines (Luzon: Ca-
marines and Sorsogon Proy.; Catanduanes; Min-
danao: Davao).
Ecol. Primary, also Dipterocarp forest at low
altitude.
Vern. Laing, Bik.
Note. Possibly related to G. quadrifida (BL.)
SLEUM.
Excluded
Gomphandra impressa Ripv. Trans. Linn. Soc.
II, Bot. 9 (1916) 29. — Stemonurus impressus
(RIDL.) SCHELLENB. Bot. Jahrb. 58 (1923) 163
= Pittosporum ramiflorum (Z. & M.) ZOLL. ex
MiQ. (Pittosporaceae).
6. RHYTICARYUM
Becc. Malesia 1 (1877) 120, t. 5, f. 18-25 (‘Ryticaryum’); ivid. (1878) 256, corr.:
Rhyticaryum; K. Sco. & Laut. FI. Schutzgeb. (1900) 415 (‘RhAytidocaryum’);
SLeuUM. Blumea 17 (1969) 249. — Pocillaria RiDL. Trans. Linn. Soc. II, Bot. 9
(1916) 28. — Fig. 12.
Dioecious shrubs or trees. Leaves spiral, entire, not rarely with sparse appressed
(strigose) hairs, midrib raised on both faces, nerves +- markedly looping before the
edge. Flowers in axillary spikes, these rarely also composed to panicles in the
same specimen, or very rarely reduced to such extent that the (few) flowers seem
to be fascicled, small, sessile, each subtended by a small bract. Ca/yx cup-shaped,
subentire or 4~5-dented to various degree. Petals 4 or 5, connate 14 to %4 their
length to a ++ open, sometimes campanulate cup, free part or lobes valvate, apex
inflexed. — 3 Flowers: Stamens 4 or 5, free, fixed to the connate part of the tube;
filaments short, anthers basifixed, oblong-elliptic to subsagittate, introrse. Ru-
diment of ovary ovoid or more cylindrical, always hairy. — 9 Flowers: Staminodes
very small, often absent, no anther cells. Ovary conical to subglobose, densely set
with erect hairs; stigma sessile, flattened. Drupe generally ovoid-ellipsoid, much
compressed laterally and almond-like, with a sharp crest, and sometimes another
less marked crest on the flattened side; exocarp fleshy, thin, red to orange; endo-
carp thin-woody, rather coarsely reticulate-alveolate or -lacunose. Seed 1, testa
thin; albumen fleshy; embryo almost of the length of the albumen, with broad
flat cordate cotyledons.
36 FLORA MALESIANA
__[ser. E vol. i
Distr. About 12 spp. in all, in NE. Australia (Queensland: Cape York Peninsula) 1 sp., Melanesia
(Solomons, Bismarcks: New Britain, New Ireland, Admiralty Is.) 1 sp., and East Malesia: New Guinea
and Moluccas, 12 spp. Fig. 11.
Ecol. Primary, rarely also secondary forests in the substage, scattered, from the lowland up to c.
2500 m.
Uses. The leaves are locally cooked and eaten by the people as a vegetable.
Note. Species are rare and still inadequately known and difficult to delimit; several types are lost.
KEY TO THE SPECIES
1. Flowers both in axillary (rather lax-flowered) spikes and Banks composed of such spikes in the
same specimen.
1. R. novoguineense
1. Flowers exclusively in axillary spikes (these sometimes very short i in 2 specimens, the flowers seemingly
fascicled then).
2. Both 3 and Q inflorescences very laxly appressedly hairy, i.e. practically glabrous to the naked eye.
3. Leaves becoming purplish when dried. j Spikes longer than 20 cm.
2. R. purpurascens
3. Leaves becoming brownish or blackish-green, or remaining green-yellowish when dried. g Spikes
usually shorter than 20 cm.
4. Base of leaves properly rounded.
4. Base of leaves attenuate, rarely almost rounded and cuneate in ‘the s same specimen.
5. Drupe 3.5-4.3 by 2.5-3 by c. 1.5 cm.
5. Drupe much smaller, rarely up to 2.6 by il. 5c cm.
3. R. rotundatum
4. R. macrocarpum
6. Leaves blackish-green and hardly reticulate above in the dry state, nerves 6—7 pairs. 2 Spikes
lax-flowered, laxly hairy.
5, Re ‘locidam
6. Leaves greenish-yellowish, or maybe greenish- greyish in "the. dry state, reticulation generally
distinct (at least slightly prominent) above.
7. Flowers laxly arranged, i.e. (much) spaced. Basal pair of nerves curved-ascendent, upper pairs
+ rectangularly from the midrib and straight.
6. R. elegans
7. Flowers (sub)densely arranged. Nerves of leaves all Sh: distinctly curved- ascendent.
8. Leaves membranous to chartaceous, with a very fine and very dense prominent reticulation on
both faces, nerves (10—)12-14 pairs.
7. R. oleraceum
8. Leaves subcoriaceous to thin-coriaceous, reticulation more ‘coarse, nerves 6—8(—10) pairs.
8. R. oxycarpum
2. Both g and Q inflorescences + densely set with brownish substrigose hairs (tomentose in juvenile
state) to the naked eye.
9. 3 Flowers + distant from each other along the rachis.
10. Rachis filiform. Leaves with 6-8 pairs of nerves. .
10. Rachis slender. Leaves with 8-10 pairs of nerves. .
. . 9. R. gracile
10. R. racemosum
9. 3 Flowers close to each other along the + stoutish rachis.
11. Spikes very short (c. 6 mm), the few flowers together almost in a fascicle. Leaves thin.
11. R. fasciculatum
11. Spikes longer to elongate (rachis at least 1 cm) and with numerous flowers. Leaves firm.
1. Rhyticaryum novyoguineense (WARB.) SLEUM.
Blumea 17 (1969) 250. — Buchanania novoguineen-
sis WARB. Bot. Jahrb. 13 (1891) 363. — Rhyticary-
um sp., LAUT. ibid. 56 (1921) 349.
Treelet or small shrub. Branchlets slender, tips
appressedly sparsely hairy, older parts early gla-
brous and covered with greyish cork and bearing
whitish-brownish elliptic prominent _lenticels.
Leaves oblong, apex slenderly cuspidate-acu-
minate (acumen 1.5 by 0.5 cm at base, tip sub-
acute or bluntish, generally slightly curved), char-
taceous, with substrigose hairs here and there on
midrib and nerves beneath, practically glabrous,
dark green when fresh, greyish-green and a little
shining when dry, 9-14(-16) by (2.5-)3-4.5(-5.5)
cm, nerves 8-10(—12) pairs, the intercalar ones
included, slightly raised above, rather sharply
prominent beneath as is the rather coarse re-
ticulation to a minor degree; petiole hairy initially,
12. R. longifolium
8-10(-13) by 1-1.5 mm, transversely lenticelled
with age. f Spikes axillary, both solitary and com-
posed to panicles (each of the pertaining spikes
with a basal subtending bract), pendent, 4-10 cm
in all, lower branches 3—5 cm; rachis + densely set
with yellowish appressed setulose hairs, slender
(0.5mm g), the flowers markedly distant from
each other. — ¢ Flowers (NGF 8386): Calyx
cupular, 1mm, 5-lobed halfway, tips sparsely
strigose. Petals 5, connate halfway, forming an
open cup, pale brownish, 2-2.5mm, lobes +
teflexed. Filaments 1-1.5mm; anthers’ sub-
sagittate, 0.8-1 mm. Rudiment of ovary hirsute.
— Q Flowers unknown. Drupe (ex WARBURG)
ovoid-rhomboid, compressed laterally, 1.7 by 1.5
by 0.5 cm; exocarp fleshy, thin, red.
Distr. Malesia: E. New Guinea (Madang &
Morobe Distr.), thrice found.
Ecol. Primary forest, 600 and 1645 m.
ICACINACEAE (Sleumer)
37
Fig. 11. Distribution of RAyticaryum. The figure
above the hyphen indicates the number of endemic
species in the island or area, the figure below the
hyphen the number of non-endemic species.
2. Rhyticaryum purpurascens SCHELLENB. Bot.
Jahrb. 58 (1923) 172; SLEUM. Blumea 17 (1969)
a
“Shrub, 2-3m. Branchlets glabrous. Leaves
elliptic, apex gradually acuminate for 1.5-2 cm,
tip obtuse, base narrowed to the petiole, perga-
maceous, glabrous, shining above, blackish-green
with pale midrib in fresh, purplish in dry speci-
mens, 15.5—20 by 5-9.5cm, nerves c. 10 pairs,
impressed above, raised beneath, reticulation very
fine; petiole glabrous, 1.5-2 cm. 3 Spikes 1 or 2
per axil, lax-flowered, 20cm long or more, pe-
duncle and rachis minutely and very laxly hairy
(lens!). Flowers yellow when fresh, purplish when
dry, 1.5 mm. Calyx glabrous, ciliolate. Petals con-
nate + halfway, minutely very laxly hairy outside.
Otherwise unknown.”
Distr. Malesia: New Guinea (Sepik Distr.:
Leonhardt Schultze R. area), once found.
Ecol. Alluvial riverine forest at 20-50 m.
Note. Translation of the original Latin descrip-
tion, as the type material is lost.
3. Rhyticaryum rotundatum SCHELLENB. Bot.
Jahrb. 58 (1923) 172; SLeum. Blumea 17 (1969)
251.
“Treelet. Branchlets glabrous or a little hairy.
Leaves oval, apex broadly and gradually acuminate,
tip bluntish, base rounded, thick-pergamaceous,
glabrous, shining, green in dry state above, paler
beneath, 10-16.5 by 4-8cm, nerves c. 6 pairs,
looping, not much prominent, reticulation fine.
3 Spikes solitary, lax-flowered, 3.5 cm; peduncle
(5 mm) and rachis minutely and very laxly hairy
(lens!). g Flowers yellow, 4—S-merous, 1.5 mm.
Calyx and petals obscurely hairy, the latter con-
nate halfway. Otherwise unknown.”
Distr. Malesia: New Guinea (Sepik R. area),
once found.
Ecol. Alluvial riverine forest, 20-30 m.
Note. Translation from the original Latin
description, as the type material is lost.
4. Rhyticaryum macrocarpum BeEcc. Malesia 1
(1878) 256; VALET. Crit. Overz. Olacin. (1886)
257; SCHELLENB. Bot. Jahrb. 58 (1923) 173;
SLEUM. Blumea 17 (1969) 251.
Erect shrub or small tree, up to 13m, up to
18 cm @; bark grey-green, with numerous brown
pustular lenticels. Tips of branchlets obtusangular,
very laxly strigose, older parts terete, glabrous.
Leaves elliptic-oblong, or elliptic, apex rather
abruptly acuminate for 1-1.5 cm, tip blunt, base
narrowed and a little decurrent to the slender
petiole, chartaceous, glabrous, with numerous
pellucid fine points against strong light, brownish
especially beneath when dry, (12—)15—20 by 4—7(-9)
cm, nerves 8-10 pairs, curved-ascending and
anastomosing, slightly raised mainly beneath, re-
ticulation very dense and finely prominent on
both faces; petiole striate lengthwise, glabrous,
1-1.5cm by c. 1mm. — ¢ Spikes (BW 11366)
10-13 cm, rachis c. 1 mm g, glabrous except some
strigose hairs, flowers for their majority distant
from each other. Calyx broadly cup-shaped, 5-
dented, hardly 0.5 mm. Petals 5, fleshy, glabrous
except some strigose hairs outside distally, con-
nate about halfway, c. 2 mm. Stamens 5; filaments
short. Rudiment of ovary hairy. — 2 Spikes very
laxly strigose, (2—)3—7 cm, lax-flowered, rachis
slender, practically glabrous in fruiting time.
2 Flowers to judge from specimen with developed
ovary (NGF 3816): Calyx and petals as in 3
flowers. Ovary subdensely hairy. Drupe almond-
like, a little oblique, broadly attenuate towards
the base, the very base obtuse, apex broadly
attenuate, much compressed laterally, the two
main crests rather sharp, with one or two less
pronounced crests on the flattish sides, 3.5-4.3
by 2.5-3 by c. 1.5 cm; exocarp red or orange red,
practically glabrous; endocarp ligneous, irreg-
ularly shallowly reticulate-costulate outside.
Seed elliptic, compressed, 2.5—3 cm.
Distr. Malesia: New Guinea.
Ecol. Second storey of primary lowland, also
swampy forest, apparently scattered, up to 475 m.
Vern. Paitjerak, Kebar.
5. Rhyticaryum lucidum SCHELLENB. Bot. Jahrb.
58 (1923) 175; Steum. Blumea 17 (1969)
Zo,
“Treelet, 4-5 m, bark greyish. Branchlets laxly
or hardly strigose. Leaves elliptic, apex gradually
attenuate to a bluntish acumen for 1.5-2 cm, base
cuneate, rigidly chartaceous, glabrous, blackish-
green above, paler beneath, shining on both faces,
9-15 by 2.7-5.5cm, nerves 6-7 pairs, curved,
distinctly raised beneath, reticulation obscure
above; petiole 4-10 mm. ¢ Flowers not known.
2 Spikes \ax-flowered, c. 6cm incl. the peduncle
(7mm), rachis laxly hairy (lens!). Flowers yel-
lowish, c. 2mm. Calyx lobes barbulate. Petals
connate less than halfway, hairy outside. Ovary
subglobose, densely set with erect hairs. Drupe
c. 2.3 by 1.5 cm; endocarp tesselate-rugose.”’
38 FLORA MALESIANA
Distr. Malesia: E. New Guinea (Hunstein
Mts), once found.
Ecol. Mossy forest at 1300 m.
Note. Description a translation of the original
Latin diagnosis, as the type specimen is lost.
6. Rhyticaryum elegans SCHELLENB. Bot. Jahrb.
58 (1923) 174; BirNIE, Nova Guinea 14, 2 (1926)
276; DAHL, J. Arn. Arb. 33 (1952) 276 (pollen);
SLEUM. Blumea 17 (1969) 251.
Shrub, 1-3 m. Branchlets slender, tips sparsely
setulose, older parts early covered with greyish
or brownish cork, lenticels elliptic. Leaves ob-
long- or elongate-elliptic, apex (rather) abruptly
narrowed to a slender bluntish or subacute, often
curved acumen (l-3cm by 5-8mm at base),
base cuneate to almost rounded, olivaceous-
brownish when dry, generally shining on both
faces, practically glabrous, 8-16 by 3-—5(—7) cm,
nerves (6—)8—-10 pairs, the lower pair curved-
ascendent, the upper ones + rectangularly from
the midrib and straight, all looping and a little
prominent above, sharply so beneath, reticulation
fine and dense, generally slightly raised on both
sides, sometimes rather obscure above; petiole
(4-)7-10 by 1mm. ¢ & 2 Spikes solitary, rarely
in twos, lax-flowered, (4~-)5—7 (in the 3 ones up
to 14) cm, rachis c. 1 mm g, very sparsely strigose,
or glabrous. — dg Flowers: Calyx low, cup-shaped,
4- or 5-dented, edge strigose, 1 mm. Petals 4 or 5,
fleshy, yellowish, sparsely strigose distally or gla-
brous, connate in the lower 14-34 to a campanu-
late cup, 3-4 mm. Filaments 1 mm; anthers sub-
ovate, 1mm. Rudiment of ovary strigose. —
© Flowers: Calyx and petals as in 3 flowers.
Ovary ovoid-conical, with large stigma, densely
strigose, 2mm. Immature drupe compressedly
ovoid, apex obtuse, laxly strigose, 1.7 by 0.8 by
0.6 cm; endocarp wrinkled.
Distr. Malesia: New Guinea.
Ecol. Montane forest undergrowth, scattered,
800-2300 m.
7. Rhyticaryum oleraceum Becc. Malesia 1 (1877)
121, t. 4, f. 18-21; VALetT. Crit. Overz. Olacin.
(1886) 257; SCHELLENB. Bot. Jahrb. 58 (1923) 173;
SLEUM. Blumea 17 (1969) 251. — R. parviflorum
PULLE, Nova Guinea 8 (1912) 659; SCHELLENB.
Bot. Jahrb. 58 (1923) 173. — R. oncocarpum
(non K. Scn. & Laut.) BirNIE, Nova Guinea 14, 2
(1926) 276.
Shrub or treelet, 0.7-5m. Branchlets laxly
strigose, early glabrescent and covered with cork.
Leaves elliptic or usually lanceolate-elliptic or
oblong-lanceolate, apex rather gradually or more
abruptly and slenderly acuminate for 1-1.5 cm,
tip bluntish, base attenuate or rarely obtuse,
thinly to firmly chartaceous green or yellowish-
green when dry, shining and glabrous on both
faces except some strigose hairs which may occur
on midrib and nerves, 15-25 by 4-7(-10) cm,
nerves (10—)12-14 pairs, curved-ascending, slight-
ly raised above, more distinctly so beneath,
looping, reticulation very fine and very dense,
conspicuously raised on both faces; petiole 0.8-1.5
[ser. I, vol. 7?
cm by c. 1.5mm. — g Spikes slender, rather
dense-flowered, 3—7(—10) cm, laxly substrigose or
glabrescent. Calyx a low cup, hardly lobed, edge
hairy. Petals 4 or 5, connate for a little more than
halfway, fleshy, greenish or yellowish, sparsely
hairy distally outside or glabrous, c. 2mm. Sta-
mens 4 or 5; filaments very short. Rudiment of
ovary hirsute. — 2 Spikes more stoutish, 1-3 cm.
Calyx and petals as in 3 flowers. Staminodes
whether or not present. Ovary ovoid, substrigose,
1.5mm. Drupe obliquely ovoid-ellipsoid, much
compressed laterally, crest bluntish, other ridges
on flat side irregular and less pronounced, orange
red, glabrous, (1.7—)2-—2.3(-2.6) by (1-)1.5 by 1
cm; endocarp coarsely reticulate-lacunose.
Distr. Malesia: Moluccas (Morotai; Sula Is.:
Mangoli I.; Ceram, Batjan, Amboina, Tanimbar
Is.: Jamdena I.; Kai Is.) and W. New Guinea
(Geelvink Bay and Southcoast).
Ecol. Primary lowland forest, rarely montane
forest, up to 1900 m, scattered.
Uses. BECCARI noted the young shoots are
eaten as a vegetable in the Kai Is. and VORDER-
MAN recorded the same for Batjan.
Vern. Dodofé, Morotai (Alfur.), sajor mam-
bos, Batjan.
8. Rhyticaryum oxycarpum K. ScuH. & Laut. FI.
Schutzgeb. (1900) 416; SCHELLENB. Bot. Jahrb.
58 (1923) 175; SLEUM. Blumea 17 (1969) 251. —
R. pulchrum SCHELLENB. Bot. Jahrb. 58 (1923)
173. — R. urophyllum SCHELLENB. /.c. 174.
Shrub or slender tree, 6—-8(—20) m; bark grey
or brownish. Branchlets sparsely strigose. Leaves
elliptic-oblong, sometimes elliptic, apex rather
abruptly acuminate for 1-1.5cm, tip bluntish,
base cuneate, slightly inequilateral, pergamaceous
or firmly chartaceous, greenish-brownish when
dry, glabrous except some scattered hairs on
midrib and nerves underneath, (7.5—)9-15(—20)
by 3.5-5.5(—7) cm, nerves 6—8(—10) pairs, patently
arcuate and ascending (though sometimes rather
straight below), looping, the lowest pair very
close to the edge, prominent beneath, reticulation
dense, usually finely raised on both faces; petiole
8-13 by 1(-1.5)mm. — 3 Spikes (LEDERMANN
7564) dense-flowered, slender, laxly strigose or
subglabrous, up to 18cm. Calyx a low cup, 0.5
mm, its 4 or 5 tooth barbulate. Petals 4 or 5,
connate + halfway, 2mm, lobes laxly strigose
outside. Stamens 4 or 5; filaments short. Rudiment
of ovary hairy. — 2 Spikes (LEDERMANN 10211),
calyx and petals as in 3 flowers. Ovary hairy.
Drupe (LAUTERBACH 2502) ovoid-ellipsoid, much
compressed laterally, 1.8—2.2 by 1.2-1.4 by 0.6-0.7
cm, coarsely reticulate-lacunose.
Distr. Malesia: New Guinea (April and Ramu
R. area).
Ecol. Lowland and montane, also swampy and
secondary forest, on alluvial soil, scattered, up to
1000 m.
9, Rhyticaryum gracile SCHELLENB. Bot. Jahrb. 58
(1923) 170; SLEUM. Blumea 17 (1969) 252.
Divaricate shrub, 1-1.5m. Branchlets very
1971]
ICACINACEAE (Sleumer) 39
slender, tips and petioles subdensely to more
sparsely strigose, older parts early covered with
greyish cork. Leaves elliptic-, or sometimes
slightly obovate-oblong, abruptly narrowed to an
acumen, 1.5—-2 cm by 3-5 mm at base, tip sub-
acute, base cuneate, thin-pergamaceous, green
when dry, a little shining, laxly covered with long
pale strigose hairs along midrib, nerves and maybe
veins on the undersurface of younger leaves,
glabrescent, 9.5—22 by (3.5—)5-7.5 cm, nerves 6-8
pairs, curved-ascending, hardly raised above,
distinctly so beneath, reticulation very dense and
finely raised above, more coarse or less prominent
beneath, petiole 5—7 by 1 mm. 3 Spikes known in
juvenile state only, solitary, the flowers spaced a
little, hirsute all over, rachis filiform, 4cm (or
maybe more in fully developed state). Flowers
5-merous, too young for description. Otherwise
unknown.
Distr. Malesia: New Guinea (Sepik-April R.
area), twice found.
Ecol. Forest at 200 and 1000 m.
10. Rhyticaryum racemosum Becc. Malesia 1
(1877) 121, t. 4, f. 22 & 23; VALET. Crit. Overz.
Olacin. (1886) 257; SCHELLENB. Bot. Jahrb. 58
(1923) 168, as to type; SLEUM. Blumea 17 (1969)
252.
Shrub or treelet, c. 1m, up to 5 cmg. Branch-
lets slender, laxly short-strigose. Leaves oblong
to subobovate-oblong, apex + abruptly acumi-
nate for 1-2 by 0.5 cm at base, tip subacute, base
cuneate and a little decurrent to the petiole,
membranous to thin-pergamaceous, greenish to
yellowish when dry, sparsely setulose at midrib
and part of the nerves beneath, otherwise gla-
brous, somewhat shining on both faces, (12-)
15-24 by (4~-)5-9 cm, nerves 8-10 pairs, curved-
ascending and looping, hardly raised above,
distinct beneath, reticulation rather coarse, finely
raised on both faces; petiole setulose, 1-1.5 cm
by 1.5 mm. —¢ Spikes (PLeyTE 574) slender, sub-
densely strigose, 3—5 cm incl. the short peduncle,
the flowers spaced. Calyx cupular, 0.5 mm, dense-
ly strigose. Petals 5, connate in the lower 7/4,
greenish, laxly strigose at apex outside, 1.5-2 mm.
Rudiment of ovary subcylindric, hairy. — 2 In-
florescence only known in fruiting stage: rachis
subdensely strigose, 3-4(—5) cm, slender (hardly
Imm @), drupes \axly arranged. Remnants of
calyx densely, of petals laxly strigose. Drupe
ovoid-ellipsoid, much compressed laterally, lateral
crests rather sharp, laxly setulose, 1.5 by 1-1.3
by 0.8-I1 cm in dry, said to measure 4 by 2cm
in fresh state; exocarp fleshy, yellow with orange
hue, or reddish.
Distr. Malesia: W. New Guinea (Vogelkop
Peninsula).
Ecol. Primary forest, also along creek in pe-
“yiodically flooded forest dominated by /nocarpus,
from sea-level up to 45 m.
11. Rhyticaryum fasciculatum Becc. Malesia |
(1877) 121, t. 4, f. 24 & 25 (fl. 2); VaALer. Crit.
Overz. Olacin. (1886) 257; SCHELLENB. Bot.
Jahrb. 58 (1923) 168; SLEUM. Blumea 17 (1969)
252.
Shrub, 1-2m. Branchlets slender, youngest
parts densely yellowish-substrigose. Leaves ob-
long to subovate-oblong, apex abruptly acuminate
for 1, rarely up to 2cm, tip rather blunt, base
broadly cuneate, membranous to chartaceous,
laxly to very laxly strigose on midrib and nerves,
rarely also on veins beneath, glabrous otherwise,
green or yellowish when dry, shining on both
faces, 12-24 by 5-9 cm, nerves 8(-10) curved-
ascending and looping pairs, prominent mainly on
the undersurface, reticulation fine and rather
dense, moderately raised on both sides; petiole
strigose, 6-12 by 1.5 mm. — ¢ Inflorescence un-
known. — 2 Flowers 3—5(—7) very close to each
other along a stout c. 6 mm long rachis, seemingly
fascicled, densely strigose in all parts. Calyx cup-
shaped, 1 mm incl. the short teeth. Petals fleshy,
connate almost halfway, 1.8mm. Ovary sub-
conical, densely strigose, 1.5 mm. Drupe + oblique-
ly subovoid-ellipsoid, much flattened laterally
and distinctly crested, laxly strigose, red, (1.3-)
1.7-2 by (1-)1.5 by 0.7-0.9cm; endocarp ir-
regularly coarsely reticulate-alveolate.
Distr. Malesia: W. New Guinea (Vogelkop
Peninsula).
Ecol. Lowland forest, apparently rare.
12. Rhyticaryum longifolium K. ScH. & Laut. FI.
Schutzgeb. (1900) 415; Nachtr. (1905) 306;
Steum. Blumea 17 (1969) 252. — R. oncocarpum
K. Sco. & Laut. Fl. Schutzgeb. (1900) 416;
SCHELLENB. Bot. Jahrb. 58 (1923) 171. — R. ?
oxycarpum (non K. Scu. & Laut.) VALET. Bull.
Dép. Agr. Ind. Néerl. 10 (1907) 30. — Pocillaria
pubescens Ripi. Trans. Linn. Soc. Il, Bot. 9
(1916) 28. — R. bullatum SCHELLENB. Bot. Jahrb.
58 (1923) 169. — R. oblongum SCHELLENB. /.c.
169. — R. elongatum SCHELLENB. l.c. 170. —
R. ovale SCHELLENB. /I.c. 171. — R. racemosum
(non BeEcc.) SCHELLENB. /.c. 168; BIRNIE, Nova
Guinea 14, 2 (1926) 276. — Antidesma megalo-
carpum Sp. Moore, J. Bot. 61 (1923) Suppl. 46.
— R. pubescens (RipL.) SLEUM. Notizbl. Berl.-
Dahl. 15 (1940) 250. — Fig. 12.
Sprawling shrub or small tree, 1-5 (rarely up
to 15) m; bark greyish-brownish. Branchlets with
laxly to subdensely strigose tips. Leaves oblan-
ceolate, or oblong, or obovate-, sometimes elliptic-
oblong, apex rather suddenly acuminate for 1-2
cm by 5 mm at base, tip acute or mostly bluntish,
base cuneate to almost rounded, firmly charta-
ceous to subcoriaceous (more stiff at higher al-
titudes), usually olivaceous-green and somewhat
shining on both faces in the dry state, glabrous
except maybe some scattered appressed hairs
on midrib and nerves of the undersurface, (6.5—)
8-17(-25) by (2.5—)3-7(-10) cm, nerves 6-8(—10)
pairs, curved-ascending and generally markedly
looping, a little raised above, more distinctly so
beneath, not rarely raised within a shallow de-
pression above (the lamina not properly bullate,
as the veins are never sunken above too), re-
ticulation rather dense, considerably or usually
40 FLORA MALESIANA [ser. I, vol. 73
Oy Ss
pee=
Fig. 12. Rhyticaryum longifolium K. Scu. & Laut. a. Habit, ¢ specimen, x 14, b. § flower bud, x20,
c. petal from inside, x 20, d. ¢ more developed flower, x20, e. stamen, x 30, f. habit of 2 specimen with
fruits, x 14, g. fruit, «114 (a-e BW 10666, f-g VAN ROYEN & SLEUMER 6136).
1971]
ICACINACEAE (Sleumer) 41
more slightly raised on both faces; petiole (4~-)6—17
by c. 1.5mm. — ¢ Spikes solitary or sometimes
few from the same axil, rather dense-flowered,
(2-)4-10(-18) cm, in juvenile state all over pale
rusty tomentose, in later stages densely short-
strigose; rachis slender to more stoutish. Calyx
cupular, 5-dented, low. Petals 4 or 5, connate
halfway, yellow or greenish, sparsely to more
densely strigose in the upper half outside, c. 2 mm.
Stamens 4 or 5; filaments short. Rudiment of
overy with erect hairs. — 2 Spikes generally
shorter than the ¢ ones, and the rachis more
stoutish. Calyx and petals as in ¢ flowers. Ovary
ovoid-conical, densely strigose. Drupe ovoid-el-
lipsoid, much compressed laterally and crested,
2-2.3(-2.5) by 1.5(-1.7) by 0.8-1 cm, red to
orange; endocarp hard, coarsely lacunose-reti-
culate.
Distr. NE. Australia (Queensland: Cape York
Peninsula), Melanesia (Solomons); in Malesia:
New Guinea.
Ecol. Substage of primary, sometimes also
secondary forest, from lowland up to 1800
(—2500?) m, scattered.
Uses. The leaves are cooked and eaten by the
natives in the Solomon Is.
Vern. Aicaruar, Wandammen, badzehkiziwo,
Orokaiva, beniros, Wagau, hahamen, Kole, kame
game, Kaugel, kamokum, Enga, namsi, Orne.
In the Solomon Is.: aiavea, Kwara’ae.
7. HARTLEYA
SLEUM. Blumea 17 (1969) 218. — Fig. 13.
Dioecious tree. Leaves spirally arranged, entire, coriaceous, penninerved,
petioled. Inflorescences axillary, 3-5 peduncled cymes arranged to a panicle. —
3 Flowers unknown. — 2 Flowers subsessile, crowded at the apex of the ultimate
branches of the inflorescence; bracts minute, caducous. Calyx cup-shaped, 4-6-
lobed halfway, lobes valvate. Petals 4, valvate, smooth inside. Staminodes 4,
of the habit of apparently normal stamina, but without pollen; filaments flattened,
glabrous; anther cells elliptic, medifixed, the connective between not produced
beyond them. Ovary with a lateral gibbosity; stigma sessile, subcapitate, papillose,
clearly set off from the ovary, subbilobed or depressed to one side. Disk unilateral,
free, opposite the gibbosity of the ovary, a semiorbicular low truncate or sub-
crenate fleshy scale. Drupe curved, ovate-acuminate when seen from one side,
exocarp thin, endocarp with numerous prominent nerves on the concave, and a
thick median rib on the convex side, the latter covered with a fleshy appendage,
the very apex excepted, and with a low semiorbicular fleshy free scale opposite the
appendage, subtended by the remnants of the calyx. Seed not yet known.
Distr. Monotypic; in Malesia: New Guinea.
Ecol. Montane forest.
Note. Closely allied to Gastrolepis TiEGH., a monotypic genus from New Caledonia, from which it
differs by the subcapitate stigma, clearly set off from the ovary (in Gastrolepis rather small and on top of
the gradually attenuate ovary), the large midrib on the concave side of the endocarp (which is not present
in Gastrolepis), the squamular disk which is adnate only to the very base of the ovary, and otherwise free
(adnate at base and both sides such as to form a pocket in Gastrolepis), and the unisexual flowers and gla-
brous filaments (bisexual and with numerous hairs in Gastrolepis).
1. Hartleya inopinata SLeum. Blumea 17 (1969)
218. — Fig. 13.
Tree, 20-35 m, spreading, 60-120 cm @; bark
grey-brown, fissured lengthwise. Branchlets rather
short, irregular, early corticate, 3-8 mm @. Leaves
elliptic or oblong-elliptic, apex short-acuminate, or
obtuse, base cuneate to the petiole, subequal,
toriaceous, brittle and brownish-blackish in dry
specimens, dark shiny green above, yellowish-
green or pale glaucous beneath in fresh state, gla-
brous, 5-8 by 2.5-4.5 cm, edge a little revolute,
midrib slightly impressed above, bold beneath,
nerves 5-6 pairs rather obscure on both faces, no
reticulation; petiole 1 cm by 2 mm. /nflorescences
panicled, branches (c. lcm) rather straight-
horizontal, each ending with a cluster of cymosely
arranged flowers, 3-4cm, the short peduncle in-
cluded, subdensely yellowish-pubescent especially
apically. Flowers subsessile, 2 only known. Calyx
c. 1 mm. Petals oblong, glabrous, yellowish green,
c. 3mm. Stamens seemingly well-developed, but
anther cells without pollen, | mm; filaments gla-
brous, c. 3mm. Ovary ovoid, glabrous, with a
lateral gibbosity, and a fleshy subtruncate semi-
orbicular free scale. Drupe curved, ovate-acu-
minate seen from one side, c. | cm by 4-5 mm;
42 FLORA MALESIANA [ser. 1, vali
Fig. 13. Hartleya inopinata SLEUM. a. Habit, x 14, b. flower bud, 7, c. ditto, with 2 petals removed,
x7, d. developed ovary, showing the vertical fleshy appendix and the basal scaly disk, 5, e. sterile
stamen from adaxial side, e’. ditto, laterally seen, «7, f. submature fruit, adaxial view with large fleshy
appendix, f’. ditto, dorsal view, f’’. ditto, laterally seen, x3 (a-e HARTLEY 12501 (type), f/ HARTLEY
11836).
exocarp thin, becoming blackish; endocarp with Distr. Malesia: New Guinea (Morobe Distr.,
numerous longitudinal prominent nerves on the twice found; possibly present in the Vogelkop
convex, and a strong mediane rib on the concave Peninsula (sterile specimens only known)).
side, the latter almost completely covered with a Ecol. Montane (also mossy) rain-forest,
fleshy flattened appendix, subtended at base by the 1830-2400 m.
lateral scale and the remnants of the calyx.
8. CODIOCARPUS
HowarbD, Brittonia 5 (1943) 53, f. 2, 3 e-f; SLEUM. Blumea 17 (1969) 188. — Fig. 14.
1971] ICACINACEAE (Sleumer) 43
Dioecious tree. Leaves spirally arranged, entire, penninerved. Jnflorescence
axillary, generally solitary, the one-bracteate peduncle with several dichotomously
branched distal branches, each branchlet with a several-flowered compact cyme
of sessile small flowers, which are articulated at the base of the calyx. Calyx
cup-shaped, shortly 5(-6)-dented. Petals 5(-6), valvate, apex minutely inflexed,
shorter in the 2 flower. — 3 Flowers: Stamens 5(—6), exserted; filaments filiform,
glabrous; anthers versatile, the cells oblong, slightly diverging below, dehiscing
introrsally. Rudiment of ovary present. — 2 Flowers: Staminodes 5(—6); filaments
shorter than ovary; anthers much reduced in size, cells empty. Ovary subcylindric
(upper part curved and contracted), with a lateral oblong swelling; stigma disk-
like, obscurely 3-lobed. Drupe oblong, attenuate upwards, flattened, slightly
curved; sarcocarp thin; endocarp thin, hard, with 3-5 (or more) ridges running
the length of the convex surface, and a longitudinal mediane furrow, i.e. the funi-
culus with a pair of woody ribs on either side of the funiculus on the concave
surface, both funiculus and ribs surmounted by an oblong fleshy appendix
extending over the whole length of the fruit, or even surpassing it a little. Seed 1,
with copious endosperm and a minute embryo.
Distr. Two very closely related spp., one in the Andaman and Nicobar Is., the other in Malesia:
Philippines (Mindoro, Palawan). Fig. 15.
Ecol. Lowland primary forest.
1. Codiocarpus' merrittii (MERR.) HOwArD,
Brittonia 5 (1943) 56, f. 2, 3 e-f (wood anat.);
Danu., J. Arn. Arb. 33 (1952) 267, f. 47 & 47 A
(pollen); SLEUM. Blumea 17 (1969) 189. — Ste-
monurus merrittii MERR. Philip. J. Sc. 3 (1908)
Bot. 240. — Tylecarpus merrittii (MERR.) SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 237. — Medu-
santhera merrittii (MERR.) SLEUM. ibid. (1941)
364. — Fig. 14.
Tree, 5—15(-30) m, up to 25cm g, sometimes
fluted. Branches horizontally spreading. Branch-
lets and petioles (1-2cm) shortly appressedly
yellowish-pubescent, glabrate. Leaves oblong-
elliptic to elliptic, apex shortly rather abruptly
acuminate, tip obtuse, base broadly cuneate to
rounded, slightly inequilateral and decurrent on
the petiole, thin-coriaceous to almost charta-
ceous, glabrous above, finely appressedly yel-
lowish-hairy beneath, ultimately glabrate, lower
surface often minutely tuberculate in dry, ap-
parently yellowish-green in fresh specimens, with
numerous fine pellucid points against strong light,
15-20(-28) by 5-10 cm, midrib grooved above,
much raised beneath, nerves 8-10 rather straight
to curved-ascending pairs, obscurely inarching
before the edge, generally prominent beneath;
petiole (1.5—)2—2.5 cm by 2-2.5 mm. Inflorescence
with a rather thick peduncle 1-2.5cm, 2-3-
chotomous, the branches short, thickish, each
with several cymosely and compactly arranged
sessile flowers, all over covered with yellowish-
greyish and + appressed hairs. Calyx c. 1.5mm,
lobes very short. Petals oblong, a little fleshy,
white, hairy outside and at the thickened inflexed
apex inside, otherwise glabrous, 4—5 mm in the 4,
c. 3mm in the & flowers. — ¢ Flowers: Filaments
ce. Smm; anthers c. | mm. Rudiment of ovary
elongate-conical, c. 2 mm. — Q Flowers: Stamino-
des a little shorter than the ovary; filaments weak;
anther cells much reduced in size, empty. Ovary
subcylindrical, glabrous. Drupe oblong, though
narrowed gradually upwards, (1.8—)2-2.5 by
0.6-0.8 cm, greenish-whitish, with 3-5 prominent
longitudinal and several oblique short ribs on the
dorsal convex, and a creamy-white fleshy oblong
appendage on the much deepened ventral surface.
Distr. Malesia: Philippines (Mindoro, Pa-
lawan). Fig. 15.
Fig. 15. Distribution of the species of Medusan-
thera ( ) and Codiocarpus (--—-). 1. M.
gracilis (KING) SLEUM., 2. M. laxiflora (Miers)
Howarp, 3. M. samoensis (REINECKE) HOWARD
and M. ovata Howarp, 4. M. vitiensis SEEM.,
5. C. andamanicus (KURZ) HOWARD, 6. C. merrittii
(Merr.) HOWARD.
Ecol. Dense lowland forest, under partial shade
up to 200 m, on clayey soil or fertile humus. FV. /r.
Jan.—Dec.
Uses. Wood dense and hard, yellowish-white,
used for house posts.
Vern. Palawan: apitong, kalas-kalas, panabd,
tibalao, Tagb.
44 FLORA MALESIANA
[ser. 1, vou 7
Fig. 14. Codiocarpus merrittii (MERR.) HowARD. a. Habit, x 4, b. flower bud, x 714, c. petal inside,
714, d. anther, dorsal side, x10, d’. anther, ventral side, x10, e. stamen, x714, f. ovary, x74,
g. drupe, dorsal side, x1, g’. drupe, lateral view, <1, g’’. drupe, ventral side with fleshy appendage,
1, h. endocarp (exocarp removed) showing the two free ribs on the ventral side, 114, i. drupe in
cross-section, showing the two free ribs of the ventral side covered by the appendage, x 114 (a—f EIMER
12622, g—i ELMER 12842).
1971] ICACINACEAE (Sleumer) 45
9. MEDUSANTHERA
SEEM. J. Bot. 2 (1864) 74: Howarp, J. Arn. Arb. 21 (1940) 469; Lloydia 6 (1943)
133; SLEUM. Blumea 17 (1969) 226. — Tylecarpus ENGL. in E. & P. Nat. Pfl. Fam.
3, 5 (1893) 247; K. Sco. & Laut. Fl. Schutzgeb. (1900) 413 (‘Tilecarpus’); SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 246; in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942)
362, f. 103, A & B. — Fig. 16.
Fig. 16. Medusanthera laxiflora (Miers) Howarp. a. Habit, x 14, b. 3 flower bud, «714, c. stamen,
dorsal view, 10, c’. ditto, ventral view, 10, d. drupe in dry state, dorsal view, showing the prominent
ribs of the endocarp, 214, d’. ditto, ventral view with large appendage, x 21/4, d’’. ditto, cross-section,
showing a low rib on either side of the median depression of the ventral side covered by the appendage,
“x24 (a-c BSIP 61 WALKER & Wuite, d-d’’ BW 3686 KALKMAN).
Dioecious trees. Leaves spirally arranged, subdistichous, entire, penninerved.
Panicles 1-2(-3) per axil, 2-3-chotomized, the branches cymose, peduncled;
pedicels short, articulated with the calyx. Calyx cupular, small, obscurely 5-
toothed. Petals 5, valvate, lanceolate-oblong, apex inflexed. — 3 Flowers: Flower
bud clavate. Stamens 5; filaments flattened and fleshy, rather abruptly narrowed
to the connective, at least the ones of the functional stamens bearing few to
numerous long-clavate hairs dorsally at the connective and ventrally below the
anther; anthers elliptic-oblong, slightly surpassing the connective, the cells di-
verging below. Ovary rudimentary, with a generally rather distinct lateral swelling.
— & Flowers: Flower bud cucullate. Staminodes with filaments reduced in size
and generally glabrous, the anther cells minute, void of pollen. Ovary subcy-
46 FLORA MALESIANA fser. I, you 7
lindrical, early curved, with a distinct lateral swelling; stigma sessile, umbilicate-
subcapitate, lobes 3—5, very short. Drupe oblong to elliptic, rounded-truncate at
both ends, or subovate and attenuate upwards, flattened and curved, the concave
side with a large oblong pulviniform fleshy appendage superficial to the sarco-
carp, the latter rather thin, also filling the groove on the concave side of the tenuous
or almost woody thin endocarp, which bears 1—3(—5) prominent longitudinal
ridges on the convex and maybe two fainter ones on the concave or flat side.
Seed 1; endosperm abundant; embryo apical, minute.
Distr. About 4 or 5 spp., 2 (or 3) in Fiji and Samoa, in Malesia 2 spp., one in Sumatra and Malaya
and one from the Philippines and Celebes to the Carolines, Bismarcks and Solomons. Fig. 15.
Ecol. Primary and secondary lowland (also coastal) and lower montane rain-forest.
KEY TO THE SPECIES
1. Leaves with (4-)5-6 (rarely —8) pairs of lateral nerves. Petals 2-2.5 mm. Drupe subovate-oblong, apex
gradually attenuate and subacute, base obtuse-truncate, 1.2—1.5 by 0.5—0.6(—0.7) cm.
1. M. gracilis
1. Leaves with (6—)8-10(-12) pairs of lateral nerves. Petals 3-4mm. Drupe broad-oblong, obtuse-
truncate at both ends, 1-1.5(-2) by 0.6—-0.8(—-1) cm.
1. Medusanthera gracilis (KING) SLEUM. Blumea 17
(1969) 226; FI. Thail. 2 (1970) 77. — Gomphandra
gracilis KING, J. As. Soc. Beng. 64, ii (1895)
115; Ripi. Fl. Mal. Pen. 1 (1922) 429, incl. var.
gracillima Rip. I.c. 430. — Gomphandra lan-
ceolata (MAST.) KING var. angustifolia KING ap.
RIpL. Fl. Mal. Pen. 1 (1922) 429, p.p. — Gomphan-
dra yatesii MERR. Pap. Mich. Ac. Sc. 19 (1934)
165, t. 28. — Stemonurus yatesii (MERR.) HOWARD,
J. Arn. Arb. 21 (1940) 469; DAHL, ibid. 33 (1952)
268 (pollen).
Shrub or small tree, rarely up to 10 m and 20 cm
@; bark smooth, grey-green. Branches hanging.
Branchlets striately-angular, early corticate, tips
slender and short-pubescent. Leaves lanceolate or
oblong-lanceolate, sometimes oblong, apex sub-
acutely acuminate and often falcate for 2—3 cm,
base much narrowed, membranous, light green,
dull, glabrous or sometimes with sparse fine
short hairs on the undersurface and/or the mid-
rib, showing numerous fine pores against strong
light, the undersurface mostly set with fine
tubercles, (5—)8-13(-20) by (1-)2-4(-8) cm, va-
riable in size, edge subentire, midrib distinctly,
nerves slightly impressed above, prominent be-
neath, nerves (4~-)5-6 (rarely -8) spreading
pairs, reticulation practically obscure; petiole
slender, (0.6—-)1-1.5cm. Peduncles axillary and
pseudoterminal, very slender, 2—-4(—7) cm, short-
hairy as are the partial inflorescences, these cy-
mose, ultimate ones umbellate and few-flowered,
pedicels very short. — g Flowers: Buds obovate.
Calyx cupular, 1 mm. Petals oblong, finally re-
flexed, glabrous, greenish-whitish, 2(—2.5) mm.
— 2 Flowers: Buds subglobose. Calyx and petals
as in g flowers. Ovary oblongoid, 2mm. Drupe
subovate-oblong, flat and curved, base obtuse-
truncate and hollow, apex gradually attenuate and
subacute, 1.2-1.5 by 0.5-0.6(-0.7) cm, glossy,
with a fleshy oblong whitish appendage on the
2. M. laxiflora
Oe ere For OM Om ery ee Poe CO, ay
much depressed ventral, and a marked longitu-
dinal distinct rib parallel with a few lateral less
distinct ones on the dorsal (convex) face.
Distr. §S. Thailand; in Malesia: Sumatra
(Eastcoast, Tapanuli), Malay Peninsula. Fig. 15.
Ecol. Dense lowland jungle or hillside, also
open bamboo forest, rarely up to 1840 m. Fi. fr.
Jan.—Dec.
Vern. Kaju si gorga, Asahan, sampin kris,
Kelantan, M.
2. Medusanthera laxiflora (Miers) Howarp, J.
Arn. Arb. 21 (1940) 470; Lloydia 6 (1943) 142;
Dan, J. Arn. Arb. 33 (1952) 269, f. 53 and 53 A
(pollen); SLEUM. Blumea 17 (1969) 227. — Platea
laxiflora Miers, Ann. Mag. Nat. Hist. IJ, 10 (1852)
111, repr. Contr. Bot. 1 (1852) 98, t. 16; VALET.
Crit. Overz. Olacin. (1886) 255. — Stemonurus
laxiflorus Miers, Ann. Mag. Nat. Hist. II, 10
(1852) 111, repr. Contr. Bot. 1 (1852) 98, nom.
event.; MERR. Philip. J. Sc. 3 (1908) Bot. 240. —
Cissus flexuosa TuRcz. Bull. Soc. Nat. Moscou
31, i (1858) 415; PLANcH. in DC. Monogr. Phan.
5 (1887) 624. — Lasianthera papuana BECC.
Malesia 1 (1877) 108, t. 3; VALET. Crit. Overz.
Olacin. (1886) 240; BorrL. Handl. 1, 1 (1890)
212. — Gomphandra laxiflora (Miers) ROoLrE, J.
Bot. 23 (1885) 211; VipAL, Phan. Cuming. Philip.
(1885) 25, 103; Rev. Pl. Vasc. Filip. (1886) 86;
MerR. En. Philip. 2 (1923) 490. — Tylecarpus
papuanus (BEcC.) ENGL. in E. & P. Nat. Pf.
Fam. 3, 5 (1893) 247, f. 138, A & B; K. Scu. &
Laut. Fl. Schutzgeb. (1900) 413; SCHELLENB.
Bot. Jahrb. 58 (1923) 159; BrrNiE, Nova Guinea
14, 2 (1926) 275. — Tylecarpus sp., PULLE, Nova
Guinea 8 (1912) 658. — Gomphandra glabra
Me_rr. Philip. J. Sc. 17 (1920) 277; En. Philip. 2
(1923) 490. — Gomphandra carolinensis KANEH.
Fl. Micron. (1933) 198 (japon.) f. 85, nom.
seminud.; Bot. Mag. Tokyo 47 (1933) 673; J. Dep.
1971]
ICACINACEAE (Sleumer) 47
Agr. Kyushu Imp. Un. 4 (1935) 358. — Tylecarpus
carolinensis (KANEH.) KANEH. & HAtTus. Bot.
Mag. Tokyo 50 (1936) 605. — M. glabra (MERR.)
Howarp, J. Arn. Arb. 21 (1940) 469, t. 2, f.
8-15; Lloydia 6 (1943) 140, t. 11, f. 1-4; Brittonia
5 (1943) 55, f. 3 a (wood anat.); Dau, J. Arn.
Arb. 33 (1952) 269, f. 55 & 55 A (pollen). — M.
carolinensis (KANEH.) HOWARD, J. Arn. Arb. 21
(1940) 469; Lloydia 6 (1943) 142; WALKER, For.
Fl. Br. Solomon Is. (1948) 127; DAHL, J. Arn.
Arb. 33 (1952) 269, f. 56 & 56 A (pollen). —
M. papuana (Becc.) Howarp, J. Arn. Arb. 21
(1940) 469; SLEuM. in E. & P. Nat. Pfi. Fam. ed.
2, 20b (1942) 363, f. 103, A & B; Howarp,
Lloydia 6 (1943) 137; Dani, J. Arn. Arb. 33
(1952) 269, f. 52 (pollen). — Tylecarpus coriifolius
SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 236. —
Tylecarpus peekelii SLEUM. I.c. 237. — M. coriifolia
(SteUM.) SLeuM. ibid. (1941) 364; Howarb,
Lloydia 6 (1943) 138. — M. peekelii (SLEUM.)
SLEUM. Notizbl. Berl.-Dahl. 15 (1941) 364;
Howarpb, Lloydia 6 (1943) 138. — Fig. 16.
Tree, rarely shrub, (3-)7-15(-27) m, bole
straight, terete, S—25(—30) cm 9, branches spread-
ing + horizontally; buttresses generally absent,
rarely up to 1.2m; bark smooth, with shallow
longitudinal cracks, greenish-grey to brownish.
Branchlets and petioles sparsely appressed-
hirsutulous to glabrate. Leaves elliptic to oblong
or obovate-elliptic, apex generally shortly and +
abruptly acuminate, rarely obtuse, base cuneate
to rounded, thin-coriaceous to chartaceous, dark
green and glossy above, greyish-green below when
fresh, with very numerous round minute pellucid
pores against strong light, lower surface minutely
granular in dried specimens, glabrous, (9—)12-21
by (3—)4.5-8 cm, midrib sulcate above, prominent
and maybe sparsely appressedly hairy beneath,
nerves (6—)8—-10(—12) pairs (with some shorter
additional sometimes between them), rather
straight to curved-ascending, + distinctly anasto-
mosing before the edge, sunken above, slightly
raised beneath, veins rather obscure; petioles
1-1.5(-2) cm. Panicles generally solitary and with
a slender peduncle (0.5—3 cm), 2-3-chotomized,
each branch bearing one to several cymes, all
over + densely appressedly hirsutulous, the calyx
and petals generally excepted. Pedicels slender,
short. Calyx cup-shaped, tube + contracted at
base, 1 mm, limb spreading, 1 mm, minutely 5-
dented. Petals oblong, 3-4 by c. 1 mm, sordid-
white to greenish. — 3 Flowers: Filaments 2-3
mm, persistently clavate-hairy to barbate in the
upper part; anthers c. 1 mm. Rudiment of ovary
conical. — 2 Flowers: Sterile stamens frequently
glabrate, not much smaller than in the ¢ flowers;
cells empty. Ovary subcylindrical, c. 3 mm, curved,
with a basal minute swelling; stigma subcapitate,
with 2-3 minute rounded lobes initially, and
numerous very short lobules in later stages. Drupe
broad-oblong obtuse at both ends, curved, base
hollow, 1-1.5(-2) by 0.6—-0.8(-1) cm, sarcocarp
thin, green to purplish, bearing on the convex
side an oblong fleshy white to pinkish (rarely
yellowish or purplish) pad for almost the entire
length, which is thin in the dry state; endocarp
thin, hard, bearing at the concave side 3(—5)
longitudinal ridges -- sharply prominent in the
dry fruit.
Distr. Micronesia (Carolines: Palau) and Me-
lanesia (Solomons); in Malesia: Philippines (S.
Luzon, Mindoro, Leyte, Samar, Mindanao, Pa-
lawan), Central & SE. Celebes, Moluccas (Hal-
mahera, Obi and Kai Is.), and all over New
Guinea. Fig. 15.
Ecol. Understorey of lowland primary and
secondary rain-forest, also in coastal plain forest
and swamp forest, on sandy and clayey soil, on
limestone and coral, 0—300(-—900) m, once found
in the Western Highlands at 1740 m. Fi. fr. Jan.—
Dec.
Disp. The fruits are reported to be favoured by
cassowary.
Uses. The wood is whitish to yellowish, me-
dium hard and heavy, with coarse prominent
rays, apparently used only as firewood. The
leaves are used orally by natives against malaria
in New Guinea (Tufi Distr.).
Vern. Celebes: tambai, Tobela; Moluccas:
modo, Halmahera; New Guinea: assi, Ramu,
balapi, Dumpu, bieroow, Kebar, bobowé, Mani-
kiong, guarom, sukumisip, Sepik, kené, Biak,
labak, Mooi, mont, Faita, nodag, Bilia, siganapa,
Orokaiwa (Mumuni), sisimal, Amele, surupa,
Baruba, turin, Onjob (Naukwate), wagewageia,
wanigela, Minufia, (Kabubu), wom, Numfur,
wugatep, Sepik (Maprik); New Britain: lamoro;
Solomon Is.: aialo, aikunu, aimaemae, aimalmal,
mae mae, Kwara’ae; Bougainville: diroka, diroma,
guam, kiana, yemollew.
Note. The species is understood here in a wide
sense as the great amount of specimens at hand
effaces differences formerly regarded as sufficient
for the discrimination of species. M. glabra was
based on a specimen with unripe fruits.
10. APODYTES
E. Meyer ex Arn. in Hook. J. Bot. 3 (1840) 155; SLeuM. Blumea 17 (1969) 184.
— Fig. 17.
Trees. Leaves spirally arranged, entire, penninerved, nigrescent. Flowers bi-
sexual, in terminal and axillary corymbs (in Mal.), small, cylindric in bud, artic-
ulate with the short pedicel. Calyx cup-shaped, (4—)5-denticulate, persistent.
Petals (4-)5, free or a little coherent at base, valvate in bud, finally spreading,
48 FLORA MALESIANA (ser. I. van?
Fig. 17. Apodytes dimidiata E. MEYER ex ARN. a. Habit, x 4, b. petal inside, x 624, c. ovary, x 6%,
d. anther, back side, x314, d’. anther, front side, 314, e. drupe, 114 (a—-d KOSTERMANS c.s. 289,
e Bon 4861).
linear, glabrous, obscurely keeled on the inner side, with a shortly inflexed apex.
Stamens 5, almost as long as the petals; filaments subulate, glabrous; anther cells
linear-oblong, sagittate below, introrse, medifixed, dehiscing by lateral slits.
Disk absent. Ovary oblique, with an unilateral swelling; style thick, + excentric,
ending ina small slightly oblique stigma. Drupe + unilaterally developed, oblique-
ly ellipsoid and + compressed, the style finally lateral to almost basal and the
lateral appendage large and succulent; endocarp crustaceous; embryo in the top
of the albumen, small. Seed 1, compressed, ovate-reniform, testa thin.
Distr. Two spp., one in Queensland, one from Malesia through SE. Asia to Africa.
1. Apodytes dimidiata E. MEYER ex ARN. in Hook.
J. Bot. 3 (1840) 155; BENTH. Trans. Linn. Soc. 18
(1841) 680, 683, t. 41, sens. lat.; SLEUM. Blumea 17
(1969) 185 (with synonyms of spp. in S. and SE.
Asia); Fl. Thail. 2 (1970) 81. — A. cambodiana
PreRRE, FI. For. Cochinch. (1892) t. 267, f. A;
GaGnep. FI. Gén. I.-C. 1 (1911) 834; Koorpb.
Atlas 1 (1913) t. 117; Merr. J. Arn. Arb. 6
(1925) 136; Crats, Fl. Siam. En. 1 (1926) 273;
SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 243;
Howarp, J. Arn. Arb. 23 (1942) 73; GAGNEP.
Fl. Gén. I.-C. Suppl. (1948) 756; Amsu. in Back.
Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 5;
Dau, J. Arn. Arb. 33 (1952) 263, f. 36 & 36 A
(pollen); BAcK. & Baku. f. Fl. Java 2 (1965) 61.
— A. javanica K. & V. Bull. Inst. Bot. Btzg 2
(1899) 3; Bijdr. 5 (1900) 159; Koorp. Nat. Tijd.
N. I. 60 (1901) 383; Jungh. Gedenkb. (1910) 175;
BACK. Schoolfl. Java (1911) 226; Koorp. Exk. FI.
Java 2 (1912) 531; Koorp.-ScHum. Syst. Verz. 1,
1971]
Fam. 162 (1912) 5. — Mappia philippinensis
Me_rr. Philip. J. Sc. 26 (1925) 467; En. Philip. 4
(1926) 249. — Neoleretia philippinensis (MERR.)
BAEHNI, Candollea 7 (1936) 180. — Nothapodytes
philippinensis (MERR.) SLEUM. Notizbl. Berl.-Dahl.
15 (1940) 247. — A. sp., ENDERT, Med.
Proefst. Boschw. Ned. Ind. 20 (1928) 211, 216.
— Fig. 17.
Tree, (10—)15—30 m, bole clear, crown high and
lax, trunk (20—)40—70 cm 9; bark rough, dark grey
to brown, thin. Branchlets with sparse oblong
small lenticels. Leaves oblong- to ovate-elliptic,
apex mostly shortly acutely acuminate, rarely
obtuse, base often unequal, acute or unilaterally
obtuse-rounded, a little decurrent on the petiole,
herbaceous to thin-coriaceous, dark green and
shining when fresh, brownish-blackish when dry,
with fine + appressed and crisped yellowish hairs
on midrib above, petioles and inflorescences, other-
wise practically glabrous, entire, (5-)6-13 by
(2.5-)3-6 cm, midrib slightly impressed above,
prominent beneath, nerves 6-10 spreading pairs,
a little raised beneath only, the finest veins mostly
ending with free ends within the network of
finer nerves; petiole 1-2cm. Corymbs many-
flowered, 3—8 cm incl. the 1-3 cm long peduncle,
their branches in a + right angle. Flower buds
oblong. Calyx cup-shaped, almost glabrous, c. 34
mm, teeth short, pubescent outside. Petals ob-
long, white to yellowish, fragrant, 5-6 mm.
Filaments 1.5mm; anther cells oblong, yellow,
3.5mm. Ovary narrow-ovoid, + densely pubes-
cent; style thick, white, 2.5mm. Drupe obliquely
ellipsoid, compressed, veined, first dark purple,
in fully mature state blackish and shining, c. 5 by
9 mm, with a lateral juicy greenish, finally scarlet
appendage, and the lateral style almost at its base.
Distr. Tropical and subtropical NE. to S.
Africa & Angola, Madagascar, Comores, Réunion,
Mauritius, Ceylon, S. India, Annam, Burma,
Thailand, Indo-China, Hainan, Yunnan; in
Malesia: Sumatra (Eastcoast), Malay Peninsula,
ICACINACEAE (Sleumer) 49
Fig. 18. Distribution of Apodytes. A. dimidiata
E. MEYER ex ARN.: approximate area in
Africa and SE. Asia, @ localities in Malesia.
x A. brachystylis F.v.M.
Java (extreme western part), Lesser Sunda Is.
(E. Sumba, Flores, W. Timor), N. Borneo (Mt
Kinabalu, Mt Trusmadi), Philippines (Mindanao:
Bukidnon), Moluccas (Batjan: Mt Sibela). Fig. 18.
Ecol. Primary rain-forest, on steep slopes or
in ravines, edge of stream, on rich maybe volcanic
soil, rare, 800-2000 m. Fi. Febr.—Sept., fr. April—
Nov.
Uses. The leaves are of a bitter and adstringent
taste. Wood light brown or yellowish, used in
Indo-China for cabinet work, described by MOLL
& JANSSONIUS, Mikr. 2 (1908) 244, f. 107 (A.
javanica).
Vern. Ki badak, S, kakomba, Sumba, simarbi-
lalang, Batak.
Note. The species is conceived here in a broad
sense. The special form occurring in Malesia,
described above, is almost matched by forms or
varieties found in Africa and Madagascar.
Pending a revision of the genus, it seems advisable
not to give a proper name on the varietal level to
the Malesian specimens.
11. MERRILLIODENDRON
KANEH. Bot. Mag. Tokyo 48 (1934) 920, f. 7; /.c. 956 (japon.); SLEUM. Blumea 17
(1969) 228. — Peekeliodendron SLeuM. Notizbl. Berl.-Dahl. 13 (1937) 509. —
Fig. 19.
Shrub or generally tree. Leaves spiral, entire, penninerved. Flowers bisexual,
arranged in lax cymes to elongate panicles, these 1-3 in foliate or defoliate axils,
or from older branches resp. from trunk. Pedicels articulated with calyx. Sepals
(3—)4-5, connate to a cup in the lower, free (and not imbricate) in the upper half.
Petals (4-)5, free, valvate, tip inflexed, villous inside, reflexed in anthesis, early
caducous. Stamens 5; filament dilated, short, glabrous; anthers elliptic, cells in-
trorsely and longitudinally dehiscent, exceeded by the connective in form of
subulate glabrous apiculus. Ovary ovoid, tapering to a thick-filiform style;
stigma punctiform. Drupe large; exocarp thin-coriaceous, smooth, early decaying;
endocarp first coriaceous and hard, becoming corky and irregularly + longi-
50 FLORA MALESIANA
[ser. 1, val?
tudinally lacunose in its outer, and woody (not hollowed) in its inner part in later
stages. Seed 1, large; endosperm copious, fibrous, starchy; embryo foliaceous.
Distr. Monotypic, scattered over Micronesia, Melanesia, and E. Malesia. Fig. 20.
1. Merrilliodendron megacarpum (HEMSL.) SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 243; in E. & P.
Nat. Pfl. Fam. ed. 2, 20b (1942) 366, f. 104 (fr.);
BALGooy in Steen. & Balgooy, Pac. Pl. Areas 2
(1966) 176 (distr. map); SLEUM. Blumea 17 (1969)
228. — Stemonurus ? megacarpus HEMSL. in Hook.
Ic. Pl. 24 (June 1895) t. 2398; Kew Bull. (June/
July 1895) 133; K. Scu. & Laur. FI. Schutzgeb.
(1900) 414; SCHELLENB. Bot. Jahrb. 58 (1923) 164.
Fig. 19. Merrilliodendron megacarpum (HEMSL.)
SLEuM. a. Inflorescence, b & b’. infructescence, the
drupes still covered with the thin-leathery exocarp,
c. old weatherbeaten fruit, showing the spongious
endocarp, d. ditto, transverse section, all x14
(a BSIP 2017, 6 BSIP 4827, b’ NGF 27224,
c-d NGF 10925).
— Mangifera xylocarpa Laut. Bot. Jahrb. 56
(1921) 354. — M. rotense KANEH. Bot. Mag.
Tokyo 48 (1934) 920, f. 7; l.c. 956 (japon.);
J. Dep. Agr. Kyushu Imp. Un. 4 (1935) 359;
KANEH. & Hatus. Bot. Mag. Tokyo 54 (1940)
435, f. 82 & 83 (phot., fl. & fr.); Hosokawa, J.
Jap. Bot. 16 (1940) 540; Merr. & Perry, J.
Arn. Arb. 27 (1946) 324; DAHL, ibid. 33 (1952)
265 (pollen). — Peekeliodendron missionariorum
SLEuM. Notizbl. Berl.-Dahl. 13 (1937) 510. —
Fig. 19.
Shrub, or generally tree, (3—)6—15(—25) m,
bole straight, sometimes fluted, up to 55 cm@; bark
green-grey to light brown, smooth, shed in ir-
regular soft flakes; blaze yellow with orange
streaks. Leaves oblong to oblong-ovate, apex
shortly attenuate, mostly blunt, rarely acutish,
base broadly cuneate to rounded, thin-coria-
ceous, glabrous, showing numerous minute pel-
lucid points against strong light, (15—)20-30 by
7-11(—16) cm, in suckers or young shoots up to
36 by 17 cm, entire, midrib strongly obtusely
prominent beneath, nerves (8—)10-12 curved-
ascendent pairs faintly looping before the edge,
veins obliquely transverse, reticulation rather in-
distinct; petiole 1.5-2 cm. Penicles composed of
short lateral cymes, laxly many-flowered, 1-3 in
foliate or already defoliate axils, or from the
wood of old branches and/or trunk, 7—20 cm,
lower branches up to 9 cm, upper ones 2-4 cm;
peduncle 2—5 cm, with a basal swelling, glabrous
or sparsely appressedly and coarsely hairy, slender
in anthesis, stout (2 mm @g) in fruiting stage as is
the rachis. Pedicels slender and 4-6 mm in anthesis,
stoutish and c. 1 cm in fruit. Sepals suborbicular,
connate in the lower half, lobes 1-1.5 mm.
Petals ovate-oblong, a little fleshy, dull cream or
yellow to purplish, 3-4 by 1.5 mm, glabrous out-
side, + densely yellowish-villous inside. Stamens
2-3 mm; filaments c. 0.7 mm; connective hairy,
exceeding the oblong anther cells (1.5mm) by
c. 0.8 mm. Ovary 2.5mm. Drupe pendulous, ob-
longoid to ellipsoid, rarely ovoid-oblongoid, slight-
ly laterally or subquadrangularly compressed,
(4-)6-10 by (2-)3-6 cm, exocarp coriaceous,
whitish or yellow to purplish when fresh and al-
ready fully ripe, becoming purplish to blackish in
older stages or when dry, smooth, thin, early dis-
soluted; endocarp remaining woody and massive
in the inner part (1-2 mm), becoming corky or
spongious and irregularly longitudinally grooved
and/or lacunose-excavated in the outer part (3-5
mm) when exposed to sea-water. Seed starchy,
4-6 by 2-2.5 cm.
Distr. Micronesia (Marianas: Rota & Guam;
Carolines: Kusaie), in Melanesia scattered on
New Ireland, New Britain, the Solomons, and the
Santa Cruz Is. (Vanikoro I.); in Malesia: Philip-
pines (once in Palawan), SE. Celebes (Muna I.),
ICACINACEAE (Sleumer) 51
Fig. 20. Distribution Merrilliodendron megacarpum
(HEMSL.) SLEUM.
and scattered in Northern New Guinea. Fig. 20.
Ecol. Generally coastal and lowland partly
said to occur at 700m), on the inland edge of
mangrove, on sandy or coral beaches, also on
volcanic soil, in moist places, often near streams
or in freshwater swamp forest, locally not rare,
though apparently scattered in general. Fi. /r.
Jan.—Dec.
Disp. The corky or spongious part of the
endocarp gives the fruit its power of buoy-
ancy, and dispersal is obviously effected or
facilitated by sea-currents and _ freshwater
streams.
Uses. Wood yellowish, straight-grained, ap-
parently not used. For wood anatomy cf. SLEUM.
Notizbl. Berl.-Dahl. 13 (1937) 511. Fruit said to
be edible.
Vern. Celebes: pake-saukatibu, Muna; New
Guinea: manggaresi, Japen (Ambai), semna wen,
Sekoli (Kemtuk); New Britain: vabilii; New Ire-
land: idalu, Pala, varunatun, Uquana; Solomon
Is.: abu, aibo, aiebo, aiembuembe, sasa su, Kwa-
ra’ae.
12, CANTLEYA
RIDL. Fl. Mal. Pen. | (1922) 436; Howarp, J. Arn. Arb. 21 (1940) 479, t. 2, f.
1-7; SLEUM. Blumea 17 (1969) 186. — Fig. 21-23.
Tree. Leaves spirally arranged, entire, inconspicuously penninerved. Cymes
axillary and solitary; peduncle completely covered with obtuse small thick spirallly
arranged bracts forming alveoli, from the upper 2 or 3 of which rise short branches,
these articulated with the peduncle and branched dichotomously, each branch
bearing at the end a cluster of 3—5(-8) sessile bisexual flowers. Calyx thick-cu-
pular, shortly subimbricately 5-parted. Petals (4—-)5, valvate, oblong, with a
longitudinal ridge and the apex inflexed inside. Stamens (4—)5; filaments fleshy,
broadened upwards, with a dense clavate-penicillate pubescence distally; anther
cells almost horseshoe-shaped, diverging at the base, introrsely and longitudinally
dehiscent, affixed at the apex. Ovary elongate-conical; stigma a fleshy reversed
cup; ovules 2, pendent from near the apex of the cavity. Fruit fusiform; exocarp
thin, pulpy; endocarp rather thin though firm, fibrous-corky, extending as a thick
longitudinal pad (similar to a second cavity) inside, finally with a lengthwise
split from the base up to the middle. Seed 1.
Distr. Monotypic; in Malesia. Fig. 24.
1. Cantleya corniculata (Becc.) Howarp, J. Arn.
Arb. 21 (1940) 479, t. 2, f. 1-7; DAHL, ibid. 33
(1952) 270, f. 61 & 61 A (pollen); Browne,
For. Tr. Sarawak & Brunei (1955) 199; SmyTuiEs,
Common Sarawak Trees (1965) 71; SLEUM.
Blumea 17 (1969) 186. — Platea corniculata BEcc.
Malesia 1 (1877) 117; VALer. Crit. Overz. Olacin.
(1886) 254. — Urandra sp., Foxw. Philip. J. Sc.
4 (1909) Bot. 492, 542, f. 47. — Urandra cor-
niculata (Becc.) Foxw. Philip. J. Sc. 6 (1911)
Bot. 179; Merr. En. Born. (1921) 356; Megan,
Emp. For. J. 4 (1925) 95; Den Bercer, Hand.
4e N. I. Wet. Congr. Batavia 1926 (1927) 401
(wood anat.); Foxw. J. Mal. Br. R. As. Soc. 5, 2
(1927) 339; Mal. For. Rec. 3 (1927) 173; ibid. 8
(1930) 13; Burx. Dict. (1935) 2205; HeEyne,
Nutt. Pl. 1 (1950) 986. — C. johorica Riv-. FI.
Mal. Pen. 1 (1922) 436. — Stemonurus cornicu-
latus (Becc.) Ripv. ibid. 5 (1925) 297; BuRGEss,
Timbers of Sabah (1966) 320. — Fig. 21-23.
Tree, 15-40 m, trunk straight, slender, terete,
30-60(—150) cm @, expanded at base, buttresses
generally not distinct, rarely up to 150 by 50 cm;
crown high, dense, with numerous small branches;
bark grey or greenish to brownish, sometimes
with brightly coloured patches, rather smooth,
2 FLORA MALESIANA
[ser. I, vane?
Fig. 21. Cantleya corniculata (BECC.) HOWARD. a. Habit, x 14, b. peduncle with bracts, «114, c. petal
inside, <x 10, d. stamen, front side, x 10, d’. ditto, back side, x 10, e. calyx and ovary, * 714 (a-e ENDERT
56 E. 2 P. 664).
finely, shallowly and rather irregularly fissured
lengthwise. Branchlets striate-angular when dry.
Leaves elliptic to subobovate-, rarely almost
oblong-elliptic, shortly rather abruptly sub-
acutely acuminate, base cuneate to rounded,
slightly inequilateral, coriaceous, glabrous, mi-
nutely papillose beneath, drying dark brown, edge
slightly revolute, 5—-9(-13) by 3-4(-5) cm, midrib
narrowly sunken above, strongly raised beneath,
nerves 8-10 pairs, straight and parallel to each
other, rather indistinct; petiole 8-13(-20) mm.
Inflorescence 1—3.5cm; flowers crowded at the
end of short dichotomous branches, these 2 or 3
on top of a 1—2(-3) cm long peduncle which is
covered all over with short obtuse thickish bracts
(each forming an alveole), sessile, small, greenish
to white or yellowish, pale puberulent except the
petals. Calyx c. 1 mm. Petals oblong, green, 4-5
by 2 mm, fragrant to almost fetid and of a bitter
taste. Stamens c. 4mm, finally a little exserted,
caducous as are the petals; filaments 3-4 mm,
glabrous below, upper part penicillate by sub-
clavate white hairs; anther cells c. 1 mm. Ovary
glabrous, 2mm. Fruit fusiform, more attenuate
distally, first green, later brown to blackish, c. 3.5
by (0.6—)0.8-1 cm; exocarp thin, pulpy, of an
acid and adstringent taste; endocarp fibrous-
corky, c. 1 mm, finally splitting from below on
one side up to the middle, extending as a thick
pad for the whole length inside. Seed 1, oblong,
Go 2-em.
Distr. Malesia: Sumatra, Malay Peninsula
(Johore, Singapore), Riouw & Lingga Arch.,
Banka, Borneo (not yet known N of the line
Labuan-Tarakan). Fig. 24.
Ecol. Drier parts of primary freshwater swamp
forest, or drier hill forest, from sea-level to c.
300 m, apparently local and scattered, on marshy
or sandy soil. Fl. fr. Jan.—Dec.
Uses. Timber highly valued, much sought and
marketed, heavy and hard, brownish-yellow, close
grained, fragrant and therefore used as a sub-
stitute for sandal wood, very durable in the ground.
Used for house and ship building, heavy con-
structions, likely to shrink and warp and suitable
for furniture or turnery only if fully seasoned. The
fruit can be eaten, but is said to be of a rather
poor quality. Fig. 22-23.
oS)
1971] ICACINACEAE (Sleumer) 3
M = on Banka known as méndaru; Borneo: kaddo,
Sy mérore, Dajak, kakal, Sampit, pérupok, Iban,
samala, Brunei (Kedayan), séranai tjéndana, Berau.
Fig. 23. Cantleya corniculata (Becc.) HOWARD.
Stem-base. S. Sumatra (Palembang) (Photogr.
ENDERT).
Fig. 22. Cantleya corniculata (Becc.) HOWARD.
Tree-shape, with buttresses. S. Sumatra (Palem-
bang) (Photogr. ENDERT).
Vern. Generally known as bédaru, daru,
dédaru, éndaru, garu, pédaru, tempilai, M; \ocal
names in Sumatra: garu bétina, g. buaja, tarai Fig. 24. Distribution of Cantleya corniculata
pahang, \Indragiri, tulang, Kedongong, tusam; (Becc.) HOWARD.
~
13. NOTHAPODYTES
Bi. Mus. Bot. Lugd.-Bat. | (1850) 248; Howarp, J. Arn. Arb. 23 (1942) 66;
SLEUM. Blumea 17 (1969) 232. — Mappia Jaco. sect. Trichocrater Miers, Ann.
Mag. Nat. Hist. Il, 9 (1852) 395, repr. Contr. Bot. 1 (1852) 64. Mappia Jaca.
54 FLORA MALESIANA [ser. I, vol. 74
sect. Nothapodytes (BL.) O. Krze in Post & Ktze, Lex. Gen. Phan. (1904) 350. —
Neoleretia BAEHNI, C. R. Soc. Phys. Hist. Nat. Geneve 53, 1 (1936) 33; Can-
dollea 7 (1936) 177. — Fig. 25.
Fig. 25. Nothapodytes montana BL. a. Habit, x 14, b. inflorescence, x1, c. flower bud, x5, d. petal, x5,
e. ovary, X714, f. stamen, adaxial side, x 714, f’. ditto, dorsal side, x 714 (a—f KOSTERMANS & KUSWATA
53).
Trees or treelets. Branches usually angular, with large leaf-scars. Leaves spirally
arranged or partly (the upper ones) subopposite, entire, appearing very finely
pellucid-punctate against strong light, penninerved. F/owers bisexual, in terminal,
rarely axillary cymes collected to corymbs, articulate with the short ebracteate
pedicels. Calyx cupular, 5-lobed or sinuate-dented to about 14, persistent. Petals
5, valvate in bud, linear-oblong, a little cohering at base, apex slightly inflexed,
short-strigose without, villose or rarely almost glabrous within. Stamens 5, free,
+ as long as the petals; filaments fleshy, somewhat flattened below, more slender
distally, glabrous; anther cells subovate- to elliptic-oblong, diverging at base, in-
trorse, with a basal abaxial reflexed pulviniform appendage + cohering to the
filament, medifixed; connective as long as the cells. Disk cup-shaped, shallowly
5- or 10-crenulate, fleshy to foliaceous, glabrous without, generally with some
hairs within and on the edge. Ovary symmetrical, hirsute, ovoid to subconical,
tapering to a short stoutish or more slender, slightly elongate, hairy or glabrous
(apparently slightly dimorphic) style; stigma subcapitate, truncate or mostly
1971] ICACINACEAE (Sleumer) 55
oblique, sometimes slightly bilobed or concave. Drupe + ellipsoid, exocarp thin,
juicy; endocarp thin-crustaceous, smooth or rugose. Seed 1, albuminous; embryo
large, almost as long as the albumen.
Distr. Four spp., 3 of which in S. India, Ceylon, Assam, Burma, Thailand, Cambodge, Tonkin, SW. to
SE. China incl. also Hainan, on Lan Yu. I. (= Botel Tobago or Kétyésyo) SE. of Formosa, Ryu Kyu
Is. (Yaeyama group); in Malesia 2 spp., of which N. foetida has a wide distribution in SE. Asia, and N.
montana is very close to N. pittosporoides (OLIv.) SLEUM. in SW. and Central China. Fig. 26.
Ecol. Primary and secondary rain-forest, also mixed deciduous forest, from sea-level to c. 2000 m.
KEY TO THE SPECIES
1. Leaves + unequal at base; petiole (2—)3-6 cm. Petals densely to more sparsely whitish-villose within.
1. N. foetida
1. Leaves + equal at base; petiole 1-2 (rarely —2.5) cm. Petals glabrous or with few hairs in the lower
a ed 8S cn) ree eee 2. N. montana
half within.
43} im5 me ‘@lle.“a” &
1. Nothapodytes foetida (WIGHT) SLEUM. Notizbl.
Berl.-Dahl. 15 (1940) 247; Howarp, J. Arn. Arb.
23 (1942) 70; DaHt, ibid. 33 (1952) 273 (pollen);
SLeuM. Blumea 17 (1969) 232 (with extra-Ma-
lesian synonyms and literature); Fl. Thail. 2
(1970) 82. — Stemonurus foetidus WiGut, Ic. 3
(1845) t. 955. — Neoleretia foetida (WiGHT)
BAEHNI, Candollea 7 (1936) 177, t. 4, f. 2, D-F.
Tree or treelet, (5S—)10-15m, 10-20cm ao.
Branchlets conspicuously angled, early covered
with straw cork (wrinkled in dry specimens), large
leaf-scars, youngest parts laxly to densely covered
with yellowish appressed + substrigose or more
softer hairs as are the petioles, the undersurface of
the leaves, the inflorescences and the fruit, the
tomentum however, variable in density or fading
early. Leaves variably elliptic-ovate to -oblong,
apex short-acuminate, base broadly cuneate to
rounded, unequal, membranous (and subglabrous
in the Philippines), or chartaceous (and tomen-
tulose beneath in Sumatra), 9-22 by 4-12 cm,
nerves (5—)6—10(-12) pairs rather steeply curved-
ascendent, raised beneath, reticulation + ob-
scure; petiole slender to stoutish, (2—)3-6 cm.
Corymbosely arranged cymes long peduncled,
many-flowered, 5-10 cm across, peduncles and axes
+ flattened or angled, all over crisped- to sub-
strigose-hairy. Pedicels 1(-2) mm. Calyx c. 1.5
mm incl. the short subacute lobes. Petals oblong-
linear, subacuminate, white, densely to more
sparsely hairy within, 4—-5(-6) by c. 1.5mm.
Filaments (2-)3-4mm, apparently accrescent
during anthesis; anther-sacs 1-1.3 mm. Ovary
ovoid to pear-shaped, hirsute, gradually narrowed
to the stouter or more slender, short or up to
3mm elongate style, which sometimes is com-
pletely glabrous. Drupe oblong-ovoid or ellipsoid,
1-2 by 0.8-l cm, black, generally short-pubes-
cent; endocarp thin-woody, shallowly rugose to
lacunose-reticulate; endosperm with a fetid odor.
- Distr. S. India, Ceylon, Assam, Burma, Cam-
bodge, Thailand, Lan Yu I. (SE. of Formosa),
Ryu Kyu Is. (Yaeyama group); in Malesia:
W. Sumatra (once on Mt Sago near Pajakumbuh),
Philippines (N. Luzon: Ilocos Norte & Isabela
Prov.). Fig. 26.
40
20
20
Fig. 26. Distribution of Nothapodytes. N. foetida
(unbroken line and dots), N. montana (squares),
N. obtusifolia (...), N. pittosporoides (—.-.-).
Ecol. Rain-forest, on Mt Sago at c. 1000 m,
in the Philippines between 100 and 300m (out-
side Malesia up to 2000m, occurring also in
mixed deciduous forest). F/. Febr.—March, /r.
June (Malesia).
2. Nothapodytes montana BL. Mus. Bot. Lugd.-
Bat. 1 (1850) 248; Steum. Notizbl. Berl.-Dahl. 15
(1940) 247; Howarp, J. Arn. Arb. 23 (1942) 67;
AMSsH. in Back. Bekn. FI. Java (em. ed.) 6 (1948)
fam. 135, p. 6; Back. & Baku. f. FI. Java 2
(1965) 61; SteumM. Blumea 17 (1969) 233; FI.
Thail. 2 (1970) 83. — Mappia montana (BL.)
Miers, Ann. Mag. Nat. Hist. II, 9 (1852) 398,
repr. Contr. Bot. 1 (1852) 67; Mia. Fl. Ind. Bat.
1 (1856) 790; Suppl. | (1860) 136; Becc. Malesia
1 (1877) 118; Vater. Crit. Overz. Olacin. (1886)
183, t. 5, f. 28 (fl.); K. & V. Bijdr. 5 (1900) 143;
Back. Schoolfl. Java (1911) 226; Koorp., Exk.
Fl. Java 2 (1912) 532. — Fig. 25.
56 FLORA MALESIANA
(ser. I, vali 7?
Tree or treelet, S—8(—20) m, 10—15(—20) cm @g;
bark smooth, light brown. Branchlets slender,
obtusangular, early covered with pale brown cork
and wrinkled in dry specimens. Leaves lanceolate
to obovate-oblong, apex obliquely acuminate (1-2
cm), subacute, base cuneate, + equilateral, her-
baceous to chartaceous, shining above, practically
glabrous, often laxly tubercled beneath, 8-16
by 3-6.5cm, midrib raised beneath, nerves in
6-9 pairs, curved or more strict, widely patent,
rather indistinctly looping before the edge, re-
ticulation of veinlets rather dense, finely raised on
both faces; petiole slender, 1-2 (rarely —2.5) cm.
Corymbs rather few-flowered, 2—4 cm across, pe-
duncle slender, 2-4 cm, densely appressedly yel-
lowish hairy in all outer parts. Calyx 1.5-2 mm
incl. lobes. Petals oblong-linear, subacuminate
white, glabrous or with few hairs in the lower half
within, 4-5 by 1.5-2(-3) mm. Filaments linear
below, filiform above, 3—4 mm; anther cells el-
liptic, 1.5mm. Ovary ovoid, gradually tapering
to a shorter or more slender style (c. 2 mm)
which at least in its lower half is appressedly hairy
as is the ovary; stigma oblique. Drupe ellipsoid,
greenish-brown when fresh, blackish when dry,
rather smooth, c. 1.5 by 1 cm.
Distr. Thailand (Prachinburi); in Malesia:
Sumatra (Eastcoast: Karo Highlands), extreme
W. and SW. Java, Lesser Sunda Is. (W. Sum-
bawa, W. Flores). Fig. 26.
Ecol. Primary and secondary forest, almost
from sea-level to c. 1650 m, also on coral lime-
stone. F/. Jan.—Aug., fr. April.
Vern. Kihadji, 8.
distally, finally recurved, a little fleshy, greenish-
14. STEMONURUS
BL. Bijdr. (1826) 648, p.p. lectotyp.; Hassk. Cat. Hort. Bog. (1844) 214 (Stemo-
nura’); BL. Mus. Bot. Lugd.-Bat. 1 (1850) 249, p.p.; em. Becc. Malesia 1 (1877)
108, 109, 111; SLEUM. Blumea 17 (1969) 255. — Urandra Tuw. in Hook. J. Bot.
Kew Misc. 7 (1855) 211. — Lasianthera (non P. Beauv.) Mia. FI. Ind. Bat. 1
(1856) 790, p.p.; KING, J. As. Soc. Beng. 64, 11 (1895) 116. — Fig. 28-31.
Trees, sometimes with buttresses or with pneumatophores, glabrous except a
fine subglandular puberulence on the inflorescences, the petals excluded. Branch-
lets and buds often vernicose. Leaves spiral, entire, coriaceous to various de-
grees, dark green to yellowish green above, paler beneath, sometimes tubercled on
the undersurface, midrib narrowly sunken above, bold beneath, nerves pinnate,
looping; petiole robust. Umbels axillary and generally solitary, peduncled, each
branch of the umbel bearing the flowers either distally and crowded, or uni- or
biseriately (scorpioid) along its upper, mostly + recurved part; bracteoles at the
base of the umbel conspicuous, sometimes persistent. Flowers bisexual, sessile,
white to yellowish, fragrant. Calyx cup-shaped, base + truncate, limb truncate
or 5-lobed to various degree, persistent. Petals (4—)5, valvate in bud, connate at
base only, oblong, apex inflexed, midrib or keel often raised on both faces, ca-
ducous. Stamens (4—)5; filaments fleshy, flat, widened upwards, shortly bearded be-
low the anther on the ventral side, and long-penicillate distally on the backside,
the penicilli inflexed in bud, at full anthesis erect and much exserted; cells ovate-
elliptic, basifixed, a little diverging downwards. Disk rim-like or patellar, or mostly
a short, whether or not shallowly lobed cup, surrounding the very base of the ovary,
the latter ovoid-conical much attenuate towards the very short slender style
and the punctiform stigma, unilocular, with 2 pendant ovules. Drupe ovoid,
ellipsoid or oblongoid-fusiform; exocarp thin, the lower 14 part dark red to purple,
the upper part white to cream or greenish; endocarp firm, coriaceous to woody,
its outer part fibrous. Seed 1; embryo small, in the top of the albumen.
Distr. About 12 spp., 2 of which in Cochinchina and Annam, 1 in Ceylon; in Malesia 9 spp., one of
which extends to the Solomon and Palau Is. Fig. 27.
Ecol. In dry hilly land or in peat swamp forest (part of the species apparently restricted to the latter),
mostly scattered, though widespread in lowland, rarely up to 1750 m.
1971] ICACINACEAE (Sleumer) 57
Disp. The ripe fruits with their fibrous outer and hard inner endocarp are apparently able to float.
This may explain the wide area of distribution of S. ammui (Palau Is., northcoast of New Guinea, New
Britain, Solomon Is.) which occurs both in lowland hillside and in coastal swamp forest, even in the
mangrove.
Uses. Wood aromatic, not durable and no commercial use known in Malesia (ENDERT, Med. Proefst.
Boschw. Ned. Ind. 20, 1928, 198, 199, 201).
Note. The genus comprises two groups of species, one with uni- or biseriately arranged flowers
around S. secundiflorus BL. (W. Malesia incl. Celebes), the other with flowers crowded at the end of the
branches of the umbel around S. umbellata Becc. (W. to E. Malesia, Solomon and Palau Is.). Within
these two groups the species must be distinguished mainly by calyx and fruit characters, as their leaf and
other flower characters overlap to rather great extent.
KEY TO THE SPECIES
1. Flowers uni- or biseriate along the whole or at least the upper (in S. grandifolius thickened) part of the
branches of the umbel, generally 5-12 per branch.
2. Bracts at the base of the umbel persistent for a while during anthesis, (3—)4—-5 by c. 1 mm.
3. Leaves (18—)26-36 by 7—11(—13) cm, very coriaceous, generally a little shining above only in the
dry state, nerves 15-20 pairs generally very iti or hardly raised on both faces, veins + obscure.
Drupe ellipsoid-ovoid, 2—2.4 by 1.2—1.5 cm.
. 1. S. grandifolius
3. Leaves 11—23(—30) by 6—9(-11) cm, coriaceous, generally shining on both “faces in the dry state,
nerves 15-20 pairs slightly though markedly raised on both faces as are the rather lax veins. Drupe
ovoid-ellipsoid, 4.5—5.5(—6.5) by 2—2.5(-3) cm. B43
2. Bracts at the base of the umbel early caducous, 1-2 by c. 0. 5 mm.
2. S. scorpioides
4. Inflorescences robust, peduncle c. 3 mm g. Calyx shallowly lobed or almost entire. Petals 5—6(—7)
mm. Drupe ovoid to oblong-ovoid, 2.5-3(-4) by (1.6-)2cm. .......
3. S. celebicus
4. Inflorescences relatively slender, peduncle 1-2 mm @. Kiba lobed + halfway. Petals c. 4 mm.
Drupe oblong-ellipsoid, 4.5-5.5 by 1-l.5 cm... .
4. S. secundiflorus
1. Flowers clustered at the end of the branches of the umbel or even at the end of the peduncle when the
branches of the umbel are very short. (Bracts at the base of the umbel generally small, fugacious or
rarely subpersistent).
5. Drupe subovoid-oblongoid to almost fusiform, at least 2 times as long as (or mostly longer than)
across. (Leaves generally smooth on the undersurface, maybe except some tubercles found close to
and along the midrib).
6. Calyx rather deeply obtusely lobed. Petals (2.5—)3(-4) mm. Disk cup-shaped. Drupe 4.5—5 by
(1-)1.5—2.2 cm.
5. S. malaccensis
6. Calyx shallowly lobed. Petals (4~)5mm. Disk low, almost patellar. Drupe (4.3—)4.5-5 by 1.5—1.7 cm.
6. S. ammui
5. Drupe ovoid to ellipsoid-ovoid, up to c. 2 times as long as across.
7. Leaves practically smooth on the undersurface; petiole (2—)2.5-3 cm. Calyx rather distinctly lobed.
7. S. gitingensis
7. Leaves generally rather + densely tubercled all over the undersurface or at least in a broad stripe
along the midrib; petiole 1-2 (very rarely up to 3) cm.
8. Calyx + distinctly obtusely lobed. Petals 3(-4, very rarely —4.5) mm.
8. Calyx truncate or very shallowly (apiculately) lobed. Petals (4~)5(—6) mm.
1. Stemonurus grandifolius Becc. Malesia 1 (1877)
114, t. 5, f. 1; VALer. Crit. Overz. Olacin. (1886)
235; Merr. En. Born. (1921) 356; SLeuM. Blumea
17 (1969) 257. — Urandra grandifolia (BECC.)
O. Krze, Rev. Gen. Pl. 1 (1891) 113. — S. doli-
chophyllus Merr. Un. Calif. Publ. Bot. 15 (1929)
171. — Urandra dolichophylla (MerRR.) HOWARD,
J. Arn. Arb. 21 (1940) 468; Dau_, ibid. 33 (1952)
270 (pollen).
Tree, (7-)25-30(-40)m, bole (6—)10—-15m,
10—30(—40) cm @; crown dense; bark greyish to
brownish, smooth or superficially fissured, yellow-
brown mottled. Branchlets robust (5-10 cm 9),
often finely warted and vernicose. Leaves lan-
ceolate or elliptic-oblong, apex very shortly acu-
minate, or merely + abruptly apiculate, base
broadly cuneate to rounded, very coriaceous, stiff,
a little shining above in the dry state, dull beneath,
8. S. monticolus
9. S. umbellatus
(18—)26-36 by 7—11(—13) cm, the extreme margin
slightly and narrowly revolute, nerves 15-20 pairs,
generally very slightly raised on both faces, veins
-+- obscure; petiole rugulose, 1.5-2(—3) by 0.3-0.5
cm. Peduncle of the umbel 3-Scm by 2-3 mm,
with 4-8(—12) shorter or longer (c. 10 mm) robust
and a little flattened branches, which bear 3-4(—5)
flowers crowded at the thickened end of each
branch, or, as normally more (up to 8) flowers
are present, these close together and uni- or
biseriate on the recurved and thickened upper
part of the respective branch. Bracteoles at base of
the umbel broad-subulate, (3—)4—-5 by | mm, +
persistent. Calyx cup-shaped, 2 mm, lobed to 1.
Petals c. 5 by 2 mm, light yellow or white. Drupe
ellipsoid-ovoid, apex gradually attenuate, obtuse,
2-2.4 by 1.2-1.5 cm.
Distr. Malesia: Borneo.
MALESIANA
[ser. I, vol. 7?
Fig. 27. Distribution of Stemonurus. The figure
above the hyphen indicates the number of en-
demic species in the island or area, the figure
below the hyphen the number of non-endemic
species.
Ecol. Primary lowland (Dipterocarp) forest,
on hills or low ridges, not rare on sandstone or
sandy soil, or coastal kerangas, never in peat
swamp, from sea-level-100(-680, a form with
narrower leaves in Tristania forest on Mt Kina-
balu at 1220) m. Fi. fr. Jan.—Dec.
Uses. The pale brown wood is apparently not
used.
Vern. Baru tuas, Bulungan, kambong, Kina-
batangan, kapal-kapal, Kedayan, katak, Dusun,
mersaut, Kutei.
2. Stemonurus scorpioides BeEcc. Malesia 1 (1877)
113, t. 6; VALET. Crit. Overz. Olacin. (1886) 234;
Hocnre. Bull. Inst. Bot. Btzg 22 (1905) 6 (var.}:
Merr. En. Born. (1921) 356 (‘scorpiurus’); RIDL.
Fl. Mal. Pen. 5 (1925) 296; Burk. Dict. (1935)
2074; HEYNE, Nutt. Pl. 1 (1950) 986; ANDERSON,
Gard. Bull. Sing. 20 (1963) 166; SLEUM. Blumea 17
(1969) 257. — Urandra scorpioides (BEcc.) O.
Ktze, Rev. Gen. Pl. 1 (1891) 113 (‘scorpiodes’);
Dau, J. Arn. Arb. 33 (1952) 270, f. 58 & 58 A
(pollen). — Lasianthera secundiflora [non (BL.)
Miq.] Kina, J. As. Soc. Beng. 64, ii (1895) 177,
p.p. — S. secundiflorus (non Bu.) RiDL. Fl. Mal.
Pen. 1 (1922) 430, p.p. — Fig. 28.
Tree 10-30(—35) m, clear bole up to 24m and
15-—30(-60) cm @; bark thin, whitish to greyish
brownish, smooth or pustular. Blunt asparagus-
like pneumatophores sometimes present. Branch-
lets robust, often vernicose as are the buds.
Leaves elliptic-oblong, apex shortly and +
abruptly acuminate, or apiculate, tip bluntish,
base broadly cuneate, coriaceous, stiff, dark green
above, paler beneath, glossy on both faces also
in the dry state, the very margin often slightly re-
volute, 11-23(-30) by 6—-9(—11) cm, nerves 15-20
subregular pairs, slightly though distinctly raised
on both faces as are the rather lax veins, veinlets
finely and prominently reticulate above only;
petiole 1.5-2cm by c. 3mm. Scorpioid cymes
4-6 in an umbel on a robust 3—-5(-6) cm long
axillary peduncle, branches of the umbel 3—5 cm,
each with 6-10 sessile flowers biseriate on each
side; bracts at base of umbel several, subulate,
IST.'69
Fig. 28. Stemonurus scorpioides Becc. Seedling,
a. with the first leaves, b. preceding stage with the
cotyledons still enclosed by the longitudinal
fibres of the endocarp, both x14 (KEP 99938).
rather persistent, (3-)4-5 by 1mm. Calyx cup-
shaped, 1.8mm, obtusely lobed in the upper
spreading 14. Petals (4-)5 by 1.5 mm, white to
cream or greenish. Drupe _ ovoid-ellipsoid,
obliquely attenuate towards the apex, 4.5—5.5(-6.5)
by 2-2.5(-3) cm, whitish above, dark green-
purplish in the lower third, with numerous slight
vertical grooves, able to float when dry.
Distr. Malesia: Sumatra (incl. also Simalur
I.), Lingga and Riouw Arch., Banka, Malay
Peninsula, Borneo.
Ecol. In Shorea albida peat swamp forest in
Borneo, generally in maybe seasonally swampy,
also in secondary vegetation, on peaty or sandy
soil, locally frequent, in lowland up to 45 m.
FI. fr. Jan.—Dec.
Uses. Timber soft, not durable, cracking,
1971]
locally used for planks and household utensils.
Pounded bark used as medicine. The fruit is
said to be edible.
Vern. S. scorpioides has remarkably numerous
native names in the Malay; part of them, how-
ever, are used also for other species of the genus,
and the name bédaru is commonly in use for
Cantleya corniculata (BEcc.) HOWARD outside
North Borneo. — Sumatra: kaju longgaha, k.
pasir, sitobal, Eastcoast, sebungku, siburuk, P.
Bengkalis, sibungkuk, Indragiri, setebal, Djambi,
bahuhu-étém, Simalur, banol, Banka, daru daru
daun lebar, P. Singkep; Malay Peninsula: bungkok,
pérépat paya, Pahang; batang torong, Manggu;
Borneo: ampasir, W. Borneo, bedaru, hantu
burok, katok; medang katuk, m. telor (Benuni);
N. Borneo & Brunei, Sarawak: entaburok, Iban,
jerumit, Milanau, katop, Dusun, semburok; kaju
malam, Kuching, pasir-pasir, Pontianak.
3. Stemonurus celebicus VALET. in Koord. Minah.
(1898) 394; Ic. Bog. 1 (1901) t. 88; Hocnr. Pl.
Bog. Exs. (1904) 45; Bull. Inst. Bot. Btzg 22
(1905) 48; SLEUM. Blumea 17 (1969) 258. —
Urandra celebica (VALET.) HOwARD, J. Arn. Arb.
21 (1940) 468.
Tree, 20-30 m tall, up to 45 cm g. Branchlets
robust, smooth or finely rugulose, vernicose, 5—12
mm 9. Leaves elliptic to obovate-elliptic, apex
broadly subacuminate-attenuate or obtusely apic-
ulate, sometimes rounded, base broadly cu-
neate, coriaceous, a little glossy above, dull be-
neath when dry, (10—)12—-16(-—22) by 5-8(—10.5)
cm, edge somewhat revolute in dry specimens,
nerves 10-14(—16) erecto-patent pairs, slightly or
hardly raised on both faces, veins -+- obscure;
petiole 1.5-2cm by c. 2.5mm. Umbels solitary,
robust, on peduncle 3-5 cm by 3 mm, branchlets
of the umbel (4~-)5-7, more slender, though
thickened distally, (2-)3-4cm, their basal bracts
ovate, obtuse, 2mm, early caducous. Flowers
sometimes only 3 or 4 and crowded at the apex,
generally 5—6(—8), close together and biseriate in
the upper half of each branchlet of the umbel,
secund. Calyx broadly cup-shaped, 2 mm, shal-
lowly lobed or almost entire. Petals 5—6(-—7) by
2.5mm, keeled, white-greenish. Drupe ovoid to
oblong-ovoid, apex gradually attenuate, obtuse,
2.5-3(—4) by (1.6—-)2 cm, darker in the lower 7.
Seed oblong, a little flattened.
Distr. Malesia: NE. and Central Celebes.
Ecol. Primary forest, rarely in swamps, 20-1000
m. Fl. April—July, fr. June—Nov.
Vern. Bilulang, Luwu, palontanggasi, Padu,
simbelan, Minahasa (Tl.), simpaga tanru, Bugin.,
woliausa, Tobela.
4. Stemonurus secundiflorus BL. Bijdr. (1826) 649;
WALP. Rep. | (1842) 378; Hassk. Cat. Hort. Bog.
(1844) 214; BL. Mus. Bot. Lugd.-Bat. 1 (1850)
fig. 45; Miers, Ann. Mag. Nat. Hist. II, 10 (1852)
36, repr. Contr. Bot. 1 (1852) 86; ScHNIZL. Ic.
3 (1866/70) t. 223; Becc. Malesia 1 (1877) 112,
t. 4, f. 16-17; VaLer. Crit. Overz. Olacin. (1886)
234; K. & V. Bijdr. 5 (1900) 145; Hocur. PI. Bog.
ICACINACEAE (Sleumer) 59
Exs. (1904) 46, incl. var. valetonii Hocur.; Bull.
Inst. Bot. Btzg 22 (1905) 41, incl. var. valetonii
Hocur.; Mott & JANss. Mikr. 2 (1908) 234, f.
105 (wood anat.); BAcK. Schoolfl. Java (1911) 226;
Koorp. Exk. Fl. Java 2 (1912) 531; Koorp.-
Scuum. Syst. Verz. 1, Fam. 162 (1912) 3; Koorp.
Atlas 1 (1913) t. 121; Rip. Fl. Mal. Pen. 1 (1922)
430, p.p.; BAKER f. J. Bot. 62 (1924) Suppl. 21;
RIp.. Fl. Mal. Pen. 5 (1925) 296; Burx. Dict.
(1935) 2074; Docr. v. LEEUWEN, Ned. Kruidk.
Arch. 51 (1941) 175, f. 48 (gall); AMsH. in Back.
Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135, p. 5;
Back. & BAKH. f. Fl. Java 2 (1965) 60; SLEUM.
Blumea 17 (1969) 258. — Croton laurifolius Nor.
Verh. Bat. Gen. 5 (1790) ed. I Art. IV, 13, nom.
nud, — Lasianthera secundiflora (BL.) Mia. FI.
Ind. Bat. 1 (1856) 792, incl. var. sumatrana
Mia. /.c. (1858) 1097; Suppl. 1 (1860) 137, incl.
var. sumatrana; ibid. (1861) 342, incl. f. sumatrana
Mia. /.c., t. 2; KING, J. As. Soc. Beng. 64, ii (1895)
177, p.p. — Urandra secundiflora (BL.) O. KTZE,
Rev. Gen. Pl. 1 (1891) 113; ENGL. in E. & P.
Nat. Pfl. Fam. 3, 5 (1893) 248; HeEyNE, Nutt.
Pl. 1 (1950) 987. — S. pauciflorus RipL. Kew
Bull. (1926) 61, non BL. 1826; Howarp, J. Arn.
Arb. 21 (1940) 481, in text. — S. ridleyanus
SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 243.
KEY TO THE VARIETIES
1. Leaves (subcoriaceous to coriaceous) generally
oblong to elliptic- or obovate-oblong, nerves
(8—)10—-12(-14) pairs, -+- distant from each
other, slightly or hardly raised, veins + obscure.
1. var. secundiflorus
1. Leaves with 14-16(—18) pairs of nerves, these
+ close together and generally a little raised on
both faces as are the lax veins.
2. Leaves elliptic, rigidly coriaceous.
2. var. hosei
2. Leaves lanceolate-oblong or -elliptic, co-
TiIaceous: :'/ Ap A ee 3. var. lanceolatus
1. var. secundiflorus.
Tree, 10-24(-32)m, trunk cylindric, up to
40cm 9, sometimes with pneumatophores in
swampy places; bark rather smooth, whitish
grey-green. Branches +- horizontal. Tips of branch-
lets and buds vernicose. Leaves rather variable in
shape and size, oblong-obovate, sometimes ellip-
tic-oblong or oblong, apex obtusely apiculate or
short-acuminate, base acutely or more rarely ob-
tusely cuneate, subcoriaceous to coriaceous,
shining above, much paler beneath, (6—)10-17
(-20) by (3-)4-8.5cm, nerves (8—)10—12(-14)
slender and + distantly subparallel, rather in-
distinctly inarching pairs, slightly or hardly
raised on both faces, veins -- obscure; petiole
1.5-2(-3) cm. Cymes arranged in an solitary
umbel, rarely in two umbels per axil, peduncle
rather slender, c. 3 cm by 1—2 mm, branchlets of
the umbel 3—5, slender, +. recurved, 2—2.5 cm,
bearing in the upper part 3-5(-7) flowers
close together in a row on _ each side;
bracts at base of the umbel ovate or oblong,
60 FLORA MALESIANA
[ser. I, vol. 7?
minute, early caducous, leaving blunt scars. Calyx
cupular, 2—2.5 mm, obtusely lobed almost half-
way. Petals c. 4 by 1.5mm, white or yellowish,
fragrant. Drupe oblongoid-ellipsoid, attenuate
towards the bluntish apex, 4.5—5.5 by 1—-1.5 cm.
Distr. Malesia: Sumatra (incl. also Simalur,
Mentawei & Enggano Is.), Banka, Billiton, Malay
Peninsula, W. Java.
Ecol. Rain-forest, 30—700(—1000) m, sometimes
in peat swamp forest or seasonal swamp forest
near sea-level, scattered. Fi. fr. Jan.—Dec.
Uses. Timber light brown, of inferior quality,
used in places under the house-roof. Medicinal
use for the cure of swellings doubtful. Seeds have
a bitter taste and are edible only after repeated
cooking.
Vern. Sumatra: lokan, Westcoast, sébungku,
Eastcoast, ganggaram, P. Bengkalis, katur, sébonku,
si bénjiet, Indragiri, séhring, Bencoolen, bahuhu
uding, suwahar uding, Simalur, mangguguk,
Mentawei, kaju bémban, Enggano, gélam kataju,
Minangkabau; sabér bubu, Banka, krikis ayér,
Billiton; Malay Peninsula: bérlat, Pahang, M,
kalapa siong, kimeong, sintok lantjang, S.
2. var. hosei SLEUM. Blumea 17 (1969) 259. —
S. labuanensis STAPF, Kew Bull. (1906) 71, p.p.;
Merr. En. Born. (1921) 356, p.p.
Leaves elliptic, very shortly acuminate, stiff-
coriaceous, nerves close together and numerous
(c. 18 pairs), slightly prominent on both faces as
is the net of veins. Inflorescence as in var. lanceo-
latus. Branchlets of the umbel 3 or 4. Fruit not
known.
Distr. Malesia: Borneo (Baram R. area in
Sarawak), once found.
3. var. lanceolatus (BEcc.) SLEUM. Blumea 17
(1969) 259. — S. lanceolatus Becc. Malesia 1
(877): 114. t. S53 fe 24S VATE Crit; Overz.
Olacin. (1886) 235; MerRR. En. Born. (1921) 356.
— Urandra lanceolata (Becc.) O. KTZE, Rev.
Gen. Pl. 1 (1891) 113; Howarp, J. Arn. Arb. 21
(1940) 468, t. 1, f. 1-6. — Urandra ‘secundiflora’,
Dau, J. Arn. Arb. 33 (1952) 270, f. 57 & 57 A
(pollen). — S. umbellatus (non BEcc.) ANDERSON,
Gard. Bull. Sing. 20 (1963) 166.
Tree; bark greyish brown. Leaves lanceolate-
oblong to narrowly elliptic, coriaceous, nerves
14-16 pairs, slightly raised on both faces as is the
lax network of veins. Umbels with 2 or 3 branches.
Drupe similar to that of var. secundiflorus, ob-
longoid, 3.5-4.5 by 1—-1.5 cm, apex attenuate and
slightly curved.
Distr. Malesia: Borneo.
Ecol. Generally mixed swamp or peat swamp
forest, or border of swamp forest with Agathis
forest, low country. Fi. fr. Jan.—Dec.
Vern. Entaburok, Iban, kepot bédjuku, Dajak,
s€mantén, Sampit, énsurok, sémburok, M.
5. Stemonurus malaccensis (MAsT.) SLEUM. No-
tizbl. Berl.-Dahl. 15 (1940) 243; Blumea 17
(1969) 260; Fl. Thail. 2 (1970) 85. — Lasianthera
apicalis (THW.) BENTH. in Mast. in Hook. f.
Fl. Br. Ind. 1 (1875) 584, as to the Labuan plant
only. — Lasianthera malaccensis Mast. in Hook.
f. Fl. Br. Ind. 1 (1875) 584; Becc. Malesia 1
(1883) 257; KiNG, J. As. Soc. Beng. 64, ii (1895)
118. — S. capitatus Becc. Malesia 1 (1877) 114,
t. 5, f. 7-11; VALET. Crit. Overz. Olacin. (1886)
236; MerRR. En. Born. (1921) 355; Rip. Fl. Mal.
Pen. 1 (1922) 431; Crain, Fl. Siam. En. 1 (1926)
272; BurRK. Dict. (1935) 2074. — Urandra ca-
pitata (BECC.) O. KTzE, Rev. Gen. PI. 1 (1891) 113.
— S. labuanensis StaPF, Kew Bull. (1906) 71,
p. p. lectotyp.; MERR. En. Born. (1921) 356, p. p.
— Urandra nitida Howarp, Lloydia 6 (1943) 147,
t. 1, f. 3-6. — Fig. 29.
Treelet or tree, (2.5—)6-15(—20) m, bole up to
10m and 20cm g; bark smooth, brownish-grey,
shallowly lengthwise fissured. Branchlets with
pustules on older parts. Leaves variable in shape
and size, oblong to subobovate-oblong, sometimes
obovate, short-acuminate or obtusely apiculate,
rarely rounded, base cuneate, subcoriaceous to
coriaceous, smooth, i.e. practically not tubercled,
a little shining above, dull beneath both in the
fresh and dry state, (6—)8-12.5(—-14) by (2—)2.5-4
(rarely —7) cm, nerves c. 15 pairs, with other
shorter ones between, rather straight and sub-
parallel, very slightly prominent on both faces,
or not rarely almost obscure beneath, similar to
those of S. secundiflorus var. secundiflorus, re-
ticulation of veins faintly visible above only, or
obscure at all; petiole 1-1.5(—1.8) cm by (1—)1.5-2
(rarely —2.5) mm. Peduncle of the umbel-like in-
florescence generally rather slender (1 mm, rarely
up to 2mm g), 1-2 cm long, with 3—5 branchlets
of the umbel (2—)3-10 mm long, each bearing
(3-)4-6 flowers crowded at the very end, the
flowers seemingly on top of the peduncle when the
branches of the umbel are very short; bracts at the
base of the umbel ovate-oblong or broadly linear,
obtuse, persisting during anthesis only, c. 2 by 2
mm. Calyx cup-shaped, c. 2mm, rather deeply
and obtusely lobed. Petals (2.5—)3(—4) mm, white-
yellowish to dull pink, fragrant. Disk cup-shaped.
Drupe subovoid-oblongoid, attenuate and some-
times a little curved distally, 4.5-5 by (1—)1.5-2.2
cm, pink or pale brown below, merging into pale
green in the upper part.
Distr. Lower Burma, southern Peninsular
Thailand; in Malesia: Malay Peninsula, Borneo.
Ecol. Primary rain-forest, also bamboo forest,
generally in lowland up to 150 m, more rarely on
hillside or ridge tops up to 1370 m, never in peat
swamp forest, apparently on fertile soil, but once
reported from Shorea materialis heath forest in
Trengganu; scattered. Fil. fr. Jan.—Dec.
Uses. Apparently not used, possibly due to
the small size of the trees, contrary to most other
species of the genus, which reach fair dimensions.
Vern. Alau daan, Murut Bokan, gigi buntol,
Selangor, baruas, pasir pasir, Kinabatangan,
péropot, Dusun, pétom, Pahang, tarung pélandok,
Perak, M.
6. Stemonurus ammui (KANEH.) SLEUM. Blumea 17
(1969) 260. — Urandra elliptica SCHELLENB. Bot.
ICACINACEAE (Sleumer) 61
Cc
Fig. 29. Stemonurus malaccensis (MAsT.) SLEUM. a. Habit, x 14, b. flower, x5, c. petal inside, x5,
d. stamen, <5, e. calyx and ovary, x71. f. fruit, x 34 (a N. BORNEO For. Dep. 7685, b-e SAN 30985,
f KEP 76124).
Jahrb. 59 (1924) 17, non MerrR. 1910. — Urandra
ammui KANEH. Bot. Mag. Tokyo 45 (1931) 293;
Fl. Micron. (1933) 199, f. 86; J. Dep. Agr. Kyushu
Imp. Un. 4 (1935) 359. —S. ellipticus (SCHELLENB).
Steum. Notizbl. Berl.-Dahl. 15 (1940) 242.
Tree, 10—24(—33) m, sometimes with buttresses
or slenderly cylindrical pneumatophores, bole
straight, 10-25(-30) cm g; bark smooth or with
fine longitudinal fissures or pustules, whitish-grey
to pale or dark brown. Leaves oblong- to obovate-
elliptic, apex shortly (1-2cm) + abruptly and
obtusely acuminate, sometimes rounded, very
rarely slightly emarginate, coriaceous, shining
above, rather dull beneath in dry specimens,
margin slightly revolute, 9-16(—19) by 4—-8(-9) cm,
smooth on both faces or practically so (a few
tubercles maybe present along the midrib under-
neath), nerves 12—14(-18) slightly curved pairs,
generally a little raised or at least visible above,
much less or hardly so beneath, reticulation of
veins faint and above only; petiole 1.5-2 cm by
ec. 2mm. Peduncles of umbel-like inflorescence
3-5 cm, stoutish, bearing 5-7 (rarely —9) branches,
these 1-1.5 cm long, each with a subulate
subpersistent basal bract c. 1 by 0.5 mm. Calyx
cup-shaped, 2mm, shallowly subacutely lobed.
Petals (4-)5 by 2mm, slightly keeled outside,
yellowish-white, with a strong lemon scent. Disk
low, almost patellar, slightly or hardly lobed.
Drupe oblongoid-subfusiform, much attenuate at
both ends, (4.3-)4.5-5 by 1.5—1.7 cm.
Distr. Micronesia (Palau Is.), Melanesia (New
Britain, Solomons); in Malesia: eastern part of
New Guinea.
Ecol. Primary forest, generally on well-drained
hilly slopes up to 200m, occasionally in fresh-
water swamp forest or mangrove along the coast.
Fl. fr. Jan.—Dec.
Vern. Aikunu, ailalo, aimarako, ainunura, mala
sata, Solomon Is. (Kwara’ae), ammui, Palau Is.
7. Stemonurus gitingensis (ELM.) SLEUM. Blumea 17
(1969) 261. — Urandra gitingensis ELM. Leafl.
Philip. Bot. 4 (1912) 1476. — S. secundiflorus
(non BL.) Merr. En. Philip. 2 (1923) 489.
Tree, 13m or higher, up to 60cm g; bark
thick, yellowish or greyish, brittle, rough with
lenticels. Leaves elliptic or obovately so, apex
rounded, base broadly attenuate and obtuse, or
sometimes rounded, dark green and shining above,
dull and much paler beneath, coriaceous and rather
stiff, smooth, i.e. practically not tubercled beneath,
edge a little revolute, 7-13 by 4-7 cm, nerves
(12—)15-17 (rarely more) pairs, rather straight,
subparallel and close together, slightly raised
above, hardly so beneath, reticulation obscure;
petiole stout, (2-)2.5-3cm. Inflorescence not
known. Infructescence an umbel of 3-4 branches
(c. 1 cm) on a robust peduncle (2-4 cm), bracts
at base of the umbel according to the scars small
and apparently early caducous. Flowers, as seen
from the left scars, 2-4 clustered at the end of
each branch of the umbel, certainly not in a row
as in S. secundiflora. Calyx on immature fruit
62 FLORA MALESIANA
cup-shaped, 2 mm, rather distinctly lobed. Drupe
ovoid-ellipsoid, short-apiculate, 3.2-3.7 by 2-2.3
cm. Seed said to be surrounded with a soft white
sourish tasting layer.
Distr. Malesia: Philippines (Sibuyan: Mt Gi-
tinggiting; after MERRILL also on Panay).
Ecol. Primary forest at 750-1750m. Fr.
March-April.
Vern. Tugbak, P. Bis.
8. Stemonurus monticolus (SCHELLENB.) SLEUM.
Notizbl. Berl.-Dahl. 15 (1940) 243 (‘monticola’);
Blumea 17 (1969) 261. — ? Urandra scorpioides
[non (Becc.) O. KTzE] PULLE, Nova Guinea 8
(1912) 659; SCHELLENB. Bot. Jahrb. 58 (1923) 165.
— Urandra monticola SCHELLENB. Bot. Jahrb. 58
(1923) 164. — Urandra umbellata (non BECC.)
PULLE, Nova Guinea 8 (1912) 658; SCHELLENB.
Bot. Jahrb. 58 (1923) 165. — Urandra brassii
Howarp, Lloydia 6 (1943) 146, t. 1, f. 1 & 2;
DauL, J. Arn. Arb. 33 (1952) 270 (pollen).
Tree, 10—-30(—40) m, bole up to 18 m and 10-30
(-60) cm @; bark grey to pale brown, smooth,
maybe with a few longitudinal fissures, a little
or not peeling; buttresses sometimes present, up
to 1.5 by 1 by 0.1 m. Leaves elliptic to obovate-,
sometimes lanceolate- or oblong-elliptic, apex
shortly obtusely acuminate, or mostly blunt, some-
times rounded and maybe a little emarginate,
base + broadly attenuate, coriaceous, dark green
to yellowish green above, paler beneath, generally
rather dull on both faces, smooth above, + dense-
ly tubercled all over the undersurface, or at least
so in a broad stripe along the midrib, very rarely
smooth beneath, the edge a little recurved in dry
specimens, (5—)6-10(-18) by (2.5—-)3-6(-9) cm,
nerves (12—)14-16 rather straight pairs, looping
before the edge, very slightly or hardly raised or
visible above and/or beneath, generally obscure
beneath; petiole (1—)1.5—2 cm by 1-2 mm. Umbel-
like inflorescences peduncled (2-4, rarely -4 cm
by 1-1.5 mm), branches 4-6 rather slender, each
bearing 3-4(—5) flowers crowded distally; bracts
at the base of the umbel ovate-subacuminate, 1-2
by 1mm, subpersistent during anthesis. Calyx
cup-shaped, 2mm, + distinctly obtusely lobed.
Petals 3(-4, rarely —4.5) by 1.5 mm. Disk a flattish
slightly 5-lobed cup. Drupe subellipsoid-ovoid,
3-3.5 by 1.6cm, apex subabruptly attenuate or
apiculate.
Distr. Malesia: New Guinea.
Ecol. Primary (and occasionally also seconda-
ry) rain-forest or forest substage, on slopes,
from sea-level to 850(-1000)m, on clayey or
sandy soil, sometimes on limestone with clay
cover, locally common. Fi. fr. Jan.—Dec.
Uses. Wood yellowish or light orange, appar-
ently not used. Scrapings of bark heated in hot
ashes are used for wounds in the Vogelkop
Peninsula.
Vern. Béram, bramsos, €mpam, keferan maper,
manséran, sandawer, sapuduper, Biak, duwili,
Kapauku, harihoh, korappi, korowapi, ukipi,
wapapi, Japen, karmuk, karmu-u, lakobiliem,
lobak, Mooi, kuwat, Amberbaken, nikoboa, Kebar,
(ser. I, vol
permeh, pirmir, Asmat, sakokohom, sakokom,
Manikiong, fafus, Berik, tifus, Mander, weijan,
Meosnum (Wanap)).
9. Stemonurus umbellatus Becc. Malesia | (1877)
115, t. 5, f. 5 & 6, incl. var. ovalifolius Becc. I.c.
t. 4, f. 14 & 15; VALET. Crit. Overz. Olacin. (1886)
235; Ript. Fl. Mal. Pen: 1 (1922) e4siherian::
Burk. Dict. (1935) 2074, p.p.; SLEUM. Blumea 17
(1969) 262. — Urandra umbellata (Becc.) O.
KTzE, Rev. Gen. Pl. 1 (1891) 113; DAHL, J. Arn.
Arb. 33 (1952) 270, f. 59 & 59 A (pollen). —
Lasianthera umbellata (BEcc.) KING, J. As. Soc.
Beng. 64, ii (1895) 117, excl. specim. cit. (= S.
malaccensis); MERR. En. Born. (1921) 355, incl.
var. ovalifolia (BECC.) MERR. — S. evenius STAPF,
Kew Bull. (1906) 71; Merr. En. Born. (1921)
355. — Urandra hallieri MeRR. Philip. J. Se. 5
(1910) Bot. 194; DAHL, J. Arn. Arb. 33 (1952) 270
(pollen). — SS. hallieri (MerR.) Merr. En.
Philip. 2 (1923) 489. — Urandra evenia (STAPF)
Howarp, J. Arn. Arb. 21 (1940) 468. — S. in-
tercedens HEINE in Fedde, Rep. 54 (1951) 237;
Pfl. d. Samml. Clemens Kinabalu (1953) 57, 114.
— Fig. 30-31.
Tree, (8-)12—-24(-37) m, (15-)20-35cm 4g;
bark smooth or finely cracked or hooped, some-
times patchy, whitish-greyish to greenish or
greyish-brownish. Branchlets dark by resinous
Fig. 30. Stemonurus umbellatus Becc. Fruiting
twig, Mt Blumut, Johore (FRI 8841) (WHITMORE,
1968).
1971]
ICACINACEAE (Sleumer) 63
excretion. Leaves oblong to obovate, sometimes
elliptic, apex generally rounded or obtusely apic-
ulate, sometimes slightly retuse, rarely shortly
Vern. Sampin kris, sampit kris, M; Borneo:
djadjar, Dajak (Tajan), éntaburok, sémburok,
Iban, kakuli, Dusun; Philippines: malatadu, Mbo.
bluntly acuminate, base cuneate, coriaceous, some-
times very stiffly so, shining above, rather dull
beneath, the whole undersurface or at least the
parts closer to the midrib set with fine epidermal
tubercles, margin a little revolute, (6—)7—12(-16)
by (3-)4-7(-9) cm, nerves numerous (20-25)
straight subparallel pairs, generally slightly raised
above only, rather obscure though visible be-
neath, reticulation, if any, remarkable above only;
petiole 1.2-2 (very rarely -3) cm by 2(-3) mm.
Cymes arranged in an umbel, its peduncle robust,
1-2(-3) cm by 2mm, the (4~-)5-6(-8) branches
(3—)5-8 mm long and c. 1.5mm 9g, each branch
bearing (1—)3—4(—S) distally crowded flowers; bract
at base of umbel ovate-subulate, c. 2 mm, sub-
persistent. Calyx broadly obconical, 3 mm, base
rugulose, margin truncate or very shallowly
(apiculately) lobed, the very edge almost pellucid
and ciliolate (lens!). Petals broad-oblong, slightly
keeled, (4-)5(—6) by 2(-—2.5) mm, white or light
brown, scented. Disk cup-shaped and shallowly
lobed. Drupe ovoid, very apex rather abruptly
attenuate, lower half glossy green, upper one dull
grey, 3—3.5(-4) by 1.5—2.2 cm, the fibrous part of
the endocarp 1 mm thick.
Distr. Malesia: Malay Peninsula, Borneo.
Ecol. Primary forest, generally on hillside or
high level alluvium, occasionally in swamp between
hills, also in peat swamp forest, in coastal and
high inland kerangas, on padang and on cliffs,
generally in lowland up to 900 m, in the Cameron
Highlands up to 1465 m, on Mt Murud in Sarawak
at 1700 m, and on Mt Pagon Periok in Brunei at
1800 m; on Mt Kinabalu up to 1220 m, on sandy,
maybe waterlogged soil (Agathis forest), sandy
peat soil or sandy loam, rarely on sandstone.
Fl. fr. Jan.—Dec.
Uses. Wood pale orange-brown or honey-
coloured, apparently not used, the value of the
timber not known.
Fig. 31. Stemonurus umbellatus Becc. Stem, Mt
Blumut, Johore (FRI 8841) (WuitTMorE, 1968).
15. WHITMOREA
SLEUM. Blumea 17 (1969) 263. — Fig. 32.
Tree. Leaves spirally arranged, entire, coriaceous, penninerved, petioled.
Flowers bisexual, generally pedicelled, uni- or biseriate towards the +. recurved
end of secondary peduncles, the latter 3 or 4 as a cross-like umbel on top of a
common axillary and solitary peduncle. Ca/yx cup-shaped, entire, persistent and
+ flattened below fruit. Petals (4—)5, oblong, valvate, free in the uppermost part,
remaining connate below at full anthesis, apex inflexed, with a conspicuous midrib
inside, caducous. Stamens (4-)5, inserted at the very base of the petals; filaments
linear, fleshy, with longish subclavate white hairs below the anther cells inside and
at the connective outside; anther cells linear, thecae diverging below, basifixed.
Disk annular, flat, rather obscure. Ovary ovoid-conical, apex truncate, i.e. inverted
there and forming a kind of cup, with the small hardly bilobed unilateral stigma
on its inner edge. Drupe oblong-ellipsoid, smooth and reddish in the lower ¥%4,
64 FLORA MALESIANA [ser. I, vol. 72
Fig. 32. Whitmorea grandiflora SLEUM. a. Habit, x14, b. inflorescence, x Y%, c. open flower, «114,
d. petal inside, x2, e. stamen, x4, f. ovary and calyx, x5, f’. upper part of ovary obliquely seen from
above to show the stigma turned inside the apical cavity of the ovary, x10, g. submature fruit, x 34
(a-f BRAss 3418A, g BSIP 5212 WHITMORE, type).
1971] ICACINACEAE (Sleumer) 65
papillose and greyish-yellowish in the upper 14 part, exocarp thin, with an outer
fibrous, and an inner crustaceous layer. Seed 1, ellipsoid-subquadrangular, with
a lengthwise depression around the middle; embryo small, in the apex of the copious
albumen.
Distr. Monotypic, hitherto only known from the Solomon Islands.
1. Whitmorea grandiflora SLEUM. Blumea 17
(1969) 264. — Urandra umbellate [non (BEcc.)
O. Krze] WALKER, For. Fl. Br. Solomon Is.
(1948) 127. — Fig. 32.
Spreading canopy tree, 6-—25(-30)m, bole
straight, up to 12 m, up to 50cm 9g, with slender
and weak pneumatophores in wet places; bark
grey to brown, smooth or cracked. Branchlets
robust, often covered with resinous exudation as
are the inflorescences. Leaves elliptic or oblong-
elliptic, apex obtuse or slightly emarginate, base
broadly cuneate to rounded, coriaceous, firm,
shining green above, dull and greyish or yellowish
green beneath, glabrous, 14-22 by 4-8(—12) cm,
nerves in 20-25 pairs, straight and parallel to
each other, slightly raised on both faces, reticu-
lation dense, visibly raised above only; petiole
2-3(-4) cm by 2(-3) mm, minutely transversely
wrinkled. Inflorescences axillary, solitary, with 3
or 4 secondary peduncles (1-3 cm) crosswise and
umbellately spreading from top of main peduncle
(2.5-4cm by 1.5—2 mm), flowers several, close
together, scorpioid, uni- or biseriate, on the upper
+ recurved part of the secondary peduncles,
thick-pedicelled for 0.5-2(—3) mm; bracts at base
of secondary peduncles knob-like, rather obscure.
Calyx obconical-cupular, maybe suddenly nar-
rowed at base for 1-2 mm to a kind of foot for
1-2 mm, rugose, persistent and more spreading
or almost patellar under the ripe fruit, 4-5 by 6-7
mm. Petals chartaceous, glabrous, veined length-
wise, becoming free only in uppermost part,
remaining connate for the rest, oblong, white with
cream centre or greenish-yellowish, with sweet
smell, (12—)13-15 by 3-4mm. Filaments linear,
fleshy, 9-10 by 1mm, with longish white hairs
below the anthers inside and at connective out-
side; anthers linear, thecae spreading below, 4 by
1 mm. Ovary ovoid-conical, truncate, hollow at
apex, its edge folded in dry specimens, 3.5—4 by
2-2.5mm. Drupe_ oblong-ellipsoid, apiculate,
reddish and smooth in the lower 34, papillate and
greyish-yellowish in the upper 14 part, 3.5-4 by
2-2.5 cm.
Distr. Melanesia: Solomon Is. (Bougainville
to San Cristobal).
Ecol. Primary and secondary rain-forest, both
on well drained slopes or ridge top and in swampy
valley bottoms, though locally not rare, from
lowland up to 950m.
Uses. Wood rather hard, moderately heavy,
cream to brownish, used for planting sticks,
suitable for not too heavy constructions.
Vern. Aialo, aikuma, aikuni, aikunu, arifanatia,
Kwara’ae, duleke, Kolombangara, porutolo,
Faro I.
16. POLYPORANDRA
Becc. Malesia | (1877) 125; SLEUM. Blumea 17 (1969) 248. — Fig. 33.
Dioecious climber, whether or not with extra-axillary or intrapetiolar tendrils.
Leaves opposite, entire, penninerved. Panicles axillary, composed of short cymes.
Flowers small, 5—6(—7)-merous. Pedicels articulated with calyx. — 3 Flowers:
Calyx cup-shaped, shortly dented. Petals valvate, inflexed appendiculate distally
inside in bud. Stamens sessile; anthers thick, clavate-subglobose, with numerous
small pollen-bearing alveoles, each of which is covered with a caducous membrane.
Rudiment of ovary conical. — 2 Flowers: Calyx deeply dented. Petals very short,
connate in the lower 14. Stamens 0. Ovary subglobose-cylindric, stigma large, disk-
shaped, the centre depressed, crenulate or very shortly many-lobed in later stages.
Fruit compressed-ellipsoid; exocarp thin; endocarp firm, irregularly foveolate-
reticulate outside when the exocarp is gone, wrinkled inside. Seed 1; cotyledons
foliaceous.
Distr. Monotypic, Solomon Is. and East Malesia. Fig. 34.
66 FLORA MALESIANA
[ser. I, vol. 72
Fig. 33. Polyporandra scandens BEcc. a. Habit, x 14, b. g inflorescence, x 14, c. § flower, x5, d. 2 flower
x10, e. anther, x20, f. drupe, <1 (a, c, e VAN ROYEN & SLEUMER 6696, b ditto 6830, d CARR 12610
f ditto 16435).
1. Polyporandra scandens Becc. Malesia 1 (1877)
125, t. 7; PULLE, Nova Guinea 8 (1912) 660;
SCHELLENB. Bot. Jahrb. 58 (1923) 176; BirNIE,
Nova Guinea 14, 2 (1926) 277; Wuite, J. Arn.
Arb. 10 (1929) 235; SLEUM. in E. & P. Nat. Pfi.
Fam. ed. 2, 20b (1942) 379, f. 109; Daut, J.
Arn. Arb. 33 (1952) 275, f. 70 (pollen); STEEN.
Nova Guinea, Bot. n. 23 (1965) 495; SLEUM.
Blumea 17 (1969) 248. — ? Iodes ovalis (non BL.)
Wars. Bot. Jahrb. 13 (1891) 299. — P. hanseman-
nii ENGL. Bot. Jahrb. 16, Beibl. 39 (1893) 13
(‘hausemanni’); in E. & P. Nat. Pfl. Fam. 3, 5
(1895) 253; K. Sco. & Laut. Fl. Schutzgeb.
(1900) 418. — Jodes sogerensis BAKER f. J. Bot.
61 (1923) Suppl. 10. — Fig. 33.
Scandent shrub or branched liana, climbing to
1971]
ICACINACEAE (Sleumer) 67
35m, stem up to 8cm @o. Branchlets slender,
divaricate, pubescent initially; nodes thickened;
internodes 4-10cm. Leaves opposite, ovate to
elliptic, apex shortly + abruptly acuminate to
cuspidate, base broadly attenuate to rounded,
slightly inequilateral, chartaceous to subcoria-
ceous, glabrous above, covered with scattered
short appressed hairs all over beneath, 7—-15(-17)
by 3.5-9 cm, margin slightly revolute, entire, mid-
rib a little impressed above, strongly prominent
beneath, nerves (4~-)6-8 curved-ascendent pairs,
prominent beneath only as is the rather dense
network of veins; petiole 8—15(—20) mm; tendrils
whether or not present at the upper axils, elon-
gate, bifurcate. Panicles 1-3 from the same axil,
few- to many-flowered, (2—)3-9 cm, ascending or
recurved, all over covered with short appressed
hairs. Peduncle 1-3cm, bearing several few-
flowered cymes. Pedicels slender in the 3, thicker
in the 2 flowers. — ¢ Flowers: Calyx tube short,
lobes ovate-acuminate, c. 1mm. Petals ovate-
caudate, + expanded, white to cream or yellow,
4-5(-6) mm, acumen abruptly narrowed, + in-
flexed, glabrous, 2mm, tip blunt. Anthers c.
1mm g. Rudiment of ovary hardly 1mm. —
° Flowers: Calyx as in 3 flowers. Petals only 1-1.5
mm. Stamens 0. Ovary densely hairy, c. 1.3 mm.
Fruit compressed-ovoid-ellipsoid, c. 2 by 1.5 by
1.5 cm, appressed-pubescent; exocarp rather soft
Fig. 34. Distribution of Polyporandra scandens
BECcc.
Distr. Solomon Is.; in Malesia: New Guinea,
Moluccas (Halmahera, Batjan). Fig. 34.
Ecol. Climber over shrubs or high liana in
primary and secondary rain-forest, from sea-level
to 700 (possibly to 1000) m. Fi. fr. Jan.—Dec.
Uses. On Bougainville young leaves are cooked
and smooth, 1 mm, apparently going early; en-
docarp hard, 0.5 mm, coarsely reticulate-foveolate.
and eaten or added to taros to give these a flavor.
Vern. Dodofu, Halmahera.
17. IODES
BL. Bijdr. (1825) 29 (‘Jédes’); SLEUM. Blumea 17 (1969) 219. — Mappianthus
HAND.-MaAzz. Anz. Ak. Wiss. Wien M.-N. KI]. 58 (1921) 150; SLEUM. Blumea 17
(1969) 225. — Fig. 36.
Dioecious climbing shrubs or lianas. Branchlets hairy, generally bearing shortly
bifid tendrils at or closely to the nodes (but not axillary!). Leaves decussate, entire,
apiculate by a somewhat projecting midrib, penninerved, with +- appressed and
strigose and/or softer, erect or obliquely spreading coarse hairs on midrib, nerves,
veins and veinlets, and sometimes besides with a tomentum of very short fine
substellate hairs which covers the intervenium beneath. /nflorescences in peduncled
cymes which may be combined to panicles, axillary, supra-axillary or a little
lateral from the nodes, sometimes terminal, solitary, the ¢ ones often elongate and
many-flowered, the ones generally shorter and rather few-flowered; lower
peduncles often sterile and becoming tendrils. Flowers small, white to cream or
yellowish, fragrant. Pedicels articulate with the calyx and often dilated distally;
bracts and bracteoles minute. Calyx cup-shaped, 4- or 5-lobed halfway or up to
8/, of its total length (Malesia), persistent in 9. Petals 4 or 5, or sometimes wanting
in °, valvate in bud, generally connate at base only, rarely and only in ¢ connate to
Y, to %, of their length into a tube, only the reflexed lobes free, a little keeled
inside, tips sometimes narrowed and inflexed already in bud, persistent in 9. —
$ Flowers: Stamens (3—)4 or 5; filaments very short and fixed either to the base of
the (then shortly connate) petals or to the end of the corolla tube between the lobes,
68 FLORA MALESIANA
or stongate and fied to the base of the corolla tube; aiittier balls almost pasifisedl
introrse. Ovary rudimentary. — 2 Flowers: Staminodes sometimes present. Ovary
sessile; stigma sessile, discoid, cut on one side, or faintly 2—5-lobed. Drupe ovoid-
ellipsoid or oblong-obovoid (Malesia), or almond-like, generally laterally com-
pressed and longitudinally grooved or crested; exocarp thin, fleshy, yellow to red;
endocarp crustaceous to thin-woody, irregularly ribbed or reticulately lacunose, or
warty outside, smooth inside. Seed 1, albuminous; cotyledons large, foliaceous.
[ser, vol. $b
Distr. About 16 spp. in tropical Africa (and Madagascar), 3 spp. in SE. Asia (Assam and Burma to
SW.-SE. China, Hainan, Indo-China, and Thailand), throughout Malesia (7 spp.), but in New Guinea
only questionably known from a single old collection; 2 Malesian spp. also known from SE. Asia. Fig. 35.
Anat. Rosrinson (Ann. Inst. Bot. Btzg 8, 1890, 111, t. 18 & 19, f. 1-5) has investigated the anatomy
of the stem of /. cirrhosa Turcz. Vascular strands are formed here also in the pith, though the bast and
wood elements are inversed.
Notes. For the determination of the various species the indumentum plays an important role. The use
of a lens is necessary, and only leaves of + the same state of maturity should be investigated or compared.
Mappianthus differs from Jodes only by rather vague characters such as filaments rather long in pro-
portion to the length of the petals and a drupe with a finely grooved and tubercled exocarp. It is regarded
here as congeneric, a point of view already expressed by BAILLON, who, however, created for J. hookeriana
a proper sect. Lasiodes (Adansonia 10, 1872, 268).
KEY TO THE SPECIES
1. Petals in the J flowers practically free, i.e. united only at the base.
2. Indument on the undersurface of -+- mature leaves consisting almost exclusively of rather sparse
to more dense manifestly appressed stiff hairs.
3. Flowers 5-merous. Pedicels of the g flowers swollen and + plate-like dilated distally when the
flower has gone. 2 Flowers without staminodes. Veinlets of leaves rather faintly reticulate.
1. I. ovalis
3. Flowers 4-merous. Pedicels of 3 flowers not swollen distally. 2 Flowers whether or not with 4
staminodes. Veinlets of leaves strongly reticulate. 2. I. yatesii
2. Indument on the undersurface of + mature leaves consisting exclusively either of numerous fine
and soft + erect hairs, or of both such hairs and more robust, stiff and generally + appressed
(rarely obliquely spreading) hairs of various density.
4. Flowers 4-merous. Hairs on undersurface of leaves rather sparse. Petals present in 2 flowers.
DI pee Sse YA A OMY ALICIN. p+ ee. . Gute Got ia, Dl. (ehh Fe 3. I. reticulata
4. Flowers 5-merous. Hairs on undersurface of leaves (rather) dense. Petals absent in 2 flowers.
Drupe up to 2 by 1.2(-1.5) by 1 cm. be Pe 52 tay ga Some: a ee 4. I. cirrhosa
1. Petals in the 3 flowers united into a tube for about their lower 24 (the tube at least 3 mm long).
5. Drupe oblong-ellipsoid, + obtuse at both ends, 1.2-1.5 by 0.8-1.2 by 0.8-1 cm. 5. I. philippinensis
5. Drupe larger.
6. Drupe oblong-ovoid, apex broadly obtuse, base narrowed, 2.8—-3.3 by 1.3-1.5 by 0.8-1.2 cm;
crustaceous endocarp laxly coarsely’ lacunose. . ..0:0 2 22.5 3 22 ae 6. I. velutina
6. Drupe almond-like, 2.3-3.5(-4.5) by 1.6-2 by 0.7-1 cm; crustaceous endocarp with several longi-
tudinal shallow furrows and numerous flattish warts... .....4.... 7. I. hookeriana
pr fet. tel «. te, ©. Jee ee
1. Iodes ovalis BL. Bijdr. (1825) 30; Mia. FI.
Ind. Bat. 1 (1856) 795, p.p.; BAILL. Adansonia 3
(1863) 364 in text, p.p.; in DC. Prod. 17 (1873)
22, incl. var. genuina BAILL. l.c. 23, p.p.; BECC.
Malesia 1 (1877) 124, p.p.; Scuerr. Ann. Jard.
Bot. Btzg 1 (1876) 14; Hai. f. Med. Rijksherb.
1 (1910) 13; Back. Schoolfl. Java (1911) 230 &
Addenda; Koorp. Exk. Fl. Java 2 (1912) 532;
Koorpb.-Scuum. Syst. Verz. 1, Fam. 162 (1912)
5; AMsH. in Back. Bekn. FI. Java (em. ed.) 6
(1948) fam. 135, p. 7; BAck. & BAKH. f. Fl. Java
2 (1965) 61; SLEUM. Blumea 17 (1969) 220. —
I. horsfieldii Batt. Adansonia 10 (1872) 267;
in DC. Prod. 17 (1873) 24. — Polyporandra jung-
huhnii Koorp. Kon. Ak. A’dam Versl. Verg.
Wis- & Natuurk. 17 (1909) 780. — J. oblonga
PLANCH. in Mast. in Hook. f. Fl. Br. Ind. 1 (1875)
597; Kina, J. As. Soc. Beng. 64, ii (1895) 128;
Rip. Fl. Mal. Pen. 1 (1922) 435.
Slender shrub, scandent with tendrils, up to
5m. Leaves oblong-lanceolate to -elliptic or
-obovate, apex acuminate to rounded, though
minutely cuspidate by the protruding midrib, base
rounded to subcordate, slightly inequilateral, thin
chartaceous, glabrous above the midrib excepted,
which is covered on both faces with a pale rusty
short villous tomentum equally found on the
branchlets and petioles, undersurface + densely
covered with stiff appressed hairs on nerves, veins
and veinlets, the proper intervenium glabrous,
(5-)6-15 by (2.5—-)3-6.5 cm, midrib and nerves
generally slightly impressed above, prominent
Ee
ICACINACEAE (Sleumer) 69
Fig. 35. Distribution of Jodes (incl. Mappianthus).
The figure above the hyphen indicates the number
of endemic species in the island or area, the figure
below the hyphen the number of non-endemic
species.
beneath, nerves 6-8 pairs, curved-ascending and
inarching, reticulation of veins rather coarse, that
of the veinlets more dense though rather faintly
raised beneath; petiole 3-15 mm. Inflorescences
all over covered with bothrusty short strigose and
softer hairs, the ¢ ones repeatedly dichotomous,
many-flowered, up to 10cm, the 2 ones con-
sisting of 8—16-flowered cymes and only 1.5—-4 cm
(incl. peduncle). Pedicels slender, swollen distally,
the very end finally plate-like dilated when the
flower has gone, 1-3 mm. Calyx 5-lobed to 14-74,
c. 1.5 mm. Petals 5, oblong, white or light yellow,
tip extended and inflexed, c. 2.5mm in the J,
lanceolate and acute (not inflexed) and c. 2 mm in
the 2 flowers, laxly appressed-strigose outside as is
the calyx. — ¢g Flowers: Stamens 5; filaments
slender, c. 0.6 mm; anthers subovate-oblong, c.
1 mm, cells with crenate-lobed margin. Rudiment
of ovary hairy. — 2 Flowers: Staminodes 0.
Ovary thick-cylindric, all over densely appressedly
hairy, 2 mm; stigma discoid, wider than the ovary,
deeply incised on one side, margin crenulate, gla-
brous. Drupe inequally ovoid-compressed, said
to be scarlet and to attain 1.5-2.5 cm in length
(seen in immature state only up to | cm); endo-
an with slightly prominent, not properly keeled
ribs.
Distr. Malesia: Sumatra (Eastcoast), Malay
Peninsula (Penang to Singapore), W. Java,
Central Java (Pekalongan: Margasari, once).
Ecol. Along border of woods, in shrubberies,
along roads or riversides, often in secondary veg-
etation, once in teak-forest (Margasari), some-
times on limestone, 50-300(-800) m. FI. /r.
Jan.—Dec.
Notes. What has been understood as J. ovalis
in numerous floras of SE. Asia and Malesia,
belongs partly to other species of the genus,
mainly to J. cirrhosa Turcz., I. philippinensis
Mera. and /. vitiginea HANCE.
I. ovalis BL. was mentioned by K. Sco. &
Laut. Fl. Schutzgeb. (1900) 417 and SCHELLENB.
Bot. Jahrb. 58 (1923) 176 for New Guinea
(Morobe Distr., once found by WARBURG), but the
specimen is lost and its identity remains uncer-
tain. There is no recent record of Jodes from New
Guinea.
2. Iodes yatesii MERR. Pap. Mich. Ac. Sc. 19
(1934) 165; SLteuM. Blumea 17 (1969) 221.
var. yatesii.
Scandent shrub. Branchlets weak, sparsely pu-
bescent; tendrils up to 15cm. Leaves ovate- or
oblong-elliptic, apex abruptly acuminate for 1—1.5
cm, tip acute, base rounded to subcordate, char-
taceous, -- shining on both sides, glabrous above,
the midrib excepted, subdensely covered beneath
with appressed strigose hairs on veins and veinlets,
and besides with softer hairs on midrib and nerves,
the proper intervenium hairless, (5—)7-12 by
3-6 cm, midrib a little sunken above, much pro-
minent beneath, nerves 5-6 curved-ascending and
inarching pairs, raised beneath, veins coarsely re-
ticulated and but little prominent beneath, re-
ticulation of veinlets dense, finely raised on both
surfaces; petiole slender, subdensely hairy, 0.5-1.2
cm. Cymes 3-8 cm incl. peduncle, few- to many-
flowered, very slender, all over covered with short
substrigose and brownish softer hairs. Pedicels
slightly dilated distally in the 9, hardly so in the
3 flowers, very slender, 2-5 mm. Calyx c. 0.8 mm,
4-lobed to 24, lobes spreading. Petals 4, oblong-
acuminate, tip not extended and hardly inflexed,
appressedly strigose outside as is the calyx, gla-
brous inside, c. 1.2 mm. —¢ Flowers: Stamens 4;
filaments 0.6 mm; anther cells ovate, margin not
crenulate, c. 0.4mm. Rudiment of ovary sub-
globular, hairy, minute. — 2 Flowers: Sepals and
petals as in g flowers, persistent. Staminodes 4.
Ovary ovoid, densely hairy; stigma flat, subgla-
brous. Drupe elongate-oblong-oveid, laterally
compressed, brownish-substrigose-tomentose, 2.5—
3 by 1.5 by 1.3 cm, slightly keeled; endocarp
shallowly and coarsely reticulate-impressed on
both faces.
Distr. Malesia: Surnatra (Eastcoast; Riouw:
Indragiri Uplands).
Ecol. Forest (also secondary) along river or
roadside. Fil. fr. March-Nov.
var. glabrescens (RIDL.) SLEUM. Blumea 17 (1969)
221. — J. reticulata KinG var. glabrescens RIDL.
Kew Bull. (1931) 35.
Appressed hairs on the undersurface of leaves
sparse. 2 Flowers without staminodes.
Distr. Malesia: Borneo (Sarawak: Kuching),
once found.
3. Iodes reticulata Kina, J. As. Soc. Beng. 64,
ii (1895) 126; Rip. Fl. Mal. Pen. 1 (1922) 434;
SieuMm. Blumea 17 (1969) 221.
Woody climber, up to 9 m. Branchlets slender,
striate, with a broad line of dilutely ferrugineous
tomentum on one side, changing sides at the nodes;
70 FLORA MALESIANA
[ser. ], voli g-
tendrils up to 15 cm. Leaves ovate-elliptic, some-
times subobovate, apex abruptly acuminate for
0.5-1.5 cm, acute, base generally slightly cordate,
a little inequilateral, thin-coriaceous, glabrous and
shining above, the hairy midrib excepted, all
over the undersurface covered with rather sparse
obliquely or patently spreading hairs, soft to the
touch, 5-12 by 3-6.5cm, midrib and nerves
slightly impressed above, boldly prominent be-
neath, nerves 4-5 curved high-ascending and
looping pairs, reticulation of veins and veinlets
finely raised above, strongly so _ beneath;
petiole densely hairy, 1-1.5 mm. Cymes lax, the
3g many-flowered and usually exceeding the leaves
in length, the 2 ones few-flowered and generally
shorter, + patently soft-rusty-pubescent in all
outer parts. Pedicels very slender in the g, thicker
and swollen distally only in the 2. —¢ Flowers:
Calyx cup-shaped, 1mm, 4-lobed to 14-2.
Petals 4, ovate-acuminate, a little fleshy, densely
hairy outside, glabrous inside, 0.8 mm. Stamens 4,
subsessile; anther cells subcordate, 0.2 mm. Ru-
diment of ovary minute, glabrous. — 92 Flowers:
Calyx as in 3 flowers. Petals 4, subovate-oblong,
united at the very base, tip acute, hairy outside as
is the calyx, reflexed, 2-2.5 mm. Staminodes 0.
Ovary thick-cylindrical or barrel-shaped, all over
densely appressedly hairy, 1.5 mm; stigma discoid,
deeply cut on one side, margin hardly crenulated.
Drupe ellipsoid, compressed, base narrowed, fer-
rugineous-velvety, 2.5-3 by 1.5 by 1 cm, on
stoutish pedicel, 5-8 mm; endocarp bony, coarsely
furrowed outside.
Distr. Malesia: Malay Peninsula (Perak, Pa-
hang, Negri Sembilan).
Ecol. Open jungle, at low altitudes up to 250 m.
Fl. May-Nov., fr. May-June.
4. Iodes cirrhosa Turcz. Bull. Soc. Nat. Moscou
27, ii (1854) 281; SLEUM. Notizbl. Berl.-Dahl. 15
(1940) 250; Herne, Pfi. d. Samml. Clemens Ki-
nabalu (1953) 58; AmsuH. in Back. Bekn. FI. Java
(em. ed.) 6 (1948) fam. 135, p. 7; HEINE in Fedde,
Rep. 54 (1951) 238; BAck. & Baku. f. Fl. Java 2
(1965) 62 (‘cirrosa’); SLEUM. Blumea 17 (1969)
221; Fl. Thail. 2 (1970) 86. — I. ovalis (non BL.)
Hassk. Cat. Hort. Bog. (1844) 172; R. Br.
in Benn. & Brown, PI. Jav. Rar. (1852) 243, t. 48;
Mig. Fl. Ind. Bat. 1 (1856) 795, p.p.; BAILL.
Adansonia 3 (1863) 364, in text, p.p.; in DC.
Prod. 17 (1873) 22, incl. var. genuina BAILL.
l.c. 23, p.p. et var. miquelii BAILL. I.c. 23; MAST.
in Hook. f. Fl. Br. Ind. 1 (1875) 596; BEcc.
Malesia 1 (1877) 124, p.p.; Kinc, J. As. Soc.
Beng. 64, ii (1895) 128; Hocnr. Pl. Bog. Exs.
(1904) 18; Bull. Inst. Bot. Btzg 19 (1904) 40;
ibid. 22 (1905) 106 (as var. genuina); BRANDIS,
Ind. Trees (1906) 154; GAGNeEP. FI. Gén. I.-C. 1
(1911) 844, incl. var. cochinchinensis PIERRE ex
GAGneP. lI.c. 845; MERR. En. Born. (1921) 356;
RIDL. Fl. Mal. Pen. 1 (1922) 435; Cras, Fl. Siam.
En. 1 (1926) 276; GaGnep. FI. Gén. I.-C. Suppl.
(1948) 757; Dan, J. Arn. Arb. 33 (1952) 274
(pollen). — Natsiatum oppositifolium PLANCH. in
Hook. Lond. J. Bot. 5 (1846) 247, nom. nud. —
I. tomentella Miq. Fl. Ind. Bat. 1 (1856) 796;
Kurz, J. As. Soc. Beng. 44, ii (1875) 137; For.
Fl. Burma 1 (1877) 243; Becc. Malesia 1 (1877)
124, p.p.; Koorp. Minah. (1898) 393; BAck.
Schoolfl. Java (1911) Addenda ad p. 229; Koorp.
Exk. Fl. Java 2 (1912) 532; Koorp.-ScuHum. Syst.
Verz. 1, Fam. 162 (1912) 5; DAHL, J. Arn. Arb. 33
(1952) 274 (pollen). — J. brandisii Kurz, J. As.
Soc. Beng. 41, ii (1872) 298; ibid. 44, ii (1875) 156;
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 596;
Kurz, For. Fl. Burma 1 (1877) 243. — J. hors-
fieldii BAiLL. Adansonia 10 (1872) 267; in DC.
Prod. 17 (1873) 24. — I. floribunda MeErRR. Pap.
Mich. Ac. Sc. 19 (1934) 166, t. 29; Damn, J.
Arn. Arb. 33 (1952) 274, f. 65 (pollen). — Fig.
36a-d.
Woody climber, 5—10(—18) m tall, stem up to
8cm g. Branchlets slender, rusty-tomentellous.
Leaves ovate to ovate-elliptic, rarely oblong- or
obovate-elliptic, apex shortly acuminate and acute
by the slightly protruding midrib, base rounded or
subcordate, firmly chartaceous, upper surface
glabrous except midrib and nerves which bear
rather dense soft yellowish erect or obliquely
spreading hairs as are found all over the under-
surface on midrib, nerves, veins and veinlets
(the proper intervenium hairless), forming a ve-
lutinous tomentum, 5—15(-18) by 3.5—8(-11.5)
cm, nerves 4-6 curved and rather steeply ascending
pairs, prominent beneath as are midrib and the
transverse veins, reticulation of veinlets dense, a
little raised beneath only; petiole 1-2(—3.5) cm,
tomentellous. Inflorescence all over covered with
a short soft rusty tomentum, the ¢ in much
branched and many-flowered, rather lax cymes,
5-15 cm incl. the peduncle (1.5—-5 cm), the 2
ones shorter and mostly rather few-flowered,
2-12 cm incl. the peduncle (1-4 cm). — 3 Flowers
not rarely transformed into globular hirsute galls
(2-3 mm @). Pedicels slender, 1-2 mm, not swollen
distally towards the end of anthesis. Calyx 5-
lobed to almost the base, densely set with sub-
strigose hairs outside, lobes narrow-lanceolate,
14-34 mm. Petals oblong-lanceolate, tip filiform
and inflexed, thin, white or yellowish, fragrant,
laxly substrigose-pubescent outside, 2-3 by0.8 mm.
Stamens 5; anthers practically sessile, ovate-
elliptic, margin entire, 0.6—-0.8 mm. Rudiment of
ovary cylindric, hairy, 0.5 mm. — 2 Flowers: Pe-
dicels thickish, c. 1mm, accrescent in fruit to
c. 3 mm, hardly dilated distally. Calyx as in 3
flowers, lobes c. 1 mm. Petals 0. Staminodes 0.
Ovary cylindric, densely substrigose-hairy, 1 mm;
stigma a dark bilobed or subreniform pad, hairy,
glabrescent in later stages. Drupe obliquely ovoid,
laterally compressed and crested, shortly + ap-
pressedly rusty- or rufous-tomentose, 1.2—1.5(-—2)
by 1—1.2(—1.5) by 0.7—1 cm; exocarp thin, reddish;
endocarp hard, coarsely lacunose-reticulate out-
side, smooth inside.
Distr. SE. Asia (Khasia, Burma, Thailand,
Indo-China); in Malesia: Sumatra (East & West-
coast, Padang), Banka, Malay Peninsula (Perak,
Selangor, Johore, Malacca), W. & E. Java,
Borneo (Sarawak, W. & N. Borneo), Philippines
1971]
ICACINACEAE (Sleumer) 71
Fig. 36. lodes cirrhosa Turcz. a. Habit, x 14, b. 3 flower, «5, c. Q flower, «5, d. drupe, x1. — J.
philippinensis Merr. e. 3 flower from the outside, <5, e’. ditto from the inside, expanded, x5 (a—b
SINCLAIR 9875, c VAN STEENIS 12717, d KING’s Coll. 6783, e-e’ SANTOS 4356).
(Palawan), Celebes, possibly in the Moluccas.
Ecol. Light primary and often secondary forest,
forest borders, shrubberies, scattered, also in open
vegetation on limestone rocks, sometimes rather
dry places, in Java in regions with at least 20 or
more rainy days in the 4 consecutive driest months,
10-1100 m. FI. fr. Jan.—Dec.
Vern. Akar mengkunyit (Selangor), M, arey
balilandak, ki kurip, 8, jagal kédang, jungetan, J,
sasir kété, Minahasa.
5. lodes philippinensis Mere. Philip. J. Sc. 3 (1908)
Bot. 241; C. B. Ros. ibid. 6 (1911) Bot. 209;
Hoitu. & H. J. Lam, Blumea 5 (1942) 205;
Daut, J. Arn. Arb. 33 (1952) 274, f. 68 (pollen);
SLteuM. Blumea 17 (1969) 224. — I. ovalis (non
BL.) BaILL. in DC. Prod. 17 (1873) 22, quoad
Philip.; F.-Vitt. Nov. App. (1880) 46; VIDAL,
Phan. Cuming. Philip. (1885) 103; Rev. Pl. Vasc.
Filip. (1886) 86; Merr. Philip. J. Sc. 30 (1926)
408; Hottu. & H. J. Lam, Blumea 5 (1942) 205.
— I. tomentella (non MiqQ.) Becc. Malesia |
(1877) 124, quoad Philip. — I. oblonga PLANCH.
ex Mast. var. moluccana Hocure. Pl. Bog. Exs.
(1904) 18; Bull. Inst. Bot. Btzg 19 (1904) 40;
ibid. 22 (1905) 107; Ann. Jard. Bot. Btzg Suppl.
Ill, 2 (1910) 855. — Fig. 36¢e-e’.
Woody climber, up to 6 m. Branchlets slender,
striate, soft villous by ferrugineous fine rather
short hairs as are the petioles; tendrils slender, up
72 FLORA MALESIANA
[ser. I, vol. 73
to 15 cm. Leaves ovate-elliptic to -oblong, apex
short acuminate, the midrib hardly projected
beyond the lamina, base rounded or mostly
slightly cordate, thin chartaceous, upper surface
finally glabrous except the midrib, undersurface
all over soft tomentose, more densely so on midrib
and nerves, the hairs being of two markedly dif-
ferent types, i.e. stoutish substrigose + appressed,
and much finer, soft and + erect ones, the latter
sometimes rather scarce, the space between the
veinlets hairless, (3—)5—9 by 2.5—5(-—7) cm, midrib
and 4-6 pairs of ascendent and + looping nerves
plain above, + strongly raised beneath, less so the
veins, reticulation of veinlets not very distinct;
petiole 0.6-1(-1.5) cm. — 3 Inflorescences: Pan-
icle; composed of several cymes, rather lax and
relatively few-flowered, 2-4(—5) cm incl. the long
peduncle, covered with + crisped and appressed
rusty substrigose hairs. Pedicels slender, 1-2 mm,
the end glabrous and a little dilated when the
flower is gone. Calyx cup-shaped, 1.5mm,
acutely 5-lobed halfway. Petals subglobose in bud,
rapidly accrescent to 6-7(-8) mm at full anthesis,
united into a narrow tube for the lower 24, free
at the 5 lobes, the latter acuminate, narrow,
tip inflexed in bud stage, ultimately refiexed or
spreading, hairy outside and at the mouth inside,
white or cream. Stamens 5, inserted at the end of
the tube, alternating with the lobes; filaments
flattened, very short; anther-sacs narrowly sub-
ovate-oblong, not crenulate, 0.8-1 mm. Rudiment
of ovary cylindric, glabrous, c. 1.5mm. — 2
Inflorescences 3-5 (rarely —12)-flowered, 2-3 cm
incl. the peduncle, hairy as are the ¢ ones. Pedicel |
slender, 3(-4) mm, not swollen distally. Calyx
cup-shaped, 1.8 mm, 5-lobed halfway or a little
more. Petals (4-)5, narrow-oblong, connate at
the base, white or cream, thin, finally reflexed,
glabrous inside, 2.5-3 by 0.6-0.8 mm. Staminodes
0. Ovary cylindric, all over clad with stiff forward
directed hairs, 2mm; stigma thick, bilobed,
glabrous. Drupe obliquely oblongoid-ellipsoid,
laterally compressed and rather sharply crested,
+ obtuse at both ends, red to orange, 1.2-1.5 by
0.8-1.2 by 0.8-1 cm, densely rusty hairy; endocarp
coarsely reticulate-lacunose; pedicels stoutish,
2-5 mm, a little swollen distally. Sepals and petals
persistent on the fruit for a long time.
Distr. Malesia: N. Borneo, Philippines (all
islands), Central E. Celebes (once found), Mo-
luccas (Talaud, Morotai, Halmahera, Ternate,
Ambon).
Ecol. Edge of forests or thickets, also seconda-
ry growths, in the lowland up to 200 m, but
ascending in the foothills of Mt Kinabalu up to
1525 m, often near rivers.
Vern. Kogopusut, Dusun (Sambunan).
6. Iodes velutina Kinc, J. As. Soc. Beng. 64, ii
(1895) 127; Ripy. Fl. Mal. Pen. 1 (1922) 434;
Burk. Dict. (1935) 1245; SLteum. Blumea 17
(1969) 224.
var. yelutina.
Climber, similar in leaves and flowers to J.
Philippinensis. Branchlets shortly rufous-pubescent
especially so on one side; tendrils glabrescent.
Leaves ovate, apex short acuminate and subacute
by a glandular tip, i.e. the protruding midrib,
base rounded or generally subcordate, firmly
chartaceous, upper surface initially + densely
short pubescent, soft to the touch, glabrescent
except the midrib, undersurface velvety by nu-
merous + appressed substrigose and even more
numerous softer + erect hairs, the tomentum
rusty-olivaceous in dry, said to be whitish in fresh
specimens, the intervenium, i.e. the proper space
between the veinlets glabrous and thus green,
(3-)4-10 by 2-5.5(-8) cm, nerves 4-5 rather
steeply curved-ascendent pairs, a little raised be-
neath only, veins and reticulation of veinlets +
hidden by the tomentum underneath; petiole
slender, tomentose, 0.5—1.5(-2) cm. — 3 IJn-
florescences: Cymes few- to rather many-flowered,
all over rusty-pubescent, 2-4 cm (incl. peduncle).
Pedicels slender, 2-3 mm, finally a little swollen
distally and glabrous at the very tip. Calyx cup-
shaped, c. 1.3 mm, 4- or 5-lobed for 14 to almost
14. Petals 4 or 5, bud club-shaped, at full anthesis
5-7 mm and connate to a tube in the lower 2/3,
lobes finally reflexed, their tip inflexed, sub-
densely + appressedly hairy outside as is the
calyx, long-hairy at the throat inside. Stamens 4
or 5, inserted at the throat; filaments laxly hairy,
c. 1mm; anther-sacs ovate-oblong, 0.8-1 mm.
Rudiment of ovary conical-cylindric, glabrous,
0.5 mm. — @ Inflorescences: Cymes 2-5-flowered,
2-3 cm incl. the rather robust peduncle. Pedicels
rather slender and 3—5 mm in anthesis, a little
accrescent and more robust in fruit. Calyx cup-
shaped, c. 1 mm, 4- or 5-lobed halfway, appressed-
ly hairy as are the petals outside. Petals 4 or 5,
subobovate-oblong, free to almost their base, 4-5
mm. Staminodes 0. Ovary cylindric, densely hairy;
stigma slightly 4- or 5-lobed. Ripe drupe oblong-
obovoid, apex broadly obtuse, base attenuate,
much compressed laterally and manifestly crested,
with a less obvious crest on each of the flat sides,
tomentose initially, finally subglabrous, 2.8-3.3
by 1.3-1.5 by 0.8—1.2 cm, the thin exocarp blackish
when dry; endocarp laxly coarsely lacunose; stigma
slightly 5-lobed, 2 mmg. Seed solitary, compressed.
Distr. Malesia: Malay Peninsula (Perak, Ne-
gri Sembilan, Malacca, Singapore).
Ecol. Woods and hedges, at low elevation.
Vern. Akar china bukit, a. salupat, M.
var. subvillosa SLEUM. Blumea 17 (1969) 224. —
I. tomentella (non MiQ.) SCHELLENB. Bot. Jahrb.
58 (1923) 176.
Undersurface of leaves covered with a felt of
small substellate hairs (and thus greyish) on the
space between the veinlets. In flower and fruit
characters not distinguishable from var. velutina.
Distr. Malesia: Sumatra (Eastcoast, Ben-
coolen), Borneo (Amai Ambit and area S. of
Kuching). Questionable for New Guinea by an
old specimen said to have been collected by Zip-
PELIUS.
Ecol. Primary forest both on sand- and lime-
1971]
ICACINACEAE (Sleumer)
73
stone in Borneo, 100-1000 m. Fi. fr. Aug.
7. Iodes hookeriana BAILL. Adansonia 10 (1872)
268; in DC. Prod. 17 (1873) 23; Mast. in Hook.
f. Fl. Br. Ind. 1 (1875) 596; Kurz, J. As. Soc.
Beng. 44, ii (1875) 157; For. Fl. Burma 1 (1877)
244; BRANDIS, Ind. Trees (1906) 154; KANJILAL
c.s. Fl. Assam 1 (1936) 254. — JI. thomsoniana
Batt. Adansonia 10 (1872) 270; in DC. Prod.
17 (1873) 25; Mast. in Hook. f. Fl. Br. Ind. 1
(1875) 596; BRanpis, Ind. Trees (1906) 154. —
L. reticulata KiNG var. glabrescens RIDL. Kew
Bull. (1931) 35. — Mappianthus borneensis MERR.
Webbia 7 (1950) 317. — Mappianthus hookerianus
(BAILL.) SLEUM. Blumea 17 (1969) 225.
Fairly large woody climber, sometimes with
tendrils, branchlets, undersurface of leaves along
nerves and midrib, petioles and inflorescences sub-
densely clothed with substrigose, appressed hairs.
Leaves elliptic- to ovate-oblong, apex abruptly
shortly (maybe bluntly) acuminate, or almost cus-
pidate, base + rounded, rarely subcordate, thin-
coriaceous, shining on both faces, margin slightly
revolute, entire (seemingly crenulate or denticulate
in dry specimens with crisped edge), (8—)10-15
(-20) by (3.5—)5-8 cm, midrib narrowly impressed
above, strongly raised beneath, nerves 7-—8(-9)
pairs, moderately curved-ascending and inarching
before the edge, faintly raised above, distinctly so
beneath as are the + transverse veins, reticulation
of veinlets dense and finely raised on both faces;
petiole 1-1.5cm. Inflorescences slightly extra-
axillary dichasia, the ¢ ones rather many-, the 2
ones few-flowered, often reduced in the lower axils
to tendrils. Pedicels c. 1 mm, slender in the 3,
stoutish in the 2 flowers. Calyx cup-shaped, c. |
mm, edge minutely 5-dented to almost entire. —
3 Flowers: Petals oblong, fleshy, c. 3 mm, con-
nate to c. 34, densely and shortly yellowish-
substrigose outside, glabrous inside. Filaments fili-
form, c. 2 mm; anther cells linear-oblong, c. 1.8
mm. Rudiment of ovary cylindric, all over shortly
strigose-tomentose. — 2 Flowers unknown. Drupe
almond-like, 2.3-3.5(-4.5) by 1.6-2 by 0.7-1 cm,
on thick pedicel c. 2mm; stigma rather small,
faintly 5-lobed; exocarp fleshy, orange red, ap-
pressedly short-strigose; endocarp crustaceous,
outside with several longitudinal shallow furrows
and numerous flattish warts, inside smooth.
Distr. SE. Asia (Assam, E. Bengal); in Malesia:
Sumatra (once found, Westcoast) and Borneo
(Sarawak: Kuching area; N. Borneo: foothills of
Mt Kinabalu, c. 900 m, twice found).
Ecol. Forests. Fr. Sept.—Nov.
Note. Closely related to and possibly con-
specific with Mappianthus iodoides HAND.-MAZZ.
(N. of Tonkin and adjacent China, eastwards to
Hunan and Fukien).
Excluded
Todes ferruginea K. Scu. & Laut. Fl. Schutzgeb.
(1900) 418; SCHELLENB. Bot. Jahrb. 58 (1923)
176, nota sub I. tomentella; SLEUM. Blumea 17
(1969) 225 = Alangium villosum (BL.) WANG.
(Alangiaceae).
18. SARCOSTIGMA
W. & A. Edinb. New Phil. J. 14 (1833) 299; SLEUM. Blumea 17 (1969) 253. —
Fig. 38.
Dioecious climbers. Leaves spirally arranged, entire, penninerved, strongly
reticulate; petiole curved or twisted, lengthwise striate and transversely wrinkled,
leaving a raised elliptic white scar. Flowers sessile or sometimes pedicellate
(2 only), clustered (or fascicled) or solitary along the rachis of long, pendulous
interrupted or spike-like racemes, these simple or composed to panicles, (4~)5(-6)-
merous. Ca/yx small, cupular, shortly dented, persistent. Peta/s shortly connate at
base, valvate in bud, afterwards recurved, oblong, persistent in 2 flowers. —
3 Flowers: Stamens 5, exserted; filaments linear; anthers small, elliptic, medifixed,
cells slightly divergent downwards. Rudiment of ovary hirsute. — 9 Flowers:
Staminodes filiform, whether or not present. Ovary ovoid-oblongoid, hirsute;
stigma almost sessile, thick, umbonate. Drupe: exocarp coriaceous, thin; endocarp
woody, thin, smooth or slightly verrucose and covered with a white membrane
inside, sometimes a little reticulate-foveolate outside in fully mature fruits. Seed 1,
testa thin; albumen none; cotyledons fleshy.
Distr. Two spp., India (Western Ghats from N. Kanara to Travancore), Lower Burma, Andamans,
Cochinchina, Annam; in Malesia: Sumatra, Malay Peninsula, Java, Borneo, Philippines. Fig. 37.
KEY TO THE SPECIES
1. Flowers in interrupted spikes; rachis stout. Petals 3-4 mm... . .
1. Flowers in slender panicles. Petals c. 2.5 mm... . .
1. S. kleinii
2. S. paniculata
o) ae @ se eee? ee es
74 FLORA MALESIANA
[ser. I, voli 74
Fig. 37. Distribution of Sarcostigma. 1 and dots S. kleinii, 2 and crosses S. paniculata.
1. Sarcostigma kleinii W. & A. Edinb. New Phil.
J. 14 (1833) 299; Miers, Ann. Mag. Nat. Hist. I,
10 (1852) 116, repr. Contr. Bot. 1 (1852) 103, t.
18; SLeEUM. Blumea 17 (1969) 254 (with further
synonyms). — S. horsfieldii R. Br. in Benn. &
Brown, Pl. Jav. Rar. (1852) 241, t. 47; Muers,
Ann. Mag. Nat. Hist. II, 10 (1852) 118, repr.
Contr. Bot. 1 (1852) 105; Miq. Fl. Ind. Bat. 1
(1856), 795: Scunize,, Tes 3) (1857/65) t.. 172,
f. 32 & 33; Hassk. Retzia ed. nov. (1858) 360;
BAILL. in DC. Prod. 17 (1873) 16; BEcc. Malesia 1
(1877) 125; Hocur. Bull. Inst. Bot. Btzg 19
(1904) 39; Pl. Bog. Exs. (1904) 16; Koorpb.
Jungh. Gedenkb. (1910) 175; Back. Schoolfi.
Java (1911) 229; Koorp. Exk. Fl. Java 2 (1912)
533; Koorpb.-ScHum. Syst. Verz. 1, Fam. 162
(1912) 5; Amsu. in Back. Bekn. Fl. Java (em. ed.)
6 (1948) fam. 135, p. 7; DAHL, J. Arn. Arb. 33
(1952) 276 (pollen); BAck. & BAKH. f. Fl. Java
2 (1965) 62.
Much branched liana, up to 26m, stem up to
8 cm 9g, sometimes leafless during anthesis; wood
hard. Branchlets slender. Leaves oblong to ob-
long-obovate or -elliptic, rarely lanceolate, apex
shortly rather abruptly and generally bluntly
acuminate, or obtuse, base broadly cuneate to
rounded, subcoriaceous to coriaceous, glabrous
on both sides, or rarely with some hairs beneath,
yellowish-green when dry, (7—)9—25 by 4-7(-11.5)
cm, midrib slightly depressed above, strongly
raised beneath, nerves (4—-)6-8 pairs, curved-
ascendent and inarching before the edge, pro-
minent beneath, reticulation dense and very
prominent (sometimes almost foveolate) on both
faces, or more distinct beneath; petiole 1—-1.5(—2)
cm, often recurved, striate as is the midrib beneath
and transversely wrinkled in dry specimens. Spikes
usually 1—3(—-5), fascicled, slightly supra-axillary
from partly already defoliate axils, or from old
wood, pendent, densely shortly brown-hairy to
-hirsutulous, 3 15-25, 2 up to 40 cm, the flowers in
numerous clusters or fascicles with flowerless
parts of the rachis between, generally sessile, rare-
ly (only in the 2) pedicelled (0.5-1.5, in fruit up
to 5mm), fetid. Calyx 1-1.5 mm, obtusely lobed
halfway, hairy below. Petals linear-oblong, sub-
acute, densely brown-hairy outside, glabrous and
with a slight median ridge inside, greenish-
yellow, 3-4 mm. — ¢ Flowers: Filaments strap-
shaped, glabrous, c. 5mm; anthers elliptic, c. 0.8
mm. Rudiment of ovary elongate-conical, hirsute.
— 2 Flowers: Staminodes whether or not present.
Ovary ellipsoid-obovoid, hirsute; stigma a reversed
subconical or flattish cup. Drupe unequally sub-
ovoid-ellipsoid or -oblongoid, laterally compressed
and almond-like, 2.5-3.5 by 1.7-2 by 1-1.5 cm,
very base suddenly contracted and hidden by the
persistent calyx and petals, apex crowned by the
thickish small (1 mm) stigma; exocarp very thin,
with numerous vertical thick lines standing out
in the dry fruit, bright orange red or yellow,
brownish-hirsute, glabrescent at length; endocarp
thin, 0.5 mm, hard, a little reticulate-lacunose out-
side, smooth or faintly verrucose and shiny white
in the fresh state inside. Seed 1, with a thin edible
sweetish pulp round the nut inside.
Distr. India (Western Ghats S. to Travancore),
Lower Burma, Andamans, Indo-China (Annam);
in Malesia: Malay Peninsula, W. &. E. Java,
Borneo (Sarawak: Semengoh F. R.). Fig. 37.
Ecol. Forests (also teak-forests), scattered, in
Java largely under seasonal climatic conditions,
up to 550m. Fi. fr. Febr.—Oct.
Uses. In India the oil of the seeds is highly
esteemed in the treatment of rheumatism; this
use is not known from Malesia.
Vern. Kamaras, tangkilan, J, buah pélay tédong,
M.
1971]
2. Sarcostigma paniculata PieRRE, Bull. Soc. Linn.
Paris 2 (1897) 1319; GAGNeEP. Fl. Gén. I.-C. 1
(1911) 841, f. 102, 1-3, incl. var. angustifolia
(PIERRE) GAGNEP. /.c. 842; Rip. Fl. Mal. Pen. 5
(1925) 298: SteEuM. Blumea 17 (1969) 254. —
S. angustifolia PIERRE, Bull. Soc. Lin. Paris 2
(1897) 1320. — S. philippinensis MERR. Philip. J.
Sc. 10 (1915) Bot. 276; En. Philip. 2 (1923) 492:
Dau, J. Arn. Arb. 33 (1952) 276, f. 71 (pollen).
— S. surigaoensis ELM. Leafl. Philip. Bot. 10
(1939) 3749; Dani, J. Arn. Arb. 33 (1952) 277
(pollen). — Fig. 38.
Climbing to 30 m, stem up to 10 cm g, crooked,
branched toward top and forming hanging masses;
wood very soft, coarsely porous, + yellow in the
centre, watery, with a large whitish pith; bark grey
to brown, smooth, rather thin. Branches divaricate,
laxly rebranched, the free ends pendent. Leaves
scattered along the slender branchlets, oblong or
ovate- to elliptic-oblong, rarely lanceolate, apex
broadly attenuate, bluntly subacuminate or obtuse,
base broadly cuneate to rounded, thin-coriaceous,
paler (greyish) green beneath, glabrous, shining
on both faces, (8—)12—20 by (2.5—)4~-9 cm, midrib
slightly raised above, prominent and longitudi-
nally striate beneath, nerves 4-6 pairs, steeply
curved-ascending, rather obscurely looping, raised
beneath, reticulation dense, finely tessellate on
both sides; petiole 1.5-2(-4) cm. — ¢ Panicles
axillary or terminal, descending, slender and
flexible, much-branched and many-flowered,
ICACINACEAE (Sleumer) 75
tomentose initially, early glabrate, 30-60 cm, the
lower branches up to 16cm; flowers 3-5 per
cluster, these spicately arranged at intervals of
5—15 mm, with brownish appressed hairs on rachis
and more densely around the clusters. Calyx
widely cup-shaped, membranous, indistinctly den-
ticulate, 0.6 mm. Petals narrowly oblong, apex
attenuate and slightly inflexed inside, thin, whitish
or yellow, practically glabrous, with 3 longi-
tudinal veins, c. 2.5mm. Filaments slender, gla-
brous, 1—1.5 mm; anthers elliptic-ovate, 0.7 mm.
Rudiment of ovary columnar, hairy. — @2 IJn-
florescences (PNH 14563) panicled, glabrate.
Flowers sessile, spaced along a rather slender
rachis. Calyx cup-shaped, hardly dented, 0.8 mm.
Petals oblong, subglabrous, 3mm. Staminodes
1 mm, no anthers. Ovary oblongoid, densely hairy,
2mm; stigma a large pad depressed in centre.
Infructescence paniculate, stoutish, hairy, c. 10 cm,
lower branches up to 4cm; pedicels c. 2mm
long and across. Drupe (subovoid-)ellipsoid, may-
be somewhat compressed, laxly hairy, glabrescent,
3-3.5 by 1.5—2 cm; exocarp blackish-brownish and
wrinkled on dry fruit; endocarp thin, woody,
smooth inside. Seed 1, white-coated.
Distr. Cochinchina; in Malesia: Sumatra
(Riouw), Malay Peninsula (Perak), Borneo,
Philippines (Leyte, Samar, Mindanao). Fig. 37.
Ecol. Primary lowland forest, from sea-level
up to 500 m (twice found at 1065 min Mindanao),
on sandy loam, scattered. Fi. fr. Jan.—Dec.
Fig. 38. Sarcostigma paniculata Pierre, a. Habit, « 14, b. f inflorescence, * y, c. d flower, x10, d.
6 flower bud, = 10, e. fructification with almost mature fruits in dry state, x 4 (a-d SAN 24459, e SAN
17107).
(ser. I, volo 7}
76 FLORA MALESIANA
19. PYRENACANTHA
Wicut in Hook. Bot. Misc. 2, 4 (1830) 107, Suppl. t. 9 & 10, nom. cons.; SLEUM.
Blumea 17 (1969) 249. — Freeria MERR. Philip. J. Sc. 7 (1912) Bot. 292.
Dioecious scandent shrubs or lianas. Leaves spirally arranged, glandular-repand-
dentate and penninerved. Flowers small, sessile, bracteate, in lax supra-axillary
spikes. Sepals or calyx 0. Petals connate at base, valvate in bud, persistent in 2
flowers. — 3 Flowers: Stamens (3—)4(-5); filaments very short; anthers minute.
Rudiment of ovary present. — 9° Flowers: Staminodes very short or absent. Ovary
sessile; stigma sessile, thick, truncate. Drupe laterally compressed; exocarp fleshy;
endocarp crustaceous, punctate outside, verrucose-papillose inside, the papillae
projecting into the interstices of the deeply ruminate and pitted albumen. Seed 1.
Distr. About 20 spp., most of them in tropical and subtropical Africa & Madagascar, 1 sp. (P. volubilis
WIGHT) in S. India and Ceylon, Indo-China and Hainan; in Malesia: 1 sp. in the Philippines.
1. Pyrenacantha repanda (MeErRR.) MErRR. En.
Philip. 2 (1923) 492; SLtEuM. Blumea 17 (1969)
249; SHaw, Kew Bull. 23 (1969) 113. — P. sp.,
CERON, Cat. Pl. Herb. Manila (1892) 46. —
Freeria repanda Mer. Philip. J. Sc. 7 (1912) Bot.
292. — Tragia luzoniensis MERR. Philip. J. Sc. 16
(1920) 564.
Woody vine, up to 8m. Branchlets slender,
minutely verrucose, the growing parts short-
pubescent. Leaves oblong to oblong-lanceolate,
apex slenderly acuminate, bluntish, base narrowed,
very base cordate, subcoriaceous, glabrous and
smooth above (though very finely papillate under
the lens), set with scattered appressed strigose hairs
(whose rounded bases are partly pellucid against
strong light) and scabrid to the touch beneath,
6-14 by 1.5-4(-5) cm, margin + repand with a
short glandular tooth at the excurrent end of each
lateral nerve, midrib and nerves prominent on
both faces, the latter in 5-6 curved-ascendent and
anastomosing pairs, reticulation rather lax and
slightly raised on both sides; petiole usually
strongly curved, glabrescent, 5—-15(-30) mm.
strigose-pubescent, (2—)4-8 cm; flowers scattered
in the upper half or less, each flower subtended by
an oblong acuminate small bracteole. Petals 4,
ovate-oblong, pubescent outside, 1.5-2 mm. —
3 Flowers: Filaments very short; anther cells ovate-
elliptic, hardly 0.5 mm. Rudiment of ovary hirsute,
subglobose. — 2 Flowers: Staminodes minute.
Ovary ellipsoid, densely pubescent, 1 mm; stigma
round, truncate, minutely papillate. Drupe elliptic-
oblong, laterally compressed to rather sharp
edges, glabrous, yellowish or red, 1.8—3 by 1-1.5
by c. 0.8 cm; exocarp pulpy or fleshy, rather thin,
wrinkled in dry fruits. Seed1; endosperm ruminate,
yellow, oily, externally deeply pitted by the pro-
tuberances on the inside of the thin endocarp.
Distr. Malesia: Philippines (Luzon: Ilocos
Norte, Bulacan, Tayabas, Sorsogon; Samar, Min-
danao).
Ecol. Primary forest at low altitudes, scattered.
Fl. March—May, fr. Sept.
Note. Apparently close to P. volubilis Hoox.
and to certain species from Madagascar in leaf-
characters, distinguishable from the first by its
Spikes supra-axillary, slender, minutely sub- larger fruits.
20. MIQUELIA
MEISN. Pl. Vasc. Gen. 1 (Sept. 1838) 152; ibid. 2 (Sept. 1838) 109; SLEUM. Taxon 17
(1968) 449 et ibid. 18 (1969) 479, nom. cons. prop.; Blumea 17 (1969) 229, non
BL. June 1838 (= Stauranthera BENTH. 1835). — Jenkinsia Grier. Calc. J. Nat.
Hist. 4 (1843) 231; ibid. (1844) t. 12, non Hook. Gen. Fil. (1842) text opp. t. 75 b.
— Fig. 39.
Dioecious climbing or twining shrubs, the wood with large porous vessels.
Leaves spirally arranged, entire or coarsely lobed (Mal.), generally minutely
papillose-tubercled on both faces, nerves palmate to pinnate. Flowers sessile,
arranged in heads or umbels; these on + elongate, distally +- thickened peduncles,
from defoliate or foliate axils; inflorescences solitary or several in lax racemes,
panicles or fascicles. Calyx small, +. deeply 4- or 5-lobed. — 3 Flowers: Petals 4
1971] ICACINACEAE (Sleumer) 77
b y
Fig. 39. Miquelia celebica Bu. a. Habit, x 4, b. 3 flower bud, x5, c. petal, x74, d. & d’. stamen,
714, e. submature infructescence, x 14, f. mature fruit, with remnants of the petals at the very base,
*1. — M. caudata KING. g. Fruit, <1 (a-d ELMER 9894, e KruyrrF 23, f KoorpDERS 19594, g CLEMENS
27608).
or 5, united into a short or mostly elongate-filiform tube adnate to the flower
axis, upper part or lobes free, valvate in bud, apex a little inflexed, patent or
+ reflexed in anthesis. Stamens 4 or 5; filaments short-linear; anthers elongate-
linear-oblong to (sub)sagittate, exceeding the filaments in length, dorsifixed a
little above the base, introrse. Rudiment of ovary minute. — 2 Flowers: Calyx
and lobes of the petals either distant from each other by the elongate flower
axis, or close together, i.e. practically no flower axis present. Staminodes flattish,
very short. Ovary thick-cylindric, hairy; stigma sessile, disk-shaped, its centre a
little depressed. Drupe oblongoid to subovoid-ellipsoid, laterally compressed,
whether or not narrowed into a stalk-like base, either sessile or on a pedicel-
like flower-axis; mesocarp thin; endocarp crustaceous, rugose or shallowly la-
cunose, ++ manifestly verrucose inside, the warts causing small pits in the flattened
ovoid seed which is surrounded by a juicy pulp; testa thin; albumen fleshy,
rugulose; cotyledons elliptic, thick-foliaceous.
Distr. About 8 spp. in SE. Asia (India: Western Ghats; Assam, Burma?, Thailand, Laos); in Malesia:
4 spp. (Sumatra, Malay Peninsula, Borneo, Philippines, Celebes). Fig. 40.
Ecol. Mainly lowland forest, rarely up to 1980 m (Mt Kinabalu).
KEY TO THE SPECIES
1. Calyx and lobes of the petals distinctly spaced both in the 2 and 3 flowers by the elongate flower-axis
"(== the basal connate tube-like part of the petals). Drupe ‘pseudopedicelled’ by this axis (and besides
much narrowed towards the base into a kind of stalk).
2. Leaves oblong to subovate-oblong, (10—-)12-17 by 4~6.5 cm; inner basal pair of nerves ascending to
the lower third of the lamina. Umbels c. 10-flowered. . . . 2. 1 6 ee ee 1. M. reticulata
2. Leaves ovate, 7-11 by 5-9 cm; inner basal pair of nerves ascending to 14 or % the length of the
ee On MS Fo Fs ie ee 2. M. philippinensis
78 FLORA MALESIANA
[ser. I, vol. 72
- Calyx and lobes of the petals close to each other in the 2 flowers and drupe, spaced in the 3 flowers.
Pig cad not “pseudopedicelled’ (but much narrowed towards the base into a kind of stalk).
- Drupe 2.3—-3.3 by 1.3-1.8 by 0.7—1 cm, narrowed at apex into a shorter or longer rather slender beak;
endocarp coarsely lacunose.
3. M. caudata
3. Drupe (0.8—)1—1.2(-1.7) by 0.7-0. 8(— i 2) by ( 0. 6 cm, shortly a and rather ‘gradually narrowed at apex;
endocarp more finely lacunose.
1. Miquelia reticulata Merr. Philip. J. Sc. 9 (1914)
Bot. 312; En. Philip. 2 (1923) 493; SLeum. Blumea
17 (1969) 230.
Scandent. Branchlets slender, sparingly hirsute
with short hairs, as are the petioles. Leaves oblong
to subovate-oblong, apex gradually attenuate or
subacuminate, base rounded and a little oblique,
very base shortly cordate, membranous, glabrous
and smooth above, set with scattered hairs all
over the undersurface and somewhat rough to the
touch, entire, (10-)12-17 by 4-6.5 cm, midrib
prominent beneath, nerves 2 basal pairs, the outer
pair short, the inner one ascending to the lower
third of the lamina, other higher nerves from the
midrib 3-4(-S) pairs curved-anastomosing, slightly
depressed above, raised beneath, reticulation
coarse and faint above, a little prominent beneath;
petiole (1—)1.5—2.5(—5.5) cm by 1-2 mm. — ¢ In-
florescences unknown. — 2 Flowers umbellately
arranged on top of a slender peduncle, the latter
either solitary and axillary, or mostly 2-4 pe-
duncles from a short thickened rachis on defoliate
stem, 3-10 cm, with about 10 flowers per umbel.
Calyx 4-lobed to almost the base, lobes hairy,
c. 1mm. Petals 4, greenish-yellow, fleshy, lobes
elongate-oblong, subacuminate, laxly puberulous
outside, glabrous inside, c. 3 by 1.5 mm, tubular
basal part of the petals 5-7 mm at full anthesis.
Stamens 4, flattened, very short. Ovary hairy,
crowned by a thick pad-like stigma. Drupe
narrowly ovoid in general, somewhat compressed,
c. 2 by 1.3 by 0.8 cm, wrinkled when dry, suddenly
attenuate at the very base into a kind of stipe
(5-8 mm), on the base of which the lobes of the
petals persist for some time; flower-axis below
these lobes 6-8 mm, slightly more slender than
the basal stipe of the drupe; exocarp thin, densely
hairy; endocarp thin, coarsely foveolate outside,
finely tubercled inside.
Distr. Malesia: Philippines (Camiguin and
Bohol Is.), twice found. Fig. 40.
Ecol. Forest at low altitudes, up to 600m.
2. Miquelia philippinensis Merr. Philip. J. Sc. 14
(1919) Bot. 416; En. Philip. 2 (1923) 493; SLEUM.
Blumea 17 (1969) 230.
Scandent shrub. Branchlets slender, laxly hir-
sute with shorter and longer hairs, as are the
petioles. Leaves ovate, apex broadly subacuminate-
attenuate, base rounded and very shallowly or
hardly cordate, membranous, glabrous and oliva-
ceous in dry state above, rather laxly and finely
short-hairy on the undersurface, entire, somewhat
repand or shortly (c. 5mm) 1-2-lobed on each
side, 7-11 by 5-9 cm, midrib and nerves faint
above, a little raised beneath, 2 pairs of basal
nerves, the outer pair rather short, the inner one
4. M. celebica
Fig. 40. Distribution of Miguelia. 1. M. kleinii,
2. M. reticulata, 3. M. philippinensis, 4. M. cau-
data, 5 and 2 dots M. celebica.
ascending to 14-34 the length of the lamina, upper
lateral ones c. 3 pairs, curved-anastomosing, re-
ticulation coarse and faint, less obvious than in
M. reticulata; petiole 2-3 cm by 0.5-1 mm. —
6 Inflorescence unknown. — 2&2 Inflorescence
axillary, solitary peduncles c. 3 cm, each bearing
c. 5 umbellately arranged flowers. Calyx lobes
ovate-acuminate, hairy, 1-1.5 mm. Petals united
below into a c. 5 mm long tube, free lobes oblong,
subglabrous, 2.5 mm. Submature drupe ob!ongoid-
ellipsoid, slightly hairy, 1-1.3 cm long, narrowed
downwards to a slender stalk as long as the fruit
proper.
Distr. Malesia: Philippines (Luzon: Ilocos
Norte Prov.), once found. Fig. 40.
Ecol. Forest at low altitude.
3. Miquelia caudata KinGc, J. As. Soc. Beng. 64,
ii (1895) 124; Ript. Fl. Mal. Pen. 1 (1922) 433;
Burk. Dict. (1935) 1478; SLEUM. Blumea 17
(1969) 230. — Fig. 39g.
A slender climber up to 6m. Branchlets pale,
striate, early glabrescent and corticate, 2-5 mm @.
Leaves oblong-lanceolate to elliptic-oblong, apex
shortly, sometimes rather abruptly acuminate,
base broadly cuneate to rounded, thin-charta-
ceous to membranaceous, glabrous above, laxly
puberulous all over the surface and soft to the
touch beneath, glabrescent, entire, (10—)12—20 by
1971]
(4-)5-8 cm, midrib and nerves moderately raised
beneath, nerves 1 or 2 basal and 5—6 upper pairs,
equally curved-ascending and rather obscurely
anastomosing, reticulation generally lax and but
slightly prominent; petiole pubescent, 1.2—3(-4)
cm. — Umbels of 3 inflorescences solitary or most-
ly several, in fascicles or along a short rachis from
hairy tubercles of foliate or defoliate axils of the
branchlets; peduncle filiform, 1.5—3 cm, the whole
inflorescence set with short almost scabrid hairs.
Calyx cup-shaped, 1mm, 4(—5)-lobed halfway.
Petals 4 (or 5) united to a filiform elongate basal
tube-like (4-10 mm), free in the upper oblong and
+ spreading part (2mm), white. Stamens 4 or
5; filaments short; anther cells elongate-linear,
subsagittate, 1.5 mm. Rudiment of ovary hirsute.
— © Flowers 8-15 together in capitules on top of
axillary, solitary stoutish peduncles, these gla-
brescent and accrescent in fruiting stage up to 11
cm. Calyx as ing flowers. Petals without a marked-
ly elongate basal part, i.e. close to the calyx, some-
what leathery and glabrous, 2.5mm. Ovary to-
mentose; stigma discoid, wider than the ovary.
Drupe broadly ovoid, laterally much compressed,
rounded at base in general, though abruptly nar-
rowed to a slender stipe for 5-7 mm, at the base of
which the remnants of the petals and calyx are
found, tapering upwards to a tail or beak for
(5—)6-10 mm, crowned by a small stigma, 2.3-3
(-3.3) by 1.3-1.8 by 0.7-1 cm; exocarp thin,
rusty-pubescent, ripening to a tomato or orange
red; endocarp bony, rather deeply (1-2 mm) and
coarsely reticulate-lacunose outside, tubercled all
over inside.
Distr. Malesia: Malay Peninsula, Borneo. Fig.
Ecol. Primary forest, also mossy forest, from
the lowland up to 1980m on Mt Kinabalu.
Uses. The stem holds a reservoir of potable
water. VAUGHAN-STEFFENS stated that the red
fruits are added to Antiaris latex in making
‘sakai (pangan)’ dart-poison (Rip. Agr. Bull.
Mal. Pen. 8, 1898, 209), but no confirmation of
this has been available.
Vern. Pisang-pisang bulu, M, sélowung, Pangan.
4. Miquelia celebica BL. Rumphia 4 (1849) 37;
Mus. Bot. Lugd.-Bat. 1 (1849) 42, f. 8; WALP.
Ann. 2 (1851) 22; Mio. FI. Ind. Bat. 1 (1856) 798;
SCHNIZL. Ic. 3 (1857/65) t. 172, f. 22-31; BAILL. in
DC. Prod. 17 (1873) 14; Becc. Malesia 1 (1877)
124; Koorp. Minah. (1898) 393; SteumM. Blumea
17 (1969) 231. — M. cumingii BAILL. Adansonia
10 (1872) 278; in DC. Prod. 17 (1873) 14; VIDAL,
Phan. Cuming Philip. (1885) 103; Rev. Pl. Vasc.
Filip. (1886) 86; Merr. En. Philip. 2 (1923) 492;
Dau_, J. Arn. Arb. 33 (1952) 275, f. 74 (pollen).
ICACINACEAE (Sleumer) 79
— M. rostrata MERR. PI. Elm. Born. (1929) 172.
— Fig. 39a-f.
High climbing shrub, much ramified. Branchlets
slender, green-olivaceous when dry, striate, sub-
glabrous. Leaves elliptic-, sometimes ovate-, rarely
lanceolate-oblong, apex shortly acuminate, tip
submucronate by the protruding midrib, base a
little oblique, broadly cuneate to truncate-rounded,
sometimes shallowly cordate, thin-chartaceous,
entire, or rarely obsoletely repand to coarsely few-
lobed, initially with scattered rather scabrid short
hairs (also found on the inflorescence) underneath,
early glabrescent, 9-15(-20) by 4~7.5(-10) cm,
midrib and nerves slightly raised beneath only,
lateral nerves 1-2 basal and 4-6 higher pairs,
all curved-ascendent and rather obscurely in-
arching before the edge, no proper reticulation;
petiole slender, 1.5—5(-7) cm. — 3 Umbels sub-
globose, many-flowered, slenderly long-peduncled,
axillary or slightly supra-axillary, rarely cauline,
either solitary or 2-3 very laxly racemosely ar-
ranged on a rachis of 1-3 cm by c. 2 mm. Calyx
shortly 4(—-5)-lobed, very small. Petals 4(-5),
united for S-7 mm to a filiform tube below, free
for the uppermost 2 mm, greenish yellow. Stamens
4(—S); filaments 1 mm; anther cells linear-oblong,
base slightly divergent, 1.5 mm. Rudiment of ovary
4-gibbous, glabrous. — 2 Flowers arranged to
many- and dense-flowered subglobose heads, these
generally solitary from the axil on a slender pe-
duncle 5-10 cm, rarely 2 or 3 laxly racemosely
arranged along a short rachis, each head then very
slenderly elongately peduncled. Calyx very small.
Petals 4, not elongate into a tube below as in the
3, oblong, thickish, 2mm. Staminodes 4. Ovary
subcylindric, densely hairy, crowned by the thick
stigmatic pad. Drupe obliquely ellipsoid-ovoid,
laterally compressed, (0.8—)1-1.2(-1.7) by 0.7-
0.8(-1.2) by 0.6 cm, apex shortly and rather grad-
ually attenuate, base suddenly narrowed to a
kind of stipe 5-7 by 1 mm (bearing the persistent
petals on its base); exocarp thin, laxly set with
short strigose hairs, yellow to reddish; endocarp
crustaceous, outside with numerous vertical low
ridges and transverse connections, shallowly
lacunose-reticulate in the dry fruit, inside with
numerous low warts which leave their mark in the
thus pitted seed.
Distr. Malesia: W. Sumatra (Padang, once
found), Borneo, Philippines (Luzon to Min-
danao), Celebes (Minahasa, Buton I.). Fig. 40.
Ecol. Primary forest, sometimes along streams,
generally in the lowland, rarely up to 700m,
in Buton I. on coralline limestone, apparently
scattered or rare.
Vern. Lalai in tasik,
Minahasa (Tt.).
sumbilan makanteh,
21. PHYTOCRENE
WALL. PI. As. Rar. 3 (1831) 11, t. 16; SLEUM. Taxon 17 (1968) 448 er ibid. 18
(1969) 479, nom. cons. prop.; Blumea 17 (1969) 234. — Gynocephalum BL. Bijdr.
(1825) 483; ENpL. Gen. (1837) 281 (‘Gynocephalium’); ReicHsB. Nom. (1841)
80 FLORA MALESIANA [ser. I, vol. "74
67 (‘Gyrocephalium’); Hassk. Cat. Hort. Bog. (1844) 78 (‘Gynaecocephalium’);
BL. Rumphia 4 (1849) 36, nota (‘Gynocephalium’); BENTH. in Benth. & Hook. f.
Gen. Pl. 1 (1862) 354 (‘Gynocephala ). — Fig. 42.
High climbing dioecious shrubs or lianas, sometimes with a robust stem,
younger parts of the latter usually hairy and prickly, youngest parts slender and
striate, sometimes rolled-in, and simulating tendrils, which are wanting. Leaves
remotely spirally arranged, entire or palmately 3—7-lobed, subpalmately 3-7-
nerved; petiole from a thickened and depressedly cup-like part of the stem, +
torted, never swollen basally or distally. — 3 Flowers in globose clusters, these
either solitary at the end of an axillary peduncle, or several spaced and racemosely
arranged then, or generally very numerous in long repeatedly branching spike-like
racemes or panicles both axillary and/or stem-borne. Ca/yx segments 3-5, + free.
Petals 3-4, +- united below, valvate, alternating with the calyx lobes. Stamens
3-4 (rarely —6), finally exserted; filaments filiform, +- connate at base into a column
which bears the hirsute rudiment of the ovary; anther cells ovate-elliptic, small. —
© Flowers in large solitary either cauline or mostly axillary peduncled globose
heads. Calyx segments 3-5, free to almost the base. Petals 3-4(—-5) as in the 3
flowers. Staminodes absent. Ovary subovoid-columnar, tapering to an equally
thick and short columnar style with several 2-lobed stigmas which form a kind of
funnel. Drupes numerous, collected in generally large, globose heads, each drupe
tipped by the style, + reversely setulose by brittle irritant hairs; exocarp thin,
larger than the endocarp and + hollow in the upper extended part at full maturity;
endocarp rather thick, ligneous, pitted to deeply lacunose outside, smooth or
warty inside. Seed 1, large, with a thin almost juicy coat which exudes a kind of
clear gum; endosperm fleshy, strongly furrowed or lobulated; cotyledons very
large, flat, foliaceous, double-folded or tortuous.
Distr. About 11 spp., of which 4 both in SE. Asia and Malesia, and 7 confined to Malesia from
Sumatra to New Guinea. Fig. 41.
Ecol. Scattered in primary and secondary lowland forest, occasionally up to 1200 m, also in open
places, as rarely found on limestone rocks.
Uses. The wood has very large porous vessels which contain water, used by the natives, hence the
generic name; for its anatomy see the comprehensive study made by TIMMERMANS, Ann. Jard. Bot.
Btzg 41 (1931) 65-104, t. 23-34, and the review given by SLEUMER in E. & P. Nat. Pfi. Fam. ed. 2, 20b
(1942) 332-336, f. 90-93. .
Note. Leaves of Phytocrene are similar in shape and nervation to those of various genera in the
Menispermaceae; besides the petiole emerges from a shallow cup-like thickening of the stem which also
occurs often in Menispermaceae. In the latter, however, generally the petiole is swollen in the uppermost
and maybe also in its basal part, which is never the case in Phytocrene.
KEY TO THE SPECIES
1. 3 Flower heads solitary, each on top of an axillary or slightly supra-axillary peduncle (as is always the
case with the 2 ones) on younger foliate parts of the branchlets... ...... 1. P. anomala
1. 3 Flower heads several together, composed to an inflorescence, the latter either from foliate or de-
foliate axils, or from tubercles on the stem.
2. $ Flower heads rather few, composed to a true raceme, i.e. the peduncled heads spaced along a
SCLIS MMMM Rect ee sw. eae See ic” REN ed ek eR OE ee 2. P. racemosa
2. § Flower heads numerous, composed to a spike-like panicle.
3. gd Flower heads in fascicles or glomerules, these markedly spaced at least in the lower half of the
rachis.
4. Leaves broadly oblong, base slightly cordate, flat in dry specimens. . . . . 3. P. interrupta
4. Leaves ovate, base rounded, -+ bullate by impressed nerves and veins in dry specimens.
4. P. malacothrix
3. § Flower heads in short-peduncled cymes, secondary racemes or panicles, these generally close
together (rarely spaced) along the rachis.
1971]
5. Bracts in the 3 inflorescences pérsistent, ‘aa, plumed by + spreading hairs.
6. Basal part of bracts hardly or not widened. St eee
6. Basal part of bracts widened up to 5 mm.
ICACINACEBAE (Sleumer)
81
5. P. trichura
6. P. borneensis
5. Bracts in the 3 inflorescences wanting (already gone in ‘full ‘anthesis), 0 or if persistent, filiform to
narrow-subulate, and not oopabere plumed.
7. 3 Inflorescences rather short,
lanceolate, early glabrescent.
2.5-5 oe (0.5—)1-1.5 cm. Leaves cals oblong or oblong-
7. P. oblonga
7. 3 Inflorescences elongate to caudate, i.e. markedly longer and at ‘least 2 2 cm g. Leaves oblong to
ovate-cordate, or palmate, generally tardily or hardly glabrescent.
8. Leaves normally palmate, i.e. deeply and narrowly (3—)5(—7)-lobed in flowering specimens.
8. P. palmata
8. Leaves normally entire or + broadly 3-lobed in flowering specimens (sometimes palmate in
sterile juvenile specimens).
9. 3 Inflorescences slender, c. 2 cm @. Bracts + persistent, numerous, much exserted beyond the
partial inflorescences. Drupe 4.5-8 by 1.5—2.3 cm incl. the hollow beak of the exocarp.
9. P. bracteata
9. 3 Inflorescences thickish, (2.5—)3 cm or more @. Bracts, if persistent, rather few and less ex-
serted.
10. Mature drupe 3-4(+4.5) by 1.5—2 cm incl. the hollow short beak of the exocarp; hard endo-
carp (2—)2.5-3(-3.5) cm long.
10. P. hirsuta
10. Mature drupe (5.5—)6—10(-13) cm long incl. the hollow elongate beak of the exocarp.
1. Phytocrene anomala Merr. J. Str. Br. R. As.
Soc. n. 86 (1922) 325; SLEUM. Blumea 17 (1969)
yx pe
Woody vine. Branchlets (slender, up to 5 mm g,
their distal part often curved and similar to a
tendril), petioles, peduncles and leaves (mainly on
the nerves of the undersurface) hirsute by short
greyish and longer (1-2 mm) rufous and more
spreading hairs, ultimately glabrescent. Leaves ob-
long to broadly oblong, or lanceolate, apex short-
acuminate, subacute, base slightly cordate, char-
taceous, olivaceous, glabrous or almost so except
the hirsute midrib above, the tomentum on the
undersurface rather soft to the touch, denticulate
by protruding veins in young, entire in mature
leaves, 10-22(-30) by (2.5—)3-10cm, nerves
6-8(-10) pairs, two of them + basal, short and
spreading, the other ones curved-ascending,
raised beneath only as is the coarse net of veins
and veinlets; petiole 1-2(-3) cm by 2-3(-4) mm.
— ¢ Inflorescences in globular peduncled heads,
solitary, from a little above the axils of subsequent
leaves of the distal part of a branchlet; peduncle
slender, c. 1 cm; heads covered with a pale fer-
rugineous tomentum of short bristly hairs, 5-8
mm 9. Calyx segments 4 or 5, linear-spathulate,
c. 2.5mm. Petals 4, elongate-obovate, connate
halfway to 34, hirsute dorsally, c. 3 mm. Stamens
4; filaments 1.5-2 mm; anther cells subovate-
elliptic, 1 mm. — @ Inflorescences as the ¢ ones
on young branchlets in axillary solitary heads, on
peduncle c. 2cm by 2mm. Calyx segments and
petals as in 3. Ovary ovoid, short-attenuate to a
conical very shortly lobed stigma, hirsute. Drupes
collected to heads of c. 4cm @, each drupe ovoid,
hirsute by simple bristly hairs all of the same type,
known up to 2.5 by 1.5cm, apparently larger at
full maturity, accrescent petals seen up to 6 mm.
Form, size and type of pits of the endocarp not yet
Distr. Malesia: Borneo (Upper Baram R. and
Sandakan area, Berau).
11. P. macrophylla s./.
Fig. 41.
above the hyphen indicates the number of en-
demic species in the island or area, the figure
below the hyphen the number of non-endemic
species.
Distribution of Phytocrene. The figure
Ecol. Primary and secondary lowland forest,
apparently rare.
2. Phytocrene racemosa SLEUM. Blumea 17 (1969)
236. — Fig. 42.
Tall woody climber, up to 40 m, stem c. 1.5 cm @
covered with thin longitudinally striate and finely
fissured brown bark which bears a few transverse
linear lenticels; branchlets and petioles with a short
fawn tomentum. Leaves oblong-obovate, apex
abruptly (c. 1.5cm) acuminate, acutish, base
broadly cuneate and shallowly cordate, charta-
ceous to subcoriaceous, glabrous except the nerves
which are hairy in their basal part above, + den-
sely set with short subappressed or more spreading
pale rusty hairs especially on nerves and veins and
soft to the touch beneath, edge slightly sinuate-
denticulate, or entire, 18-32 by 7-20 cm, nerves
MALESIANA
[ser. I, vol, 72
Fig. 42. Phytocrene racemosa SLEUM. a. Habit, with 3 inflorescence, x YA, b. fruit
the exocarp removed, showing the endocarp, x1, d. pitted endocarp, x1, e. cross
d JT69
» X%, c. ditto, part of
~section of stem, x | y,
(a, e MurTHYy & ASHTON § 22325, b Smytuigs § 17155, c-d§ 19513).
flat above, much Prominent beneath as is the rather by 2-4 mm. — & Racemes from tubercles of the
Coarse reticulation of veins and veinlets, nerves 2 stem or from Slightly above foliate or defoliate
basal pairs, the outer pair short, the inner one axils, all over fawny, tomentose, peduncled (1 cm)
rather straight-ascending to the middle of the heads (5-10, ¢. 5 mm 9g)
, these laxly arranged along
lamina, upper c. 4 pairs from the midrib also the 6-12 cm long rachis. Calyx segments 5, linear
rather steeply curved-ascendent; petiole 1.5-3 cm spathulate, long-haired, 2 mm. Petals 4, linear-
1971]
ICACINACEAE (Sleumer) 83
oblanceolate, free to almost the base, long-haired
dorsally, 2.5 mm. Stamens 4; anther cells 0.4 mm.
— 2 Racemes from c. 1 cm above defoliate axils,
with about 5 laxly arranged heads, fulvous-hirsute
at peduncles and the c. 4cm long rachis. Flower
heads globose, c. 5mm @g, on thick peduncles
1.5-2 cm. Calyx segments and petals as in ¢
flowers. Ovary cylindric, hirsute, c. 2mm. De-
veloped ovary or immature drupe (c. 1 cm long)
densely covered with short and shortly hirsute
emergences in the upper, and longer and longish-
hirsute ones in the lower half. Mature drupe ar-
ranged to a globose head of c. 8 cm g, each drupe
showing the same shorter and longer emergences of
the exocarp as already found in the very young
state, subelliptic-oblongoid, attenuate at both ends,
4-5 by 2-3 cm; endocarp oblongoid-ellipsoid, ob-
tuse at both ends, much compressed laterally,
smooth, with numerous almost linear rather deep
pits, which appear on the inner side as flat warts,
c. 3 by 1.5 by 1.3 cm.
Distr. Malesia:
lections.
Ecol. Mixed Dipterocarp forest, from low
elevation up to 250m, apparently very scattered.
Vern. Buah pisang munsang, Kapit.
Borneo (Sarawak), 3 col-
3. Phytocrene interrupta SLEUM. Notizbl. Berl.-
Dahl. 15 (1940) 252; Blumea 17 (1969)
236.
Climber. Branchlets slender (5-7 mm 9), deeply
striate and brownish-hirsute, older parts laxly set
with hardly pungent small emergences. Leaves
broadly oblong, apex for 3-10 mm subabruptly
acuminate, base slightly cordate, firmly charta-
ceous, glabrous above except nerves, covered
with soft hairs all over the undersurface, and with
stiffer ones on midrib and nerves, soft to the
touch, entire, 7-17 by 4-8(-9) cm, midrib and
nerves flat or a little impressed above, markedly
prominent beneath, nerves 2 basal pairs (of which
the outer pair is short, the inner one ascending to
the lower 14 of the length of the lamina), other
pinnate nerves 6-8 pairs, rather straight and sub-
parallel to each other and the inner basal pair,
reticulation of veins and veinlets fine and hardly
raised above, more coarse and much raised
beneath; petiole 2-3.5 cm by 2-3 mm, hirsutulous.
— ¢ Inflorescences spike-like, several in a fascicle
from a tubercle of the stem (c. 1.5 cm @), pendent,
the heads of flowers sessile or very shortly pe-
duncled and (2—)3-6 of them fascicled, these fasci-
cles spaced in the lower, less or not so in the upper
half of the rachis (10-30 cm by 2 mm), the whole
inflorescence covered with shorter soft and longer
stiff-brownish hairs; peduncle of flower heads
slender, up to 4mm, the basal bract subulate,
2-3 mm, apparently early going. Calyx segments
4, ovate, fleshy, concave, hirsute, 0.8 mm. Petals
3 or 4, connate to the lower 4 to 4, ovate-
oblong, subacuminate, green, -+- densely substrigose
dorsally, c. 1.5mm. Stamens 3 or 4; anther cells
ovate-elliptic, 0.4mm. — @& Inflorescences and
drupe unknown yet.
Distr. Malesia: New Guinea (Central Distr.
in ascent from Port Moresby to Mt Victoria),
once found.
Ecol. Forest at c. 450 m.
4. Phytocrene malacothrix SLEUM. Notizbl. Berl.-
Dahl. 15 (1941) 361; Blumea 17 (1969) 237.
*“Scandent. Branchlets soft-olivaceous-tomentose
in the younger parts, glabrescent below. Leaves
ovate, apex subabruptly acuminate for | cm, tip
obtuse, base rounded, thin-chartaceous, glabrous
above, softly brownish-tomentose all over beneath
especially on midrib and nerves, 16-18 by 8-10 cm,
midrib raised on both faces, nerves curved-as-
cendent, + distinctly impressed as are the veins
above (the lamina bullate in dry specimens),
much raised beneath, reticulation rather dense,
slightly prominent beneath only; petiole tomentose,
3-4 cm. 3 Spikes only known ina poorly developed
state, brownish-tomentose, solitary from the axils
of the upper leaves, the flower heads composed to
glomerules, the latter spaced on a rachis c. 5cm
long.”
Distr. Malesia: New Guinea (Morobe Disttr.:
Boana), at 1030 m, once found.
Note. Imperfectly known species, said to be
related to P. interrupta SLEUM. in the original
diagnosis, from which the description given above
was made; type material lost.
5. Phytocrene trichura RipLi. Fl. Mal. Pen. 1
(1922) 433; SLEUM. Blumea 17 (1969) 237.
Tall woody climber. Stem of branchlets shallow-
ly striate, patently rufous-setose distally as are the
petioles, corky and with numerous roundish len-
ticels (no prickles) downwards, 4-6 mm @. Leaves
ovate in circumference, whether or not 3-lobed to
14-24, lateral lobes acutish, base shallowly cor-
date, thin-coriaceous, subdensely short-tomentu-
lose on both faces initially, glabrescent and a
little rough above, soft to the touch beneath,
12-19 by 8-ll cm, palmately 5-nerved, midrib
and inner pair of basal nerves more prominent
than the other basal and the few upper lateral ones
beneath, reticulation coarse, strongly raised be-
neath; petiole 3-6 cm by 2-3 mm. — ¢ Panicles
from bare lower part of stem (1.5 cm Q), sessile,
pendulous, grey-green, squirreltail-like, 12-23 by
3.5-4 cm, formed by numerous peduncled dicha-
sia, the ultimate pedicels (3—6 mm) bearing globular
heads consisting of numerous flowers, all densely
shortly greyish-pubescent; bract of each globule
partly connate with pedicel, slender, almost fili-
form, persistent, spreading, its basal part hardly
or not widened, short-pubescent, upper part
plumose by spreading rufous 2—3 mm long hairs.
Calyx segments 4 or 5, irregularly obcuneate,
1 mm, free, apex fringed with long hairs. Petals
4, ovate-oblong, connate to the lower 4, sparsely
short-hairy or papillose outside, 1.5 mm. Stamens
4, slightly exserted; anther cells elliptic, 0.5 mm.
— 2 Inflorescences and drupe not yet known.
Distr. Malesia: Malay Peninsula (Perak, Pa-
hang, Selangor).
Ecol. Dense forest, ascending the top of lofty
trees, apparently very rare.
84 FLORA MALESIANA
[ser. I, voliay?
6. Phytocrene borneensis Becc. Malesia 1 (1877)
128; Merr. En. Born. (1921) 356; SLEUM. Blumea
17 (1969) 237. — P. porphyrea STAPF, Kew Bull.
(1906) 72.
Big woody climber. Stem robust (seen up to
2.5cm @), longitudinally ribbed (2-3 mm) and
with distant transverse linear lenticellar cracks.
Branchlets -- patently rufous-setose and striate
at distal part (3-6 mm@). Leaves ovate in cir-
cumference, simple, or subtrilobed, or deeply
obtusely 3-lobed, base cordate, edge very shortly
denticulate by protruding veins, chartaceous to
subcoriaceous, young ones soft-tomentulose all
over both faces, mature ones finally glabrous above,
+ densely covered with appressed to suberect
rather stiff hairs especially on nerves, and soft to
the touch beneath, 15—25(—30) by (8—)10-18(—22)
cm, nerves slightly sunken above, strongly
raised beneath, basal palmate nerves 7-9 pairs,
branched mainly outward, upper ones fromthe
midrib 3-4 pairs of entire, and 4-6 ones in the
central lobe of lobed leaves, all these nerves
curved-ascending, connected by numerous straight
and + transverse veins and a coarse reticulation
markedly prominent underneath only; petiole
subappressedly rufous to ferrugineous-setulose,
(3-)4-8(-12) by 24mm. — ¢ Inflorescences
pendent from old wood, tail-like as in P. trichura,
sessile, compact, consisting of numerous repeatedly
branched and shortly pedicelled dichasia, all over
covered with greyish to brownish softer and coarse,
in part hooked hairs, (12—)15-—25(—50) by 3-4(-5)
cm, most of the ultimate pedicels bearing a
narrowly to broadly subulate-caudate, (1.2—)2-3.5
cm long and at base (1—)2—5 mm broad persistent
bract, the latter covered with the same short to-
mentum and besides with longer, spreading, often
dark red or brownish hairs which give the bracts
a + plumose appearance; flowers in globular pe-
dicelled heads (3-4 mm @). Calyx segments 4 or
5, irregularly obcuneate to spathulate, distally
densely hairy outside, glabrous inside, 1 mm.
Petals 4, oblong-obovate, acutish, fleshy, slightly
to densely hairy outside only especially below,
1.5mm. Stamens 4; anther cells oblong-elliptic,
0.5mm. — 2 Inflorescence and drupe unknown.
Distr. Malesia: Borneo.
Ecol. Lowland forest, up to 1065 m.
Uses. Water from the stem is used for drinking.
Vern. Bilis akar, S. Kinabatangan.
7. Phytocrene oblonga WALL. Pl. As. Rar. 3
(1831) 12; Cat. (1832) n. 4948; WaLp. Rep. 1
(1842) 98; Bait. in DC. Prod. 17 (1873) 13;
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 592;
Kina, J. As. Soc. Beng. 64, ii (1895) 122; GAGNEP.
Fl. Gén. I.-C. 1 (1911) 839, f. 102, 4-8; RipL. FI.
Mal. Pen. 1 (1922) 432; Crars, Fl. Siam. En. 1
(1926) 275; GAGNepP. Fl. Gén. I.-C. Suppl. (1948)
757; SLEUM. Blumea 17 (1969) 237; Fl. Thail. 2
(1970) 90. — Gynocephalum oblongum (WALL.)
TrEcuL, Ann. Sc. Nat. III, 8 (1847) 149. —
Miquelia cancellata Kurz, J. As. Soc. Beng. 44,
ii (1875) 201.
Climber to top of tall trees, with rather slender
stem (up to 1.5cm g@); bark brownish, rather
rough, striate, not prickly. Branchlets densely set
with fine retrorse prickly lenticels, tip short-
pubescent. Leaves oblong or oblong-lanceolate,
-+- abruptly and shortly acuminate, acumen rather
slender and slightly curved, subacute, base cu-
neate to rounded, coriaceous, glabrous and
shining above, rather densely stiffly short-hairy
and a little rough to the touch beneath, early
glabrescent, entire or undulate-subdentate, (10-)
12-22 by (4-)5-9 cm, nerves 5-7 pairs, one of
which basal, ascending and + manifestly in-
arching before the edge, prominent as is the rather
dense reticulation beneath; petiole rugose, 1—2.5
cm by 2-3 mm. — 3 Panicles axillary, or mostly
clustered on woody tubercles on the stem or larger
branches, consisting of numerous pedicelled few-
flowered, more laxly or densely arranged um-
bellules, all over patently rusty to rufous-tomen-
tose, 2.5—-5 cm by (0.5—)1—1.5 cm; pedicels slender
to more stoutish, 2-5 mm, each subtended by a
subulate bract as long as or shorter than the
pedicel, which is generally hidden in the cluster
of flowers. Calyx segments 4 or 5, oblong-ovate,
free, 1 mm. Petals 4, almost free, oblong-ovate to
obovate, glabrous and yellow-green inside, rufous-
sericeous outside, c. 1.8mm. Stamens 4; anther
cells 0.3-0.5mm. — 2 Flowers (often diseased)
in peduncled globose heads (c. 8mm @), borne
on stem and branches, peduncle stout, set with
subulate bracts, c. 1 cm, all parts of the inflores-
cence brownish-rufous-tomentoseas are the 3 ones.
Calyx segments linear, 2 mm. Petals subspathulate,
base slightly connate, 3mm. Ovary cylindric,
apex somewhat contracted into a very short
thick style, tomentose; stigmas (2—)3—4, flattish,
glabrous, spreading. Drupes in globular orange
coloured clusters 6-9(-11)cm g, each drupe
ovoid to ovoid-oblongoid, obtuse, the very apex
only + apiculate by the style, covered especially
in the upper part with stout straight reddish-
brown subulate prickles (5-8 mm) which after
falling leave numerous circular pits (c. 1 mm g) in
the endocarp, (3—)3.5(-4) by 2-2.5 cm incl. the
prickles; exocarp coriaceous; endocarp crusta-
ceous, slightly tubercled inside, c. 2.5 by 1.5 cm.
Distr. Lower Burma, Thailand (Pattani),
Indo-China (Annam, Cochinchina); in Malesia:
Malay Peninsula (Penang to Malacca and Johore).
Ecol. Lowland forests, up to 150 m.
Uses. The leaves are used by Malay natives
for fever headache. The caustic seed is reported
to be edible.
Vern. Akar chumprai, M.
8. Phytocrene palmata WALL. PI. As. Rar. 3 (1831)
12; Cat. (1832) n. 4949; WaLp. Rep. 1 (1842) 98;
Mia. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 248;
BAILL. in DC. Prod. 17 (1873) 11; MaAst. in
Hook. f. Fl. Br. Ind. 1 (1875) 592; Becc. Malesia 1
(1877) 127; Kina, J. As. Soc. Beng. 64, ii (1895)
123; RIDL. Fl. Mal. Pen. 1 (1922) 433; Burk. Dict.
(1935) 1722; SLEuM. Blumea 17 (1969) 237; FI.
Thail. 2 (1970) 91. — Gynocephalum palmatum
(WALL.) TRECUL, Ann. Sc. Nat. III, 8 (1847) 149.
1971]
ICACINACEAE (Sleumer) 85
— P. stylocarpa GriFF. Not. (1854) 320; Ic.
(1854) t. 489, I.
Liana; stem generally not branched, younger
parts with prickles (1 mm), striate, tips rufous-
hispid. Leaves limited to the upper part of the
branches, orbicular to reniform in outline, deeply
palmately (3—)5—-(rarely —7)-lobed, lobes oblong
to oblong-obovate, acute or acuminate, base of
lamina deeply cordate, entire or shallowly wavy,
upper surface glabrous and shining, lower sur-
face densely covered with longish coarse sub-
appressed rufous to yellowish hairs especially on
midrib and nerves, soft to the touch, 15-30 cm
long, breadth about the same, main nerves 5(—7),
palmate, lateral nerves per lobe 34 pairs, curved-
ascending and looping, with more numerous
coarse transverse veins in a + right angle from
the main nerve, reticulation coarse, + deeply
impressed above, much raised beneath as are all
nerves, the lamina not rarely almost bullate in
dry specimens; petiole stout, often twisted,
rufous- to rusty-tomentose, 7—-12.5 cm. — Panicles
of 3 flowers subcylindric, axillary or from lower
part of stem, (6-)15-40 by c. 3.5 cm, all over
covered with a tomentum of short soft rufous to
ferrugineous hairs, the ultimate branches (c. 1 cm)
bearing small, 12-—15-flowered, pedicelled (2-4
mm) umbellules or heads (c. 2.5mm 9), part of
them subtended by a + caducous subulate bract
(24mm). Calyx segments 3 or 4, obovate,
hairy, 0.8mm. Petals (3—)4, oblong-lanceolate,
pale green, glabrous inside, hairy outside, c. 1.5
mm. Stamens 4; anther cells linear-oblong, 1 mm.
— @ Flowers in peduncled (c. 6mm) axillary
ovoid heads, c. lcm go. Ovary ovoid, hirsute.
Drupes numerous, collected in subglobular heads
7-10 cm 9, elongate-ellipsoid, or slightly obovoid,
much tapering to the base, apiculate by the
conical style, all over + densely covered with
yellowish shining stiff hairs, (3.5-)4-5.5 by
1.6-2(-2.5) cm; exocarp thin; endocarp bony,
rather smooth outside where the hairs are gone,
laxly pitted outside, low-papillate inside respec-
tively.
Distr. Thailand (Pattani); in Malesia: Malay
Peninsula, Sumatra (Palembang; Riouw Arch.:
P. Bintan).
Ecol. Hilly lowland forest, up to 500m.
Vern. Akar pisang pisang bulu, Malay Penin-
sula, képajang akar, Palembang, M.
9. Phytocrene bracteata Wat. PI. As. Rar. 3
(1831) 12; Cat. (1832) n. 4947; Wap. Rep. 1
(1842) 98; Batt. in DC. Prod. 17 (1873) 12;
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 592;
Becc. Malesia 1 (1877) 127; Merr. En. Born.
(1921) 357, p.p.; Ript. Fl. Mal. Pen. 1 (1922)
432; Baker /. J. Bot. 62 (1924) Suppl. 21; Merr.
PI. Elm. Born. (1929) 172; Daut, J. Arn. Arb.
23 (1952) 275 (pollen); Steum. Blumea 17 (1969)
238; Fl. Thail. 2 (1970) 91. — Gynocephalum
bracteatum (WALL.) TrécuL, Ann. Sc. Nat. III,
8 (1847) 149. — P. macrocarpa Grirr. Not.
(1854) 322; Ic. (1854) t. 487 (fl. 9) & t. 488 (fr.).
High climbing liana; stem up to 2.5cm 9,
striate, setose and minutely prickly in younger
parts. Leaves broadly ovate, often 3-lobed and
obscurely dentate then, apex acute, base cordate,
coriaceous, glabrous and a little asperulous above,
the nerves excepted, densely covered all over the
undersurface with short brownish hairs and soft
to the touch, and the nerves moreover set with
bristle-like stiffer hairs beneath, 10—-20(—25) by
7-15(-20) cm, 7 pairs of basal palmate nerves
and 2-3 pairs of upper nerves, curved-ascending,
slightly sunken above, prominent beneath, re-
ticulation low though distinct on the undersurface;
petiole hairy and bristly, + torted, 3—-7(—9) cm by
2-3 mm. — Panicles of 3 flowers mainly axillary,
rarely from older branches, solitary or 2-3, spike-
like, all over shortly greyish to brownish tomentose,
10-20 by c. 2cm, rachis and peduncles of main
branches slender, ultimate branches consisting of
minute clustered, (3—)6—10-flowered pedicellate
(2-5 mm) umbellules, each pedicel subtended by a
filiform appressed-hispid, not properly plumed +
persistent bract (1 cm), which is partly connate
to the pedicel and exserted from the mass of
flower heads. Calyx segments 3—5, free, obcuneate,
villous outside, hardly 0.6 mm. Petals generally 3
(rarely 4), ovate-oblong or lanceolate, slightly
hairy outside only, practically free, finally re-
flexed, 1-1.2mm. Stamens 3(-4); anther cells
oblong-elliptic, 0.6 mm. — 2 Flowers in oblong to
spherical peduncled heads, either solitary or 2-3
per axil. Calyx segments 4, spathulate. Petals 4,
short-strigose outside as are the calyx segments.
Ovary strigose; style very short, 3-—4-partite.
Drupes in pendulous globose clusters up to 20 cm
@, each drupe ovoid-oblongoid, tapering to both
ends, densely covered with appressed long soft
yellowish bristles, 4.5-8 by 1.5-2.3 cm at full
maturity; exocarp fleshy especially in the upper
part of the drupe which bears a short beak of the
former style; endocarp ligneous, ovoid-oblongoid,
up to 4 by 2 by 1.5 cm, rather smooth and minutely
pitted outside, faintly tubercled inside. Seed testa
sparsely and shallowly pitted.
Distr. Thailand (Surat, Nakawn Sritamarat),
in Malesia: Sumatra (Eastcoast, Palembang),
Malay Peninsula, Lingga Arch., possibly in
Borneo.
Ecol. Lowland hillside woods and hedges.
Vern. Akar pisang pisang, M (as P. palmata).
Note. All investigated Bornean specimens ap-
parently belonging to P. bracteata bear immature
inflorescences, or are sterile, and could possibly
belong to P. borneensis.
with spiny tubercles 1 mm. Branchlets + deeply
10. Phytocrene hirsuta BL. Rumphia 4 (1849) 37;
Mus. Bot. Lugd.-Bat. | (1849) 42; WALP. Ann. 2
(1851) 22; Mio. Fl. Ind. Bat. | (1856) 797; Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 247; BAILL. in DC.
Prod. 17 (1873) 12; Becc. Malesia 1 (1877) 127;
Koorp. Minah. (1898) 629; Hatt. f Med.
Rijksherb. 1 (1910) 14; Steum. Blumea 17 (1969)
238, non TeysM. & BINN. Hort. Bog. (1866) 207.
— P. crinipes Batt. ex Bureau, Mon. Bign.
(1864) 146, in text, nom. nud. — P. minahassae
86 FLORA MALESIANA
{ser. I, vol. 73
Koorpb. Minah. (1898) 394, nom. nud. — P. dasy-
carpa (non MiqQ.) DAHL, J. Arn. Arb. 33 (1952)
275 (pollen).
Climber; stem (lcm or more g) smooth.
Branchlets slender, set with longish rufous sub-
setose and + spreading hairs, older parts also
laxly with very short pale retrorse prickles. Leaves
ovate, or ovate- or elliptic-oblong, apex acuminate,
the midrib characteristically protruding up to 8
mm beyond the leaf margin as a hirsute breakable
thread in young leaves, base rounded to slightly
cordate, firmly chartaceous, finally glabrous above
except the nerves, remaining hirsute or tomentose
by shorter and longer rufous to ferrugineous
hairs and soft to the touch beneath, entire or
shallowly sinuate-dented, or obscurely repand
especially in the lower half, 12-17 by 6-9 cm,
midrib and nerves very slightly immersed or flat
above, prominent beneath, nerves 2 pairs palmate-
ly from the base of the lamina (outer pair short,
inner one rather straight ascending to + half
the length of the blade), other 3(-4) pairs of nerves
from the upper 24 of the midrib generally more
curved, reticulation dense and finely raised above,
coarser and more prominent beneath; petiole
thickened at base, rufous-hirsute, 3.5-4 cm by c.
2mm. — ¢ Inflorescences in spike-like panicles,
solitary in the axils of lower leaves, up to 25 by 3
cm, composed of numerous shortly peduncled
cymes, the ultimate peduncle (2-3 mm) bearing
numerous flowers in a head (c. 4mm g) and
provided at base with a subulate + persistent
bract (1-1.5mm), the whole inflorescence cov-
ered with shorter greyish and longer rufous,
bristle-like and + spreading hairs. Calyx seg-
ments 3 or 4, connate below, obcuneate, hirsute,
1.3mm. Petals 3 or 4, almost free, oblong, dirty
yellow, with a few hairs inside, hirsute outside,
c. 2mm. Stamens 3 or 4; anther cells 0.5 mm. —
2 Flowers in heads c. 8 mmg, on stout peduncle
2-3 cm, generally from old wood. Calyx segments
and petals as in 3 flowers. Ovary glabrous below,
hirsute above; stigmas 4 or 5, short, acuminate.
Drupes collected in subglobular masses, each
drupe ovoid-oblongoid to -ellipsoid, laterally
much compressed, apex + broadly attenuate,
densely set with retrorse setose bristle-like stiff
hairs, 3-4(—4.5) by 1.5-2 by 1.5 cm, incl. the short
hollow beak of the exocarp; endocarp hard, with
minute scattered roundish pits, (2-)2.5-3.5 by
1.5-2 cm.
Distr. Malesia: Celebes (incl. Buton I.), Mo-
luccas (Buru).
Ecol. Primary and secondary lowland forest,
up to 800m, also in rather dry places, even on
limestone rocks.
Vern. Kunét ri sasap, Minahasa (TI.), pitji pitji
lantjeh, Makassar, sapét kulo, Minahassa (Tt.).
11. Phytocrene macrophylla (BL.) BL. Rumphia 4
(1849) 36; Mus. Bot. Lugd.-Bat. 1 (1849) 41, f. 7;
WALP. Ann. 2 (1851) 22; ZoLi. Syst. Verz. 2
(1854) 112; Mig. Fl. Ind. Bat. 1 (1856) 796;
Suppl. 1 (1860) 137; Ann. Mus. Bot. Lugd.-Bat. 3
(1867) 247; BaiLt. in DC. Prod. 17 (1873) 10;
Hist. Pl. 5 (1874) 282 (fl.); Becc. Malesia 1 (1877)
127; C. B. Ros. Bot. Z. 47 (1889) 645 (wood anat.);
ENGL. in E. & P. Nat. Pfl. Fam. 3, 5 (1893) 255,
f. 140; RaciB. Flora 87 (1900) 13 (foliar biology);
Hocur. Pl. Bog. Exs. (1904) 17; Bull. Inst.
Bot. Btzg 19 (1904) 39; ibid. 22 (1905) 114;
BAcK. Schoolfi. Java (1911) 229 (‘macrocarpa’);
Koorp. Exk. Fl. Java 2 (1912) 533; Koorp.-
ScHum. Syst. Verz. 1, Fam. 162 (1912) 6; WaArB.
Pflanzenwelt 2 (1916) 348, f. 231 A, 1-9, f. 232
(phot.); Doct. v. LEEUWEN, Zoocecidia (1926) 332
(galls); UittT1eN, Réc. Trav. Bot. Néerl. 25 (1929)
464 (veg.); AMSH. in Back. Bekn. FI. Java (em.
ed.) 6 (1948) fam. 135, p. 8; Back. & BAKH. f.
Fl. Java 2 (1965) 62; SLEuM. Blumea 17 (1969)
239. — Gynocephalum macrophyllum Bu. Bijdr.
(1825) 483; TRECUL, Ann. Sc. Nat. III, 8 (1847)
148. — P. gigantea WALL. Phil. Mag. N.S. 3 (1828)
223, nom. nud.; Pl. As. Rar. 3 (1831) 11, t. 216;
Cat. (1832) n. 4946; WaALP. Rep. 1 (1842) 98; BL.
Rumphia 4 (1849) 36; Ann. Mus. Bot. Lugd.-Bat.
1 (1849) 41; Zoxt. Syst. Verz. 2 (1854) 112;
GrirF. Not. (1854) 329; Ic. (1854) t. 490, f. II
(fl. ¢ & anat.); Mig. Fl. Ind. Bat. 1 (1856) 796;
SCHNIZL. Ic. 3 (1857/65) t. 172, f. 1-21; BAILL. in
DC. Prod. 17 (1873) 9; Mast. in Hook. f. FI. Br.
Ind. 1 (1875) 591; Kurz, J. As. Soc. Beng. 44,
ji (1875) 156; For. Fl. Burma 1 (1877) 241;
Becc. Malesia 1 (1877) 127; BRANpis, Ind. Trees
(1906) 152. — Kadsura blancoi AZAOLA in Blanco,
Fl. Filip. ed. 2 (1845) 594; ed. 3, 3 (1879) 118. —
Gynocephalum giganteum (WALL.) TRECUL, Ann.
Sc. Nat. III, 8 (1847) 149. — P. calicarpa GrirF.
Not. (1854) 327; Ic. (1854) t. 490, f. I (fr.). —
Gynocephalum luzoniense LLANOS, Rev. Progr.
Cienc. 15 (1865) 55. — P. dasycarpa Mig. Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 248, t. 7, as to
fruits only. — P. luzoniensis (LLANOS) BAILL.
Adansonia 10 (1872) 281; in DC. Prod. 17 (1873)
10; Hist. Pl. 5 (1874) 283 (fr.); F.-VILL. & NAVEs
in Blanco, FI. Filip. ed. 3, 4 (1880) 86; F.-VILL.
Nov. App. (1880) 46; CERON, Cat. Pl. Herb.
Manila (1892) 46. — P. blancoi (AZAOLA) MERR.
Philip. J. Sc. 2 (1907) Bot. 432; Spec. Blanc.
(1918) 237; En. Philip. 2 (1923) 492; Daun, J.
Arn. Arb. 33 (1952) 275, f. 75 (pollen). — P.
obovoidea MerrR. Philip. J. Sc. 10 (1915) Bot.
322; En. Philip. 2 (1923) 492. — P. forbesii
BAKER f. J. Bot. 62 (1924) Suppl. 21.
See for further synonyms under the varieties.
KEY TO THE VARIETIES
1. Drupe (7—)7.5-10(-13) by 3-3.5 cm; endocarp
(4-)5-6 by 2-2.5 cm. . 1. var. macrophylla
1. Drupe (5.5—)6-6.5(-7) by 1.5—2 cm.
2. Endocarp 44.5 by c. 1.5 cm. 2. var. caudigera
2. Endocarp c. 2.5 by 1cm. . 3. var. dasycarpa
1. var. macrophylla.
High climbing, often much branched shrub or
liana, up to 25 m; stem up to 10 cm g, not spiny.
Branchlets striate, younger parts with appressed
or somewhat spreading shorter and longer rufous
or ferrugineous hairs, and besides + densely set
1971]
with pale conical prickles (0.5 mm), older parts
glabrescent and smooth, the cork with transverse
lenticellar cracks. Leaves of young shoots often
orbicular with rather deeply cordate base, and
broadly and + obtusely 3—5-lobed, such of older
parts generally ovate-oblong from a subcordate
base and subacuminate, tip obtuse, firmly sub-
coriaceous, entire, glabrous above, the nerves
excepted, the whole undersurface with a tomentum
of short, weak rather pale rusty (partly almost
stellate) hairs, and less numerous longer strigose
rufous hairs especially on nerves and veins, soft
to the touch, + glabrescent at full age, 14~30(—33)
by 9-12(-20) cm, palmately 5—7-nerved, outer
pair(s) of nerves short, inner one(s) ascending to
14 and even 34 of the length of the lamina, other
3-4 pairs from upper part of the midrib, all nerves
almost flat above, well raised beneath, reticula-
tion dense and very fine above, moderately
raised beneath; petiole 5-15 cm by 2-3 mm. —
3 Inflorescences spike-like panicles, from old
branches or stem, generally several together,
cylindric, dense or rarely more loose, all over short-
ly greyish-brownish-tomentose, up to 25 cm long
and (2.5—)3(—5) cm 9, composed of lateral racemes
or panicles up to 4 cm long which bear the flowers
in heads +6 mm @. Calyx segments 3-5, + nar-
rowly obcuneate, apex truncate, hirsute outside,
0.7-1.2 mm. Petals 3 or 4, light green to yellowish,
connate — halfway, lobes recurved, densely ap-
pressedly hairy outside, 1.5-2 mm. Stamens 3 or
4 (in the same inflorescence!); anther cells ovate-
elliptic, c. 0.4mm, often devoid of pollen. —
2 Inflorescences collected in large, solitary or
fascicled heads, each head on a thick peduncle
(3-10 cm)and c. 1.5 cmg, all over short-tomentose.
Calyx segments as in g flower. Petals 2.5-3.5
mm. Ovary columnar, tomentose with forward
directed hairs, c. 4mm, stigma 2—4-lobed. Fruit
heads 20-25cm go. Drupe obovoid-oblongoid,
(7—)7.5—10(-13) cm long incl. the hollow beak and
the basa] narrowed part of the exocarp, 3-3.5 cm
wide, slightly variously compressed or round in 9,
rusty-tomentose by normally forward directed
hairs (sometimes reversely so in specimens from
Sumatra and the Philippines), these hairs + ap-
pressed, stiff, shining, easily going; endocarp
oblongoid, bony, laterally flattened, the crests
less marked on the flattened side, with numerous
roundish pits (1mm), (4-)5-6 by 2-2.5 by
(1.5—)2 cm.
Distr. Burma; in Malesia: Sumatra, Java,
Philippines (Luzon, Polillo, Samar, Biliran, Leyte,
Boho!, Mindanao, Palawan).
Ecol. Lowland thickets and forests from sea-
level to c. 1200 m, also in secondary and in
swamp forest, in the Philippines also in Diptero-
carp forest, scattered.
Galls. Docrers vAN LEEUWEN mentions a
ICACINACEAE (Sleumer) 87
leaf-gall caused by a gall-midge.
Uses. Fruits said to be used on Palawan as
‘talang’, i.e. fish poison. In Java used against
stomach pain and tumors.
Vern. Akar pisang, Sumatra Westcoast, M,
labu kumbung silai, olor delos, Simalur, areuj
pitjung tjéléng, hondjé bu-ut areuj, undjéwut, S;
Philippines: olo-olo, Bag.
2. var. caudigera (SLEUM.) Blumea 17 (1969) 240.
— P. caudigera SLEUM. Notizbl. Berl.-Dahl. 15
(1940) 253; Herne, Pfl. d. Samml. Clemens
Kinabalu (1953) 57.
Differs from var. macrophylla by generally
smaller leaves 7-15 by 5-8 cm, and mainly by the
oblongoid, slightly 4~-6-angled minor drupes,
(5.5—)6-6.5(-7) by c. 2cm; endocarp 44.5 by
c. 1.5cm, the pits much smaller than in var.
macrophylla. The hairs on the drupe are strigose
and retrorse.
Distr. Malesia: Borneo, only known from the
Mt Kinabalu area, in forest around Dallas at
915-1370 m.
3. var. dasycarpa (Mriq.) SLEUM. Blumea 17
(1969) 240. — P. dasycarpa Miq. Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 248, t. 7, the fruit excluded
(which is var. macrocarpa); BAILL. in DC. Prod.
17 (1873) 11; BEcc. Malesia 1 (1877) 127; Koorpb.
Minah. (1898) 394; Hocnure. Pl. Bog. Exs. (1904)
n. 92; Bull. Inst. Bot. Btzg 22 (1905) 114; Ic. Bog.
2 (1906) t. 108; Koorp.-ScHuM. Syst. Verz. 3
(1914) 74; Koorp. Minah. Suppl. 2 (1922) t. 49;
Suppl. 3 (1922) 25. — P. hirsuta (non BL. 1849)
TeysM. & BINN. Hort. Bog. (1866) 207. — P.
ovalifolia KoorD. Minah. (1898) 394, nom. nud.,
629, descr.; KOORD.-SCHUM. Syst. Verz. 3 (1914)
74; Koorp. Minah. Suppl. 2 (1922) t. 50; Suppl. 3
(1922) 26. — P. tinosporifolia Koorp. Minah.
(1898) 394, 629; Koorp.-ScHuUM. Syst. Verz. 3
(1914) 74; Koorp. Minah. Suppl. 2 (1922) t. 51;
Suppl. 3 (1922) 26.
Differs from var. macrophylla by its smaller
drupes, (5.5—)6—6.5 by 1.5-2 cm; endocarp c. 2.5
by 1 cm.
Distr. Malesia: Celebes.
Ecol. Lowland forest, up to 350 m.
Vern. Apo, Tontembuan, samadan, Tonsea,
sapet kulo, Tt., s. putih, Tl., tali tali utan, Bt.
Excluded
Phytocrene loheri Merr. Philip. J. Sc. 7 (1912)
Bot. 293; En. Philip. 2 (1923) 492, mixtum, nom.
illeg.
The leaves belong to a Menispermacea (probably
Hypserpa), and the loose fruits to Phytocrene
macrophylla var. macrophylla, cf. SLeuM. Blumea
17 (1969) 241.
Excluded
Leucocorema Rip. Trans. Linn. Soc. I, Bot. 9 (1916) 29 = Trichadenia Tuw. (Flacourtiaceae), cf.
Steen. Bull. Jard. Bot. Brux. 27 (1957) 114.
Pentastira Riv. Trans. Linn. Soc. Il, Bot. 9 (1916) 27 = Dichapetalum Tuou. (Dichapetalaceae),
cf. Leennouts, Fl. Mal. I, 5 (1957) 305.
are pene
Tn a
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i wry,
LOPHOPYXIDACEAE (H. Sleumer, Leyden)
1. LOPHOPYXIS
Hook. f. Ic. Pl. 18 (1887) t. 1714; SLEUM. Blumea 16 (1969) 322. — Combretopsis
K. Scu. in K. Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 69. — Treubia PIERRE ex
BoerL. Handl. 1, 2 (1890) 445. — Fig. 1.
Scandent or liana-like shrubs or small trees, with interxylary bast elements,
watch-spring tendrils (modified leaves or subtending leaves of inflorescences),
and umbrella-like branching. Leaves spiral, simple, serrulate to crenulate, stip-
ulate. Flowers small, regular, monoecious, in glomerules on the branches of
loose axillary panicles. Sepals 5, valvate, shortly united basally, persistent. Petals
5(-6), much smaller than the sepals, free. Stamens 5(-6), opposite to the sepals,
with filiform filaments and subglobose introrse, almost basifixed anthers, alter-
nating with 5(—6) oppositipetalous cordate glands, these in the ¢ + adnate to the
subtending petal, and in the 2 + concrescent into a 5(—6)-lobed disk (the glands
or lobes opposite the ovary cells); pollen grains + ellipsoidal, tricolporate, exine
reticulate. Ovary superior, conical, shallowly 5-ribbed, 5(—4)-celled, with 5(-4)
sessile subulate stigmas; ovules 2 per cell, pendulous, apical, axile, anatropous,
epitropous, bitegmic, each surmounted at the micropyle by a small obturator-
like appendage coming from the funicle. Fruit obovoid or ellipsoid, indehiscent,
fusiform, 1-locular, 1-seeded, with 5 broad stramineous wings. Seed oblong, with
endosperm; embryo erect, with oblong cotyledons and a short erect radicle.
Distribution. Monotypic, in Malesia, Melanesia, and Micronesia. Fig. 2.
Anatomy. Stem. Anomalous secondary growth: ENGLER, Sitz. Ber. Preuss. Ak. Wiss. Berl. 18 (1893)
265-266; HANDA, Bot. Mag. Tokyo 54 (1940) 41-47; PFEIFFER, Rev. Sudam. Bot. 10 (1951) 3-6; SLEUMER
in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 393, f. 118.
The young stem has five ribs with a continuous xylem cylinder enclosing a pentagonal pith with a
central portion of thick-walled parenchyma cells and a marginal area of thin-walled cells, erroneously
referred to by ENGLER (/.c.) as intraxylary phloem. This thin-walled tissue may become caducous, at least
in herbarium specimens. Between the ribs the secondary xylem has numerous vessels; in the ribs the
vessels are narrower and scarcer. Vessel perforations are simple. Through anomalous activity of the
cambium, 5 phloem strands become enclosed within xylem in the young shoot. Later interxylary phloem
is formed as continuous bands alternating with secondary xylem. The phloem is stratified into soft and fi-
brous portions. Axial xylem parenchyma is scarce and paratracheal (only seen in young twigs). The
ground tissue of the xylem is composed of fibres with numerous minutely bordered pits. Rays vary from
1—6-seriate in the young stem. The outer phloem is surrounded by a cylinder of fibres and stone cells.
Cork arises in the layer below the epidermis.
The anatomy of the leaf has hitherto never been described. The petiole shows a strongly incurved arc
of separate vascular bundles and two additional latero-dorsal bundles as seen in transverse section
through the distal end. The vascular system forms a closed flattened cylinder in the midrib. The stomata,
confined to the abaxial surface, are paracytic. Crystals are present as solitary rhomboids and clusters.
The hairs are unicellular.
To evaluate the taxonomic significance of the vegetative anatomy of Lophopyxis with regard to the
affinities of the genus more research is still needed. The only straightforward conclusion to be drawn at
present is that Lophopyxis is anatomically entirely different from Gouania (Rhamnaceae), to which AIRY
SHaw (in Willis Dict. ed. 7, 1966, 668) related it. Gouania differs e.g. in having anomocytic stomata, sty-
loids, unlignified perivascular fibres and exclusively narrow rays. Dr. C. R. METCALFE (Kew) kindly put
slides and anatomical information of Lophopyxis and Gouania at my disposal. — P. BAAs.
Taxonomy. This genus was tentatively ascribed to the Euphorbiaceae by Hooker/., but removed from
this family by Pax (1890). ENGLER (1893) accommodated it as a distinct subfamily Lophopyxidoideae
within /cacinaceae, from which I rejected it in 1942. HUTCHINSON (Fam. Flow. PI. 1959) placed it in the
Celastraceae.
Its gross morphology, wood anatomy, embryology and pollen morphology is well known and it is
(89)
90 FLORA MALESIANA [ser. I, vol. 7#
Fig. 1. Lophopyxis maingayi Hook. f.a. Habit, x 14, a’. showing domatia in the axils of lateral nerves,
x 214, b. 2 flower, <7, c. ovary, cross-section, 7, d. ovary, longit. section, x7, e. f flower buds, x2YV,
J. 3 flower, 5, g. stamen, front view, x 15, g’. stamen, back view, 15, A. infructescence, x 3/5, i. fruit,
x 8/5 (a-d, h-i KANEHIRA (1938), partly altered, e-g SLEUMER (1942)).
1971]
LOPHOPYXIDACEAE (Sleumer) 91
now apparent that it should be placed within the Geraniales-Sapindales-Celastrales. It seems, however,
that it does not fit in any of the established families of these orders. Its relation to Rhamnaceae, suggested
by SHAW (1966) rests on a superficial habit similarity with Gouania, as shown by BAKHUIZEN & VAN
STEENIS, Fl. Mal. Bull. 21 (1966) 1426 (see also sub Anatomy).
The best solution is to regard it as the type of a family of its own, as has been casually proposed by
VAN TIEGHEM (1897) and PIERRE (1897), and formally by PFEIFFER (1951). I have this more fully explained
in my precursory paper in Blumea 16 (1969) 320.
1. Lophopyxis maingayi Hook. f. Ic. Pl. 18 (1887)
t. 1714; Fl. Br. Ind. 5 (1888) 673; PAx in E. & P.
Nat. Pfi. Fam. III, 5 (1890) 117; ENGL. ibid.
(1893) 238, 257; Hari. f. Med. Rijksherb. 1
(1910) 9; Riot. FI. Mal. Pen. 1 (1922) 435;
Stoor. Bull. Jard. Bot. Btzg III, 7 (1925) 364;
Hottn. & H. J. Lam, Blumea 5 (1942) 205, f. 7;
SLeum. in E. & P. Nat. Pfil. Fam. ed. 2, 20b (1942)
393; Fl. Males. I, 5 (1954) 63; Blumea 16 (1969)
322, with extensive bibliogr. — Combretopsis
pentaptera K. Scu. in K. Sch. & Hollr. Fl. Kais.
Wilh. Land (1889) 69. — Treubia combretocarpa
PIERRE ex BoERL. Handl. 1, 2 (1890) 445. — L.
pierrei BOERL. l.c. 673, nom. ill.; HEYNE, Nutt.
Pl. 1 (1950) 987. — L. schumannii BoERL. Handl.
1, 2 (1890) 674, nom. ill. — L. combretocarpa
(BoERL.) ENGL. in E. & P. Nat. Pfl. Fam. III, 5
(1893) 257. — L. pentaptera (K. SCH.) ENGL.
Sitz. Ber. K6n. Preuss. Ak. Wiss. (1893) 265,
t. 2, f. 6 & 7; SLEUM. in E. & P. Nat. Pfl. Fam. ed.
2, 20b (1942) 393, f. 117 (habit) & 118 (wood anat.,
embryol.); DAHL, J. Arn. Arb. 36 (1955) 160, 161
f. 2 & 2A (pollen); PEEKEL, Fl. Bismarck Arch.
MS p. 1083, fig. — Homalium gilgianum Laut. in
K. Sch. & Laut. Nachtr. (1905) 320. — Sinapistrum
RumpH. Herb. Amb. 5 (1747) 73, t. 39, f. 1. —
Fig. 1.
Vigorous woody vine or climbing shrub,
3-8(-30) m; stem up to 7cm @g, bark whitish.
Branches virgate, youngest parts longitudinally
grooved, with elliptic lenticels in the grooves, pu-
berulent. Leaves ovate to oblong, apex + acu-
minate, acutish, base cuneate to obtuse, rarely
rounded or subcordate, subequal, chartaceous to
subcoriaceous, caducous-puberulous especially
at the nerves beneath, the hairs persisting in the
nerve axils as domatia, serrulate-crenulate or
crenate, sometimes subentire, 8—18(—24) by
4-8(-10) cm, nerves 1(-2) basal or slightly supra-
basal, and 3—4 upper pairs, all arched and steeply
ascending, veins and veinlets densely reticulate,
slender but prominent on both faces; petiole c. 10
by Imm. Stipules small, knob-like. Axillary
branchlets metamorphosed into strong woody
tendrils coiled only at the end, often bearing a bud.
Panicles \oose, composed of a few axillary or
terminal! spike-like racemes, pendent, puberulous,
10-25 cm. Flowers solitary, or mostly crowded into
glomerules, these spaced along the slender to
filiform rachis. Pedicels very slender, up to 2 mm.
Bract at the base of the inflorescences often meta-
morphosed into a weak flat completely coiled
tendril. Sepals ovate, greenish white or yellowish,
hairy on both sides, c. 1.5 mm. Petals ovate, thin,
c. 1mm. Disk yellowish. — 3 Flowers: filaments
hairy, 2mm; anthers subglobose, 0.5mm; ru-
diment of ovary subglobose, shallowly 5-ribbed,
hairy. — 2 Flowers: ovary ovoid, whitish-yellowish
puberulent, 2mm. Capsule obovoid-ellipsoid in
outline, 5-winged, green, later dark brown,
caducous-pubescent, 2.2-3(-3.5) by 1.3-1.8 cm;
wings chartaceous, 5-8 mm wide, with irregularly
crenulate margin; pedicel short, subtended by the
non-accrescent calyx. Seed 1, subcylindrical,
acuminate, lengthwise grooved, 12-15 by 5-6 mm.
Fig. 2.
Distribution of Lophopyxis maingayi
Hook. f.
Distr. Micronesia (Palau Is.), Melanesia (New
Ireland, Duke of York I., New Britain, Solomon
Is.), in Malesia: Malay Peninsula (Penang to
Malacca), North Borneo, W. Central and N.
Celebes, Moluccas (Talaud Is., Ceram, Sula Is.,
Ambon), New Guinea. Fig. 2.
Ecol. Straggling climber in canopy or edge of
primary lowland forest, both in well drained and
in swampy riverine forest, in littoral forest and
even sometimes in the mangrove, also in disturbed
gully forest or forest regrowths, seaside scrub;
scattered, though locally not too rare, from sea-
level up to c. 300 m, often on alluvial soil.
Uses. The rather hard stem splits easily into
pieces, and apparently for this reason is used in
New Britain for tying thatch. RUMpHIus under
his ‘Sinapistrum’ says, that crumpled leaves give
a strong smell of mustard, and are used to cure
ulcers.
Vern. Akar bélimbing hutan, Brunei (Kina-
batangan), simpuru, Celebes, taburuh, Talaud, tali
sasawi, Ambon (sec, RUMPHIUS).
REP owe as
_ 4
CARDIOPTERIDACEAE (H. Sleumer, Leyden)
1. CARDIOPTERIS
[WALL. ex] Roy Le, Ill. Bot. Himal. Mts (1834) 136, in text; em. BL. Rumphia 3
(1847 or 1849) 205; ibid. 4 (1849) t. 177; HAssk. Flora 30 (1847) 110, in adnot.;
Nat. Tijd. N. I. 10 (1855) 64; ENGL. in E. & P. Nat. Pfl. Fam. 3, 5 (1893) 239,
257 (‘Cardiopteryx’); BAKH. f. & STEEN. FI. Mal. Bull. 15 (1960) 725; Taxon 11
(1962) 28; BACK. & BAKH. f. FI. Java 2 (1965) 62. — Sioja BUCH.-HAM. ex LINDL.
Nat. Syst. ed. 2 (1836) 82, nom. nud. — Peripterygium Hassk. Tijd. Nat. Gesch.
Phys. 10 (1843) 142; Cat. Hort. Bog. (1844) 235; SLEuM. in E. & P. Nat. Pfl. Fam.
ed. 2, 20b (1942) 400; AMsH. in Back. Bekn. Fl. Java (em. ed.) 6 (1948) fam. 135,
p. 9. — Fig. 1.
Sinistrorsely twining herbs with white milky juice. Leaves spirally arranged,
simple or lobed to varying degrees, cordate, palmatinerved, long-petioled, glabrous
as is the stem, exstipulate. Flowers bisexual, or polygamous (andromonoecious),
small, subsessile, in unilateral repeatedly forked cincinni, composed of loose
axillary panicles, ebracteate. Calyx deeply (4~-)5-partite, lobes imbricate in bud,
whether or not slightly accrescent in fruit, persistent. Petals (4-)5, caducous,
lower half forming a widely funnel-shaped corolla, lobes imbricate in bud.
Stamens (4-)5 inserted on the upper part of the corolla tube, alternate with its
lobes; filaments very short, glabrous; anthers medifixed, introrse; pollen grains
oblate, triangular in polar view, tricolporate. Disk absent. Ovary oblong-ovoid,
subquadrangular (rudimentary in 3), 1-celled; styles 2, one with a deep longitudinal
groove, and two inequal ovate rather obtuse distal divisions, accrescent in fruit and
becoming linear and succulent then, the other short-curved, capitate at apex,
deciduous; ovules 2 (often one of them aborted), pendent from apex of cavity,
anatropous, with dorsal raphe. Fruit indehiscent, compressed, with 2 longitudinal
broad and transversely striate stramineous wings, obovate-elliptic to orbicular in
outline, apex emarginate, crowned by the columnar accrescent soft and green
stigma, base very shortly or hardly (Mal.), sometimes elongately contracted into
a kind of stipe. Seed 1, linear, sulcate; testa thin; embryo minute, conical, in top
of granular fleshy albumen.
Distr. Two spp., one in SE. Asia and W. Malesia, and one in E. Malesia.
Ecol. Climber on edge of (mostly secondary) forest and thickets, in open places as limestone rocks.
Uses. The leaves are eaten as a vegetable.
Taxon. Cardiopteris was conceived as type of a monogeneric family by BLUME (1847 or 1849) and
R. Brown (1852); it was considered to constitute a subfamily of the Jcacinaceae by ENGLER (1893) and
Post & Kuntze (1904), and again as a distinct family within the Celastrales by KiNG (1893), WILLIAMS
(1915, as Peripterygiaceae), GAGNEPAIN (1910, 1911), SLEUMER (1942, as Peripterygiaceae), HUTCHINSON
(1959) and TAKHTAJAN (1966).
The pollen of the genus shows no distinctive features against [cacinaceae and resembles very much that
of the Afro-Malagasian genus Cassinopsis SONDER.
Note. The family name Cardiopteridaceae is derived from the original spelling Cardiopteris used by
WALLICH and later by BLUMe and others. ENGLer changed the name to Cardiopteryx, which would be
more correct as an allusion to the winged fruit, a substitute name, however, which cannot be used ac-
cording to the Code; for this reason, the family name ‘Cardiopterygaceae’, proposed recently, has not
been used.
(93)
94 FLORA MALESIANA [ser. I, vol. 73
Fig. 1. Cardiopteris moluccana BL. a. Habit, with 3 inflorescence, x 14, b. 3 flower, corolla and stamens,
15, c. 2 flower, corolla and ovary, x15, d. flower, lateral view with calyx and corolla, x7, e. flower
calyx, x10, f. infructescence, x 14 (a—f BLUME).
1971]
CARDIOPTERIDACEAE (Sleumer) 95
KEY TO THE SPECIES
1. Leaves membranous, generally + deeply and + acutely 3—5(-9)-lobed, very rarely (sub)entire. In-
florescence 1—2(-3)-forked, cincinni rather few-flowered.
1. C. quinqueloba
1. Leaves firmly chartaceous, entire. Inflorescences with more numerous forks, cincinni rather many-
flowered. ......
1. Cardiopteris quinqueloba (Hassk.) HAssk. Nat.
Tijd. N. I. 10 (1855) 64. — Peripterygium quin-
quelobum Hassk. Tijd. Nat. Gesch. Phys. 10
(1843) 142; Cat. Hort. Bog. (1844) 235; SLEUM.
in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 400, f.
120 F—G; Amsu. in Back. Bekn. FI. Java (em. ed.)
6 (1948) fam. 135, p. 9. — C. javanica BL.Rumphia
3 (1847 or 1849) 206, nom. illeg.; ibid. 4 (1849) t.
177, f. 1A; BACK. & BAKH. f. Fl. Java 2 (1965) 63.
— C. lobata R. Br. [in WALL. Cat. (1847) n. 8033]
in Benn. & Br. Pl. Jav. Rar. (1852) 246, t. 49,
nom. illeg.; Mig. Fl. Ind. Bat. 1 (1856) 799;
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 597;
Kurz, J. As. Soc. Beng. 44, ii (1875) 157; Becc.
Malesia 1 (1877) 131; Kinc, J. As. Soc. Beng.
64, ii (1895) 131; GaAGNep. FI. Gén. I.-C. 1
(1912) 849; Back. Schoolfl. Java (1911) 230;
Koorp.-ScHum. Syst. Verz. 1, Fam. 162 (1912)
6; Merr. En. Born. (1921) 357; Ripi. Fl. Mal.
Pen. 2 (1923) 464; Crars, Fl. Siam. En. 1 (1926)
276; Burk. Dict. (1935) 456; KANJILAL c.s. FI.
Assam 1, 2 (1936) 254; HENDeERS. J. Mal. Br. R.
As. Soc. 17 (1939) 59; GAGNeEP. FI. Gén. I.-C.
Suppl. (1948) 759; HeyNe, Nutt. Pl. 1 (1950) 987;
Wu & WAnG, Act. Phytotax. Sin. 6 (1957) 284;
LarsEN, Dansk Bot. Ark. 23 (1963) 71. — C.
rumphii BatLt. Adansonia 10 (1872) 280, nom.
illeg., incl. var. lobata BAILL. et var. subhamata
BAILL. /.c. 281; DC. Prod. 17 (1873) 26; SCHEFF.
Ann. Jard. Bot. Btzg 1 (1876) 14. — C. platycarpa
GAGNneEP. Not. Syst. 1 (1910) 198; Fl. Gén. I.-C. 1
(1911) 847, f. 104, p.p., f. 105. — Peripterygium
platycarpum (GAGNEP.) SLEUM. Notizbl. Berl.-
Dahl. 15 (1940) 257; in E. & P. Nat. Pfl. Fam. ed.
2, 20b (1942) 400. — C. moluccana (non BL.)
Wu & Wana, Act. Phytotax. Sin. 6 (1957) 284.
Much branched with twining terete stems, which
often preserve their freshness after the fall of the
leaves and are loaded with fruits then, 2—5(-9) m.
Leaves widely spaced, polymorphous, broadly
ovate in outline, with a cordate base, generally +
deeply or irregularly 3—5(—9)-lobed, apex generally
acutish, lateral lobes acute or obtuse, very rarely
entire or almost so, herbaceous, caducous, pale
green, (4-)6-12(-17) by (3—)4~-7(-16) cm, main
nerves 7-9 radiating from apex of the petiole,
slightly raised beneath, veins rather obscure;
petiole (3—)5—12 cm. Panicles solitary on 3—5(—10)
cm long peduncles, 1—2(—3)-forked, cincinni rather
few-flowered, glandular-puberulous or almost gla-
brous, 5-10 cm, accrescent in fruiting stage.
Flowers distant from each other along rachis, on
short pedicels (1 mm, accrescent to 5 mm in fruit).
» Calyx lobes 2mm. Petals white, 2.5(-3.5) mm.
Fruit obovate-elliptic in outline, the wings in-
cluded, apex emarginate, base cuneate and sudden-
ly contracted to a kind of foot up to 2mm, on
the very base of which the persistent calyx is found,
2. C. moluccana
wings glossy yellow or light stramineous colour,
tipped by the accrescent style which remains
green and fleshy for a long time.
Distr. SE. Asia (W. Bengal and Assam to
Burma, Thailand, Indo-China and Yunnan);
in Malesia: N. & NE. Sumatra, Malay Peninsula
(Kedah, Perlis, Perak), SE. Borneo (twice found),
Java (incl. also Bawean & Kangean Is. and Ma-
dura), Lesser Sunda Is. (Lombok, Sumbawa,
Sumba, Alor), Celebes (incl. also Saleijer Is.),
Moluccas (Tenimber Is.).
Ecol. Primary and secondary (also seasonal,
deciduous) forest edge, scrub jungle, bamboo
forest, hedges around villages, open waste places,
even on open limestone rock, generally at low
elevations, rarely up to 1000 m, scattered.
Uses. Locally the leaves are used as a vegetable.
Vern. Angi anginan, iri iri, kanjar kawang,
parianom, rindengan, serintil, sobowengi, wiwi wiwo,
J, oke oke, Mad., gambas kawaong, riru ruruan,
saburung, S, kokrah, Alor.
Fig. 2. Distribution of Cardiopteris. 1. C. quinque-
loba (Hassk.) HAssk., 2. C. moluccana BL.
2. Cardiopteris moluccana BL. Rumphia 3 (1847
or 1849) 207; ibid. 4 (1849) t. 177, f. 1 B, f.2 A & B,
incl. var.; Mig. Fl. Ind. Bat. 1 (1856) 799; Becc.
Malesia 1 (1877) 131; K. Scu. Notizbl. Berl. 2
(1898) 130; K. Scu. & Laut. FI. Schutzgeb. (1900)
418; Merr. Philip. J. Sc. 2 (1907) Bot. 423;
SCHELLENB. Bot. Jahrb. 58 (1923) 176; Merr. En.
Philip. 2 (1923) 493; Heyne, Nutt. Pl. 1 (1950) 987.
— Dioscorea sativa L. Amoen. Ac. 4 (1759) 133,
pr. pl. Amboin. — C. rumphii BALL. var. blumeana
BaiLt. Adansonia 10 (1872) 281, incl. var. inte-
grifolia Baw. Lc. 280; DC. Prod. 17 (1873)
26. — C. lobata R. Br. var. moluccana (BL.)
Mast. in Hook. f. Fl. Br. Ind. 1 (1875) 597. — C.
lobata (non R. Br.) Becc. Nuov. Giorn. Bot. Ital.
9 (1877) 100, t. 8; F.-ViLL. Nov. App. (1880) 46;
96 FLORA MALESIANA
[ser. I, vol. 74]
F. M. BarLey, Queensl. Agric. J. 24 (1910) 20.
— C. celebica R. BR. ex Koorpd. Minah. (1898)
392, nom. nud. — Aspidocarya kelidophylla K. SCu.
& Laut. Fl. Schutzgeb. (1900) 313, sec. DIELS,
Menisp. (1910) 320. — Peripterygium moluccanum
(BL.) SLEUM. Notizbl. Berl.-Dahl. 15 (1940) 257;
in E. & P. Nat. Pfl. Fam. ed. 2, 20b (1942) 400,
f. 120 A-E; Dant, J. Arn. Arb. 36 (1955) 161
(pollen). — Olus sanguinis RuMPH. Herb. Amb. 5
(1747) 482, t. 180. — Fig. 1.
Leaves ovate-cordate, entire, acutely acuminate,
base -- deeply cordate, firmly chartaceous, edge
often slightly recurved in dry specimens, up to
24 by 22cm. Panicles generally 3- or more-
forked, cincinni rather rich-flowered, flowers
usually close together. Otherwise as C. quinqueloba.
Distr. Malesia: Celebes, Philippines (Luzon,
Panay?, Negros, Samar, Bohol, Catanduanes,
Mindanao), Moluccas (Buru, Ambon (type),
Ternate, Ceram), New Guinea, and New Britain
(Gazelle Peninsula).
Ecol. Climber in tall rain-forest or forest edge,
also in secondary vegetation and in native gardens,
generally in the lowland, ascending to 1460 m in
New Guinea, scattered.
Uses. A decoction of the stem is used against
hepatitis in Ternate. The leaves are eaten as a
vegetable.
Vern. Matta matta, tamatta, Makassar, telin
teli, Menado, uge jabba, Ternate, uta lala, u. turi,
Ambon; Philippines: bangogan, Bik., gurisan,
sagumati, Bag., lila, Buk., tagulauai, P. Bis.;
New Guinea: foyomangeni, Wapi (Miwaute),
kehunghe, Mekeo (Maipa), swalow, Buang.
Note. According to Mique. also in Bali,
which is apparently erroneous.
OCHNACEAE (A. Kanis, Leyden)
Woody plants, very small undershrubs to tall trees. Leaves distichous or spirally
arranged, stipulate, simple, glabrous; midrib prominent on either side. /nflorescen-
ces 1- to many-flowered, cymose, racemose, or thyrsoid, bracteate; pedicels
articulate. Flowers actinomorphic, bisexual (rarely functionally polygamous).
Sepals 5, free or a little connate at base, quincuncial, persistent. Peta/s 5—10, free,
contort, caducous. Staminodes 0-~. Stamens 5—10—~; anthers basifix, + latrorse
and dehiscing lengthwise, or with 1-2 apical pores. Carpe/s 2-S—10(—15), superior,
free with 1 ovule, or fused with 2—~ ovules per carpel; styles fused, basigynous or
epigynous; stigmas free or + fused. Fruit(s) a drupe(s), berry, or capsule. Seeds
l—-~, small or large, sometimes winged, with or without albumen.
Taxonomy. There is little doubt that the family of the Ochnaceae represents a natural one among the
more primitive in the Guttiferales (= Clusiales or Theales s. 1.). Nonetheless, there are striking differences
between the genera, even at first sight. It is not difficult to arrange them in a few distinct, supra-generic
taxa. A supposed natural system, as far as relevant to the Malesian genera, is as follows:
Subfamily Ochnoideae
eeCONaAR 1s (hw aed! Ae vd oe aa Sabtribe Ochninge 25 Sogine
2. Brackenridgea
Subtribe Ouratinae 3. Gomphia
Subfamily Sauvagesioideae
EO a ay a rrr Perm
Mribe SAUVAGESIFAE.. 2. 2 5 iw ww ew cl et we eo ) Subtribe Sauvagesiinae.5. Neckia
6. Indovethia
7. Schuurmansiella
8. Schuurmansia
Distribution. About 30 genera and c. 250 spp. through the tropical, rarely subtropical countries
(S. Africa, N. India), chiefly in S. America and Africa. In Malesia 8 genera and 13 spp.; absent from Java
and the Lesser Sunda Islands.
In subfam. Ochnoideae, the monogeneric tribe Elvasieae and the genus Ouratea are restricted to S.
America. Of the 3 genera of the Ochneae mentioned above, only Brackenridgea is chiefly Malesian.
Ochna and Gomphia are principally African, both reaching the western part of Malesia with one species.
In subfam. Sauvagesioideae, the monogeneric tribe Lophireae occurs only in Africa, whereas the sub-
tribe Luxemburgiinae is restricted to S. America. The 5 relevant genera mentioned above are almost
purely Malesian. Of the subtribe Sauvagesiinae, Sinia Dies and Indosinia VIDAL are found in continental
SE. Asia, whereas the majority of the genera is restricted to S. America, only Sauvagesia L. occurring
both in S. America and Africa.
There is an interesting parallelism between some of the Sauvagesioideae, which are more or less restric-
ted to sandstone areas in W. Malesia, and related genera found in similar areas in northern S. America
(Roraima flora).
Ecology. The Malesian genera are restricted to everwet areas, except Ochna which is adapted to a
seasonally dry climate and is deciduous. They are more or less adapted to poorer, sandy or peaty soils
in relatively undisturbed areas, except Schuurmansia which is a genus of pioneer species.
Pollination. There is no literature on this subject. Pollination probably takes place by insects,
because of the brightly coloured and (sometimes?) scented flowers. The colours of the petals are yellow
in Ochna and Brackenridgea sect. Notochnella, mostly white, creamy, or tinged purple in other genera,
sometimes dark purplish red in Schuurmansia. Only Ochna has flowers c. 3 cm across. Flowers of other
genera usually do not exceed 1 cm, but they are often combined to conspicuous inflorescences. The
undershrubs of Neckia, however, have solitary flowers with relatively small, early caducous petals.
Dispersal. The bluish or black, 1-seeded fruits of the Malesian Ochneae are probably mainly dis-
persed by birds. In Ochna and Brackenridgea they are contrasting with a purple calyx and torus. The
Euthemideae have red or white berries on a dark red calyx, which are probably also dispersed by birds
(see Riptey, Disp. 1930). The Malesian Sauvagesieae have many-seeded capsules. Their seeds do not
show adaptations to a special mode of dispersal, except those of Schuurmansia with two wings like
propeller blades. The latter characteristic points to wind dispersal, which fits the pioneer-like nature of
the genus concerned. The sepals in Neckia are also turning dark purple in fruit; the meaning of this
phenomenon is not understood.
(97)
98 FLORA MALESIANA [ser. I, vol. 72
Morphology. From the description of the family and the key to the genera it will be clear that the
morphological differences between subfamilies and tribes are considerable. A discussion on these dif-
ferences I gave in my thesis (Blumea 16, 1968, 8-15). A short note should be made here on the inflores-
cence types, as these may not always be easily understood.
The inflorescences in subfam. Ochnoideae are all considered to be of a thyrsoid nature, viz racemes with
cymose branches. Those of Gomphia serrata are lateral and terminal, bearing terminal flowers which
makes sympodial growth of the vegetative branches necessary. Those of Ochna integerrima have also
terminal flowers, but they are terminal on short side branches and monopodial growth of the main
branches remains possible. The inflorescences of Brackenridgea spp. are terminal, sometimes also lateral.
A terminal flower is lacking and monopodia! growth of the rachis to a vegetative shoot is still possible,
although not equally frequent in all species. The cymose branches are very much shortened here, espe-
cially in sect. Brackenridgea where flowers are almost sessile on the rachis. In some species there are
several branches per inflorescence, each bearing 3(—5) flowers. In other species there are only a few
branches, each bearing 7 or more flowers in pseudo-umbels. In sect. Notochnella the cymose branches
are not shortened so much, whereas the bracts on the rachis sometimes have a more leaf-like appearance.
The inflorescences in subfam. Sauvagesioideae are more of a paniculate nature. They are profusely branch-
ing in Schuurmansia, but in the other genera most branches are very much shortened, flowers and
fruits of different age standing closely together. In Neckia the rachis is bearing several bracts, but only
one (terminal?) flower.
Anatomy. The Ochnaceae are characterised by the presence of cortical bundles without resin canals
(GILG, Ber. Deut. Bot. Ges. 11, 1893, 20-25). The subfam. Ochnoideae shows typical ‘cristarque’ cells in
branches and leaves (VAN TIEGHEM, Bull. Mus. Hist. Nat. Paris 8, 1902, 266-273). For other data, see
METCALFE & CHALK, Anat. Dicot. 1 (1950) 104, 108, 333-338, 340, f. 76, 77, and DECKER, Phytomor-
phology 16 (1966) 39-45.
Palynology. The pollen of the Ochnaceae only show interesting differences at generic or higher level,
but they do not differ fundamentally. See ERDTMAN, Pollen Morph. Pl. Tax. (1952) 290, and MULLER,
Rey. Palaeobot. Palynol. 9 (1969) 149-173.
Phytochemistry. No data available. Some species are used locally for medicinal purposes, because
of bitter components of unknown nature.
Uses. No economically important applications of Malesian spp. have been recorded. For properties
of the wood of some species, see under Ochna integerrima and Gomphia serrata.
Note. The present revision is based on my precursory treatment of Indo-Pacific Ochnaceae in Blumea
16 (1968) 1-83.
KEY TO THE GENERA
1. Stamens 10-ov. Carpels (3—)5-10(-15), free. Fruits 1-5, 1-seeded drupes on a swollen torus. Leaves
distichous.
2. Anthers opening with 2 apical pores. Stipules intrapetiolarly united. Inflorescences with (1—)3-cy,
++ remote flowers, usually thyrsoid, sometimes simple cymes.
3. Stamens 12-0; ovaries (3—)5—10(-15); embryo straight. Leaves without an intra-marginal nerve.
1. Ochna
3. Stamens 10; ovaries 5; embryo curved. Leaves with a distinct intra-marginal nerve. 3. Gomphia
2. Anthers opening with 2 longitudinal slits. Stipules free. Inflorescences of umbelloid appearance with
© flowers in conferted, cymose clusters of 3 or more... . eee = Brackenridgea
1. Stamens 5. Carpels 2—5, fused. Fruit a more-seeded berry or capsule; ‘torus not distinctly enlarged in
fruit. Leaves alternate, not distichous.
5. Ovary 5-carpelled, S-celled. Fruit a berry. Anthers opening by 1 apical pore. Leaf margin always
stiffly, though sometimes very finely, denticulate. . . . . . 4, Euthemis
5. Ovary 3-carpelled, 1-celled. Fruit a capsule. Anthers opening by 2 longitudinal slits. Leaf margin
entire or (bi)serrulate.
6. Seeds not winged. Fruit opening with 3 valves. Inflorescences simple, or compound and (nearly)
all branches shortened. Leaves evenly spaced. Leaf margin (bi)serrulate. Shrubs or undershrubs.
7. Inflorescences axillary; the rachis bearing a varying number of bracts, but only 1 flower. Under-
shrubs, up to 1 m, but often much smaller... . . > eeSiNeckia
7. Inflorescences terminal (or pseudo-axillary by sympodial growth), many-flowered. Shrublets or
shrubs, up to 7 m high.
8. Staminodes 10, in 1 whorl. Fruit subglobose. Leaf blades oblanceolate, up to 35 cm long.
6. Indovethia
8. Staminodes ov, in more than 1 whorl, those of the inner whorl larger. Fruit fusiform. Leaf blades
linear oblong, up to 17 cmlong.. . . . . . . 7. Schuurmansiella
6. Seeds winged. Fruit opening with 3 longitudinal slits ‘under the persistent style. Inflorescences
much-branched panicles. Leaves distinctly tufted. Leaf margin entire, glandular dotted. Treelets,
Haually, NOL exccedine 15m... s 2 2 sels swe eee Bk ee ee alee
1971] OCHNACEAE (Kanis) 99
1. OCHNA
LinnE, Gen. Pl. ed. 5 (1754) 229; Sp. Pl. 1 (1753) 513; KANIs, Blumea 16 (1968)
22, 83. — Diporidium WENDL. f. in Bartl. & Wendl. Beitr. Bot. 2 (1825) 24; O.K.
Rev. Gen. Pl. 1 (1891) 104, emend. illeg., incl. typ. Ochna. — Discladium TieGH.
Bull. Mus. Hist. Nat. Paris 8 (1902) 214, nom. illeg. — Polythecium TieGH. Ann.
Sc. Nat. Bot. VIII, 16 (1902) 196, 366. — Pleopetalum TieGH. Bull. Mus. Hist.
Nat. Paris 9 (1903) 163. — Polythecanthum TieGH. Ann. Sc. Nat. Bot. IX, 5
(1907) 160, 175. — Fig. 1.
Shrubs or treelets, sometimes undershrubs. Stipules small, intrapetiolarly united,
caducous. Leaves shortly petioled, chartaceous or subcoriaceous; nerves curved
upward, especially near the margin, not joining; veinlets + at right angles to the
nerves near the midrib and joining in irregular secondary nerves, + transverse
near the margin. /nflorescences lateral or terminal thyrses with a terminal flower;
peduncle + persistent, bearing many, small, distichously conferted, caducous
bracts at base, leaving a distinct annulus of scars; pedicels filiform, articulate.
Flowers with + hemispherical torus, distinctly tumid and red in fruit. Sepals 5,
greenish, accrescent and turning red in fruit. Peta/s 5-10, in 1-2 whorls, yellow.
Stamens ~ in 2 or more whorls; filaments subterete; anthers opening with 2
apical pores. Ovaries 5-10(-15), obovoid; ovule atropous; stigmas as many as Ova-
ries, on short branches or + united. Fruits 1-3(—5), greenish, turning black when
ripe.
Distr. The majority of the species is found in Africa, south of the Sahara, and in Madagascar. In
India and Ceylon 4 spp. occur, one of these ranging from Assam to Indo-China, Hainan, the Nicobar Is.,
and Malesia: in the North of the Malay Peninsula.
Ecol. Adapted to a seasonal climate, on poorer soils, below 1500 m. Dispersal by birds because of
conspicuous black fruits on red torus and calyx (RIDLEY, Disp. 1930, 419).
1. Ochna integerrima (Lour.) MErRR. Trans. Am.
Phil. Soc. n.s. 24, 2 (1935) 265, emend.; KANIS,
Blumea 16 (1968) 36; Fl. Thail. 2 (1970) 25. —
Elaeocarpus integerrimus Lour. Fl. Cochinch.
(1790) 338. — O. wallichii PLANCH. in Hook.
Lond. J. Bot. 5 (1846) 650; KiNG, J. As. Soc.
Beng. 62, ii (1893) 231; Rip. J. Str. Br. R. As.
Soc. nm. 59 (1911) 83; Crais, FI. Siam. En. 1
(1931) 244; Burk. Dict. 2 (1935) 1569. — O.
crocea GriFF. Not. Pl. As. 4 (1854) 463; Kurz,
J. As. Soc. Beng. 40, ii (1871) 49, err. in syn.
Gomphia sumatrana JACK; KING, ibid. 62, ii (1893)
233. — O. grandis Riv. J. Str. Br. R. As. Soc.
n. 59 (1911) 83; FI. Mal. Pen. 1 (1922) 365. —
Ouratea crocea (GrirF.) Burk. Dict. 2 (1935)
1614; Kew Bull. (1935) 318, p.p., quoad typus. —
Fig. 1.
Deciduous undershrub, shrub, or treelet up to
12m and 45cmg. Stipules 5-8 by 2-3 mm.
Leaves mostly obovate-oblong or (obovate-)
lanceolate, rarely obovate or linear-lanceolate,
6-20(-25) by 2-7 cm, mostly acuminate, some-
times acute or obtuse at apex, mostly acute,
sometimes obtuse at base, margin finely denticu-
late; petiole 2-5 mm. Inflorescences compound,
many-flowered; rachis 14—114(-4) cm; branches
1-3-flowered, monochasial; pedicels 2-4 cm, in
fruit up to S5cm, the basal 2-8 mm persistent.
Torus -1 mm high, 114-244 mm 4g, in fruit up
to 6mm high, 10 mmg@. Sepals 5, ovate to ovate-
oblong, 10-16 by 4-9 mm. Petals 5S—6(-10), obo-
vate, 15-25 by 8-15 mm, tapering at base or
subunguiculate. Stamens (25—)30-60(-75); fila-
ments 214~7 mm, unequal, the outermost longest;
anthers 4-6 by 0.4-0.8 mm. Ovaries 6-10(-15),
0.7-1.1 by 0.5-0.7 mm; style 10-15 by c. 4% mm,
in fruit up to 20 mm; stigmas sometimes on up to
1 mm long branches. Fruits mostly 2-3, up to 11
by 8 mm.
Distr. NE. India, E. Pakistan, Burma, Anda-
man and Nicobar Is., Thailand, Laos, Cambodia,
Vietnam, Hainan, in Malesia: Malay Peninsula
(Peninsular Thailand, Perlis, Kedah, Langkawi
Is.).
Ecol. From sea-level up to 1200m in hilly
country, in moist or dry, deciduous forests, often
of a mixed Dipterocarp type, on loamy, sandy, or
rocky soils. Tall specimens are found near river
banks, small shrubs near seashores. Flowering
shortly before or during development of new leaves:
in the northern part of the area mainly in Febr.
and March, in the southern part generally a
little earlier, but less restricted, especially in the
Malay Peninsula.
100 FLORA MALESIANA [ser. I, volege
7
‘4
QJ
Ja
Fig. 1. Ochna integerrima (Lour.) MERR. a. Fruiting twig, x 7, b. inflorescence, x 24, c. stamen, x4,
d. gynoecium, x4, e. fruit, x2 (a & e RipLey 15746, b-d GARRETT 1346).
1971]
Uses. The bark tastes bitter and yields a diges-
tive tonic (Cochinchina). The wood is recorded
as used for huts in the Andamans.
Properties of wood. Light brown, hard, close-
grained and brittle.
Excluded
Ochna decaisnei TrEGH. Bull. Mus. Hist. Nat.
Paris 8 (1902) 47-49. — Diporidium decaisnei
OCHNACEAE (Kanis)
101
TieGu. Ann. Sc. Nat. Bot. VIII, 16 (1902) 356 =
O. mauritiana LAMK, fide KaANis, Blumea 16
(1968) 80.
A species based on one collection by RIEDLE,
erroneously recorded for Timor. The specimen
must have been collected during Capt. BAUDIN’s
expedition on I’Ile de France (Mauritius) on the
way to Timor. It must have been mislocalized later
on.
2. BRACKENRIDGEA
A. Gray, New Gen. Pl. (1853) 5, preprint of Proc. Am. Ac. Arts Sc. 3 (1857) 51;
KANIs, Blumea 16 (1968) 41. — Campylopora TigGH. Bull. Mus. Hist. Nat. Paris 8
(1902) 547. — Notochnella T1EGH. /.c. 549. — Fig. 2.
Trees or treelets. Stipules small, free, often more or less laciniate, caducous.
Leaves shortly petioled, chartaceous, glossy above, nerves strongly curved to the
apex, often some of the lower ones partly parallel to the margin, the higher ones
joining successively, veinlets branching, -++ transverse. Inflorescence thyrsoid, but
of umbelloid appearance, made up of simple or compound, distichously arranged,
shortened cymes, the rachis often growing on vegetatively after flowering; bracts
small, broadly linguiform to triangular, + laciniate, + caducous, often many at
base of inflorescence, leaving a distinct annulus of scars; pedicels filiform, + ac-
crescent and turning red in fruit, articulate at base. Flowers with + hemispherical
torus, distinctly tumid and red in fruit. Sepals 5, accrescent, fleshy and red in fruit.
Petals 5(-10), white or yellow. Stamens 10 (or ~); filaments subterete; anthers
dehiscing from the apex downwards by longitudinal slits. Ovaries 5(—10), obovoid;
ovule camptotropous, epitropous, -_ annularly curved around 2 connecting
intrusions of the endocarp; stigma small. Fruits 1-2(—5), greenish, turning black
or almost so when ripe.
Distr. 2 or 3 spp. in tropical eastern Africa and Madagascar, probably forming a distinct section;
in Malesia 4 spp. in 2 sections are found; a related species occurs in NE. Queensland and Fiji.
Ecol. Confined to the everwet tropical areas, up to c. 1000 m. Dispersal mainly by birds because of
conspicuous, black fruits on red torus and calyx (Guppy, Obs. Nat. Pacif. 2, 1906, 569; RipLey, Disp.
1930, 265). The fruits are also capable of floating because of two air-filled spaces between exocarp and
endocarp; this was recorded from the Kapuas R. (BEccARI, Wand. Borneo 1904, 187) and from the New
Guinea seadrift (HEMSLEY, Bot. Chall. Exp. 3, 1885, 289, t. 54). No specimen has ever been collected in
beach forest.
KEY TO THE SPECIES
1. Petals 5. Stamens 10. Ovaries 5. (Sect. Brackenridgea).
2. Inflorescences never axillary, but sometimes terminal on short side-branches.
3. Inflorescences made up of many-flowered cymes, the pedicels in 2 or more tiers. Leaves mostly
Pae-eOvemiongs:. 2 i 5. .edne .ote oho ee fe ie ina. 1. B. hookeri
3. Inflorescences made up of 3(—5)-flowered cymes, the pedicels in 1 tier. Leaves mostly 4-12!4 cm long
2. B. palustris
2. Axillary inflorescences always present when in fertile state... .....4..-. 3. B. forbesii
1. Petals 5—7-(10). Stamens 10-45. Ovaries 5-10. (Sect. Notochnella)...... .
1. Section Brackenridgea
Brackenridgea A. Gray. — Campylopora TieGH.
Cymes much conferted; peduncle and branches much shortened. Flowers of |
102
FLORA MALESIANA
[ser. L vol. bs
cyme flowering atdlontbotety: Corolla regularly 5-merous, white. Stamens 10,
in | whorl. Ovaries 5.
Distr.
Chiefly Malesian with 2 spp. in the Malay Peninsula and Borneo, one reaching Sumatra,
Palawan, and Celebes, the other reaching the Andamans and Ko Chang; another sp. in New Guinea.
Also 1 sp. in NE. Queensland and Fiji.
1. Brackenridgea hookeri (PLANCH.) A. GRAY,
New Gen. PI. (1853) 6, preprint of Proc. Am. Ac.
Arts Sc. 3 (1857) 51; Furtapo, Gard. Bull. Sing.
19 (1962) 182: KaANis, Blumea 16 (1968) 45;
Fl. Thail. 2 (1970) 27. — Gomphia hookeri
PLANCH. in Hook. Lond. J. Bot. 6 (1847) 3;
Mia. FI. Ind. Bat. 1, 2 (1859) 675, err. ‘G. gla-
berrima PLANCH.’; BENN. in Hook. f. Fl. Br. Ind.
1 (1875) 525; KING, J. As. Soc. Beng. 62, ii (1893)
233, excl. var. corymbosa; RipL. Fl. Mal. Pen. 1
(1922) 366. — Ochna hookeri (PLANCH.) O.K.
Rey. Gen. Pl. 1 (1891) 106. — B. perakensis
TieGcH. Ann. Sc. Nat. Bot. VIII, 16 (1902) 396.
— Gomphia corymbosa (KING) RiDL. J. Str. Br.
R. As. Soc. n. 54 (1910) 33, p.p. excl. typus;
FI. Mal. Pen. 1 (1922) 367. — Ouratea hookeri
(PLANCH.) BuRK. Kew Bull. (1935) 318; Dict. 2
(1935) 1615. — B. denticulata FURTADO, Gard.
Bull. Sing. 19 (1962) 183. — Fig. 2b1.
Tree, up to 33m, 1m g. Leaves oblong to
lanceolate, 714-20 by 214-6 cm, obtuse to acute,
sometimes acuminate at apex, acute, often +
tapering at base, margin entire, + acicular
denticulate in young treelets; petiole 5-12 mm.
Inflorescences terminal, made up of many-flowered
shortened, sessile cymes, flowering successively or
simultaneously, the rachis often not growing on
vegetatively after flowering; pedicels 10-15 mm,
up to 20mm in fruit, those of one cyme fixed
close together in c. 3, + distinct tiers. Torus c. 1
mm high, 1 mm g, in fruit up to 3 mm high, 5mm
9. Sepals ovate to obovate, 4-5 by 114-2 mm.
Petals ovate to obovate, 314-5 by 114-134 mm.
Filaments 1-114 mm; anthers 114-2 by c. 4% mm.
Ovaries c. 0.7 by 0.5mm; style 114-3 mm, in
fruit up to 5mm. Fruits up to 614 by 514 mm.
Distr. India: Andamans, and Thailand: Ko
Chang (NE. Gulf of Thailand); in Malesia: Malay
Peninsula and Borneo.
Ecol. From sea-level up to 750 m, in kerangas
forests on sandy soils, on dry hillocks in swampy
forests, in primary lowland Dipterocarp forests
and in hill forests.
Vern. Mal. Pen.: bunga kélat mérah, bunga
maskam, kayu luru, Malacca; Borneo: émpodat,
Sarawak, sémukau, P. Madijang.
2. Brackenridgea palustris BARTELL. Malpighia 15
(1901) 165, t. 10; FurtTaApDo, Gard. Bull. Sing. 19
(1962) 183; Kanis, Blumea 16 (1968) 46; FI.
Thail. 2 (1970) 27. — B. hookeri (PLANCH.) A.
Gray var. leucocarpa SCHEFF. Nat. Tijd. N. I. 32
(1873) 411. — Gomphia hookeri PLANCH. var.
corymbosa KiNG, J. As. Soc. Beng. 62, ii (1893)
233. — B. serrulata BARTELL. Malpighia 15 (1901)
163, t. 9; FURTADO, Gard. Bull. Sing. 19 (1962)
184. — B. kingii TieGH. Ann. Sc. Nat. Bot. VIII,
16 (1902) 395. — B. corymbosa (KING) TIEGH.
lic. 395. — B. rubescens TiEGH. lI.c. 396. — Gom-
phia corymbosa (KiNG) RIDL. J. Str. Br. R. As.
Soc. n. 54 (1910) 33, p.p.; Fl. Mal. Pen. 1 (1922)
367. — Ochna forworthyi ELMER, Leafl. Philip.
Bot. 5 (1913) 1823; Merr. En. Philip. 3 (1923)
68. — B. foxworthyi (ELMER) FURTADO, Gard.
Bull. Sing. 19 (1962) 184. — Fig. 2.
Tree, up to 30m, 1.2mg. Leaves (ovate-)
oblong to (ovate-)lanceolate, 4-1214 by 114-5 cm
(up to 20 cm long on young treelets), mostly acute
to acuminate, sometimes obtuse at apex, rounded
to acute, often + tapering at base, margin entire,
+ acicular denticulate in young treelets; petiole
3-10 mm. Inflorescences terminal, made up of
3(—5)-flowered, shortened cymes, sessile, or with
up to 14 cm long peduncle, flowering simul-
taneously, sometimes 2 inflorescences of different
stages in close succession, the 5—10(—25) mm long
rachis growing on vegetatively, sometimes branch-
ing in the lower parts; pedicels 8-15 mm, up
to 20 mm in fruit, those of one cyme fixed close
together in one tier. Torus c. % mm high, 1 mm g,
in fruit up to 4mm high, 6 mm g. Sepals ovate to
oblong, 314-6 by 1144-3mm. Petals ovate to
obovate-lanceolate, 314-7 by 114-3 mm. Filaments
(i4-)1-2(-214) mm; anthers 114-3 by c. 4% mm
Ovaries c. 0.7 by 0.5 mm; style (1-)24%4-4 mm, +
accrescent. Fruits up to 8 by 6 mm.
Distr. Malesia: Sumatra, Malay Peninsula,
Borneo, Philippines (Palawan), and Celebes.
KEY TO THE SUBSPECIES
1. Anthers 114-2 mm long. Style 244-4 mm long
during anthesis.. . . . 1. ssp. palustris
1. Anthers 2-3 mm long. Style 114-244 mm long
during anthesis.
2. Filaments c. 2 mm long during anthesis.
Ovaries c. 0.7 by 0.5mm.. 2. ssp. foxworthyi
2. Filaments c. 14 mm long during anthesis.
Ovaries c. 0.5 by 0.3mm. . 3. ssp. kjellbergii
1. ssp. palustris. — All synonyms except Ochna
foxworthyi ELMER.
Cymes 3-flowered, sessile, up to 5-flowered and
with up to 14 cm long peduncle in young treelets.
Sepals 34-514 by 1144-3 mm. Petals 344-6 by
114-3 mm. Stamens with 1-2 mm long filaments;
anthers 114-2 mm long, up to 3mm in young
treelets. Ovaries c. 0.7 by 0.5 mm; style 24444 mm
during anthesis. Fruits up to 8 by 6 mm.
Distr. Malesia: Sumatra, Mentawai Is.,
Banka, Billiton, Malay Peninsula, Borneo.
Ecol. Usually found in the lowlands, but oc-
1971]
OCHNACEAE (Kanis)
103
Fig. 2. Brackenridgea palustris BARTELL. a. Flowering twig, < 24, 5. part of twig with scars of flowers and
bracts arranged in simple cymes, x 4 (b’. ditto of B. hookeri (PLANCH.) A. GRAY, scars of many flowers
arranged in shortened, cymose, partial inflorescences, « 4), c. petal from bud, d. flowerbud, sepals and
petals removed, e. flower, petals and anthers shed, sepals removed, all x 6, f. fruiting flower, x2 (a-b & e
RAHMAT SI TOROES 4187, c—d ditto 4162, f IBOET 214; b’ PAlE 13589).
casionally up to 1000m, reported from peat-
swamp forests and from kerangas forests on sandy,
sometimes rather rocky soils, with humic podsols.
Uses. The wood is reported twice as being
used in house-building (Malacca, Sarawak).
Vern. Sumatra: majang majang, mampat,
rampat dahan, séniang, M, kaju barat laut, k.
galugus badak, k. ludé, k. saholat, k. topa topa,
Kota Pinang Distr.; madu luai, mensulung kaju,
Banka; ménsolongang, Billiton; Mal. Pen.: lidah
mura, Pahang, péndorah, chéharahan, Malacca;
Borneo: mata undang, timur bésih, P. Madjang.
2. ssp. foxworthyi (ELmMerR) KaANis, Blumea 16
(1968) 48. — Ochna foxworthyi ELMER.
Cymes 3-flowered, sessile. Sepals 5-6'4 by 2-3
mm. Petals 5-7 by 114-2mm. Stamens with
134-214 mm long filaments; anthers 2-3 mm long.
Ovaries c. 0. 7 by 0.5 mm; style 1'4-2'14 mm long
during anthesis. Fruits up to 5 by 4mm.
Distr. Malesia: Philippines (Palawan).
Ecol. At sea-level, along river in forest, and
from 150m on rocky hillside near river-bank.
3. ssp. kjellbergii KANnis, Blumea 16 (1968) 48.
Cymes 3-flowered, sessile. Sepals 4-514 by
114-2 mm. Petals 4-414 by 134-214 mm. Stamens
with 14-34 mm long filaments; anthers 214-3 mm
long. Ovaries c. 0.5 by 0.3 mm; style 1-2 mm long
during anthesis. Fruits up to 5 by 4mm.
Distr. Malesia: Celebes.
Ecol. At sea-level in swamp and from 400 m
at the edge of a lake.
3. Brackenridgea forbesii TieGH. in Morot, J.
Bot. 16 (1902) 46, nom. nud.; Ann. Sc. Nat. Bot.
VIII, 16 (1902) 395, descr.; PULLE, Nova Guinea 8
(1912) 667; ReENDLE, J. Bot. (1923) Suppl. 7;
KANIS, Blumea 16 (1968) 49.
Tree, up to 30m, 55cm @. Leaves oblong to
lanceolate, 5-15 by 1'14—5 cm, acute to acuminate
at apex, acute or a little tapering at base, entire;
petiole 3-Smm. Jnflorescences terminal and
axillary, made up of a varying number of mostly
3-to 5- flowered, shortened, sessile cymes, flowering
simultaneously, the 2-S(-10) mm long rachis
mostly growing on vegetatively when terminal,
sometimes when axillary; pedicels c. 1 cm, up to
lcm in fruit, those of one cyme fixed close to
each other in one tier. Torus c. 1 mm high,
4-4 mm @, in fruit up to 244 mm high, 4 mm @.
Sepals ovate to elliptic, 3-444 by 14%4-14%4 mm.
104
Petals obovate-lanceolate, 3-414 by 1-114 mm,
acute at apex. Stamens with c. 144 mm long
filaments; anthers c. 114 by 14 mm. Ovaries c.
0.6 by 0.5 mm; style 114-2mm, in fruit up
to 3 mm. Fruits up to 6 by 5 mm.
Distr. Malesia: New Guinea.
FLORA MALESIANA
[ser. I, vol. 7?
Ecol. Primary rain-forest on flat country to
steep slopes, up to 750m, on clay, sand, or peat,
in sites which may be inundated during the wettest
season.
Vern. W. New Guinea: jobias, Je, obaisang,
Mooi, serukdeho, Manikiong.
2. Section Notochnella
(TieGH.) KANis, Blumea 16 (1968) 43. — Notochnella T1EGH.
Cymes ++ remote; peduncle + distinct; branches + shortened, sometimes
unequal in length, the longer overtopping the central flower. Flowers of 1 cyme
flowering successively. Corolla irregular, yellow. Stamens ~, in more than |
whorl. Ovaries 5-10.
Distr. Malesia: Philippines (excl. Palawan), monotypic.
4. Brackenridgea fascicularis (BLANCO) F.-VILL.
Noy. App. (1880) 40; KAnis, Blumea 16 (1968) 43.
— Ochna fascicularis BLANCO, Fl. Filip. ed. 2
(1845) 245; ibid. ed. 3, 2 (1878) 92; VIDAL, Sinopsis
(1883) 19, t. 27A; Rev. Pl. Vasc. Filip. (1886)
79; BARTELL. Malpighia 15 (1901) 162; Merr.
Goy. Lab. Publ. Philip. 27 (1905) 29; Sp. Blanc.
(1918) 263; En. Philip. 3 (1923) 68. — Diporidium
fasciculare (BLANCO) O.K. Rev. Gen. Pl. 1 (1891)
105; TreGH. in Morot, J. Bot. 16 (1902) 203.
— Notochnella fascicularis (BLANCO) TIEGH. Bull.
Mus. Hist. Nat. Paris 8 (1902) 549; Ann. Sc. Nat.
Bot. VIII, 16 (1902) 403. — Ouratea mindanaensis
Merk. Philip. J. Sc. 17 (1920) 287; En. Philip. 3
(1923) 68.
Tree up to 25m, 30cm g. Leaves oblong to
lanceolate, 5-15 by 2-5 cm, sometimes obtuse,
mostly acute to + acuminate at apex, obtuse to
acute, sometimes a little attenuate at base, margin
entire or + finely denticulate; petiole 4-8 mm.
Inflorescences terminal, made up of a varying
number of many-flowered, -+_ shortened cymes,
sometimes separate cymes in the axils of normal
leaves; pedicels 1-2 cm, up to 3cm in fruit,
the basal 1-5 mm persistent. Torus c. 1 mm high,
1144 mm g, in fruit up to 4mm high, 7mmg.
Sepals elliptic to obovate, 6-8 by 3-4 mm.
Petals obovate to obovate-lanceolate, 6-8 by
244-4144 mm. Filaments 114-244 mm; anthers
214-3 by c. Y% mm. Ovaries 0.7-0.8 by 0.5—0.7
mm; style c. 4mm, in fruit up to 7mm. Fruits
up to 7 by 6mm.
Distr. Malesia: Philippines.
Ecol. In primary forests at low and medium
altitudes, once reported from logged Dipterocarp
forest and once along a stream.
KEY TO THE SUBSPECIES
1. Stamens 20-45 . Cymes with 2-5 mm long
PEGUNCIES Hy en eh eo ante 1. ssp. fascicularis
1. Stamens 10-15. Cymes with 5—10(-20) mm
long peduncles. . . . 2. ssp. mindanaensis
1. ssp. fascicularis. — Ochna fascicularis BLANCO.
Inflorescences with 1-214 cm long rachis; cymes
with 2-5 mm long peduncles, up to 15-flowered.
Sepals 5. Petals 5—7(-10). Stamens 20-45. Ovaries
7-10.
Distr. Malesia: Philippines (Luzon, N. Visayas).
Vern. Philip.: aniatan, dirigkalin, bitas, mala-
kiting kiting, masalisi, Luzon, Tag., bansilai,
Visayas, Bis.
2. ssp. mindanaensis (MERR.) KANIS, Blumea 16
(1968) 44. — Ouratea mindanaensis MERR.
Inflorescences with 2-6 cm long rachis, often
indistinct by development of bracts to normal
leaves; cymes with 5—10(—20) mm long peduncles,
up to 7-flowered. Sepals (3-)5. Petals (3-)5.
Stamens (8—)10-15. Ovaries 6-7.
Distr. Malesia: Philippines (Mindanao).
Doubtful
Brackenridgea elegantissima (WALL.) KANIS,
Blumea 16 (1968) 50. — Euthemis elegantissima
WALL. in Roxb. FI. Ind. 2 (1824) 305; in Hook.
Bot. Misc. 2 (1830) 77, note; PLANCH. in Hook.
Lond. J. Bot. 5 (1846) 647; ibid. 6 (1847) 2, err.
in syn. Gomphia sumatrana JACK; BENN. in Hook.
f. Fl. Br. Ind. 1 (1875) 526; Kina, J. As. Soc. Beng.
52, ii (1893) 235; BARTELL. Malpighia 15 (1901)
160; RipL. FI. Mal. Pen. 1 (1922) 368. — Euthemis
? pulcherrima WALL. ex BENN. in Hook. f. FI.
Br. Ind. 1 (1875) 526, err. in syn. Gomphia suma-
trana JACK; KING, J. As. Soc. Beng. 52, ii (1893)
233; BARTELL. Malpighia 15 (1901) 160.
Based on one sterile, juvenile specimen from
Singapore I. It can not be decided whether it
belongs to B. hookeri (PLANCH.) A. GRAY or to
B. palustris BARTELL., although it should be cer-
tainly one of these species. WALLICH’S epithet
is the oldest available for any of the species in
Brackenridgea.
1971] OCHNACEAE (Kanis) 105
3. GOMPHIA
ScHREB. Gen. Pl. ed. 8 (1789) 291, p.p.; KANis, Taxon 16 (1967) 420, 422; Blumea
16 (1968) 51. — Ochna LINNE, Sp. Pl. 1 (1753) 513; Gen. Pl. ed. 5 (1754) 229, p.p.
excl. typ. — Ouratea [non AvBL. Hist. Pl. Gui. Fr. 1 (1775) 397] BAILL. Hist.
Pl. 4 (1873) 367, emend., p.p. excl. typ. — Meesia GAERTN. Fruct. | (1788) 344,
nom. rej., non HeDw. Sp. Musc. (1801) 173, nom. cons. — Campylospermum
TreGH. in Morot, J. Bot. 16 (1902) 40, 194, 197, nom. superfl. — Cercinia TrEGH.
l.c. 198. — Campylocercum TiEGH. Bull. Mus. Hist. Nat. Paris 8 (1902) 546. —
Fig. 3.
Shrubs or treelets. Stipules small, intrapetiolarly united, caducous. Leaves
shortly petioled, chartaceous, nerves close, parallel, + straight, curving upward
near the margin, forming an inconspicuous marginal nerve and a conspicuous,
somewhat wavy nerve parallel to the margin at some distance, veinlets reticulate,
joining in irregular secondary nerves parallel to the primary ones. Inflorescences
lateral and/or terminal thyrses with + reduced branches; peduncle ++ persistent
with sometimes a few small bracts at base, not leaving a distinct annulus of
scars; pedicels filiform, articulate at base. Flowers with a short, columnar, 5-
ribbed gynophore, enlarging and sometimes turning subglobular in fruit. Sepals 5,
tinged pinkish, accrescent. Petals 5, yellow, creamy or white. Stamens 10 in 1
whorl; filaments terete, very short; anthers opening with 2 apical pores. Ovaries 5,
obovoid; ovule camptotropous, epitropous; stigma punctiform. Fruits 1—2(-5),
yellowish green, turning dark purple or blue-black when ripe.
Distr. The majority of the species is found in Africa, S. of the Sahara, and in Madagascar. One sp.
in SW. Peninsular India, Ceylon, E. Thailand, Indo-China, Hainan, and W. Malesia.
Ecol. Confined to tropical areas with an everwet climate or with a moderately dry monsoon, up to
1500 m. Dispersal possibly by birds, but the fruits are not as conspicuous as in Ochna and Brackenridgea,
as calyx and torus are not coloured.
1. Gomphia serrata (GAERTN.) KANIS, Taxon 16
(1967) 422; Blumea 16 (1968) 53; FI. Thail. 2
(1970) 28. — Meesia serrata GAERTN. Fruct. 1
(1788) 344, t. 70, f. 6. — G. angustifolia VAHL,
Symb. Bot. Upsal. 2 (1791) 49; ScHEFF. Nat.
Tijd. N. I. 32 (1873) 411; Benn. in Hook. f. FI.
Br. Ind. 1 (1875) 525; F.-ViLL. Nov. App. (1880)
39; VIDAL, Sinopsis (1883) 19; Rev. Pl. Vasc.
Filip. (1886) 79; Phan. Cuming. (1895) 101;
Backer, Schoolfl. (1911) 194. — G. sumatrana
Jack, Mal. Misc. 1, 5 (1821) 29; in Hook. Bot.
Misc. 2 (1830) 77; PLANCH. in Hook. Ic. PI. II, 4
(1845) t. 712; in Hook. Lond. J. Bot. 6 (1847) 2;
Mio. FI. Ind. Bat. 1, 2 (1859) 675; Sumatra (1860)
209, 534; Scuerr. Nat. Tijd. N. I. 32 (1873) 411;
Benn. in Hook. f. FI. Br. Ind. 1 (1875) 525, err.
‘sumatrensis’; KiNG, J. As. Soc. Beng. 62, ii (1893)
232; Rip. Trans. Linn. Soc. II, Bot. 3 (1893)
285; Fl. Mal. Pen. 1 (1922) 365; RENDLE, J. Bot.
(1924) Suppl. 16; Ript. Kew Bull. (1925) 79;
Fl. Mal. Pen. 5 (1925) 296; Merr. J. Arn. Arb.
33 (1952) 226. — Ouratea angustifolia (VAHL)
BAILL. ex Laness. PI. Util. Col. Fr. (1886) 607;
Koorp. Exk. Fl. Java 2 (1912) 107; HALL. f. Beih.
Bot. Centralbl. 34, 2 (1916) 34; Merr. J. Str.
Br. R. As. Soc. n. 86 (1921) 387; En. Philip. 3
(1923) 68; RipLt. Kew Bull. (1930) 76. — Ochna
angustifolia (VAHL) O.K. Rev. Gen. PI. 1 (1891)
106. — Ochna sumatrana (Jack) O.K. lc. 106.
— Ouratea sumatrana (JACK) GILG in E. & P.
Nat. Pfl. Fam. 3, 6 (1895) 142; BARTELL. Mal-
pighia 15 (1901) 160; HALL. f. Beih. Bot. Centralbl.
34, 2 (1916) 35; Merr. J. Str. Br. R. As. Soc. n. 86
(1921) 387; Rip. Kew Bull. (1930) 76. — Ouratea
borneensis BARTELL. Malpighia 15 (1901) 156, t. 6;
Me_rr. J. Str. Br. R. As. Soc. n. 86 (1921) 387;
Ripv. Kew Bull. (1930) 76. — Ouratea neriifolia
BARTELL. Malpighia 15 (1901) 158, t. 7, sphalm.
‘neerifolia’; Merr. J. Str. Br. R. As. Soc. n. 86
(1921) 387; Airy SHAW, Kew Bull. (1940) 249.
— Ouratea beccariana BARTELL. Malpighia 15
(1901) 159, t. 9; Merr. J. Str. Br. R. As. Soc. n.
86 (1921) 387; Airy SHAW, Kew Bull. (1940) 249.
— Campylospermum sumatranum (JACK) TIEGH.
in Morot, J. Bot. 16 (1902) 197; Ann. Sc. Nat. Bot.
VIII, 16 (1902) 298; ibid. 18 (1903) 21. — Campylo-
spermum borneense (BARTELL.) T1EGH. in Morot,
J. Bot. 16 (1902) 197; Ann. Sc. Nat. Bot. VIII, 16
(1902) 301. —- Campylospermum beccarianum
(BARTELL.) TieGu. Ann. Sc. Nat. Bot. VIII, 16
(1902) 301. ——- Campylocercum _ neriifolium
(BARTELL.) TireGu. Lc. 304. — Campylocercum
(ser. I, vol. 7?
106
FLORA MALESIANA
S
v7 cee
Q o
ml ON
DY) Mew my \ y) —
SAN |S
LP
GTS
i 2
ro di
h gynophore,
f, x 114, c. flower, x2,
ls removed, f. stamen, abaxial view, g. gynoecium wit
KANISs. a. Flowering twig, 2, b. part of lea
%2 (a-g AmpuRIA 41444, h SAN 25040).
all «6, h. fruiting flower,
Fig. 3. Gomphia serrata (GAERTN.)
d. petal, x4, e. flower, sepals and peta
1971]
OCHNACEAE (Kanis)
107
zollingeri TYEGH. l.c. 305. — Campylospermum
wallichianum TieEGH. Bull. Mus. Hist. Nat. Paris
9 (1903) 76; Ann. Sc. Nat. Bot. VIII, 18 (1903) 17.
— Campylospermum plicatum TieGH. Bull. Lc.
78; Ann. /.c. 19. — Campylospermum strictum
TreGH. Bull. /.c. 79; Ann. /.c. 20. — Campylo-
spermum kingii T1EGH. Bull. /.c. 79; Ann. /.c. 21. —
Campylospermum perakense TieGH. Bull. /.c. 80;
Ann. /.c. 21. — Campylospermum abbreviatum
TieGuH. Bull. /.c. 80; Ann. /.c. 21. — Campylo-
spermum cumingii TIEGH. Bull. /.c. 80; Ann. /.c. 22.
— G. microphylla Rib. Fl. Mal. Pen. 1 (1922) 365,
f. 38. — G. oblongifolia RipL. Kew Bull. (1925)
281; Fl. Mal. Pen. 5 (1925) 296. — Ouratea arcta
Crais, Kew Bull. (1926) 341; BACKER & BAKH. f.
Fl. Java 1 (1963) 327. — Curatea megacarpa
Rip. Kew Bull. (1930) 76. — Ouratea micro-
phylla (RiDL.) CRAB, Fl. Siam. En. 1 (1931) 245.
— QOuratea sumatrana (JACK) GILG var. nervosa
Cras, /.c. 245. — Ouratea crocea (GRIFF.)
Burk. Kew Bull. (1935) 318, p.p. excl. typ. —
Fig. 3.
Shrub or tree, up to 25 m, 40 cmg. Branchlets of
young plants sometimes scrambling. Leaves ovate-
to obovate-lanceolate, 6-20 by 2-6 cm (up to 35
by 10cm in young plants), chartaceous, mostly
acute to acuminate, sometimes obtuse at apex,
acute or a little tapering at base, margin finely
denticulate; petiole 214-714 mm. Inflorescences
many-flowered; rachis (2%—)5-20(-35) cm, pri-
mary branches of terminal inflorescences usually
up to 10(-15) cm long, secondary branches or
primary ones of lateral inflorescences usually
reduced or up to 1 cm long, cymose, with (1-)
3-7(—~~), + conferted flowers; pedicels 14-34 cm,
up to 1 cm in fruit. Torus 0.7-1 mm high, 0.7-1
mm @, in fruit up to 5mm high, 5mm g. Sepals
mostly ovate to elliptic, sometimes obovate,
4-7 by 2!4-4mm, mostly acute to obtuse, some-
times rounded. Petals obliquely obovate to broad-
spatulate, 414-8 by 214-6 mm, obtuse, rounded
or truncate, mostly yellow, sometimes creamy or
white. Stamens subsessile or with up to 14 mm
long filaments; anthers 2!4-—5(—6) by 0.5—0.8 mm.
Ovaries 0.7—1 by 0.4-0.6 mm; style 3-5 mm long,
up to 7 mm in fruit; stigma minute. Fruits up to
8(-10) by 6(-8) mm.
Distr. SW. Peninsular India, Ceylon, E. Thai-
land, Indo-China, Hainan; in Malesia: Sumatra,
Banka, Malay Peninsula, Karimundjawa _Is.,
Borneo, Philippines, and Celebes.
Ecol]. From sea-level up to 1200 m, up to 1500
m on Mt Kinabalu, in primary and secondary
vegetation: lowland and submontane, mixed
Dipterocarp forests, peat swamp forests, high
kerangas forests, ridge forests, and open scrub veg-
etation; on level land to steep slopes, also near
river-banks and on cliffs near the sea. Soils are
rocky, sandy, loamy or clayish, but usually acid;
limestone is reported occasionally, but it is very
probable that the soils in the localities concerned
have been leached out.
The flower buds are often galled by unidentified
insects, especially in the Malay Peninsula, Su-
matra, and the smaller islands east of Sumatra.
These buds are about ovoid, with small sepals and
petals somewhat sunken in an enlarged torus,
whereas gynoecium and androecium are strongly
suppressed.
Ripe fruits in normal flowers are often empty,
which is probably also caused by insect damage,
as perforations of the pericarp are sometimes
clearly visible. It is recorded once that ants did
attack a collector’s press for these fruits (CLEMENS,
Indo-China). Seeds are rare in herbarium speci-
mens of Malesian collections.
Uses. The wood is reported as being used as
poles or planks for the construction of houses in
Ceylon, Vietnam, Sumatra, Malay Peninsula,
and Sarawak. The leaves are chewed by jungle
tribes in Malaya. Roots and leaves are bitter and
are decocted in S. India for a stomachic and anti-
emetic tonic. Young branches are used against
tooth-ache in Cambodia.
Properties of wood: dull red or red-brown,
hard and strong, apt to split in drying.
Vern. Sumatra, North: kaju ndolak, Batak:
West: kalék djambak, sébalusi, Minangkabau;
South: kaju mat, k. sépah, lakodjong, madjang
madjang; Banka: méntungging, mésulung putih:
Mal. Pen., Thailand: rong pling; Malaya: bunga
kélimbing bési, chinta mula, lidah mura, mata
kétam batu, mémbatu, ménarah, méndapor,
méramong, murmagong, pénarah, péngling, pokok
lébah, p. luas, tampang bési; several compound
names were recorded occasionally with géronggang
(= Cratoxylon), jambu and kélat (= Eugenia);
Borneo, Sarawak: kéladang, kélutak, Iban, aam,
Kenyah; compound names with ubah or ubar
(= Eugenia) were recorded occasionally; Brunei:
chénaga lampong, Iban, pinis, Malay; Sabah:
majang majang, quintalai, Kadazan, antimagas
gimbaan, Murut, kolambang, Rungus Dusun,
posoon, tulangkara, Kinabatangan, alas, tampala-
nuk, Tidong, bakan bitanag, biobi, malatangor,
ondogong, Malay; West Borneo: mélindingan, Da-
yak; East Borneo: amir burgang, Dayak, batu batu,
mulak, téngkédjing kéring, Malay; Philip., Pa-
lawan: anduyong, bibingo, huisac; Luzon: salactoc,
Zambales, simahima, Bicol, sasahit, Tagalog;
Visayas: bulocauan, caranan, Panay, minsaray,
Mindoro; Celebes: /ébani, parasinga tjila,
Bugi, mampa, Rangkas, morosisio, wulisi mapute,
Tobela.
Notes. As this species grows under rather dif-
ferent ecological conditions, the variation in its
morphological characters is not surprising. Apart
from these variations, some geographically more
or less separated ‘races’ can be recognized. This
is the reason why in certain Floras two or three
different species have been recognized although the
distinctive characters can hardly be used at a
specific level. Over a large area it is impossible to
make a key to these ‘races’, because the same dif-
ferences occur in remote parts of the specific area.
In Sumatra and the Malay Peninsula plants are
relatively uniform, the greatest variation occurring
in Borneo, especially in Sarawak. In more ex-
posed habitats, on cliffs and on poor kerangas
soils, specimens often have small leaves. On
108
limestone in the Langkawi Islands a race ‘micro-
phylla is found with small leaves and very reduced
inflorescences. Similar, but less reduced forms are
found elsewhere along the coasts of the Malay
FLORA MALESIANA
[ser. I, vol. 7}
Excluded
Gomphia magnoliaefolia Zirp. ex SPAN. Linnaea
15 (1841) 186, nom. nud. = Pycnarrhena longifolia
DIELS,
(DECNE) Becc. (Menispermaceae), fide
Peninsula. Pflanzenreich, Heft 46 (1910) 51.
4. EUTHEMIS
JACK, Mal. Misc. 1, 5 (1821) 15; KANIs, Blumea 16 (1968) 62. — Fig. 4.
Shrubs or shrublets, sparsely branched. Stipules free, caducous. Leaves coria-
ceous, glabrous, denticulate, nerves numerous, parallel, from the midrib curving
sidewards, straightly ascending to the marginal veins at an angle of c. 80°; petiole
+ winged. Inflorescences terminal, many-flowered, compound racemes; bracts
small, caducous. Flowers % or polygamous. Sepals 5, turning purplish red in fruit.
Petals 5, white or pinkish. Staminodes 0(-S), filamentous. Stamens 5, free; anthers
subsessile, rostrate. Ovary 5-celled; ovules 2 per cell, pendulous, axile; stigma
minute. Fruit a berry with 5 pyrenes. Seeds 1 (2) per cell.
Distr. SW. Cambodia, in Malesia: Sumatra, Malay Peninsula, Borneo.
Ecol. Everwet tropical forest below 1250 m, in kerangas forests, on low ridges in peat-swamp forests,
and in open ridge forests, on poor, mostly sandy soils. Dispersal probably by birds because of conspic-
uous, white, rose-pink or red berries (RIDLEY, Disp. 1930, 410).
KEY TO THE SPECIES
1. Inflorescence a panicle, branches well developed with scattered flowers. Leaves 8-40 cm long, margin
distinctly denticulate. Mature fruit white.
1. E. leucocarpa
1. Inflorescence a very slender, often cernuous raceme, nearly all branches reduced with conferted
flowers. Leaves 4-15 cm long, margin faintly denticulate. Mature fruit red.
1. Euthemis leucocarpa JAcK, Mal. Misc. 1, 5
(1821) 16; WALL. in Roxb. FI. Ind. 2 (1824) 303;
JACK in Hook. Bot. Misc. 2 (1830) 69; PLANCH. in
Hook. Ic. Pl. II, 4 (1845) t. 711; Mra. FI. Ind.
Bat. 1, 2 (1859) 675; Sumatra (1860) 208, 533,
incl. var. latifolia; SCHEFF. Nat. Tijd. N. I. 32
(1873) 411; BENN. in Hook. f. FI. Br. Ind. 1
(1875) 526; Kina, J. As. Soc. Beng. 62, ii (1893)
234; Rip_. Trans. Linn. Soc. II, Bot. 3 (1893)
285; BARTELL. Malpighia 15 (1901) 167; HALL.
f. Beih. Bot. Centralbl. 34, 2 (1917) 30; Merr.
J. Str. Br. R. As. Soc. n. 86 (1921) 388; Ript. FI.
Mal. Pen. 1 (1922) 368; Diets, Bot. Jahrb. 60
(1926) 311; Burk. Dict. 1 (1935) 987; Airy SHAW,
Kew Bull. (1940) 249; Merr. J. Arn. Arb. 33
(1952) 224; VipaL, Adansonia 1 (1961) 60;
KANIs, Blumea 16 (1968) 62; FI. Thail. 2 (1970)
29. — E. robusta Hook. f. Trans. Linn. Soc. 23
(1862) 163; BARTELL. Malpighia 15 (1901) 168;
HALL. f. Beih. Bot. Centralbl. 34, 2 (1917) 32;
RIDL. Fl. Mal. Pen. 1 (1922) 368.
Shrub up to 6m. Branchlets stout, green.
Stipules ovate, 4-6 by c. 2 mm, acute to acuminate,
ciliate. Leaves oblong to linear oblong, 8-40 by
2-10 cm, acute at apex, tapering at base, margin
distinctly and irregularly denticulate, nerves
1-2 mm apart; petiole 2-5 cm. Panicles erect,
8-20 cm; pedicels 4-10 mm, articulate at base;
bracts 8-10 by 2-4 mm, lanceolate, acute. Flowers
2. E. minor
%, erect, often in pairs. Sepals obliquely ovate to
elliptic, unequal, 4-7 by 2-344 mm, ciliate.
Petals obliquely obovate to spatulate, 4-10 by
214-5 mm. Anthers 3-5 by c. 1 mm, yellow. Ovary
ovoid to bottle-shaped, 2-4 by c. | mm, style
114-3 mm. Fruit globular, up to 1 cm g, fleshy,
via red turning white. Seeds like sectors of a
sphere, c. 4 by 2 mm.
Distr. SW. Cambodia;
Riouw & Lingga Is., Banka; Billiton,
Peninsula, Anambas Is., Borneo.
Ecol. From sea-level up to 1000 m, on poor
soils, preferably in moist, shady places (see also
under the genus).
Uses. Medical applications of the roots is
reported from Malaya. In Brunei the fruits are
used against eye-diseases.
Vern. Sumatra: bélusung putih, kaju padang,
mata pélanduk, Banka; balong, Billiton; Malaya:
pélawan bérok; Borneo: tambu, Sarawak, ranggas
hutan, Sabah, iur iur, W. Borneo.
in Malesia: Sumatra,
Malay
2. Euthemis minor Jack, Mal. Misc. 1, 5 (1821) 18;
WALL. in Roxb. FI. Ind. 2 (1824) 304; Jack in
Hook. Bot. Misc. 2 (1830) 69; Mra. FI. Ind. Bat.
1, 2 (1859) 675; Sumatra (1860) 209, 534; SCHEFF.
Nat. Tijd. N. I. 32 (1873) 412; BENN. in Hook.
f- Fl. Br. Ind. t (1875) 526; KiNG; J-Asiesee-:
Beng. 62, ii (1893) 235; BARTELL. Malpighia 15
1971] OCHNACEAE (Kanis) 109
Fig. 4. Euthemis minor Jack. a. Fertile twig, x %/, 6. detail of leaf margin, «4, c. flowerbud, d. open
- 3 flower, both x4, e. ditto, stamen, ad- and abaxial view, f. ditto, reduced pistil, both x6, g. pistil of 2
or % flower, x4, A. fruit, «2, i. seed, x4 (a-c MevER 21807, d-f KosrerMans & ANTA 1256, g Hans
WINKLER 1421, A-i HALLIER f. 1445).
110
(1901) 168; Ripi. J. Str. Br. R. As. Soc. 1. 54
(1910) 34; HALL. f. Beih. Bot. Centralbl. 34, 2
(1917) 32; Merr. J. Str. Br. R. As. Soc. n. 86
(1921) 388; Ript. Fl. Mal. Pen. 1 (1922) 368;
ArrY SHAW, Kew Bull. (1940) 250; Merr. J. Arn.
Arb. 33 (1952) 224; KaANis, Blumea 16 (1968)
65. — E. obtusifolia Hook. f. Trans. Linn. Soc. 23
(1862) 163; BARTELL. Malpighia 15 (1901) 169;
HALL. f. Beih. Bot. Centralbl. 34, 2 (1917) 33;
Merr. J. Str. Br. R. As. Soc. n. 86 (1921) 389;
Ary SHAW, Kew Bull. (1940) 250. — E. engleri
Gitc in E. & P. Nat. Pfl. Fam. 3, 6 (1895) 152,
f. 78; BARTELL. Malpighia 15 (1901) 169; HALL.
f. Bern. Bot. Centralbl. 34, 2 (1917) 33; Atry
SHAW, Kew Bull. (1940) 250. — E. ciliata PEARSON,
Kew Bull. (1906) 3; HALL. f. Beih. Bot. Centralbl.
34, 2 (1917) 33. — E. hackenbergii DiELs, Bot.
Jahrb. 60 (1926) 310; Atry SHAW, Kew Bull.
(1940) 250. — Fig. 4.
Shrublet up to 3(—5S?) m. Branchlets slender,
blackish. Stipules c. 3 by 1 mm, acuminate, ser-
rulate. Leaves oblong to oblanceolate, 4-15 by
114-4 cm, obtuse at apex, mucronate, tapering at
base, margin faintly denticulate, nerves c. 1 mm
apart; petiole to 1144 cm. Racemes lax, 5-30 cm
long, often pseudolateral, sometimes drooping,
rachis slender, branches ++ shortened; bracts on
the rachis 5-12 by 1-2 mm, caducous, on branches
trigonous, c. 1mm long, acuminate. Flowers
functionally polygamous, complete, often 2 or
more conferted. Sepals obovate, unequal, 4-414
FLORA MALESIANA
[ser. I, vol. 71
by 214-314 mm, ciliate near the apex, acuminate.
Petals lanceolate, 6-8 by 214-3mm, distinctly
reflexed in 3 flowers. Anthers 3'\4-414 by c. 1 mm,
yellow. Ovary in § or $ flowers obovoid, 5-ribbed,
c. 244 by 114 mm; style c. 214 mm long, widening
into the ovary; ovary in g flowers very much
reduced, depressed-globular, shallowly 5-lobed,
c. 0.2 by 0.4 mm; style c. 0.2 mm, broadly conical
at base. Fruit globular, up to 6 mm g, acuminate,
5-ribbed, red. Seeds semi-annular, c. 4 by
214 mm.
Distr. Malesia: Central E. Sumatra, Riouw &
Lingga Is., Banka, Billiton, S. Malay Peninsula
(?), Borneo.
Ecol. From sea-level up to 1250m, often
found with the previous species, but generally in
drier and more exposed places (see also under the
genus).
Vern. Sumatra: bélusung mérah, kétjing pé-
landuk, Banka; Borneo: buah iték, Sarawak,
pétikawo, Sabah, Kedayan, mata pélandok, W.
Borneo.
Notes. Many specimens are doubtlessly
functionally male. I am not certain whether fruiting
specimens generally have bisexual or func-
tionally female flowers, but these possibilities are
probably both realised.
This species was never recorded from the
mainland of the Malay Peninsula. Three older col-
lections are known to me from Singapore L.,
where it is probably now extinct.
5. NECKIA
KortTH. Ned. Kruidk. Arch. 1 (1848) 358; KANis, Blumea 16 (1968) 69. — Fig. 5.
Shrublets or undershrubs. Stipules pectinate. Leaves with short petioles, nerves
ascending at an angle of 70-80°, slightly curved. Inflorescences axillary, much
reduced, the rachis bearing some small bracts, but only 1 terminal flower; pedicels
articulate. Flowers 3, 5-merous. Staminodes ~, of 2 types, the inner ones forming a
tube at base. Stamens adnate to the tube. Ovary 3-carpelled; stigma 1. Style of ripe
fruit splitting into 3 parts. Seeds not winged.
Distr. Malesia: Sumatra, Malay Peninsula, Borneo, Philippines. Monotypic.
1. Neckia serrata KortTH. Ned. Kruidk. Arch. 1
(1848) 358; Mia. Fl. Ind. Bat. 1, 2 (1859) 118;
Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 218; Bogr-.
& Koorp. Ic. Bog. 1, 4 (1901) 1, t. 76; Ripv. J.
Str. Br. R. As. Soc. n. 49 (1908) 13, 14; Merr.
ibid. n. 86 (1921) 388; KaNnis, Blumea 16 (1968)
69. — N. lancifolia Hoox. f. Trans. Linn. Soc. 23
(1862) 158; Rip. J. Str. Br. R. As. Soc. n. 49
(1908) 13; Airy SHAW, Kew Bull. (1940) 252, incl.
f. major (RIDL.) Airy SHAW. — N. humilis Hook. f.
Trans. Linn. Soc. 23 (1862) 159; RipL. J. Str. Br.
R. As. Soc. n. 49 (1908) 14. — N. malayana
RIDL. /.c. 11, incl. var. angustifolia RiDL.; Fl. Mal.
Pen. 1 (1922) 134, f. 13, incl. f. major RIDL. et
f. minor Riv_. — N. distans Rie. J. Str. Br. R.
As. Soc. n. 49 (1908) 12; Merr. ibid. n. 86 (1921)
388; Airy SHAW, Kew Bull. (1940) 250. — N.
klossii Rip. J. Str. Br. R. As. Soc. n. 49 (1908)
13; Airy SHAW, Kew Bull. (1940) 251, incl. var.
borneensis Airy SHAW. — N. parviflora RIDL. J.
Str. Br. R. As. Soc. n. 49 (1908) 14; Merr. ibid. n.
86 (1921) 388. — N. ovalifolia Carit. Bull. Soc.
Bot. Fr. 57 (1910) 397, t. 13; Merr. J. Str. Br. R.
As. Soc. n. 86 (1921) 388; Airy SHAW, Kew Bull.
(1940) 252. — Sauvagesia jaheriana Capit. Bull.
Soc. Bot. Fr. 57 (1910) 397, t. 13; 3MiEnRE
Str. Br. R. As. Soc. n. 86 (1921) 388. — N. gran-
difolia RipL. Kew Bull. (1925) 77. — N. obovata
Airy SHAW, Kew Bull. (1940) 251. — N. philip-
pinensis MERR. & Quis. Philip. J. Sc. 82 (1954)
329, t. 2. — Fig. 5.
Shrublet up to 1 m, often much smaller, often
unbranched. Stipules compound, 1-3 mm wide,
the segments unequal, acicular, 14-1 cm long,
1971] OCHNACEAE (Kanis) 111
oe a nn ees aee
ST
LCS
SS
LTR are awe
TH
—aor TT
_ Fig. 5. Neckia serrata Kortu. a. Habit, x 2/4, b. stipule, x4, ¢. flower, x4, d. androecium, inner view of
tube, 3 anthers removed, e. ditto, outer view, complete, both «8, f. pistil, x6, g. fruit, x2, h.
seed, X20, i. habit of a dwarfed form, * %, j. leaf of a narrow type, x %, k. leaf of a rounded type,
x%%,, I. leaf of a large type, x2 (a-f Mever 6854, g-h Brooke 9613, i KOSTERMANS 9087, / BROOKE
10610, k Brooke 10006, / PosrHumus 600).
112
FLORA MALESIANA
{ser. I, vol. 73
lacerate. Leaves obovate to long obovate-lanceo-
late, 3-15(-25) by 34-2'4(-714) cm, mostly
acute to acuminate, sometimes obtuse, tapering
at base, biserrulate, membranous to chartaceous,
nerves 114-3 mm apart. /nflorescences with fili-
form rachis, 1-5 cm; bracts up to 10, linear, c.
2 mm; sometimes with small stipules; pedicels fili-
form 8-15 mm. Flowers pendulous. Sepals lan-
ceolate, 4-714 by 1-3 mm, acute, margins dentate.
Petals elliptic, 244-5 by 144-2 mm. Outer sta-
minodes ~\, unequal, long spatulate or gland-like,
14-1 mm long; inner staminodes 15-25, 1-134 mm
long, + half connate, the free lobes spatulate, the
5 lobes alternating with the stamens longer.
Stamens with terete filaments, the free part up to
1mm; anthers lanceolate, 1-144 by c. 4% mm,
yellow. Ovary ovoid, 1-1.2 by 0.5-0.6 mm, style
c. 1mm long; stigma clavate. Fruit subtrigonous
in cross-section, c. 5 by 214 mm. Seeds ellipsoid,
c. 0.5 by 0.3 mm, areolate.
Distr. Malesia: Sumatra, Mentawei Is.,
Riouw & Lingga Is., SE. Malay Peninsula (S.
Trengganu, Pahang, Johore), Borneo, and Philip-
pines (Samar). Monotypic.
Ecol. Found up to 1200m, in moist, shady
places in kerangas forests as well as in richer rain-
forests, especially on boulders, cliffs and wet
slopes, along brooklets and small rivers, sometimes
near waterfalls or subject to occasional inundation.
Soils are reported as sandstone, acid sand, sandy
loam, loam, or tuff.
Notes. The variability in shape and size of
this species must primarily be due to variation in
ecological conditions. The plants need moist,
shady places and will be capable to germinate and
to produce a few flowers even while growing in
a moss carpet over boulders. The smaller speci-
mens are dwarfed by lack of nutrition. Shade,
water-supply, occasional inundation, or other
ecological circumstances may also exert influence
upon the habit of the plants.
There will probably be genetic differences
between local populations, but it seems impractic-
able to distinguish infra-specific taxa, since no
useful differences are found in the geographical
distribution of morphological characters. The
greatest variability is found among Bornean
specimens, whereas the plants with the largest
leaves are found in Sumatra.
6. INDOVETHIA
BOERL. Feestbundel P. J. Veth (1894) 89; Ic. Bog. 1 (1897) 9; KANIs, Blumea 16
(1968) 72. — Fig. 6.
Shrublets. Stipules pectinate. Leaves subsessile, nerves ascending at an angle
of 60-70°, slightly curved. Inflorescences terminal, many-flowered, compound ra-
cemes; mostly some lower branches well developed, the other ones much shortened
with a few flowers; bracts small. Flowers %. Sepals 5. Petals 5, white. Staminodes
10 in 2 alternating whorls, mutually and with the stamens connate at base. Ovary
3-carpelled; stigma 1. Style of ripe fruits splitting into 3 parts. Seeds not winged.
Distr. Malesia: Central E. Sumatra and NW. Borneo. Monotypic.
1. Indovethia calophylla BorrL. Feestbundel P.
J. Veth (1894) 90, plate; Ic. Bog. 1 (1897) 10, t. 1;
BARTELL. Malpighia 15 (1901) 173; Merr. J. Str. Br.
R. As. Soc. n. 86 (1921) 388; KANis, Blumea 16
(1968) 72. — J. beccariana BARTELL. Malpi-
ghia 15 (1901) 172, t. 11; Merr. J. Str. Br. R. As.
Soc. n. 86 (1921) 388. — Fig. 6.
Shrublet c. 1m. Stipules compound, the seg-
ments unequal, lanceolate, 5-10 by 2-4 mm,
margin lJacerate. Leaves oblanceolate, 10-35 by
214-814 cm, acute to acuminate at apex, tapering
at base, biserrulate, membranous, stouter nerves
3-714 mm apart. Inflorescences 5-20 cm long;
rachis c. 2mm 9; pedicels filiform, mostly short,
up to 12mm when fruiting; bracts with small
stipules, lanceolate, the lower ones up to 8 by2 mm,
the higher ones smaller, shortly pubescent under-
neath. Flowers erect. Sepals suborbicular to obo-
vate, 314-5 by 3-314 mm, margins denticulate.
Petals obovate, 3-4 by 2-3mm. Staminodes
spatulate, those opposite to the stamens 134-214
by c. % mm, with 3 + distinct, parallel nerves,
the alternating ones 114-2 by c. 4% mm, with 1
distinct nerve. Stamens with flattened filaments
0.2-0.4 mm long; anthers 114-134 by 0.5-0.8 mm,
the anther-cells basally separated by the cuneate
connective provided with a c. 4 mm long mucro.
Ovary subglobose, c. 1 mm g; style c. 1 mm long;
stigma minute, trigonous. Fruit subglobose, up
to 12 mm g, papillate. Seeds c. 0.8 by 0.2 mm,
areolate.
Distr. Malesia: Central E. Sumatra (Riouw
Prov.), NW. Borneo (Sarawak, Ist Div.; W.
Borneo, NE. of Pontianak).
Ecol. A rare, local lowland species, reported
from moist, shady places, in primary or secondary
(Dipterocarp) forests, on richer (loamy) soils.
Occasionally reported from the edge of a lake and
above a torrent in the forest. Also once collected
under rubber.
1971] OCHNACEAE (Kanis) L&3
Fig. 6. Indovethia calophylla BoERL. a. Fertile twig,
d. ditto, part of androecium, inner view, e. ditto, outer view, both x20, f. inner staminode from flower,
g. ditto, outer staminode, h. ditto, stamen, i. ditto, pistil, all x6, j. fruit, x2, k. seed, x10 (a after Ic.
Bog. 1, t. 1, b-g BUWALDA 6256, h—-j TEYSMANN HB 10893, k BROOKE 9455).
7. SCHUURMANSIELLA
HALL. f. Rec. Trav. Bot. Néerl. 10 (1913) 344; KANis, Blumea 16 (1968) 73. —
Fig. 7.
Shrubs with slender branchlets. Stipules acicular. Leaves short-petioled, nerves
ascending at an angle of c. 80°. Inflorescences terminal, many-flowered, compound
racemes; branches much shortened, with flowers of successive age; bracts small,
those of one branch conferted. Flowers %. Sepals 5. Petals 5, white or pinkish,
_ purplish at base, in anthesis soon surpassing the sepals in size. Staminodes ~,
of 2 types, the inner ones mutually and with the stamens connate at base. Ovary
3-carpelled; stigma 1. Style of ripe fruits also splitting into 3 parts. Seeds not winged.
Distr. Malesia: NW. Borneo. Monotypic.
114
FLORA MALESIANA
[ser. 1, vol. 74
Fig. 7. Schuurmansiella angustifolia (HOOK. f.) HALL. f. a. Fertile twig, x 24, b. flowerbud, ~ 4, c. flower,
petals shed, x4, d. part of androecium, inner view, x 6, e. pistil, «8, f. fruit, g. open fruit, both x2,
h. seed, 6 (a-f ABANG Muas 5139, g ANDERSON 8368, 4 BUJANG 12990).
1. Schuurmansiella angustifolia (Hook. f.) HALL. f.
Rec. Trav. Bot. Néerl. 10 (1913) 345, t. 7; MERR.
J. Str. Br. R. As. Soc. 1. 86 (1921) 387; KAnis,
Blumea 16 (1968) 73. — Schuurmansia angustifolia
Hook. f. Trans. Linn. Soc. 23 (1862) 157. —
Fig. 7.
Shrub up to 7 m high. Stipules up to 12 by c. 14
mm. Leaves linear-oblong, 8-17 by 4-114 cm,
long acuminate at apex, - tapering at base,
chartaceous, margin serrulate, nerves 14-1 mm
apart. Inflorescences c.15cm long; rachis c. 1 mmg;
branches with up to 5 flowers; pedicels filiform, up
to 12 mm in fruit; bracts acicular, 2-3 mm under
the branches, smaller at base of pedicels. Flowers
1971]
OCHNACEAE (Kanis)
115
erect. Sepals + linguiform, 1-114 by 34-1 mm.
Petals ovate, 3144-414 by 134-214 mm, distinctly
reflexed. Outer staminodes ~, small; inner ones
25-30, linear, 2-3 mm, purplish. Stamens with
flattened filaments 44-1 mm long, purplish;
anthers lanceolate, 144°-24%4 by 0.3-0.6 mm.
Ovary ovoid, c. 1.2 by 0.7 mm; style c. 0.3 mm,
purplish; stigma capitate. Fruit ellipsoid, sub-
trigonous in cross-section, c. 814 by 3 mm. Seeds
0.5-1 by 0.2-0.5 mm, tomentose.
Distr. Malesia: NW. Borneo (Sarawak, Ist
and 4th Div.).
Ecol. A lowland species, reported from up to
600 m, in moist places, especially in kerangas
forests on poor soils, and on sandstone cliffs,
even near the sea.
8. SCHUURMANSIA
BLuME, Mus. Bot. Lugd. Bat. | (1850) 177, t. 32; KANis, Nova Guinea, Bot. 6
(1961) 63; Blumea 16 (1968) 74. — Fig. 8-10.
Trees or treelets with stout, often hollow branches. Stipules entire. Leaves
conferted at intervals (fig. 10), glandular along the margins, nerves almost straight-
ly ascending at an angle of 60~-70°. Inflorescences terminal, many-flowered,
panicles often with many hypsophylls at base; peduncle and branches distinctly
ribbed, often with con- or recaulescences. Flowers 3 or polygamous. Sepals 5.
Petals 5, white, creamy or purplish. Staminodes ~, in 1 or 2 whorls, the inner ones
mutually and with the stamens connate at base. Ovary 3-carpelled; stigmas | or 3.
Fruit opening with 3 longitudinal slits, the style usually not splitting. Seeds winged
like a propeller with 2 blades.
Distr. Solomons and Bismarcks; in Malesia: Borneo, Philippines, Celebes, Moluccas, and New
Guinea.
Ecol. Pioneer plants, from sea-level up to 3000 m, especially in natural or anthropogenous secondary
vegetation.
KEY TO THE SPECIES
1. Style as long as the ovary or longer; stigma capitate or punctiform. . . .. . - - 41. S. elegans
1. Style half as long as the ovary or shorter; stigma 3-lobed.
2. Filaments + twice the length of the anthers. ...........++ee68 2. S. vidalii
2. Filaments as long as the anthers or shorter... ........2-++-ee-8 . 3. 8S. henningsii
1. Schuurmansia elegans BLUME, Mus. Bot. Lugd.
Bat. 1 (1850) 178, t. 32; Mig. FI. Ind. Bat. 1, 2
(1859) 118; Illustr. 1 (1871) 66, t. 29; Capit. Bull.
Soc. Bot. Fr. 57 (1910) 397; HALL. f. Rec. Trav.
Bot. Néerl. 10 (1913) 346; Merr. Philip. J. Sc. 11
(1916) Bot. 19; En. Philip. 3 (1923) 68; Herne in
Fedde, Rep. 54 (1951) 240; Pfl. Samml. Clemens
Kinab. (1953) 63; KaNnis, Nova Guinea, Bot. 6
(1961) 64, f. la; Blumea 16 (1968) 75. — S. par-
viflora Rip. Trans. Linn. Soc. II, Bot. 9 (1916) 18.
— S. borneensis Rip_. Kew Bull. (1930) 77.
Treelet or tree, up to 15(—30?) m, sometimes
with low stiltroots. Stipules linguiform, 114-4 by
114-24%4 mm, sometimes shortly ciliate. Leaves
obovate-oblong to -lanceolate, 10-30 by 2!4-10cm,
rounded or somewhat acuminate at apex, + tape-
ring at base, margins + involute, chartaceous,
nerves I-1'4 mm apart; surface of dried leaves
very finely reticulate by protruding intercellular
walls of epidermal cells, giving an impression of
Striation parallel to the nerves; petiole 114-6
em. Inflorescences 10-25 cm; peduncle 2-5 mm 9;
pedicels filiform, 3-6 mm; bracts broadly lin-
guiform, up to 2mm long, sometimes shortly
ciliate. Flowers bisexual, erect. Sepals elliptic to
obovate, 3-6 by 114-4 mm. Petals obovate, 4-8
by 2-S mm. Outer staminodes 0-—cy, linear, 1-114
mm long; inner ones 15-25, linear to spatulate,
2144-5 mm long, with 1 distinct nerve. Filaments
1-2 mm; anthers 114-2 by c. 4 mm, connective
distinctly protruding. Ovary subovoid, 114-3 by
1144-2 mm, papillate, sometimes with a few glan-
dular-capitate hairs; style 1144-3 mm, growing in
fruit, widening into the ovary; stigma small,
capitate or punctiform. Fruit fusiform, up to
24 by 4 cm, acuminate. Seeds c. ¥% by 4% mm,
with c. 3 mm long, slender wings.
Distr. Malesia: Borneo, Philippines (Min-
danao, only 1 collection of uncertain identity
reported from Agusan), Celebes, Moluccas, and
New Guinea.
Ecol. From sea-level up to 2000 m, in primary
and secondary forests, on level land to steep
slopes, sometimes near river-banks or in swampy
localities; on clay or more sandy or rocky soils.
Probably not uncommon locally, but rarely
flowering.
Vern. Philip., Mindanao: ftanang, Manobo;
FLORA MALESIANA [ser. I, vol. 73
Mf
M//- “
, 1
(hae OT)
BO,
i
A
LLILITITLA
Vi
rer tne
front ne
/
Meh
LLL
LIT] LLY
Yipee
vy, .
is
a
Fig. 8. Schuurmansia henningsii K. Scu. a. Fertile branch, x 14, b. 3 flower, <5, c. ditto, part of androe-
cium, inner view, d. ditto, reduced pistil, both x8, e. 2 flower, x5, f. ditto, sepals and petals removed,
x8, g. fruit, h. open fruit, both «114, i. seed, x5 (a-d HOOGLAND & PULLEN 5356, e—f BRASS 29335,
g HOOGLAND 3507, h-i BW 4987).
Celebes: labo labo, Toradja; Ambon: wat lopu; 229, plate. — S. parvifolia MERR. Philip. J. Sc. 11
New Guinea, Vogelkop Pen.: hikselah, Tehid. (1916) Bot. 19.
Treelet up to 7m. Stipules 1-214 by 1-2 mm,
2. Schuurmansia vidalii (F.-VILL.) MerR. Philip. shortly ciliate. Leaves obovate-lanceolate, 5—11
J. Sc. 11 (1916) Bot. 19; En. Philip. 3 (1923) 69; by 114-4 cm, obtuse to obtusely acuminate at
KANiIs, Nova Guinea, Bot. 6 (1961) 69, f. 1c; apex, tapering at base, margins somewhat involute,
Blumea 16 (1968) 76. — Calophyllum vidalii strongest nerves c. 1 mm apart, papyraceous to
F.-VILL. in Ceron, Cat. Pl. Herb. Manila (1892) _chartaceous; petiole 14-2 cm. Inflorescences 7-15
75 Se Pe
= Y
Fig. 9. Schuurmansia henningsii K. Scu. After anthesis, Cycloop Mts (Photogr. SLeuMER, July 1961).
cm; peduncle 114-314 mm @; pedicels 1-2 mm;
bracts very small, semi-annular to triangular.
Flowers %, erect. Sepals obovate, 2-274 by 144-2
mm. Petals obovate, c. 314 by 244 mm, white or
pink. Outer staminodes 10—-\, filiform, 44-1 mm
long; inner staminodes c. 25, linear, 114-2 mm,
c. Ya mm connate at base. Filaments c. 134 mm;
anthers 34-1 by 0.4-0.5 mm. Ovary subglobular,
3-lobed, 0.4-0.5 mm @, glabrous; stigma 3-
lobed, subsessile. Fruit fusiform, c. 10 by 3 mm,
acuminate. Seeds c. 0.6 by 0.4mm, with c. 2mm
long, basally connate wings.
Distr. Malesia: Philippines (Luzon: Aurora,
Camarines Sur & Sorsogon Prov., 3 collections).
Ecol. In (mossy) forest, from 500 to well over
1000 m, possibly also at sea-level.
Trav. Bot. Néerl.
3. Schuurmansia henningsii K. Scu. Bot. Jahrb. 9
(1888) 210; K. Scu. & Hoxie. FI. Kais. Wilh.
Land (1889) 50; WARB. Bot. Jahrb. 13 (1891) 283;
E. & P. Nat. Pfl. Fam. 3, 6 (1895) f. 75; K. Scu. &
Laut. Fl. Schutzgeb. (1901) 448; PuULLE, Nova
Guinea 8 (1912) 667; HALL. f. Rec. Trav. Bot.
Néerl. 10 (1913) 346; E. & P. Nat. Pfl. Fam. ed. 2,
21 (1925) f. 41; LANE Poo.e, For. Res. (1925) 116;
Wuite & FRANCIS, Proc. R. Soc. Queens]. 38
(1927) 247; Wuitr, J. Arn. Arb. 10 (1929) 241;
KaANIs, Nova Guinea, Bot. 6 (1961) 65, f. 1b;
Blumea 16 (1968) 76. — S. bamleri K. Scu. &
Laut. Fl. Schutzgeb. (1901) 448; Nachtr. (1905)
318, incl. var. longifolia Laut.; HALL. f. Rec.
10 (1913) 350. S. gilgiana
Laut. in K. Sch. & Laut. Fl. Schutzgeb. Nachtr,
118
FLORA MALESIANA
[ser. I, vol. 7!
(1905) 319; HALL. f. Rec. Trav. Bot. Néerl. 10
(1913) 348. — S. microcarpa Capit. Bull. Soc.
Bot. Bre57) 910) 398; t. 112-12.) 17. BAKERS J.
Bot. 61 (1923) Suppl. 4. — S. theophrasta HALL. f.
Rec. Trav. Bot. Néerl. 10 (1913) 346; HoLTH. &
Lam, Blumea 5 (1942) 213. — S. pseudopalma
HALL. f. Rec. Trav. Bot. Néerl. 10 (1913) 347.
— §. rauwolfioides HALL. f. l.c. 349; K. Scu.
Bot. Jahrb. 9 (1888) 211, sine nomen. —S. longifolia
(LAuT.) GiLG in E. & P. Nat. Pfl. Fam. ed. 2, 21
(1925) 80; A. C. Smitn, J. Arn. Arb. 22 (1941) 524.
— ? S. lophiroides Gia, I.c. 80. — ? S. oreophila
GILG, /.c. 80. — ? S. schlechteri GiLG, lI.c. 80. —
? S. crassinervia GILG, l.c. 80. — S. coriacea A. C.
Smitu, J. Arn. Arb. 22 (1941) 525. — S. montana
A. C. SmiTH, /. c. 526. — S. grandiflora A. C.
SMITH, /.c. 527. — Fig. 8-10.
Treelet or tree, up to 15(—-20?) m, sometimes
with stiltroots up to 1 m high. Stipules 14-5 by
14-3 mm, often up to 3 mm long, ciliate. Leaves
obovate-lanceolate, 6-85 by 114-15 cm, obtuse to
acuminate at apex, tapering at base, margins some-
what involute, nerves 2-7 mm apart, chartaceous
or subcoriaceous; petiole up to 4 cm. Inflorescen-
ces 7—65cm; peduncle 114-8 mm 9; pedicels filiform,
2-5 mm; bracts usually very small, + triangular,
sometimes larger and transitional to leaves.
Flowers % or functionally unisexual, erect. Sepals
obovate to elliptic, 3-5 by 114-3 mm, greenish,
sometimes purplish. Petals obovate-oblong, 4-714
by 114-4 mm, white, creamy, pink or purplish red.
Outer staminodes 0-c\, filiform, c. 1mm long;
inner ones 5-30, filiform to linear, 114-2 mm
long. Stamens in 3 flowers with 34-214 mm long
filaments; anthers 1144-214 by 14-34 mm; in 92
flowers with 14-14 mm long filaments; anthers
c. 1 by 14 mm. Ovary subglobular to ovoid, -+
3-lobed, glabrous, in g flowers 14-1 by 14-34 mm,
in 2 flowers c. 3 by 2 mm; style cylindric, up to
1mm; stigma 3-lobed. Fruit fusiform, up to
144 by 34 cm, acuminate. Seeds c. 1 by 14 mm
with c. 214 mm long, slender wings.
Distr. Solomons and Bismarcks; in Malesia:
Moluccas (Talaud Is., Halmahera, Ceram),
New Guinea.
Ecol. From sea-level up to 3000 m, in primary
and secondary forests, on landslides and in
secondary grasslands, on level land to steep slopes,
sometimes near river-banks or in swampy lo-
calities, on clay or more sandy or rocky soils;
scattered or locally common in open _habi-
tats.
Vern. Moluccas, Talaud Is.: arisusu uruné;
Tidore: malétopé; Ambon: ut lapu, was lapu;
W. New Guinea, Ajamaru: batsjevak, Maibrat;
Biak I.: rambuan; NE. New Guinea, Wabag:
opaga, orpach; Hagen: pappai, popai, porkai,
pupai; Minj: bubar, bubus; Chimbu: akeéssa,
a ulareh, hahéssa, ménmén; Aiyura: arebi; Budemu:
sipulund; Huon Pen., Sambui: pelip; Sattelberg:
(m)beli; Andarora: haiwinge, Nauti, yatsiga,
Manki; SE. New Guinea, Tari: obbo; Kutubu:
karadéwa; Mendi: op; Kairuku: engefukenge:
Buna: kembusa; Bausa: wérawera.
Notes. The striking variation in vegetative
characters is for the greater part due to differences
in ecological conditions. The greatest influence
is exercised by the altitude. Papuan material, ar-
ranged according to increasing altitude, shows a
very regular decrease in leaf size, especially be-
tween 1000 and 3000 m. At the same time the
leaves become more coriaceous and more distinct-
ly petioled. Exposition and age of the plants play
a less important role in the determination of the
leaf size.
Fig. 10. Schuurmansia henningsii K. Scu. Small-
leaved, young plant, Edie Creek, Morobe Distr.,
c. 2000 m, MICHAEL holding the twig (NGF 13949)
(Photogr. SLEUMER, 1961).
The dimensions and colours of flower parts are
also rather variable, but less easy to correlate
with other data. These variations probably indicate
genetical differences between local populations.
More detailed local studies will be necessary to
decide whether any infraspecific taxa can be dis-
tinguished.
A majority of the specimens has relatively large
anthers and small pistils, whereas a minority
has relatively sma!l anthers and large pistils.
It was observed once, that most specimens in a
profusely flowering stand had proportionally large
anthers and did not set fruit. It is not certain, how-
ever, that the flowers are always functionally uni-
sexual. Polygamy is likely to occur, as the reduc-
tion of either stamens or ovary is never complete.
Excluded
Sauvagesia erecta LINNE, Sp. Pl. 1 (1753) 203; Mia. Fl. Ind. Bat. 1, 2 (1857) 118; Koorp. Exk. Fl. Java
2 (1912) 607; BACKER & Baku. f. FI. Java 1 (1963) 327; KANis, Blumea 16 (1968) 80.
1971] OCHNACEAE (Kanis) 119
SND gum we a eee
Erroneously recorded for Java by MIQUEL without reference to material or other source of information.
His error might be caused by a collection of Sauvagesia erecta L. in G, which is labelled: ‘Perrottet, Java,
1819. Although Perrottet visited the eastern part of Java in that particular year, it is clear that the speci-
men concerned was mislabelled and in fact was collected in Cayenne on the same voyage.
Tetramerista Mig. Fl. Ind. Bat. Suppl. (1860) 534 belongs to the Theaceae, fide GiLG, Ber. Deut.
Bot. Ges. 11 (1893) 22, et auct. div.
al
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ily Piglet. Pitas ate dr
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2
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4 - @ 7 ‘>
- =
ian ' : :
Ssremcig se t
7 : od =
PORTULACACEAE (R. Geesink, Leyden)
Annual to perennial, erect or creeping, mostly branched herbs or shrubs, oc-
casionally woody at the base, often with a tuberous or swollen main root, oc-
casionally rooting at the nodes. Leaves spirally arranged to opposite, subsessile,
occasionally with axillary hairs or scales (in Mal. only in Portulaca), nervation
pinnate or reticulate. Flowers bisexual, actinomorphous (occasionally cleistoga-
mous), in axillary and/or terminal thyrsi, dichasia, in terminal capitules or solitary
(terminal or axillary). Bracts leaf-like or membranous. Sepals 2 (4-8 in extra-Mal.
Lewisia and Grahamia), boat-shaped, deltoid to obovate at base shortly connate
and confluent with petals and stamens. Peta/s (3—)4-6(-8), mostly obovate and
unequal, shortly connate. Stamens (1-)3-~, in 1-~ + distinct whorls; filaments
basally shortly connate; anthers 2- or 4-celled, dorsifixed, dehiscing lengthwise.
Ovary superior or half-inferior, originally 2—20-celled, soon becoming 1-celled;
style with 2-20 mostly papillous arms. Ovu/es 4-~ on a central, dendroid placenta,
campylotropous. Capsule 3—7-valved or with a caducous operculum, occasionally
surrounded by the persistent calyx. Seeds 1-~, smooth or ornamented, kidney-
shaped to + globular, laterally compressed, mostly with a caruncle. Embryo
curved, almost filling the ripe seed.
Distribution. About 15 genera with possibly 200 spp. Cosmopolitan, with some tropical species
occurring as adventives in temperate regions. In Malesia 4 genera with 11 spp.
Ecology. The Malesian species are all more or less succulent herbs or semi-shrubs, the aquatic
Montia fontana excepted. Only two genera are native, Portulaca and Montia.
All Portulacas occur at low altitude, preferably in disturbed vegetation, waste places, or on the coast.
Most are indifferent to climate, only P. macrorhiza and P. pilosa ssp. sundaensis are restricted to the
seasonal climate of the Lesser Sunda Is. The sandy beach is tolerated by P. oleracea, P. lutea, and P.
pilosa ssp. pilosa (race tuberosa).
The occurrence of Montia fontana in the high mountain bogs of New Guinea is singularly interesting,
as it is the only tropical montane occurrence in the Old World between its range in the temperate northern
hemisphere and the southern counterpart in the SE. Australian Alps (at c. 36° SL), SE. Australia (Mt
Lofty Ranges), Tasmania, and New Zealand, an almost bipolar type of distribution.
Flower-biology. As far as known all Portulacaceae are self-pollinating.
Whether cross-pollination occurs, deserves further study. Self-pollination results in genetically constant
local populations, representing pure lines in nature.
The production of seeds is profuse; seeds can usually stand a long time of drought.
Man has doubtless been responsible for the dispersal of several Portulacaceae. Seed can be transported
with cargo; some species are used as food or as ornamentals, and they easily escape as adventives and
maintain in warm regions. For these reasons the native range of certain species cannot any longer be
ascertained, notably of Portulaca oleracea, P. pilosa, P. quadrifida, Talinum paniculatum, and T. triangulare.
The structure of the inflorescence in this family is mostly, possibly always, of a cymose nature. Through
contraction and reduction several modifications are represented; some derivations are schematically
depicted in figures 1 & 2. Talinum has in principle a thyrse (fig. la, 1b & 2) of which a full explanation is
given in the notes under the genus. At first sight Calandrinia would seem to possess a true raceme (fig. 1d)
but this could also well be a reduced thyrse. Montia has also probably a reduced cymose inflorescence
(fig. 1c). The inflorescences in Portulaca are in subg. Portulacella (Australia) the most primitive and are
compound dichasia from which the structure in the capituliform inflorescences of the other subgenus
can be derived, as I have explained in my precursor (Blumea 17, 1969, 277).
Phytochemistry. Red betacyanins (replacing anthocyanins) and yellow betaxanthins form an
outstanding chemical character of the family. Such nitrogen-containing pigments (chromoalkaloids)
were demonstrated to be present in members of the genera Anacampseros, Calandrinia, Claytonia,
Montia, Portulaca, Spraguea, and Talinum. Saponins seem to occur frequently in the family; their chem-
istry, however, is not yet known. The seeds store starch in the perisperm and fatty oil in embryos;
at the same time they contain appreciable amounts of protein. The chemical features known at present
~ from this family confirm its intimate relationships with other members of the centrospermous alliance.
A summary of phytochemical literature is to be found in HEGNAUER, Chemotax. d. Pfl. 5 (1969) 383-387.
— R. HEGNAUER.
(121)
he? FLORA MALESIANA (ser. I, volar
d
Fig. 1. Scheme of the inflorescences of some Portulacaceae and their morphological relations. a. Thyrsus
in Talinum paniculatum, b. corymboid thyrsus, resembling a pleiochasium, in Talinum triangulare, c.
raceme-like inflorescence, cymose or racemose, in Montia fontana, d. raceme with terminal flower in
Calandrinia grandiflora.
Morphology. VON PoELLNITZ (in Fedde, Rep. 37, 1934, 240, and other papers) called the sepals
“Involucralblatter’’, suggesting their homology with the involucral leaves surrounding the capituli, the
latter being called by him ‘‘falsche Involucralblatter’. Also LEGRAND (in Com. Bot. Mus. Hist. Nat.
Montevideo 31, 1, 1953, 1, and other papers) called the calyx lobes ‘‘pseudosepalos”’. |
Taking into account, that the calyx normally is considered as (metamorphosed) leaves and that in
Portulacaceae the phyllotaxal position of the sepals is so distinctly set off against the bracts, it seems to
me that there is no morphological argument to accept the flower as being monochlamydeous.
Anatomy. See also my precursory paper (Blumea 17, 1969, 276-279) on the axillary hairs; METCALFE
& CHALK, Anat. Dicot. 1 (1950) 153; KowaL, Monogr. Bot. 12 (1961); CHorINSKI, Oest. Bot. Z. 80
(1931) 308.
Uses. See Heyne, Nutt. Pl. (1927); OcHsE & BAKH. Ind. Groent. (1931) 615; BURKILL, Dict. Ec.
Prod. Mal. Pen. (1935); Carus, J. Bomb. Nat. Hist. Soc. 41 (1939) 369. See also under the species.
KEY TO THE GENERA
1. Fruit dehiscing circumscissile with an operculum. Ovary half-inferior. . . . .. . . 4. Portulaca
1. Fruit valved or irregularly dehiscing. Ovary superior.
DECAY CSG uAtII ONS vOPDOSILG tiie (0 a) bos oh to ie al) des ye Le eee « by we elie aaeenie:
2. Leaves more than 6 mm long, the middle cauline leaves spirally arranged.
3. Seed glabrous. Fruit globular, 3-5 mm g. Calyx caducous ornot. ........ 2. Talinum
3. Seed hairy. Fruit obpyriform, c. 10 by 8 mm. Calyx persistent, distinctly accrescent.
1. Calandrinia
1971] PORTULACACEAE (Geesink) 123
1. CALANDRINIA
H.B.K. Nov. Gen. Sp. 6 (1823) 77, nom. cons.; POELLN. in Fedde, Rep. 35 (1934)
161-173; D.C. ANON SUAREZ DE CULLEN, Bol. Soc. Arg. Bot. 5 (1953) 1-29.
Mostly succulents, occasionally with short stems, occasionally unbranched,
glabrous, pubescent or glandular. Leaves spirally arranged and/or in a basal
rosette, occasionally with axillary hairs. Flowers arranged in mostly terminal,
occasionally axillary thyrsi (corymboid or not), glomeruli or dichasia, or solitary.
Bracts and bracteoles subulate or leaf-like, partly without an axillary axis. Calyx
accrescent (caducous in the American sp. C. punae). Sepals glabrous, pubescent,
spiny or glandular, occasionally with dentate, ciliate or glandular margin. Petals
5-7, persistent, twisted after anthesis. Stamens 3—~, anthers linear to elliptic.
Ovary superior; style 1 with 3-5 arms (seldom 3 simple styles). Fruit pyriform,
globular, obpyriform or elleptic, 3-7-valved. Seeds at least occasionally without
a caruncle.
Distr. About 150 spp., native in tropical and subtropical America and Australia.
1. Calandrinia grandiflora LINDL. Bot. Reg. (1828) smaller. Sepals roundish, up to c. 14% cm @,
t. 1194: G. Don, Gard. Dict. 3 (1834) 80, t. 18; | acuminate in fruit. Peta/s 5-6, + orbicular, up to
L. H. Bamey, Man. Cult. Pl. (1949) 336; BACKER c. 2 cma, purplish. Stamens c. 75; filaments up to
& BAKH. f. Fl. Java 1 (1963) 217. — Fig. 1d. 1 cm; anthers c. 2 by 0.8mm. Style c. 24% mm,
Erect, up to c. 30cm. Leaves spirally arranged, with several short, thick lobes. Fruit obpyriform,
obovate, up to 13 by 5 cm, without axillary hairs; c. 10 by 8 mm, 3-valved. Seeds ~, c. 1mm 9;
petiole semi-amplexicaulous; apex acute to acu- testa cells about hexangular, with hair-like ap-
minate. Flowers in compound terminal racemes at _pendages.
least occasionally with a terminal flower; each Distr. Native of S. America, sometimes escaped
raceme 1—6-fiowered. Bracts and bracteoles ovate from gardens.
to elliptic, acute, up to c. 1 by 14 cm, apical ones
2. TALINUM
Jussieu, Gen. (1789) 312, nom. cons. prop.; POELLN. in Fedde, Rep. 35 (1934)
1; Danby, Taxon 18 (1969) 464. — Fig. 1a, b, 2.
Herbs or semi-shrubs (in Malesia glabrous). Leaves spirally arranged (the lower-
most sometimes opposite), linear to obovate. Flowers in terminal whether or not
corymboid thyrsi, or cymosely arranged, seldom axillary or solitary. Sepals
mostly caducous. Petals mostly 5. Stamens 5-~. Ovary superior; style mostly
with 3 arms. Fruit globular; mostly 3-valved or irregularly caducous.
Distr. About 50 spp., native in S. and Central America and S. Africa, the two treated species now
pantropically naturalized.
Morph. The inflorescences of the Talinums studied are explained in comparison with the vegetative
ramifications, thus in agreement with C. TROLL, who assumes that there is no essential difference between
the vegetative and the inflorescential ramification. Their leaves are spirally arranged, the lowermost
sometimes excepted. Specimens of T. paniculatum grown at Leyden in winter under low light intensity
and short day light produced all opposite leaves and opposite primary inflorescential branches. The
flowers of T. paniculatum are arranged in a wide thyrse with a terminal flower (fig. 1a). A change in
phyllotaxis, in contrast with the rule of TRoLL mentioned, is found in the ultimate dichasial ramifications.
Superficially, 7. triangulare is trichasial in the first ramification of the inflorescence (fig. 1b, 2b). I
have tried to bring this in agreement with the inflorescence of T. paniculatum by the following argu-
mentation: the vegetative branches always have 2 basal cataphylls, each with a dormant axillary bud
b (fig. 2a C, B). The inflorescences lack these cataphylls and buds. The cataphylls easily fall, and mostly
| the scars are hardly to be found. The cataphylls are also present if the axillary axis is not developed
(fig. 2a’). In the cultivated specimens these dormant buds developed after the axillary axis was pinched
<i
124 FLORA MALESIANA [ser. I, vol. 7?
= A
Sat at
Fig. 2. Talinum triangulare (JACQ.) WILLD. Schematic demonstration of the homology of the vegetative
ramifications and those of the inflorescence. In all three figures L means leaf bearing the lateral axis A.
The main axis is indicated by M. Each lateral axis carries 2 cataphylls at the base indicated by C, each
cataphyll having an axillary bud indicated by B, which remains normally dormant in the vegetative part
(fig. a), but develops into an axis in the inflorescence (fig. b), but can by removal of the lateral axis also
artificially be produced (as in fig. c). For further explanation see the text.
out (fig. 2c, A), but this only succeeded near the inflorescence. Now the inflorescence may be explained
as follows: The main axis (fig. 2a, b, c M) forms the central flower. The axillary axis (fig. 2a, c A; fig. 2b A)
of the leaf (fig. 2a, c L; fig. 2b L, not visible, cf. arrow), is branched cymosely. From the dormant buds
(fig. 2a, c B) branches are developed, which are also branched cymosely, and are only slightly poorer than
the axillary axis. So the axillary axis (fig. 2b A) is supposed to represent the first and single racemous
ramification of a thyrsus.
KEY TO THE SPECIES
1. Flowering axes sharply triangular in cross-section. Branches with 2 lateral, basal buds (scars!).
1. T. triangulare
2. T. paniculatum
1. Talinum triangulare (JAcQ.) WILLD. Sp. Pl. 2
(1799) 862; HEYNE, Nutt. Pl. (1927) 612; PoELLN.
in Fedde, Rep. 35 (1935) 15; W. H. Brown,
Useful P]. Philip. (1950) 520; BACKER & BAKH. f.
Fl. Java 1 (1963) 217. — Portulaca triangularis
JAcQ. Enum. Pl. Carib. (1760) 22. — Portulaca
racemosa L. Mant. (1771) 242. — JT. racemosum
(L.) Rowers. in Mart. FI. Bras. 14, 2 (1872) 297;
DEN BERGER, Trop. Natuur 7 (1918) 28, t. 1-3. —
Fig. 1b, 2.
Erect semi-shrub up to c. 1m. Leaves elliptic
to obovate, up to 15 by 5 cm, acute to acuminate;
nervation pinnate. Axillary buds with 2 subulate,
small cataphylls; the latter always with a dormant
axillary bud up to 114 cm in the side-axes (caducous
in dried specimens leaving a scar). Thyrsi terminal,
corymboid, up to c. 12 cm @, the axes sharply
triangular with 8—c. 28 flowers. Bracts and brac-
teoles subulate. Sepals deltoid, c. 4.6 by 314 mm,
acuminate. Petals 5, obovate, up to 10 by 4mm,
emarginate, pink. Stamens 20-40; filaments up
to 5 mm; anthers c. 0.7 by 14 mm. Style c. 2.7 mm,
2-3-fid. Fruit up to 5mm g, 2-3-valved, yellow.
Seeds ~, c. 1.2 mmg@; testa cells radially elongated,
smooth, tubercled at the edge.
Distr. Pantropic weed, still extending its range,
native of tropical America. According to Dr.
LEEUWENBERG it extends enormously along new-
made ways in Africa.
Ecol. Waysides, waste places, edges of forests.
Uses. A commonly used vegetable, easily
propagated by cuttings, profusely used in war
prison camp gardens, in Java imported from
Surinam (cf. HEYNE, Nutt. Pl. 1927, 612).
Vern. Surinam purslane. Malesia:
blanda, posleén, 8S.
krokot
2. Talinum paniculatum (JACQ.) GAERT. Fruct. 2
(1791) 219, t. 128; HeyNe, Nutt. Pl. (1927) 612;
POELLN. in Fedde, Rep. 35 (1935) 10; A. C. Smit,
Bull. Torr. Bot. Club 70 (1943) 537; GREENWOOD,
J. Arn. Arb. 30 (1949) 75; BACKER & BAKH. f. FI.
Java 1 (1963) 217. — Portulaca paniculatum
Jacq. Enum. PI. Carib. (1760) 22. — Portulaca
1971]
PORTULACACEAE (Geesink)
125
patens L. Mant. (1771) 242. — T. patens (L.)
WILLD. Sp. Pl. 2 (1799) 863; Hassk. Hort. Bog.
(1858) 74; DRAKE DEL Cast. Ill. Fl. Ins. Mar.
Pac. (1890) 111; F. M. BartLey, Queensl. Fl. 1
(1899) 95; Backer, Ann. Jard. Bot. Btzg Suppl. 3
(1910) 416; Schoolfl. (1911) 84; Crars, FI.
Siam. Enum. 1 (1925) 110; Merr. & CHUN,
Sunyatsenia 1 (1934) 56; F. B. H. BRown, Bern.
P. Bish. Mus. Bull. 130 (1935) 77. — Fig. 1a.
Erect semi-shrub up to c. 70 cm. Leaves elliptic
to obovate, up to 11 by 5 cm, acute to acuminate;
nervation pinnate. Axillary buds with 2 subulate
cataphylls. Terminal thyrsi up to c. 18 by 15 cm,
with up to c. 10 dichasia, each with up to 30
flowers. Bracts and bracteoles subulate. Sepals
suborbicular, c. 1.1 mm, acute. Petals (4-)5(-6),
obovate, c. 4 by 2mm, pink, apex emarginate.
Stamens (4—)15; filament c. 2.7mm; anthers
c. 0.4mm. Style c. 1.6 mm, 3-fid. Fruit c. 3 mm g,
yellow or pink, 3-valved. Seeds c. 1.2mm 9g;
testa cells radially elongated, shortly tubercled or
not, with small pits between the cells.
Distr. Pantropic weed, native of tropical
America, still extending its range.
Ecol. Waysides, waste places, edges of forests.
Uses. Cultivated as an ornamental, and with
edible leaves. According to HEYNE (Nutt. Pl. 1927,
611) a decoction of the roots is used by Chinese
as a surrogate aphrodisiac.
Notes. 1. My cultivated specimens had pink
sepals, petals, filaments, and style. According to
field-labels, there are in America also forms with
white and yellow petals.
2. I received at the Leyden Hortus seed of ob-
viously two strains (pure lines?) recognizable by
the seed-coat; the one has small tubercles on the
cells of the testa, the other has no such tubercles.
Specimens raised from these two seed types in the
glass-house of the Leyden Hortus on vegetable
mould showed different characters, as tabulated
below:
Seed with tubercles:
Leaves up to 7 by 4 cm.
Panicle c. 8 cm wide.
Dichasia with up to 15 flowers.
Fruit reddish.
Seed without tubercles:
Leaves up to 11 by 5 cm.
Panicle c. 15 cm wide.
Dichasia with up to 30 flowers.
Fruit yellowish to olivegreen.
From the herbarium it appears that these 2 seed
types occur almost throughout the range of the
species. The correlating characters are difficult to
observe in the herbarium, and besides are liable
to be influenced by the ecology of the locality
(leaves and panicle). There are no transitions and
obviously these two strains keep constant as a
sort of pure lines.
3. MONTIA
LINNE, Sp. Pl. (1753) 87; SWANSON, Brittonia 18 (1966) 229. — Fig. Ic, 3.
Copiously branched, succulent herbs or waterplants. Basal rosette mostly
absent. Leaves ovate to linear, occasionally parallel-veined, without axillary hairs.
Ue
Fig. 3. Montia fontana L. a. Flowering branch, x 114, b. ditto, «5, c. flower, «10, d. sepals, «10,
e. corolla, * 10, f. stamens, * 10, g. young fruit with style, «10, /. burst fruit with 1 seed, » 20 (a-h
Brass & Mever Drees 9972, L).
126
FLORA MALESIANA
[ser: I, vols?
Flowers in axillary and/or terminal cymes or solitary. Calyx persistent. Petals
5. Stamens 3 or 5; anthers 4-celled. Ovary superior; style arms 3. Fruit pyriform to
globular, 3-valved, after dehiscence occasionally twisted. Seeds 1-S.
Distr. About 50 sp». Temperate and warm-temperate in Europe, Northern Asia, N. & S. America,
Central Africa, and in Tasmania and New Zealand (the present species only!); the sole tropical localities
in the Old World are on Mt Kilimanjaro in Central Africa and the high mountains of New Guinea.
Generic delimitation, e.g. against Claytonia, is not very satisfactory.
1. Montia fontana LINNE, Sp. Pl. (1753) 87. —
M. lamprosperma CHAM. Linnaea 6 (1831) 565;
MERR. & Perry, J. Arn. Arb. 23 (1942) 386;
GEESINK, Pac. Pl. Areas map 184 (inedit.). — Fig.
Ics.
Aquatic herb, in patches up to c. 6cm high.
Leaves opposite, elliptic, up to 4 by 2mm, not
caducous but decaying. Flowers solitary or 2-3
together with a membranaceous bract, inserted
in the axil of a leaf. Sepals suborbicular, c. 1.2
mm g, acute to mucronate. Petals 5, obovate to
spathulate, 2 larger, up to 1.6 by 1 mm, 3 smaller,
up to 1.6 by 0.6mm, the latter each with an
epipetalous stamen. Filaments up to c. 1mm;
anthers c. 0.34 by 0.17 mm. Style arms 3, sub-
@; the 3 valves twisted after dehiscence. Seeds
2-3, c. 1.2mm 4g; testa cells radially elongated.
Distr. Northern hemisphere in temperate lo-
calities, Americas (Rocky Mts, Andes), Central
Africa (Mt Kilimanjaro), SE. Australia (Alps and
Mt Lofty Ranges, south of 36° SL), Tasmania,
and New Zealand; in Malesia: West New Guinea
(Mt Wilhelmina) and East New Guinea (between
Mt Dickson and Kuputivava). This peculiar dis-
tribution in Australasia reminds of that of
Hydrocotyle vulgaris L. (cf. Fl. Mal. 1, 4, 1949, 116).
Ecol. Stream banks, 3500-3650 m.
Note. A variable species of which several in-
fraspecific taxa are distinguished, which to some
have the status of species.
sessile, c. 0.1 mm long. Fruit globular, c. 1.2 mm
4. PORTULACA
LInneE, Sp. Pl. (1753) 445; Pax & HorrM. in E. & P. Nat. Pfl. Fam. ed. 2, 16a
(1934) 246; PoELLN. in Fedde, Rep. 37 (1934) 240; LEGRAND, Com. Bot. Mus.
Hist. Nat. Montevideo 31, 1 (1953) 1; ibid. 34, 1 (1958) 1; GEESINK, Blumea 17
(1969) 275. — Fig. 4-6.
Mostly succulent, copiously branched herbs. Leaves linear to orbicular, in most
species with axillary hairs. Flowers in (1—)2-30-flowered, terminal capituli; re-
ceptacle infundibular, mostly with hairs or scales in the axils of the bracts (and
bracteoles), surrounded by a whorl of c. 3-30 involucral leaves (or in some Austra-
lian spp. in axillary and/or terminal dichasia). Sepals occasionally keeled or hooded,
persistent or caducous with the petals, stamens and style. Petals 4-6(—8), mostly
obovate, occasionally emarginate or mucronate. Stamens 4—~, in 1 whorl. Ovary
half-inferior, occasionally apparently inferior; style with 2-18 arms. Capsule with
a caducous circumscissile operculum. Seeds ~ (in some Australian spp. 1-4).
Distr. Mainly tropical and subtropical all over the world, not more than c. 40 spp., possibly several
very polymorphous, a few worldwide anthropochorous weeds.
Ecol. Savannahs, shores, in pastures and ruderal places, along waysides, etc.
Uses. P. oleracea is a well known vegetable with cultivars, P. guadrifida and other species are used as
medicine (see Caius, J. Bomb. Nat. Hist. Soc. 41, 1939, 369), P. pilosa sens. lat. cultivated for ornamental
purpose.
Morph. In my opinion (Blumea 17, 1969, 277) the capitulum is a condensed cyme. It is surrounded
by 3-18 involucral leaves, not to be confused with the “‘Involukralblatter”’ in the sense of VON POELLNITZ.
The nerved scales between the flowers are considered bracts and bracteoles and have, like the vegetative
leaves, axillary hairs. In P. quadrifida the small white tubercles at the base of a tuft of hairs are supposed
to represent the bracts and bracteoles.
The axillary hairs are mostly homologized with stipulae, but in my opinion (/.c.) they are probably
not of stipular nature. They are mostly found in 2 tufts in the leaf-axils. In Malesian spp. they are con-
fluent at the base. In P. quadrifida they are present in a whorl around each node.
Taxon. In my precursor (/.c.) I have modified and simplified the infrageneric subdivision of the genus.
Of the two subgenera, one is confined to Australia (subg. Portulacella).
1971] PORTULACACEAE (Geesink) 127
As to the species concept, the one adopted here is distinctly larger than currently accepted, including
that used by VON POELLNITZ and LEGRAND. As I have shown, and experimentally checked in all four
genera, selfing in bud seems the rule in the family, which explains that in nature pure lines are formed,
and that populations may consist of several pure lines keeping distinct for the small characters in which
they differ. For those who want to give a name and rank to any constant difference there is hence ample
opportunity for distinguishing varieties, subspecies, and even species, especially in variable complexes
such as those of P. oleracea and P. pilosa. In view of this genetical situation it seems that according to a
normal specific concept there are far less good species than distinguished by VON POELLNITZ and LE-
GRAND who recorded some 130 for the world and c. 62 (+21 varieties) for the Americas only respectively.
KEY TO THE SPECIES’
1. Allleaves opposite. Hairs intra- and interpetiolar. 1. Sect. NEOSSIA . ...... 1. P. quadrifida
1. At least the middle-cauline leaves spirally arranged. Hairs if present only axillary. 2. Sect. PORTULACA.
2. The largest leaves obovate to spathulate. Axillary hairs inconspicuous. Sepals distinctly carinate
(20 magn.). 1. Subsect. Portulaca.
3. Mostly 1-3 involucral leaves with an axillary axis. Capituli (2—)3—30-flowered. Stamens 7-15.
Fruit c. 4 mm long. Se 2. P. oleracea
3. Involucral leaves mostly without axillary axes. Capituli (1—)2—3(-6)-flowered. Stamens 18-50.
Fruit c. 7 mm long. SE Te TTR £8 ee ee See chy A ie is ty 3. P. lutea
2. The largest leaves linear to elliptic ‘(obovate to spathulate in P. macrorhiza from Timor). Hairs
mostly conspicuous. Sepals not carinate (20 magn.), occasionally with an apical, dorsal, about
dome-shaped hood. 2. Subsect. Stellulato-tuberculatae.
4. Largest leaves obovate to spathulate, c. 8mm wide. Timor..... .
4. Largest leaves linear to elliptic, less than 4 mm wide. Pantropic . .... .
4 ah Se ake ee ei ee DER ee a ves Veta 1é
4. P. macrorhiza
5. P. pilosa
1. Subgenus Portulaca
GEESINK, Blumea 17 (1969) 288. — Subg. Euportulaca SPEGAZZINI, Ann. Soc.
Ci. Argent. 82 (1917) 17; POELLN. in Fedde, Rep. 37 (1934) 242.
Leaves opposite or spirally arranged. Flowers sessile, solitary or in glomeruli.
Note. The single other subgenus Portulacella is exclusively Australian, and has peduncled flowers in
cymes.
1. Section Neossia
LEGRAND, Com. Bot. Mus. Hist. Nat. Montevideo 34, 1 (1958) 3; GEESINK, Blumea
17 (1969) 289. — Subsect. Tuberculatae POELLN. in Fedde, Rep. 37 (1934) 243. —
Subg. Enantiophylla LEGRAND, Com. Bot. Mus. Hist. Nat. Montevideo 31,
(1953) 5.
Leaves opposite. Hairs or scales intra- and interpetiolar. Bracts and bracteoles
reduced to small, white tuberculi at the base of a tuft of hairs or scales (in the
capitulum).
Distr. About 7 spp., pantropic, 5 in Africa, 1 in South India and Ceylon.
1. Portulaca quadrifida L. Mant. PI. 1 (1767) 73;
DC. Prod. 3 (1828) 354; Decne, Herb. Timor.
Descr. (1835) 120; Hassk. Pl. Jav. Rar. (1848)
437; Mig. PI. Jungh. (1855) 396; Fl. Ind. Bat. 1, 1
(1858) 1061; Dyer, FI. Br. Ind. 1 (1874) 246;
KING, J. As. Soc. Beng. 59, ii (1890) 145; BACKER,
Pl. (1927) 613; Backer, Onkr. Suiker. (1930)
242, Atlas t. 255; PogELLN. in Fedde, Rep. 37
(1934) 275; BACKER & Baku. f. FI. Java | (1963)
216; GEESINK, Blumea 17 (1969) 290. — P.
meridiana L. f. Suppl. (1781) 248; BL. Bijdr.
(1826) 1136; BLANco, FI. Filip. ed. 3, 2 (1878)
Fl. Bat. 1 (1907) 79; Voorl. (1908) 18; Schoolfl.
~ (1911) 84; Merr. FI. Manila (1912) 200; Int.
Rumph. (1917) 217; Gisss, Arfak (1917) 211;
Mere. Sp. Blanc. (1918) 142; Ripv. FI. Mal. Pen.
1 (1922) 151; Merr. En. Philip. 2 (1923) 136;
Rip. Fl. Mal. Pen. 5 (1925) 289; Heyne, Nutt.
162. — P. quadrifida var. meridiana DC. Prod. 3
(1828) 354; Decne, Herb. Timor. Descr, (1835)
120; Boer. in Veth. Midd. Sumatra 4 (1884) 17.
— Fig. 4.
Creeping herb, rooting at the nodes; nodes
with a whorl of hairs. Leaves elliptic to cordate,
FLORA MALESIANA
{ser. I, vOILa-
F. VIJISMA "69
cz XY Zi
Fig. 4. Portulaca quadrifida L. a. Habit of part of a plant, «144, b. flowering end of a branch, x5,
c. flower, x 5,d. corolla, <7, e. sepals, x7, f. stamens, <7, g. young fruit, operculum partly removed,
<7, h. node with a whorl of hairs, x 5 (a—h WEBER S.n.).
—
2-20 by 0.8—7 mm with c. 5 mm long axillary hairs.
Flowers terminal, 1(-3) on an infundibuliform,
profusely hairy stem-apex, at edge with 4, seldom
more leaves. Sepals c. 3 mm long. Petals 4, obo-
vate, up to 5 by 4mm, yellow. Stamens 8 or 12;
filaments up to 314 mm; anthers c. 0.30 by 0.30
mm. Style up to c. 4mm with (3—)4(-5) arms.
Fruit + obovate, up to c. 2-314 by 3 mm; oper-
culum c. 74 the height, shining, straw-yellow. Seeds
cw, 0.8-1 mm g, dull; testa cells elliptic, radially
arranged, the surface either convex or with a
pyramidal tubercle.
Distr. Pantropic, except Australia and the
Pacific east of Samoa, throughout Malesia.
Ecol. Waysides, deforested land, ruderal places,
often between gravel or in sand, not on the sandy
beach, below c. 300m. Fi. fr. Jan.—Dec.
Uses. A native vegetable and used as a medi-
cine. See HEYNE, Nutt. Pl. (1927) 613; Carus, J.
Bomb. Nat. Hist. Soc. 41 (1939) 369; QuiIsUM-
BING, Med. PI. Philip. (1951) 284; BurKILL, Dict.
ed. 2, 2 (1966) 1833.
Vern. Rumput ségan, Mal. Pen.; gélang pasir,
krémi, J; Philip.: maravigalok, Ilk., Ib., sayikan,
Tag.; djalu djalu bubudo, Ternate.
Note. Several authors have determined speci-
mens of P. pilosa as P. quadrifida, especially in
New Caledonia and in the rest of the Pacific.
2. Section Portulaca
1. Subsection Portulaca
GEESINK, Blumea 17 (1969) 291. — Sect. Carinatae POELLN. in Fedde, Rep. 37
1971]
PORTULACACEAE (Geesink)
129
(1934) 242. — Sect. Portulaphiton LEGRAND, Com. Bot. Mus. Hist. Nat. Monte-
video 31, 1 (1953) 6.
At least the middle-cauline leaves spirally arranged. Hairs, if present, only
axillary. Bracts and bracteoles membranous.
2. Portulaca oleracea LINNE, Sp. Pl. (1753) 445;
BL. Bijdr. (1826) 1136; DC. Prod. 3 (1828) 353;
Mia. Pl. Jungh. (1855) 396; Fl. Ind. Bat. 1, 1
(1858) 1061; Sum. (1860) 150; BENTH. & F. v. M.
Fl. Austr. 1 (1863) 169; Dyer, FI. Br. Ind. 1
(1874) 247; BLANco, FI. Filip. ed. 3, 2 (1878) 162,
t. 164; Kina, J. As. Soc. Beng. 59, ii (1890) 144;
Backer, FI. Bat. 1 (1907) 80: Voorl. (1908) 18;
Schoolfi. (1911) 84; Merr. Fl. Manila (1912) 200;
Int. Rumph. (1917) 217; Sp. Blanc. (1918) 142;
Ripi. Fl. Mal. Pen. 1 (1922) 151; Merr. En.
Philip. 2 (1923) 136; BACKER & SLooT. Theeonkr.
(1924) 110, t. 110; HeyNe, Nutt. Pl. (1927) 612;
BACKER, Onkr. Suiker. (1930) 243, Atlas t. 254;
POELLN. in Fedde, Rep. 37 (1934) 258; BACKER &
BAKH. f. Fl. Java 1 (1963) 216; GeESINK, Blumea
17 (1969) 292. — P. diptera Zipp. ex SPAN.
Linnaea 15 (1841) 207, nom. nud.
Herb, up to c. 40 cm. Leaves spirally arranged
to subopposite, obovate to spathulate, up to
40 by 20 mm, with inconspicuous, up to c. 1 mm
long axillary hairs. Capituli 2—30-flowered. Mostly
1-2(-3) of the 2-8 involucral leaves with an
axillary axis. Flowers surrounded by up to c. 5
by 6 mm long bracteoles and inconspicuous hairs.
Sepals up to c. 6 by 6 mm, carina up to c. 3 by 2
mm. Petals (4-)5, broadly obovate, up to 7 by 6
mm, yellow. Stamens 7—10(-15); filaments up to
4 mm; anthers 0.2-0.5 by 0.2-0.4 mm. Style up to
c. 5 mm with (4-)5 arms. Fruit ovate, c. 4 by 3 mm;
operculum 14-14, as high as the fruit, shining,
straw-yellow. Seeds ~, 1,-1.2 mm 9g, granulate;
testa cells stellulate, with many fine tubercles.
Distr. Pantropic, throughout Malesia.
Ecol. Waysides, deforested and ruderal places,
sandy shores, even on exposed rocks, up to 1800 m.
FI. fr. Jan.—Dec.
Uses. Eaten as a vegetable and used as a med-
icine. See HEYNE, Nutt. Pl. (1927) 612; OCHsE &
Baku. Ind. Groent. (1930) 615; BURKILL, Dict.
(1935) 1833; Carus, J. Bomb. Nat. Hist. Soc. 41
(1939) 369; W. H. Brown, Useful Pl. Philip. 1
(1950) 520.
Vern. Purslane, E, postelein, porselein, D;
gélang, S, M, J, gélang pasir, sesegan, M, krokot,
J, ré-séréjan, Mad., silobar pinggan, M (Sum.),
segan jantan, rumput béremi, M (Mal. Pen.);
Philip.: alusiman, ausiman, galusiman, Bik.,
olasiman, sahikan, ulisiman, Bik., Tag., golasiman,
kolasiman, makabling, Tag., bakbakad, luigum,
If., dubdupil, Bon., vigalug, Ilk., kantatdba, Pan-
gasinan; djalu djalu kiki, Ternate.
Notes. Several authors distinguished the sub-
species (or varieties) oleracea (syn. sylvatica) and
sativa, the latter is the cultivated form. The dif-
ference is probably caused by a different chromo-
some number (according to a letter of G. Tiscu-
LER to O. DEGENER) added to a Hawaiian specimen
in the Bishop Museum: “‘sativa’’ would be a
hexaploid with 2n = 54, “‘sylvatica’’ a diploid
with 2n = 18. In the herbarium these two taxa
cannot sharply be separated: “‘sativa” is generally
tall, with large seeds, ‘sylvatica’ is smaller,
with smaller seeds, but they are connected by
transitional specimens also.
For an anatomical comparison of the seeds of
these two forms, see KOwAL, Monogr. Bot. 12
(1961) 1, who also found the characters overlapping .
3. Portulaca lutea Forster [PI. Esc. (1786) 72,
nomen] ex SEEMANN, FI. Vit. (1865) 9; GEESINK,
Blumea 17 (1969) 291.
Herb, up to 70(?) cm. Leaves spirally arranged
to subopposite, obovate to orbicular, up toc. 30 mm
long, not translucent in the dried state, with
up to 6mm long axillary hairs. Capituli 1-6-
flowered. Flowers surrounded by up to 4 by 2 mm
long bracteoles and up to c. 2mm long hairs.
Sepals suborbicular up to c. 9 mm long, fleshy in
centre, with an up to 4 by 114 mm long, dorsal,
apical keel. Petals 5, broadly obovate to obovate,
up to c. 10 mm, emarginate to mucronate, yellow.
Stamens 18—c. 50; filaments up to c. 4 mm; anthers
up to c. 0.7 by 14 mm, at least occasionally red.
Style up to c. 5mm with (4-)S arms. Fruit +
ovate, c. 7 by 5mm; operculum 7 as high as the
fruit, shining, straw-yellow. Seeds o., up to c. |
mm @g; testa cells stellulate, flat, convex, whether
or not with tubercles or spines.
Distr. Pacific Islands, from Samoa and New
Caledonia to the Marquesas, not yet recorded
from Malesia.
2. Subsection Stellulato-tuberculatae
POELLN. in Fedde, Rep. 37 (1934) 242; Geesink, Blumea 17 (1969) 293. — Sect.
_Rotundatae PorLin. in Fedde, Rep. 37 (1934) 242. — Sect. Pseudohipsoclasis
LEGRAND, Com. Bot. Mus. Hist. Nat. Montevideo 31, | (1953) 7. — Sect. Catocla-
sis LEGRAND, Com. Bot. Mus. Hist. Nat. Montevideo 31, | (1953) 8; ibid. 34, 1
(1958) 10, excl. subsect. Squamosae.
130
FLORA MALESIANA
[ser. I, vol. 71
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E.VITSMA ‘69
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Fig. 5. Portulaca macrorhiza GEESINK. a. Branch, nat. size, b. flowering end of a branch, x2, c. flower,
x5, d. sepals, «5, e. corolla, the frontal petal halved, x5, f. stamens, <7, g. young fruit, the operculum
partly removed, 7, h. node with axillary hairs, x5 (a-h ZIPPELIUs s.n., type, HLB 200, 850, L).
4. Portulaca macrorhiza [Zipp. ex SPAN. Linnaea
15 (1841) 207; Mia. FI. Ind. Bat. 1, 1 (1858) 1060;
Pax & HorFrM. in E. & P. Nat. Pfl. Fam. ed. 2,
16c (1934) 247, nom. nud.] GEESINK, Blumea 17
(1969) 293. — Fig. 5.
Herb, up to 10cm. Leaves spirally arranged,
obovate to spathulate, up to c. 30 by 8mm,
obtuse to truncate; axillary hairs up to c. 4mm.
Capituli 2—-3-flowered. Flowers surrounded by up
to c. 3mm long hairs and c. 114 by 1.7 mm long
bracteoles. Sepals «<. 5.3 mm. Petals broadly
obovate, c. 7 (by 5?) mm, yellow. Stamens c. 60;
filaments up to c. 4mm; anthers c. 0.6 by 0.4 mm.
Style c. 4mm with 5 arms. Fruit globose, c. 3 mm
@; operculum about half as high as the fruit,
shining, straw-yellow. Seeds ~, elliptic, c. 0.7 by
14 mm; testa cells elliptic, + stellulate, shining.
Distr. Malesia: Lesser Sunda Islands (W. and
E. Timor).
Ecol. On limestone karst, c. 350 m, in E. Timor
(Baucau), only once noted. Fi. fr. Dec. (one
record).
1971]
PORTULACACEAE (Geesink)
131
Uses. As a medicine against gonorrhoea.
Vern. Nati biti, Timor.
5. Portulaca pilosa LINNE, Sp. PI. (1753) 445;
BACKER, Schoolfl. (1911) 84; Merr. FI. Manila
(1912) 235; En. Philip. 2 (1923) 136; Rip. FI.
Mal. Pen. 5 (1925) 289; HEYNE, Nutt. Pl. (1927)
612; PoELLN. in Fedde, Rep. 37 (1934) 261;
HENDERSON, Mal. Wild FI. 4 (1949) 33; BACKER
& BAKH. f. FI. Java 1 (1963) 216; GEESINK,
Blumea 17 (1969) 294.
Variable herb. Leaves spirally arranged, obovate
to linear, 2-30mm long, with axillary hairs.
Capituli 1-12-flowered. Flowers surrounded by
bracteoles and hairs. Petals 4—6. Stamens (6—-)10—
75. Style arms 4-8. Fruit ovate, globular or obo-
vate; operculum 14 to 34 as high as the fruit.
Seeds ~; testa cells elliptic to stellulate.
KEY TO THE SUBSPECIES
1. Ripe seeds bluish, grey, or dull black.
2. Cells of the testa convex to pyramidal, without
a nipple. Petals 214-12 by 1.8-11 mm.
1. ssp. pilosa
2. Cells of the testa flat, but at seed-edge nippled.
Petals larger, up to c. 25 mm g.
2. ssp. grandiflora
1. Ripe seeds dark brown or shining black.
3. Cells of the testa stellate, in shape orbicular,
sometimes nippled. Flowers orange to red.
3. ssp. sundaensis
1. ssp. pilosa. — GEESINK, Blumea 17 (1969) 295.
— P. tuberosa Roxs. FI. Ind. ed. Carey 2 (1832)
464; Dyer, FI. Br. Ind. 1 (1874) 246; BACKER,
Voorl. (1908) 19; Schoolfl. (1911) 85; PoELLN.
in Fedde, Rep. 37 (1934) 312; BACKER & BAKH. f.
FI. Java 1 (1963) 216 (race tuberosa). — P. australis
ENDL. Atakta Bot. (1833) 7, t. 6; ? J. J. Smitn,
Teysmannia 10 (1899) 92 (race australis). —
P. helianthemoides Zipp. ex SPAN. Linnaea 15
(1841) 207, nomen (race tuberosa). — P. cincta
FENZL, Nat. Tijd. N. I. 14 (1857) 162; Mia. FI.
Ind. Bat. 1, 1 (1858) 1061; BoeRL. Handl. 1
(1890) 85; PoELLN. in Fedde, Rep. 37 (1934) 312
(race tuberosa). — P. filifolia F. vy. M. Fragm. 1
(1859) 169 (race filifolia). — P. sclerocarpa [non
Gray, Bot. U.S. Expl. Exp. 1 (1854) 141] Koorp.
Minah. (1898) 345; Koorp.-ScHum. Syst. Verz.
3 (1914) 40, record corrected to P. pilosa (race
pilosa). — P. pachyrhiza [non GAGN. Bull. Soc.
Bot. Fr. 56 (1909) 41] Merr. En. Philip. 2 (1923)
136 (race tuberosa). — P. samoensis POELLN.
in Fedde, Rep. 33 (1933) 163; ibid. 37 (1934) 300
(race tuberosa). — P. javanensis POELLN. Rev.
Sudamer. Bot. 7 (1943) 273, incl. var. grisea
POELLN. (race tuberosa).
Herb, 10-c. 30cm. Leaves elliptic to linear,
4-28 by 14-4 mm; axillary hairs 1-18 mm long.
Capituli (1—)2—10-flowered; more or less congested
with adjacent quasi-axillary capituli. Flowers sur-
rounded by 0.7—214 by 0.7—2.2 mm long bracteoles
and 3-18 mm long hairs. Sepals 2-6 by 1-4 mm,
occasionally inconspicuously hooded at apex.
Petals 4-6, obovate, 214-12 by 1.8-11 mm,
yellow or pink. Stamens (7—)20—30(—35?); filament
1-5 mm; anthers globose to elliptic, 0.35-0.7 mm.
Style 2-8 mm, 3-7-fid. Fruit + globose, c. 2—3(-4)
mm @; operculum 14 to % as high as the fruit,
shiny, straw-yellow to olive green. Seeds 0.4—0.7
mm @, dull light to dark grey, or bluish; testa
cells elliptic, elliptic with lobes or stellate; pyra-
midal all over the seed, or not.
Distr. Pantropical, throughout Malesia.
Taxon. A _ subspecies consisting of several
‘races’, which are easy to distinguish as shown in
the following table.
Race Range Petals Stamens Length Seed Colour Shape Surface
anthers rs) seed testa cells testa cells
pilosa pantrop., pink (7-)20-30 0.5 0.4-0.6 blue stellate pyramidal
excl. Austr.
tuberosa_ E. Asia, yellow (17-)25-30 0.7 0.6-0.7. dark elliptic, lobed, convex
Malesia, grey stellate at seed
W. Pacific edge
filifolia Australia, yellow 12-30 0.4 0.6-0.7 bright stellate pyramidal
Java: in grey
Hort. Bogor
P. australis ENDL. is entered in the synonymy,
but it remains uncertain whether this race was
indeed represented by the material J. J. Smitu
identified from the island Groot Kombuis (Bay
- of Djakarta), as no specimen is available. The
Australian race filifolia is only represented by
specimens collected in the Botanic Gardens at
Bogor.
Uses. A native vegetable and used as a medi-
cine. See Ocuse & BAKH. Ind. Groent. (1931) 615,
and Catus, J. Bomb. Nat. Hist. Soc. 41 (1939) 369.
Vern. Rose-flowered purslane, E, pénawar, M
(Mal. Pen.), rebha ledkah, Mad.; Philip.: romruki,
Ilk.; njalé njalé wolanda, Ternate.
2. ssp. grandiflora (HooK.) Gresink, Blumea 17
(1969) 297. — P. grandiflora Hoox. Bot. Mag.
n.s. 3 (1829) t. 2885; Backer, Voorl. (1908) 19;
FLORA MALESIANA
E VITSMA’69
Fig. 6. Portulaca pilosa L. ssp. sundaensis (POELLN.) GEESINK. a. Habit, x 114, b. flowering end of a branch
x5, c. flower, <5, d. sepals, x7, e. corolla, x7, f. stamens, <7, g. young fruit, 7, A. node with axillary
hairs, <5 (a-h JaaG 799, L).
Schoolfl. (1911) 85; BRUGGEMAN, Ind. Tuinb.
(1938) 84, t. 55; STEEN. FI. Sch. Indon. (1949)
176; BACKER & BAKH. f. Fl. Java 1 (1963)
2G:
Herb, up to c. 30 cm? Leaves linear, up to 25
by 3 mm, with c. 5 mm long axillary hairs. Flowers
in each capitulum up to c. 5, flowering succedane-
ously, each surrounded by up to 10 mm long hairs
and deltoid, up to 4.7 by 2 mm long bracteoles;
transitions between leaves and bracteoles present.
Sepals c. 8mm long, with a very small apical
keel. Petals + obovate, c. 25mm @g, pink, red,
orange or yellow (cultivated in many colours).
Stamens c. 40-75; filaments up to c. 6mm;
anthers elliptic, c. 1.4 by 0.3 mm, 2- and 4-celled.
Style up to c. 13 mm, with 5-18 arms. Fruit +
globose, c. 5 mm @; operculum 7% to 14 as high as
the fruit, shining, straw-yellow. Seeds c. 0.7
mm @g, shining; testa cells + stellulate, those
at the edge of the seed with a central tubercle.
Distr. Native of tropical America, cultivated
as an ornamental and occasionally escaped
elsewhere.
3. ssp. sundaensis (POELLN.) GEESINK, Blumea 17
(1969) 298. — P. sundaensis POELLN. in Fedde,
Rep. 50 (1941) 105. — Fig. 6.
Herb, up to c. 10 cm. Leaves oblong to obovate,
up to 7 by 3mm, with up to 6 mm long hairs.
Capituli 2—3-flowered. Flowers surrounded by up
to 0.7 by 1 mm long bracteoles and c. 6 mm long
hairs. Sepals 5, c. 2.6 by 2.2 mm. Petals obovate,
subacute, c. 2.3 by 1.7mm, orange. Stamens
(6—)10-15; filaments c. 114 mm; anthers c. 0.4 by
0.35 mm. Style c. 1.6 mm, with 5 arms. Fruit +
globose, c. 244mm @; operculum 4 as high as
1971] PORTULACACEAE (Geesink) 133
the fruit, dull, grey-brown. Seeds c. 0.55 mm og, +
shining; testa cells stellulate.
Distr. Malesia: Lesser Sunda Islands (Sum-
bawa, Flores, Wetar, Alor), from each island one
specimen.
Ecol. Close to the coasts, along roads, between
rocks.
Vern. Tamadsiang ata, Alor.
Excluded
Portulaca axilliflora (non PERS.) BLANCO, FI.
Filip. ed. 2, 2 (1845) 285 and P. toston BLANCO,
Fl. Filip. (1837) 408 are both, according to MErR-
RILL, En. Philip. 2 (1923) 136, Trianthema portu-
lacastrum L. (Aizoac.).
Portulaca teretifolia L.; F.-VILL. Nov. App.
(1880) 15, an American species, probably an in-
correct identification.
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BYBLIDACEAE (C. G. G. J. van Steenis, Leyden)
In the former century Byblis was mostly included in the Droseraceae, for example
by BENTHAM & Hooker. f. (Gen. Pl. 1, 1859, 220); even ENGLER had it in that
position in 1912 (Syllabus ed. 7, 329). PLANCHON had in 1848 (Ann. Sc. Nat.
III, 9, 1848, 80, 90) already pointed to affinity with Cheiranthera of the Pitto-
sporaceae; HALLIER f. merged Bybdlis and Roridula with Tremandraceae, curiously
referring this to an Ochnaceous assemblage (Abh. Gebiete Naturw. Hamburg 18,
1903, 53). About the same time LANG argued (Flora 88, 1901, 179) that on mor-
phological and anatomical grounds Byblis cannot belong to Droseraceae, but should
be referred to Lentibulariaceae.
Diets (Pfl. R. Heft 26, 1907, 51) and Domin (Act. Bot. Bohem. 1, 1922, 1)
definitely concluded to the alliance with Pittosporaceae, and so did HUTCHINSON
(1926, 1959) and SCHULTZE-MENZ (Syllabus 1964): resemblance with Drosera
is superficial, sympetaly unimportant. HALLIER f. and HUTCHINSON include the
S. African genus Roridula also in the family Byblidaceae, but others regard this as
an allied family.
BYBLIS
SALIsB. Parad. Lond. (1808) t. 95; Bru. Fl. Austr. 2 (1864) 469; LANG, Flora 88
(1901) 179; Domin, Act. Bot. Bohem. 1 (1922) 1; Bibl. Bot. 22 (1929) 702; Diets
in E. & P. Nat. Pfl. Fam. ed. 2, 18a (1930) 288; Vester, Bot. Arch. 41 (1940) 563,
map 192. — Fig. 1.
Erect herbs, viscid, by longitudinal rows of minute, sessile glands and capitate-
glandular hairs. Leaves linear to filiform, involute-coiled in vernation, exstipulate,
spirally arranged. Flowers axillary, solitary, without bracteoles, 5-merous. Sepals
imbricate, persistent, short-connate at the base. Petals larger than sepals, contorted,
with a broad, dentate apex and cuneate base, oblique, -+ connate at the very
base. Stamens 5, sometimes unequal; anthers basifixed, alternipetalous, cells
opening with a very short pore-like slit. Disk none. Ovary superior, 2-celled, with
~ ovules attached to the axis of the dissepiment about the middle; integument
1; style 1, simple, with a faintly 2-lobed stigma. Capsule + compressed, 2-celled,
loculicid with 2 valves, sometimes the valves later splitting, the dissepiment
splitting + halfway. Seeds dark, rugose; embryo elongate, cylindric; cotyledons
short, fleshy; albumen present.
Distr. Two spp., one in SW. Australia, the other from NW. to NE. Australia, in Malesia: the N.
Australian species in South New Guinea.
Ecol. This is the fifth genus of insectivorous plants in Malesia, the others being Nepenthes, Utricularia,
Drosera and Aldrovanda. Both species grow in depressions which are swampy on poor soils or which
become swampy or water-logged in the wet season. Often gregarious.
The way of catching insects (small flies, mosquitoes, moths and ants) superficially resembles that in
Drosera, but differs in that the capitate-glandular hairs make no movement towards the prey. RICA
Erickson (Austr. PI. 3, 1966, 319, 321) calls it a ‘flypaper trap of the passive type’. According to GRIEVE
(ibid. 1, n. 9, 1961, 23) “insects are usually first caught by the sticky mucilage exuded from the gland-
tipped hairs and these tend to collapse or bend as they pour out secretion. The insect is thus also brought
into contact with the numerous, minute sessile glands and becomes enveloped in additional secreted
fluid. The process of secretion and absorption continues until all of the soft parts of the insect are dis-
solved and absorbed, and only the hard, indigestible parts remain. The glands then stop secretion and
the stalked ones commence to recover to their normal position. In due course the hard parts of the insect
which are left dry out and fall off.”
(135)
FLORA MALESIANA [ser. I, voli 7
Fig. 1. Byblis liniflora SALIsB. a. Habit, x 24, b. stem tip, 4, c. leaf tip, 12, d. flower, corolla removed
x4, e. anther, x12, f-g. fruit, x4, h. seed, x8, i. flower, x3, j. flower, corolla removed, x4, k. anther
x 12, 1. lengthwise section of ovary (a-—h HOOGERWERF 273, i-/ BYRNES 230).
It has been suggested that the capitate-glandular hairs secrete a sticky mucilage, but that the secretion
of the sessile ones is less sticky and would serve mainly for digesting proteins, but I have no pertinent
data to sustain this opinion.
The large West Australian species, B. gigantea, is well-known as the ‘rainbow plant’, a name ‘‘believed
to be derived from the fact that on looking through the plant towards the setting sun, one can see a
spectrum of colours where the edges of the leaves are bordered by the shining drops of liquid on the
glands.”
Anat. FENNER (Flora 93, 1904, 382-388) gave a detailed account of the anatomy of the glands of
B. gigantea LINDL.
1971]
BYBLIDACEAE (van Steenis)
137
1. Byblis liniflora SALisB. Parad. Lond. (1808) t.
95; DC. Prod. 1 (1824) 319; ENpDL. Iconogr.
(1841) t. 113; Bru. FI. Austr. 2 (1864) 470;
F. v. M. Fragm. 10 (1876) 81; BAILEY, Queensl.
Fl. (1900) 551; Compr. Cat. (1913) 174, f. 145;
BRITTEN, Ill. Pl. Banks 1 (1900) 30, t. 96; Ewart
& Davies, Fl. North. Terr. (1917) 117; Domin,
Act. Bot. Bohem. 1 (1922) 4; Bibl. Bot. 22
(1929) 703; SpecutT, Rec. Am.-Austr. Exp.
Arnhem Land 3 (1958) 231; STEEN. Blumea 16
(1968) 355. — Fig. 1.
Unbranched, weak herb, c. 15-40 cm. Rhizome
thin. Leaves filiform, very thin, blunt, c. 4-6 cm,
spreading. Pedicels already from the base of the
plant, usually exceeding the leaves, in fruit up to
c. 10cm, patent, the lower ones reflexed. Pistil
and stamens somewhat zygomorphic. Sepals
ovate-lanceolate, acute, erect, 3-4 mm, with
scarious margin, glabrous, c. 3-4 by 114 mm.
Petals oblanceolate, acute, with dentate upper
margin, 4-8(-11) mm. Stamens glabrous, anthers
varying from 114-3 mm, filaments longest where
anthers are shortest. Ovary glandular; style c. 3
mm. Capsule transversely elliptic, with 2 shallow
grooves, c. 2 by 4mm. Seeds ellipsoid, at one
end + pointed, black, 1 mm, lengthwise ribbed,
ladder-like tessellate between the ribs.
Distr. Queensland to NW. Australia, in Ma-
lesia: South New Guinea (near Merauke: Hoo-
GERWEREF 273), one collection.
Ecol. Marsh herb of shallow, seasonal swamps in
Melaleuca-Acacia-Tristania-Eucalyptus savannah,
at very low altitude. Fr. March.
CLETHRACEAE (H. Sleumer, Leyden)
1. CLETHRA
LINNE, Gen. Pl. ed. 5 (1754) 188; Sp. Pl. (1753) 396; SLEUM. Bot. Jahrb. 87 (1967)
55. — Fig. 1-5.
Trees or shrubs, evergreen (Mal. spp.); leaf-scars large. Leaves crowded towards
the end of the shoots, spiral, simple, exstipulate, serrate with glandular teeth,
often with an apical gland, more rarely entire; nerves a little decurrent along the
midrib, both midrib and nerves + impressed above, + prominent beneath. In-
dumentum of branchlets, leaves and inflorescences consisting of simple, and/or
long, fascicled and + patent, and/or minor, -+ depressed stellate hairs. Flowers
bisexual, regular, 5(—6)-merous. Inflorescences sometimes simple solitary terminal
racemes, but mostly consisting of a terminal raceme and several lower approx-
imate racemes, each of the latter from the axil of a + reduced or caducous leaf,
thus forming together a panicle-, fascicle- or umbel-like inflorescence; bracts mostly
caducous during anthesis, rarely subpersistent. Calyx lobes 5(-6), persistent,
quincuncially imbricate, united at the base only. Petals 5(-6), generally free, some-
times cohering to some degree, alternate with the calyx lobes, rather early caducous,
generally sweet-scented. Stamens 10(—12) in 2 whorls of 5(—6), the outer whorl
opposite the petals, the inner one opposite the calyx lobes; filaments adnate to the
corolla at the extreme base; anthers dorsifixed, overturned outwards in bud, erect
in anthesis, introrse, upper part of cells -- divergent, opening with apical, slit-
like pores; pollen grains single, tricolporate, psilate. Ovary superior, 3-celled, with
axile placentation; ovules ~, small, anatropous; style simple, mostly shortly,
very rarely hardly divided into three apical lobes, sometimes more deeply so and
trifid, each lobe stigmatic at the top. Fruit a 3-valved, loculicidal capsule, the
septae of which become loose from the persistent central axis, subtended or + en-
closed at maturity by the persistent calyx. Seeds ~, small, subovoid to irregularly
angular or subtrigonous, with a foveolate-reticulate testa (all Mal. spp.). Endo-
sperm fleshy. Embryo cylindrical.
Distribution. A small, monogeneric family in the Ericales, of (sub)tropical Asiatic-Malesian, and
temperate and tropical American distribution, and with 1 sp. in Macaronesia (Madeira, and formerly in
Teneriffe).
Of the total of 64 spp., 2 temperate spp. are found in North America (C. alnifolia L. and its var. pu-
bescens Ait., and C. acuminata MicHx), 38 spp. inhabit (sub)tropical America (Cuba-Jamaica-Central
America, in S. America in the Mt Roraima area and in Trinidad, in the Andes from Venezuela to SE.
Bolivia and NW. Argentina, and a small separate area in SE. Brazil), 10 spp. are found in SE. Asia
(Upper Burma, SW., Central and SE. China, Japan, Indo-China).
In Malesia 13 spp., of which 2 in the Malay Peninsula, 1 in Sumatra, | in Java, 3 in the Lesser Sunda
Islands, 4 in the Philippines, 3 in Borneo, 2 in Celebes, 1 in the Moluccas, and 4 in New Guinea. Fig. 2.
Ecology. Clethra occurs mainly in the lower and upper montane primary (and also secondary)
forest. As to climate, the genus prefers the wet montane tropical and subtropical zone, though it extends
far into the northern temperate zone, in North America to about 45°, in Japan to about 42° N, whilst its
southern limit is about 29° S in SE. Brazil. As the majority of the Ericales, Clethras have preference for
acid soils. They occur in Malesia from sea-level to about 3000 m; and up to c. 3800 m in Szechuan and
Yunnan, and in the Andes. They are light-demanding, remain sterile in dense forest, and flower in forest
borders with open vegetation or exclusively in the latter, also in light secondary plant communities.
Dispersal. Nothing is known about the dispersal of Clethras. The rather small, deeply impressed-
reticulate and light seeds of the Malesian spp. point to dispersal by wind.
Anatomy. Den Bercer, Determinatietabel van Malesié, Veenman, Wageningen (1941) 31 (hand
(139)
140 FLORA MALESIANA [ser. I, vol. 7?
——s
Ss
=& = yy
= pW FSS —— :
———SSS AL SLS)
SSS JL OARS = —s >.>
ee BSia= So Ia z
Se QZ: LS ?}
= Fes PLUMS AS ~—
=n C2. aS\Sis Ga
SS
—S—
—
———— SSS
=e = : SS g
SANE
=>
~———>
——-
=
\ \h\ ( N
\ VN OR AES » Ai
AY WEEN |
bit (
i .
=
SS
—S
=
es
wk
\y
a
Fig. 1. Clethra pachyphylla MERR. a. Habit, x 7, b. flower, <4, c. petal, x4, d. stamens in bud stage,
4, e. stamens at full anthesis, 4, f. longitudinal section of flower, <4, g. fruit, x4, h. seeds, «8
(a-f CHEW, CORNER & STAINTON 1845, g—h SINCLAIR 9082).
1971] CLETHRACEAE (Sleumer) 141
lens, wood); DEN BERGER & BIANCHI, Tectona 24 (1931) 894-903 (wood); HARA, Bot. aE? Tokyo 70
(1957) 108-114 and J. Fac. Sc. Un. Tokyo, Sect. 3, Bot. 7 (1958) 367-450 (shoot apex and leaf histo-
genesis); LEMESLE, Bull. Soc. Pharm. Bordeaux 104 (1965) 37-48 (localization of tanniferous substances);
LeMESLE & Dupuy, C. R. Ac. Sc. Paris 263 (1966) 250-253 (tannins and mucilagenous substances);
Supo, Identification of Japanese Hardwoods, Bull. Govt. Exp. Stat. Meguro 118 (1959); UpHor, Hum-
MEL & STAESCHE, Plant Hairs, in Handb. Pflanzenanat. IV, 5 (1962). For general surveys of older lit-
erature see SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899) 541-551 and ibid. (1908) 195-197 (under
Ericaceae), and METCALFE & CHALK, Anat. Dicot. 2, Oxford (1950) 836-839.
The wood shows primitive characters such as narrow solitary vessels with scalariform vessel perfo-
rations, fibre-tracheids, apotracheal parenchyma which is diffuse or arranged in short uniseriate bands and
heterogeneous rays of two sizes. The petiole is supplied with a strongly incurved to closed vascular
strand. Hairs occur as multicellular stellate and uniseriate types. Stomata mainly paracytic. Crystals soli-
tary and clustered. Secretory tanniferous cells are present in the parenchymatous tissues of petiole and
stem. METCALFE & CHALK (/.c.) state that Clethra has many anatomical features in common with
Ericaceae. — P. BAAS.
Pollination. The flowers are protandrous. The pollen is shed at least partly before the flowers are
open and the stigmas are receptive.
Galls. DocTers VAN LEEUWEN (Zoocecid. Neth. Ind. 1926, 439, fig. 825 & 826) mentions leaf-galls on
C. sumatrana from the Petani ravine near Medan, caused by a Psyllid and a gall mite. On the leaves of
C. canescens in N. Celebes (Minahasa) globular galls are found.
Uses. Temperate species of Clethra, mainly from North America, Madeira and Japan, much less
from China, are cultivated as ornamentals in many parts of the world. Not a single species from Malesia
so far has been introduced in Botanic Gardens.
Phytochemistry. Leucoanthocyanins, caffeic acid, flavonols, taraxeron and ursolic acid are known
to occur in species of Clethra (HEGNAUER, Chemotax. d. Pfl. 3, 1964, 432-433; TANABE c.s. J. Pharm.
Soc. Japan 86, 1966, 441). This spectrum of phytoconstituents is compatible with the generally accepted
relationship between Clethraceae and Ericaceae. — R. HEGNAUER.
Taxonomy. A monograph of the genus has been published by the author (Bot. Jahrb. 87, 1967,
36-175, pl. 1). In this work it was shown, that the North American and the Asiatic-Malesian species of
Clethra belong to sect. Clethra by their subovoid to subtrigonous and impressedly reticulate seeds,
whilst the Central and South American species and C. arborea in Madeira form a second section, Cuel-
laria, distinguished by flat and variously winged seeds.
For the determination of Clethras close observation of the indumentum with a lens is necessary.
There are (i) simple, mostly rather long, appressed or patent hairs, (ii) fascicled, +- obliquely erect,
generally elongated hairs, (iii) stellate hairs of generally small size and + flattish. On most Clethras a
combination of 2 or all 3 types of hairs is found, but their density may differ greatly on the various
organs, resulting in a fine tomentellous, a thicker tomentulose or tomentose to villous tomentum of
white to greyish, pale ferrugineous to dark rusty and even rufous colour. Besides the various kinds of
hairs, the discrimination of species is based on floral characters. Collectors should try to collect flowering
material and (sterile) branchlets with young, not yet glabrate leaves from the same specimen, and abstain
from collecting sterile or fruiting material.
KEY TO THE SPECIES
1. Filaments hairy.
2. Filaments hirsutulous. Style elongate in fruit (up to 12mm)... . . . 1. C. sumatrana
2. Filaments laxly set with longish hairs. Style a little or hardly elongate i in fruit (up to 7 mm).
3. Leaves subcoriaceous, practically glabrous in mature state. Inflorescence dilutely rusty-tomentellous
in all parts... . . . 2. C. hendersonii
3. Leaves coriaceous, ‘still densely stellate-hairy along midrib. and primary nerves in mature state
beneath. Inflorescence rusty-villous in all parts. . . .....4. 4.4... . . 3. C. symingtonii
1. Filaments glabrous.
4. Younger, not yet glabrate leaves (which still show the full quality and quantity of the tomentum)
laxly to more densely hairy only on midrib, nerves and veins, and glabrous on the intervenium be-
neath, mature ones glabrescent, or leaves glabrous from the beginning.
5. Lateral nerves (15—)17—20(—23) pairs, + approximate toeach other..; .°.. . 4. C. javanica
5. Lateral nerves 10—12(—14, rarely up to 15) pairs, more distant from each other. 5. C. canescens
4. Younger, not yet glabrate leaves tomentellous to tomentose all over the undersurface, mature ones
but slightly or tardily glabrescent.
6. Younger leaves + appressedly tomentellous on the intervenium beneath, i.e. covered there ex-
clusively with minute, + flattish stellate hairs.
7. Younger leaves densely covered with minute stellate hairs on the whole undersurface in a coherent
layer, and besides more laxly set with stellate-fascicled and darker (generally rusty) hairs on nerves,
veins and veinlets ... . . . 6. C. kebarensis
7. Younger leaves on the undersurface set with dense minute ‘stellate hairs, which form a coherent
pale layer, other larger and/or darker stellate-fascicled hairs very few, or mostly absent.
142
FLORA MALESIANA
[ser. I, vol. 73
8. Leaves regularly serrate in the upper 24, with (15—)17—20 pairs of lateral nerves.
8. Leaves subserrate to entire, with 10-14 (rarely up to 15) pairs of lateral nerves.
9. Petals 2—2.2(—2.5) mm.
9. Petals (3.5—)4(—5) mm.
6. Younger leaves +-
7. C. sumbawaensis
8. C. tomentella
9. C. arfakana
erectly tomentulose or tomentose on the intervenium beneath, i.e. covered there
with minute and more coarse stellate (suberect), or with minute stellate and coarse (+ erect)
fascicled hairs.
10. Racemes generally more slender and elongate (flowers + laxly arranged), and tomentellous by
minor stellate, rather flattish hairs. .
10. C. longispicata
10. Racemes generally more robust and shorter (flowers -t =. ’ densely arranged) and tomentulose to
tomentose by a combination of minor stellate and robust, + erect fascicled hairs.
11. Pedicels 1-2 mm at anthesis. Style 1-1.5 mm at anthesis, hardly 2 mm in fruit. Lateral nerves of
leaves -- inarching before the margin .
. 11. C. papuana
. Pedicels | 2-3(—5) mm at anthesis. Style 2.5- 3 mm 1 at anthesis, 3-4 mm in fruit. Lateral nerves of
leaves -- excurrent along the margin.
12. Ovary tomentose by long erect hairs.
12. Ovary tomentulose by short hairs. .
1. Clethra sumatrana J. J. S. Ic. Bog. 4, 1 (1910)
t. 319; MerrR. Philip. J. Sc. 14 (1919) 248; Doct.
v. LEEUWEN, Zoocecid. Neth. Ind. (1926) 439,
f. 825 & 826; METCALFE & CHALK, Anat. Dicot.
2 (1950) 838, f. 192, F-G (anat.); SLEUM. Bot.
Jahrb. 87 (1967) 77. — C. pulcherrima RiwL. J.
Mal. Br. R. As. Soc. n. 87 (1923) 75; MEeErR.
Contr. Arn. Arb. 8 (1934) 121. — C. longipedi-
cellata MERR. Pap. Mich. Ac. Sc. 19 (1934) 179.
— C. pubifolia Merr. I.c. 180; BURRET, Notizbl.
Berl.-Dahl. 15 (1940) 188, in text.
Much-branched shrub, or tree with erect or
crooked trunk and a dense crown, (3—)5-15
(rarely up to 25)m; bark whitish. Branchlets
stoutish, younger parts covered with a rusty stel-
late tomentum maybe mixed with simple hairs,
early glabrescent. Leaves lanceolate to oblong- or
obovate-elliptic, apex shortly acutely acuminate,
base cuneate, thin-coriaceous, young ones covered
with a thick tomentum of rusty to rufous shorter
stellate and longer fascicled hairs at least on mid-
rib and nerves, older ones mostly rather persistent-
ly so by both or but one type of these hairs on the
entire undersurface or only on midrib, nerves
and maybe veins, generally distantly serrate with
callose teeth especially in the upper part of the
lamina, rarely subentire or entire, reddish when
young, dark green at maturity, 4-11(-15) by
(1.5—)2-4.5(-8) cm, nerves in 8-13 pairs, curved-
spreading, slightly raised beneath as are the trans-
verse veins, reticulation not conspicuous in gen-
eral; petiole 6—-12(-15, rarely up to 30) mm.
Racemes elongate, erect-ascendent, 8-30 cm, usu-
ally solitary and terminal, more rarely also from
the axil of 2 or 3 upper leaves, the latter mostly
normal in size and + persistent, rarely reduced
and caducous during anthesis, covered by the
same, but generally more dense and more per-
sistent indumentum found in the branchlets and
leaves. Pedicels rather slender, 3-10 mm (in fruit
10-20 mm), bracts lanceolate-linear, caducous,
8-15 by 1-2.5 mm. Calyx lobes subovate or ovate-
oblong, subacute, rusty-tomentellous or -tomen-
tose in the upper part inside and all over outside,
the fascicled hairs few to numerous, (4~-)5—6(-8)
12. C. pulgarensis
13. C. pachyphylla
by 2-3mm. Petals obovate, erose-crenulate at
apex, glabrous, white-creamy to pinkish, (S—)6—7
(-9) by 4-5(-6) mm. Filaments linear, glabrous
at both ends, longish and subappressedly hirsutu-
lous between, 45mm; anthers obsagittate,
1.5-2(-2.5) mm, base extended into an acute
triangular appendix. Ovary subappressed-hirsute;
style slender, glabrous or with a few hairs at the
very base, 6-7 mm at anthesis, 8-12 mm in later
stages, apex thickened and very shortly trilobed
at anthesis, rarely more deeply so in fruit. Capsule
subglobose, hirsutulous, 4-Smmg. Seeds sub-
obovoid-oblong or subtrigonous, variously com-
pressed, c. 1.5 mm.
Distr. Malesia: Sumatra (Atjeh, Eastcoast,
Tapanuli, in the Westcoast Res. only at Brani
near Bukit Tinggi).
Ecol. In primary and secondary forest, also
in Pinus merkusii forest, (390—)900-—3000(—33007)
m. Fl. fr. Jan.—Dec.
Fig. 2. Distribution of Clethra in Malesia and
adjacent SE. Asia. The number above the hyphen
indicates the number of endemic species, that
below the hyphen the number of non-endemic
species in each island or district.
CLETHRACEAE (Sleumer)
143
Fig. 3.
6a
Clethra hendersonii SLEUM., Mt Beremban, 1720m, Malay Peninsula, Cameron Highlands
(SLEUMER 4675) (SLEUMER, 1963).
Vern. Kumbawang, Karo, bodi-bodi, boli-boli,
darik-darik, hau si martadjau, simar darih-darih,
Toba.
2. Clethra hendersonii Steum. Bot. Jahrb. 87
(1967) 79. C. canescens (non REINW. ex BL.)
Ript. J. Fed. Mal. St. Mus. 6 (1915) 158. —
Fig. 3.
Slender, small or medium-sized, few-branched
tree; bark rather smooth, greyish, minutely
longitudinally fissured. Branchlets rather slender,
dilutely rusty-stellate-tomentellous distally. Leaves
elliptic-oblong or elliptic, sometimes subovate-
elliptic, apex shortly acuminate, base cuneate and
slightly inequilateral, subcoriaceous, dark green and
dull above, paler and shining below, flush reddish,
denticulate-serrulate, 8—10(—12) by 4-5.5(—6) cm,
young ones stellate-puberulous, early glabrate
above, set with sparse simple hairs at midrib and
moreover laxly with small stellate hairs at
midrib, nerves and veins below, mature ones
finally glabrous, nerves (10—)11-13 pairs, curved-
ascendent, raised beneath, reticulation faint;
petiole rather slender, 1.5-2.2cm. Racemes 2
144
FLORA MALESIANA
(ser. I, vol
or 3(—5), subdensely set with flowers, up to 22 cm,
tomentellous by rusty-brownish simple and paler
stellate hairs in all parts. Pedicels rather robust and
3-4 mm (in fruit 6-7 mm), bracts subulate, 3-4
mm, caducous. Calyx lobes oblong, 4-5 by 2 mm.
Petals obovate, apex erose, spreading, white,
slightly fragrant, glabrous on both faces, sparsely
fimbriate, (5—)6-7 by (3-)4-5 mm. Filaments
laxly set with longish hairs, 5(—-6) mm; anthers
narrowly obcordate, base acute, 1.8mm. Ovary
subsericeous-tomentose; style glabrous, shortly
trilobed, 5-6 mm in anthesis, 7mm in fruit.
Capsule depressed-globose, 4 by 5mm, sub-
tended by the spreading sepals, Seeds oblong-
ovoid, c. 1.5 mm.
Distr. Malesia: Malay Peninsula (Pahang: Cam-
eron Highlands and MtTahan; _ Selangor:
Mt Nuang).
Ecol. Scattered in forest, often on ridges or
summits, 1340-1740 m. Fi. April—July.
3. Clethra symingtonii SLEUM. Bot. Jahrb. 87
(1967) 80.
Shrub. Tips of branchlets and innovations short-
ly rufous-villous. Leaves elliptic-oblong, apex
shortly subacutely attenuate, base cuneate, very
base sometimes obtuse, slightly inequilateral, co-
riaceous, young ones early glabrate above, +
densely set with rather large stellate hairs on mid-
rib, nerves and veins beneath as are the petioles,
mature ones persistently so, 6-9 by (2.5—)3-4
cm, glandular-subserrate-denticulate in the upper,
entire in the lower part, nerves in 9-10 pairs
excurrent along the edge, well-raised beneath,
veins transverse and prominent beneath, retic-
ulation visible; petiole 2-3 cm. Racemes terminal
and from the upper 3 or 4 axils (which are mostly
defoliate in flowering time) close together forming
a panicle, all over with a rusty, almost villous
tomentum of numerous longer crisp fascicled and
shorter stellate hairs; rachis robust. Pedicels
thickish, 2-3 mm (in fruit 5-6 mm), bracts subu-
late, caducous, 3mm. Calyx lobes oblong, c. 5
by 2.5mm. Petals obovate, apparently cup-like
converging, quite glabrous, white, scented, 6 by
3 mm. Filaments laxly long-hairy; anthers narrow-
obcordate, base acute, 2mm. Ovary densely set
with erect hairs; style subappressedly hairy in its
lower third, glabrous for the rest, 5 mm (in fruit
6mm), stigma shortly 3-lobed. Capsule sub-
globose, c. 4mm 9g.
Distr. Malesia: Malay Peninsula (Perak: G.
Kerbau For. Res.), once found.
Ecol. On open ridge top, 1830 m, not common.
FI. fr. July.
4. Clethra javanica Turcz. Bull. Soc. Nat.
Moscou 36, ii (1863) 232; Koorp. Album Na-
tuurmonum. Ned. Ind. ser. 1 (1918) 3, pl. 9
(phot.); Bull. Jard. Bot. Btzg III, 1 (1919) 148, t.
10; BAck. Bekn. FI. Java (em. ed.) 7 (1948) fam.
161, p. 1; BAck. & BAKH. f. Fl. Java 2 (1965) 178;
SLEuUM. Bot. Jahrb. 87 (1967) 83. — C. canescens
(non REINW. ex BL.) Miq. FI. Ind. Bat. 2 (1859)
1056, p.p.; Koorp. Jungh. Gedenkb. (1910) 184;
Exk. Fl. Java 3 (1912) 1, p.p.; BACK. Bull. Jard.
Bot. Btzg II, 12 (1913) 11;J.J.S.in K. & V. Bijdr.
13 (1914) 85, as to descr. — C. sp. ZOLL. Syst.
Verz. 2 (1854) 138, p.p.
var. javanica.
Small tree, 3—6(—10) m, trunk up to 15cm @,
often crooked, or much branched shrub, 2-5 m;
branches up to 5cmg. Branchlets rather robust,
covered distally with a rusty hirsute or almost
lanate tomentum as are the petioles. Leaves
lanceolate to oblong-elliptic, sometimes partly
obovate in the same specimen, apex shortly
acuminate, base cuneate, subcoriaceous, dark
green, shining and glabrous above, much paler
beneath, and set: there still towards the maturity
with both longish simple or fascicled rufous and
fewer minor stellate hairs on midrib and nerves,
whereas the veins bear only scattered stellate hairs
and the intervenium is glabrous, finally glabres-
cent, (5—)6-10(-13) by 2.5-4(-6) cm, rather reg-
ularly and deeply (1.5-2 mm) serrate, nerves
(15—-)17—20(-23) pairs, spreading, -+ approx-
imate and subparallel, rather straight from the
midrib and partly forked before the edge, +
sharply prominent beneath, veins transverse, +
raised beneath too; petiole robust, 1(—1.5) cm.
Racemes panicled or subumbellate, rachis and
pedicels covered with a floccose and rusty, almost
hirsutulous tomentum, 5—-10(-14) cm, flowers
rather densely arranged, subpatent. Pedicels rather
thickish, 3—5(-6) mm, bracts subulate, 2—4(-6)
mm, caducous. Calyx lobes ovate-oblong, sub-
acute, dorsally a little keeled and stellate-tomen-
tellous, fimbriate, (4~-)4.5-6 by 2.5mm. Petals
obovate, apex slightly erose, thin, glabrous,
white or pinkish, (5—)6—7 by c. 3 mm. Filaments
glabrous, 2.5mm; anthers obcordate, base sub-
acute, 1.3 mm. Ovary shortly rufous-hirsutulous;
style glabrous, 3—-3.5 mm, apex shortly trilobed.
Capsule subglobose, 2.5—3 mm @, pedicel elongate
to 10 mm, style slender, 3.5-4mm. Seeds ovoid,
subtrigonous or variously compressed, 0.5-0.8
mm.
Distr. Malesia: East Java (Mt Jang).
Ecol. In mixed primary forest edges, 1830-2300
m. FI. fr. April—Oct.
var. lombokensis SLEUM. Bot. Jahrb. 87 (1967) 84.
— C. canescens (non REINW. ex BL.) Mia. FI.
Ind. Bat. (1859) 1056, p.p. — C. sp. ZOLL. Syst.
Verz. 2 (1854) 138, p.p.
Leaves very similar to those of var. javanica,
but thinner, young ones on the undersurface
sparsely clothed with fascicled hairs at midrib
and nerves, and with stellate ones at the veins,
becoming almost completely glabrous at maturity,
up to 15 by 5cm, more deeply (up to 4 mm) and
more irregularly serrate.
Distr. Malesia: Lesser Sunda Islands (Lombok),
in montane forest and Casuarina bush of Mt Rind-
jani, 1700-2400 m.
Ecol. On dry soil of volcanic debris. Fi.
June—Aug.
1971]
CLETHRACEAE (Sleumer)
5. Clethra canescens REINW. ex BL. Bijdr. (1826)
863; G. Don, Gen. Syst. 3 (1834) 842; Mia. FI.
Ind. Bat. 2 (1859) 1056, p.p.; Ann. Mus. Bot.
Lugd.-Bat. 1 (1863) 41; Koorp. Minah. (1898)
515; SARASIN, Reisen Celebes 1 (1905) 29, in
text.; J. J. S. Ic. Bog. 4 (1910) 61, t. 318; Koorp.-
Scuum. Syst. Verz. 3 (1914) 100; Koorp. Bull.
Jard. Bot. Btzg III, 1 (1919) 148, in text.; ibid. LI,
2 (1920) 255; voN MALMin Fedde, Rep. 34 (1934)
284; SLeuM. Bot. Jahrb. 87 (1967) 85.
KEY TO THE VARIETIES
1. Racemes and pedicels + robust. Petals 3-4 mm.
2. Leaves early glabrescent; nerves starting from
the midrib at an + acute angle.
1. var. canescens
2. Leaves with more persistent minute rusty
stellate hairs underneath; nerves starting from
the midrib at + right angles.
2. var. luzonica
1. Racemes and pedicels slender or almost so.
3. Leaves underneath on midrib, nerves and
veins more laxly to rather densely clothed
with subpersistent rather large stellate hairs.
Petals 3-4 mm. : 3. var. clementis
3. Leaves underneath on midrib and nerves,
rarely on veins, laxly set with fascicled and/or
minute stellate hairs, early glabrescent, or
young ones already subglabrous.
4. Racemes generally elongate, rather dense-
flowered. Petals 2—3 mm. 4. var. novoguineensis
4. Racemes generally short, rather lax-flowered.
Petals (3-)4 mm. . . 5. var. ledermannii
1. var. canescens.
Tree up to 20m, bole up to 40 cm g, covered
with greyish bark; at higher altitudes a shrub,
2-5 m. Branchlets covered with a woolly-floccose
rufescent, later greyish tomentum at the tips.
Leaves elliptic or obovate-elliptic, apex shortly
subacutely acuminate, base cuneate, thin-coria-
ceous, entire below, regularly serrate with callose
teeth c. 1 mm long, younger ones glabrous above,
+ densely set with minute stellate + rusty hairs
on midrib, nerves, veins and veinlets, and with
sparse, longer simple or fascicled hairs on the
midrib beneath, glabrous on the intervenium
beneath, mature ones glabrescent, generally
(4-)5-8(—11) by (2—)2.5-4 cm, at higher altitudes
constantly smaller, (2.5—)3-5(-6) by (1-)1.5-2
cm, nerves 10-12 (rarely up to 15) curved-
ascendent pairs, which rather distinctly inarch
before the edge, veins slightly prominent, veinlets
rather inconspicuous beneath; petiole +- robust,
0.6—-1.2(—1.5) cm. Racemes few to several, forming
an erect panicle, all over covered with a floccose-
lanate, + rusty or finally greyish tomentum,
6-12(-16) cm, the numerous flowers rather laxly
set along the thickish rachis. Pedicels robust,
43-)4-5 mm at anthesis, up to 6mm in fruit,
bract subulate, up to 6mm, caducous. Calyx
lobes ovate-oblong, rather abruptly acuminate,
long-ciliate, pale rusty-floccose dorsally, 3-4 mm.
Petals obovate, crenulate all along the margin,
145
glabrous, white (or suffused with red initially),
with a rather unpleasant smell, 3-4 by c. 2 mm.
Filaments glabrous, c. 2.5mm; anthers broad-
obovate, base almost mucronate, 0.8 mm. Ovary
villous: style glabrous, 3—3.5 mm (in fruit 3.5—4
mm), very shortly trilobed. Capsule subglobose,
tomentulose, 2.5mm. Seeds irregularly ovoid-
angular, 0.8-1 mm.
Distr. Malesia: Celebes, Lesser Sunda Islands
(Flores); slightly different forms in the Moluccas
(Buru, Ceram and Ambon).
Ecol. A tree in mountain forest or forest edge,
ascending into more open summit vegetation and
shrub-like there, sometimes on volcanic tuff,
rather common locally, (1160—)1400—2800(—3000)
m. Fl. fr. Jan.—Dec.
Vern. Kaju parang, M, madausiy, Ts. (Mina-
hasa), pitjisan djabong, Makassar.
2. var. luzonica (MERR.) SLEUM. Bot. Jahrb. 87
(1967) 88. — C. luzonica MERR. Publ. Gov. Lab.
Philip. n. 29 (1905) 38; Philip. J. Sc. 5 (1910) Bot.
371; En. Philip. 3 (1923) 242.
Leaves elliptic, more rarely oblong-elliptic,
apex shortly subacutely acuminate, sometimes
more obtuse, base broadly cuneate to obtuse, sub-
coriaceous, younger ones laxly set with minute
stellate hairs above, densely clothed with rather
small floccose stellate hairs on the prominent
midrib and nerves, and more laxly so on the
veins beneath, glabrous on the intervenium,
moreover sparsely set with longer simple or
fascicled hairs on midrib and nerves beneath,
mature ones glabrescent first above, tardily
beneath, subentire or minutely (rarely more
deeply) serrate, (3.5—)4-7(-10) by (1.5—)2—4(—5)
cm, nerves 10-12(—-14) pairs, curved, rather ap-
proximate, + parallel and anastomosing; petiole
rather robust, 0.3-1(-1.3) cm. Racemes panicled
or almost umbellate, 3—6(—11) cm, covered with a
tomentum of small dark-rusty stellate floccose
hairs; flowers -+- dense along the thickish rachis.
Pedicels robust, 1(—2) mm, slightly accrescent in
fruit, bract subulate, caducous, 3-4mm. Petals
white or cream, irregularly erose at apex, 3—3.5 mm,
Distr. Malesia: Philippines (Luzon: Mountain
Prov.; Isabela Prov.).
Ecol. In montane Pinus insularis, Quercus or
Podocarpus forest, also in mossy forest or in
shrubby summit vegetation, 1220—2250(—2700?) m,
rather common locally. Fi. fr. Jan.—Dec.
Vern. Amog, kamueg, \g., apiit, Bon.
3. var. clementis (MeRR.) SLEUM. Bot. Jahrb. 87
(1967) 86. — C. clementis Merr. Philip. J. Sc. 13
(1918) Bot. 104; En. Born. (1921) 460. — C. ca-
nescens (non REINW. ex BL. s. str.) STAPF, Trans.
Linn. Soc. Bot. 4 (1894) 198; Gisss, J. Linn. Soc.
Bot. 42 (1914) 105.
Leaves oblong-lanceolate to oblong, apex
acutely acuminate, base cuneate-attenuate, thinly
subcoriaceous, younger ones first stellate-hairy,
early glabrescent above, -- densely set with fas-
cicled and stellate, rather coarse hairs at midrib
and nerves beneath, laxly or hardly so at the veins,
146
FLORA MALESIANA
[ser. I, vol. 7?
intervenium glabrous, mature ones quite gla-
brous and a little shining above, glabrescent first
at the veins, more tardily so at midrib and nerves
beneath, subentire or shallowly glandular-serrate,
(6-)7-14 by (1.5—)2—3(—3.5—-4) cm, nerves (10-)
12-14 pairs, curved-ascending towards the edge,
rather indistinctly inarching there, prominent
beneath, veins but slightly raised beneath; petiole
slender, 1—-1.5(—2) cm. Racemes panicled, slender,
10-20 cm, covered with a dilutely rusty tomentum
of both smaller and larger stellate and/or fascicled
hairs; flowers rather densely arranged along the
robust rachis. Pedicels slender, 1-2 mm (in fruit
c.3 mm). Petals erose-fimbriate at apex, 3-4 mm.
Distr. Malesia: Borneo.
Ecol. Generally in mountain (also mossy)
forest or on forest edges, rarely in lowland (ap-
parently secondary) forest, locally common,
especially on Mt Kinabalu, (30—)700—1200(—1830)
m. Fi. fr. Jan.—Dec.
Vern. Kalintuhan, mitindike, Dusun.
4. var. novoguineensis (KANEH. & HATUS.) SLEUM.
Bot. Jahrb. 87 (1967) 90. — C. longispicata J. J. S.
var. novoguineensis KANEH. & HATus. Bot. Mag.
Tokyo 56 (1942) 474, f. 3. — C. lancifolia Turcz.
Bull. Soc. Nat. Moscou 36, ii (1863) 231; MerrrR.
Philip. J. Sc. 1 (1906) Suppl. 111; ibid. 2 (1907)
Bot. 292; En. Philip. 3 (1923) 242. — C. alnifolia
(non L.) BLANCO, FI. Filip. ed. 2 (1845) 259;
ed. 3, 2 (1878) 117; MeErR. Sp. Blanc. (1918) 297.
— C. canescens (non REINW. ex BL. s. str.) F.-VILL.
Novis. App. (1880) 121; VIDAL, Sinopsis Atlas
(1883) 30, t. 60, f. B; Phan. Cuming. Philip.
(1885) 123; Rev. Pl. Vasc. Filip. (1886) 172;
Merr. En Philip. 3 (1923) 242. — C. williamsii
C. B. Ros. Bull. Torr. Bot. Cl. 35 (1908) 73, 75.
— C. castanea ELM. Leafl. Philip. Bot. 9 (1934)
3182.
Leaves lanceolate to oblong, apex shortly sub-
acutely acuminate, base + cuneate, younger ones
glabrous above, more densely or laxly set with
longish simple or fascicled, -+- appressed pale rusty
hairs on midrib and nerves, -+ laxly and minutely
stellate-hairy on veins beneath, mature ones gradu-
ally glabrescent, finally sometimes quite glabrous,
shortly or rarely more deeply and distantly serrate
in the upper part, occasionally subentire, (4~-)5-10
(-14, rarely -17) by (1.5—)2-3.5(-4.5, very rarely
-8) cm, nerves (8—)10-14 (rarely -16) pairs,
curved-ascending, -+ joined before the edge,
raised beneath, veins often obscure though a fine
reticulation is visible; petiole rather slender,
0.5—1(—1.8) cm. Racemes panicled, (4—)6—15(—24)
cm, flowers rather dense along the + slender
rachis, covered with a tomentum of longer, fas-
cicled rusty and minor pale stellate hairs (rarely
only with the latter ones). Pedicels generally rather
slender, rarely more robust, 1-2 mm (in fruit 2-3
mm). Petals irregularly erose-crenulate mainly at
apex, (2—)2.5(-3) mm.
Distr. Malesia: SE. Celebes, Philippines (Batan
Is, Luzon, Mindoro, Samar, Leyte, Biliran,
Negros, Mindanao, Catanduanes), NW. New
Guinea.
Ecol. In the Philippines and in Celebes in
mountain (also mossy) forest, mainly on ridges and
peaks, (400—)700—2100 m, in New Guinea in open
beach forest and in savannah-like thickets on dry
hill (300 m) near the coast, presumably mainly in
secondary vegetation. Fil. fr. Jan.—Dec.
Vern. Alibuiigog, C. Bis., kalyapi, kayat-
buntot, Bag., kamog, Uk., kamueg, Ig., malaklak,
Tag., maratunrgau, Ibn., tagobahi, P. Bis., mayaorin,
Batan.
5. var. ledermannii (SCHLTR) SLEUM. Bot. Jahrb. 87
(1967) 87. — C. papuana SCHLTR, Bot. Jahrb. 52
(1915) 219, non J. J. S. 1914. — C. ledermannii
Scu tr, ibid. 55 (1918) 194.
Leaves lanceolate-elliptic to lanceolate, apex
acutely acuminate, base cuneate, thin-subcoria-
ceous, younger and submature ones glabrous
above, set with sparse rather coarse rufescent
simple or fascicled and/or finer stellate hairs on
midrib and nerves, hardly at veins beneath, in-
tervenium glabrous, mature ones almost com-
pletely glabrous, rather sharply serrate in the
upper part, 5—8(-11) by 2-3(-4) cm, nerves
12-14(-15) pairs, -- curved, subparallel, rather
indistinctly inarching before the edge, prominent
beneath, veins + obscure; petiole rather slender,
(0.7—)1-1.5 cm. Racemes panicled, all over cov-
ered with a very short tomentum of brownish to
greyish stellate hairs, in which a few larger fas-
cicled hairs occur, rather lax-flowered, 5-12 cm.
Pedicels rather slender, 2—3 mm (in fruit 3-3.5 mm).
Petals erose-crenulate, (3—)4 mm.
Distr. Malesia: E. New Guinea (Sepik
Distr.; Western Highlands).
Ecol. In mountain (also mossy) forest, and in
regrowths, (200—)400-1830 m. F/. June—Sept.
Vern. Talellpi, Wapi (Torricelli Mts).
6. Clethra kebarensis SLEUM. Bot. Jahrb. 87 (1967)
92. — Fig. 4.
Laxly branched shrub or treelet, 2—3(—6) m,
trunk often blackish, 4-10cm 9g. Branchlets
distally ferrugineous-tomentellous. Leaves obo-
vate-oblong or -elliptic, more rarely oblong, apex
-+- shortly subacutely acuminate, base cuneate,
rarely obtuse or even rounded in the same spec-
imen, subcoriaceous to firmly chartaceous,
young ones very laxly stellate-hairy above, sub-
densely covered beneath with short, fascicled
rusty hairs at midrib, and all over the under-
surface with very dense minute stellate hairs,
which form a pale coherent layer, moreover
nerves and especially veins and veinlets set with
darker rusty stellate-fascicled, tardily evanescent
hairs so to say on top of the pale layer mentioned
above, mature leaves dark green above, yellowish
rusty beneath, early glabrescent above, tardily so
beneath, generally shortly to very shortly sub-
serrate- or crenate-dentate, (4-)6-9 by (2—)2.5-4(-
5.5) cm, nerves in 14-16(-17) pairs, slightly
curved or rather straight from the midrib, rather
obscurely inarching, prominent below, veins but
slightly raised; petiole 9-14 by c. 1 mm. Racemes
3-6, -- fascicled, partly branched below, erect,
1971]
CLETHRACEAE (Sleumer)
Att
7 ;
Fig. 4. Clethra kebarensis SteuM., Kebar Valley, Nertoi, 700 m, NW. New Guinea (VAN ROYEN &
SLEUMER 6841) (SLEUMER, 1961).
dense-flowered, 6-12 cm, covered with a short,
almost villous, rusty (finally greyish) pubescence
of fascicied and stellate hairs; rachis rather robust.
Pedicels thickish, 1—-1.5 mm at anthesis, subtending
bracts subovate-subulate, 2 mm, persistent for a
while. Calyx lobes ovate, subacuminate, 2-—2.5
mm. Petals obovate-spathulate, often connate or
coherent in the lower part, crenulate, glabrous,
white or cream, (2.5—)3—3.5 by c. 1.5mm. Fila-
ments dilated downwards, glabrous, 2(—2.5) mm;
anthers obcordate, c. 0.6mm. Ovary rusty-
tomentulose; style glabrous, 1.2—1.5 mm (in fruit
to 2.5mm). Capsule subglobose, c. 3mm @.
Seeds subtrigonous-ovoid, 0.8 mm.
Distr. Malesia: New Guinea (Vogelkop
Peninsula).
Ecol. In patches of shrubberies within grass-
land or fern thickets of Gleichenia, or in forest
edges, not rarely in periodically burned vegetation,
fairly common locally, sometimes forming pure
loose stands on sandy or clayey soil, 700-1200
(-1950) m. Fi. fr. Jan.—Dec.
7. Clethra sumbawaensis Steum. Bot. Jahrb. 87
(1967) 94.
Treelet, 4-8 m. Branchlets distally clad with a
rusty rather scabrid pubescence. Leaves elliptic-
oblong or elliptic, apex +. shortly and acutely
acuminate, base cuneate, thin-coriaceous, younger
and submature ones glabrous above, subdensely
set with longish simple and fascicled (or stellate)
rusty hairs along midrib and nerves, sparsely so
148
on the veins beneath, moreover beneath all over
the undersurface with a dense thin layer of minute
stellate and pale (or here and there rusty), finally
greyish hairs, apparently very tardily glabrescent,
serrate in the upper 274, (5-)6-9 by 2-4 cm,
nerves (15—)17—20 pairs, rather straight and sub-
parallel, in part forked distally, reticulately dis-
solving before the edge, raised beneath, veins
transverse, slightly prominent beneath, veinlets
obscure; petiole 1—1.5(—2) cm. Racemes rather
numerous, condensed to a panicle, 5—8(—10) cm;
rachis stoutish, covered with a floccose rusty kind
of wool. Pedicels rather slender, with a short
stellate pubescence as are the sepals, 3-4 mm,
bracts subulate, tomentulose, 8-10 mm, caducous
after anthesis. Calyx lobes ovate-acuminate, less
hairy at the fimbriate margin, glabrous inside, 3
mm. Petals obovate, thin, white, fragrant, gla-
brous, minutely fimbriate-crenulate + all along
the margin, 5 by c. 2.5 mm. Filaments glabrous, 2
mm; anthers broadly obovate, base apiculate,
1 mm. Ovary tomentose; style slenderly columnar,
glabrous, 4—4.5 mm, shortly 3-lobed.
Distr. Malesia: Lesser Sunda Islands (Sum-
bawa: Mt Batulanteh; Flores: Mt Ranaka).
Ecol. In forest, (900—)1600—2400 m, on an-
desite soil. F/. April, Oct.
8. Clethra tomentella ROLFE ex DUNN, Kew Bull.
(1922) 185; Merr. En Philip. 3 (1923) 243;
SLEUM. Bot. Jahrb. 87 (1967) 94.
Small tree or shrub. Branchlets rather slender,
tips grey-tomentellous. Leaves lanceolate to
elliptic-oblong, or oblanceolate, apex + shortly
subacutely acuminate, base attenuate, firmly
chartaceous, subentire or sometimes sparsely very
shortly serrate-dentate in the upper part, younger
and submature ones glabrous above, appressedly
greyish-stellate-tomentellous all over beneath,
mature ones glabrescent tardily, first on the inter-
venium, finally also almost completely along
midrib and nerves, 6-9(—11) by (2—)2.5—3(—3.5) cm,
nerves (10—)12-14(-15) pairs, prominent beneath,
reticulation visible only in the already glabrate
parts; petiole rather slender, (5—)6-10(—12) mm.
Racemes several, panicled, laxly or more densely
set with flowers, 7—-14(-20) cm, covered with a
short, greyish or rusty-stellate tomentum; rachis
slender. Pedicels slender, c. 2mm (c. 3mm in
fruit), bracts subulate-linear, 3-5 mm, caducous
in the beginning of anthesis. Calyx lobes ovate-
acuminate, glabrous inside in part, 1.5—2 mm.
Petals obovate-spathulate, minutely erose apically,
glabrous, white, 2-2.2(-2.5) mm. Filaments
dilated towards the base, glabrous, 1 mm; anthers
obovate, hardly 44mm. Ovary tomentellous;
style thickish, glabrous, 1mm, very shortly 3-
lobed. Capsule subglobose, 2.5mm @. Seeds sub-
trigonous-ovoid, 0.7—1 mm.
Distr. Malesia: Philippines (Luzon: Mountain
Prov. to Batangas).
Ecol. Scattered in thickets or forests on slopes,
chiefly on ridges in mossy forest, 600-1800 m.
Fl. fr. May—Sept.
Vern. Amog, kamug, kamung, Ig., ayusan, Tag.
FLORA MALESIANA
[ser. I, vol. 71
9. Clethra arfakana SLeum. Bot. Jahrb. 87 (1967)
95.
Shrub or erect treelet, 2-3 m, much branched.
Branchlets slender, tips rusty-tomentellous. Leaves
obovate-oblong or -elliptic, apex shortly acutely
acuminate, base cuneate, slightly inequilateral,
subcoriaceous to firmly chartaceous, flush red-
dish, younger and mature ones dark green and
glabrous above, whitish greyish below, i.e. except
the sparsely hairy or subglabrous midrib and
nerves covered by a short tomentum of minute
stellate hairs, very tardily glabrescent, (sub)entire
or shallowly dentate, (2—)3-4.5 by (1—)1.3—2 cm,
more deeply dentate and up to 7.5 by 3 cm in not
flowering new shoots, nerves in 10-12 pairs, +
straight from the midrib, curved upwards, slightly
prominent beneath; petiole 5—8(—10) mm. Racemes
several, panicled, 5—7(—8) cm, rather lax-flowered,
pale-rusty stellate-tomentellous in all outer parts;
rachis slender. Pedicels slender, 2-3 mm, basal
bracts early fugacious. Calyx lobes ovate-ob-
long, long-ciliate, 3 by 1.5mm. Petals obovate-
spathulate, sometimes coherent at the base,
glabrous, white, shortly erose-fimbriate all along
the edge, (3.5—)4(—5) by 2—2.5 mm. Filaments di-
lated towards the base, glabrous, 2.5 mm; anthers
obcordate, about 0.5mm. Ovary almost hirsute;
style glabrous, 1.8mm at anthesis, hardly ac-
crescent in fruit. Capsule subglobose, 2.5—3 mm @.
Seeds oval to subtrigonous, c. 1 mm.
Distr. Malesia: New Guinea (Vogelkop
Peninsula: Arfak Mts around the Anggi Lakes).
Ecol. In edge of Nothofagus-Myrtaceae-
forest or in more open heath formation within
devastated montane forest, 2000-2600 m, common
locally. Fl. fr. Jan.
10. Clethra longispicata J. J. S. Bull. Jard. Bot.
Btzg III, 4 (1922) 240; SLteum. Bot. Jahrb. 87
(1967) 96. — C. elongata J. J. S. Bull. Jard. Bot.
Btzg ILI, 1 (1920) 398, t. 47, non Russy, 1907.
Shrub or treelet, up to 5(-12) m, trunk up to
10cm g, bark green, with shallow pale brown
flakes. Branchlets and petioles rusty to greyish
tomentellous. Leaves oblong-lanceolate to elliptic-
oblong, more rarely elliptic, apex rather shortly
acuminate and subacute, base cuneate, rarely
almost obtuse, inequilateral, subcoriaceous, young-
er ones glabrous above, covered all over beneath
by a coherent tomentum of minor stellate cin-
namomeous to greyish hairs, and of larger stel-
late hairs so to speak on top of that layer, mature
ones tardily glabrescent beneath, furthermore
sometimes with sparse fascicled hairs on the mid-
rib beneath, (4.5-)6-11 by (1.7—)2-3.5(-4) cm,
callose-denticulate or subentire, nerves 10—12(—14)
pairs, generally starting at an acute angle, curved-
ascending towards the edge, a little impressed
above, prominent beneath, veins distinctly,
veinlets rather inconspicuously raised beneath;
petiole 0.6—-1.3(-1.6) cm. Racemes panicled,
rather slender, (15—)20-—30(—35) cm, laxly many-
flowered, with a short stellate tomentum, in which
also some fascicled hairs may be found. Pedicels
slender or almost so, 2-3 mm (to 4 mm in fruit),
1971]
CLETHRACEAE (Sleumer)
149
Fig. 5. Clethra papuana J. J. S., Kebar Valley, Mt Nettoti, 1900 m, NW. New Guinea (VAN Royen &
SLEUMER 7449) (SLEUMER, 1961).
bracts subulate, caducous. Calyx lobes narrowly
ovate-subdeltoid, 1.5—2(-2.5) mm. Petals partly
connate at base, broadly spathulate, crenulate,
glabrous, white, scented, c. 3 by 1.5 mm. Filaments
glabrous, 1.5mm; anthers obcordate, 0.5 mm.
Ovary appressedly hairy; style glabrous, c. 1.7 mm
(to 2mm in fruit), very shortly 3-lobed. Capsule
subglobose, c. 2.5 mm @o@. Seeds convex-
ovoid, 0.7 mm.
Distr. Malesia: Borneo, Philippines (Palawan)
and Central Celebes.
Ecol. Generally in mountain forest, also in
secondary vegetation, on clayey ground, (60-)
500-1525 m. Fl. Jan.—Dec.
Vern. Pongo, Toradja, lang,
Dusun.
Iban, garapoi,
11. Clethra papuana J. J. S. Nova Guinea 12
(1914) 169, t. 53; KANeH. & Hatus. Bot. Mag.
Tokyo 56 (1942) 476; SLeuM. Bot. Jahrb. 87
(1967) 97. — Fig. 5.
var. papuana.
Shrub or treelet, (1—)3—6(—8) m, more rarely a
tree up to 15m, bole up to 10m by 30cm.
Branchlets at tips rusty subvillous-tomentose.
Leaves elliptic or obovate-elliptic, sometimes ob-
long-elliptic, rarely elliptic-lanceolate, apex short-
ly subacutely acuminate, rarely rather blunt
(more acutely acuminate in sterile shoots), base
inequilateral, broadly cuneate to subtruncate-
obtuse, coriaceous, firm, younger ones with
stellate and fascicled hairs mainly along midrib
150
and nerves above, clothed all over beneath by a
dark rusty subvillous tomentum formed by
stellate and fascicled hairs, mature ones glabrate
above, dark green in the fresh state there, under-
surface rusty- to greyish- or (almost silvery-)
tomentulose (the large fascicled hairs having
mostly disappeared by then), entire or shortly ir-
regularly serrate-dentate (more deeply and sharply
serrate in sterile shoots), (4-)5-8(-12) by 2-4
(-5.5) cm, variable in size even in the same
specimen, sometimes microphyllous, nerves 12-14
pairs, much spreading, + inarching at or a little
before the edge, well raised beneath, veins not
much conspicuous beneath, reticulation faint;
petiole rusty-tomentulose, (1—)1.2-1.6(-1.8) cm.
Racemes panicled, erect, all over rusty-tomentose,
flowers generally densely arranged, rarely up to
15cm; rachis robust. Pedicels thickish, 1-2 mm
(up to 4mm in fruit), bracts thickish, subulate,
caducous. Calyx lobes oblong-ovate, 3—3.5(—4) mm.
Petals not rarely connate or coherent at base,
obovate-spathulate, edge slightly erose, white.
sometimes pale cream or suffused with red, gla-
brous, slightly scented, c. 3.5 by 2 mm. Filaments
glabrous, hardly 2mm; anthers + obcordate,
0.8mm. Ovary almost villous; style glabrous,
shortly trilobed, (1—-)1.2-1.5mm (hardly 2mm
in fruit). Capsule subglobose, c. 3mm g@. Seeds
ovoid-oblong, irregularly angular, 0.8-1 mm.
Distr. Malesia: NW. New Guinea (Vogelkop
Peninsula: Kebar Valley and Arfak Mts).
Ecol. In (mossy) Nothofagus-Conifer-forest
edge or in open heath vegetation on crest, 1800—
2600 m. Fi. Oct.—Jan.
var. trichostyla SLEUM. Bot. Jahrb. 87 (1967) 98.
Petals (3mm) laxly hairy along the edge.
Style set with a few appressed hairs at base.
Otherwise as var. papuana.
Distr. Malesia: W. New Guinea (Wissel
Lakes area), once found.
12. Clethra pulgarensis Erm. Leafi. Philip. Bot. 5
(1913) 1757; Merr. En. Philip. 3 (1923) 243;
Sieum. Bot. Jahrb. 87 (1967) 99.
Low, stunted tree, or shrub-like; branches as-
cending, rigid, covered with brown bark. Branch-
lets short, rather robust, short-hairy. Leaves ob-
long to subobovate-elliptic, apex shortly at-
tenuate or subacuminate, base broadly cuneate,
the very base often obtuse, subcoriaceous, firm,
younger ones very densely shortly stellate-hairy
above, covered all over beneath with a tomentum
of minor and larger stellate, brownish greyish
hairs, large fascicled hairs on petiole and midrib
below few or absent, mature ones glabrous and
a little shining above, rather persistently greyish
tomentulose beneath, finely serrate-dentate in the
upper part, tips of the serratures usually callose,
5-8 by (2—)2.5-4 cm, midrib bold beneath, nerves
in 9-11(-12) pairs, curved, excurrent or divided
FLORA MALESIANA
[ser. I, voli, 74]
along or before the edge, raised beneath, reticula-
tion rather faint; petiole 0.5-1 cm by c. 2mm.
Racemes panicled, (8—)10-15 cm, covered with a
fulvous-greyish tomentum of stellate and fascicled
hairs; rachis stoutish. Pedicels robust, 2—3(—5)
mm, bracts subulate, caducous. Calyx lobes
ovate-oblong, 4-5 by 2(—2.5) mm. Petals obovate-
spathulate, minutely crenulate at apex, white,
glabrous, (4—)5(—6) by c. 2.5 mm. Filaments gla-
brous, 3mm; anthers obcordate, 1mm. Ovary
longish erect-tomentose; style thickish, glabrous,
(2.5-)3 mm (3-4mm in fruit), shortly 3-lobed.
Capsule depressed-globose, c. 3mm g. Seeds
oval, 1 mm.
Distr. Malesia: Philippines (Palawan), in low
woody vegetation on summit of Mt Pulgar near
Puerto Princesa, 1200 m. Fi. fr. May.
13. Clethra pachyphylla Merr. Philip. J. Sc. 13
(1918) Bot. 103; En. Born. (1921) 460; SLEUM.
Bot. Jahrb. 87 (1967) 99. — Fig. 1.
Shrub or mostly small tree, 2-4(-10) m high,
trunk up to 20cm g, laxly branched, branches
obliquely ascending to almost horizontal. Branch-
lets robust, rufous-ferrugineous-tomentose at apex.
Leaves oblong or obovate-oblong, more rarely
oblong-elliptic or elliptic, apex shortly subacutely
acuminate, base cuneate, coriaceous, firm, younger
ones subdensely floccose- or subvillous-stellate-
hairy above, all over beneath with a tomentum of
minor pale stellate hairs and a more coarse one
of major stellate and fascicled, initially rusty,
finally greyish hairs, midrib and nerves mainly
with major fascicled hairs, entire or more rarely
and but in part irregularly serrulate, 4~-9(-14)
by (2-)2.5-3.5(-4, -5) cm, midrib bold beneath,
nerves (9—)10-12(—13) pairs, prominent beneath,
curved and -+ excurrent at the edge, veins slightly
raised beneath, reticulation more distinct but in +
mature leaves; petiole 1.2-1.5(—-1.8) by 1.5—2 mm.
Racemes panicled, rather dense-flowered, robust,
all over villous or almost so by minor and major
stellate and coarse fascicled rusty hairs, 5-10(—12)
cm; rachis stout. Pedicels thickish, 2-3(-4) mm,
bracts subulate, 4-5(-8) mm, caducous. Calyx
lobes oblong-ovate, subacuminate, (3.5—)4—-5 mm.
Petals spathulate to spathulate-oblong, irregularly
erose or crenulate at apex, white to cream, scented,
glabrous, (3—)4(—-5) by 2.5-3 mm. Filaments gla-
brous, 2—2.5 mm; anthers subobovate, 0.8—1 mm.
Ovary short-tomentulose; style thickish, glabrous,
2.5-3 mm (3.5-4 mm in fruit), shortly 3-lobed.
Capsule depressed-globose, 3 mm @. Seeds sub-
trigonous, 1—1.2 mm.
Distr. Malesia: NE. Sarawak (Mt Murud
area) and North Borneo (Mt Kinabalu).
Ecol. Scattered in Quercus havilandii and Tris-
tania forest, still not too rare in mossy Lepto-
spermum-Dacrydium forest, also in secondary
vegetation, (870—)1200-2500(-3050) m, on poor
soil. Fl. fr. March—Oct.
OXALIDACEAE (J. F. Veldkamp, Leyden)
Herbs, sometimes with scaly rhizomes, bulbs, bulbils or stolons, or woody
perennials, shrubs, lianas or trees. Leaves penninerved, digitately or pinnately
trifoliolate, imparipinnate or paripinnate, basal, alternate, subopposite or apically
tufted. Stipules sometimes present. Petioles with basal joint, petiolules articulated.
Inflorescences basal, axillary or pseudoterminal, cymose to pseudumbellate, rarely
racemose, l1-many-flowered, bracteate and bracteolate. Flowers 3, very rarely also
$ specimens (Dapania), actinomorphic, 5-merous, hetero-tri-, -di-, or homostylous,
sometimes cleistogamous. Pedicels articulate. Sepa/s imbricate, free or connate at
base, sometimes with apical calli (Oxalis), persistent. Petals contort, quincuncial
or cochlear, free but usually cohesive above the base (“‘pseudosympetal’), clawed
(sometimes minutely so), glabrous or inside sometimes with minute papillae or
pilose. Filaments 10, obdiplostemonous, connate at base into an annulus, persistent,
the epipetalous (shorter) sometimes with a basal gland near the insertion of
the petals, or sometimes with 2 scales or dark lines on the annulus (Dapania),
rarely without anthers; the episepalous (longer) with a dorsal tooth (Oxalis) or
hunchbacked; anthers dorsifixed, versatile, 2-celled, dehiscing extrorsely by
longitudinal slits. No disk. Ovary 5-celled, superior; styles 5, terminal, persistent,
free, in LF' and MF erect, in SF patent to recurved, rarely reduced (3 flowers);
ovules 1—2-several per cell in 1-2 rows, epi- and anatropous, pendulous, super-
posed, bitegmic. Fruit capsular, loculicid, 5-celled, dry, rarely fleshy and indehis-
cent. Seeds usually with an aril; endosperm copious, fleshy, rarely absent; embryo
straight.
Distribution. 6(7?) genera with c. 850 spp. Of the Malesian representatives Oxalis, the largest genus,
is most numerous in S. America and S. Africa and Biophytum in S. America and Madagascar; Dapania
has 2 spp. in Malesia and | in Madagascar; Sarcotheca (11 spp.) is endemic in Malesia, while Averrhoa
(2 spp.) assumedly also originated here; it is now cultivated pantropically.
In Malesia there are 5 genera with 29 species, of which 14 endemic.
The generic distribution of the family offers in the Old World two remarkable patterns. First, that of
Dapania which shows the characteristic disjunction between West Malesia and Madagascar. Second,
that of Oxalis sect. Acetosellae which almost resembles that of Euphrasia, that is, a temperate genus with
two stepping stones across the tropics (Luzon, New Guinea), otherwise bipolar. Fig. 2.
Ecology. Many members of the family have sensitive leaves and show sleeping movements, some being
also seismonastic which is conspicuous in Biophytum.
Dispersal. The fleshy fruits of Averrhoa, Dapania, and Sarcotheca are no doubt eaten by various animals
as bats, birds, and monkeys, and dispersed by them. The seeds of Oxalis and Biophytum have a peculiar
ejaculative aril originally enveloping the entire seed which at maturity shoots them away for some distance
(ZieGLeR, Ber. Bayer. Bot. Ges. 36, 1963, 61). The cultivated species of Oxalis which set no fruit in
Malesia, are locally tenacious weeds through their bulbils. The native Oxalis species propagate also
vegetatively, by stolons and root-stocks.
All Oxalidaceae have an arillate seed except the species of Sarcotheca and Averrhoa bilimbi.
On germination some remarks are made under the genus Biophytum.
Morphology. Episeptal rimae are found in some species of Oxalis, in most of Dapania and all
Sarcothecas; in Averrhoa they are only inconspicuous, apical furrows. The most primitive state is probably
represented in Oxalis corniculata. The septs fail to enlarge towards fructification, whereby the cells of the
fruit are only united by their attachment to the central axis. The walls of the cells are pressed together
and slits are formed, especially conspicuous by transparent ridges. These ridges are wide apart in the
Malesian Dapanias and are slightly bent inwards. In Sarcotheca the septs are developed, at least in the
lower half; in the upper half the rimae may be open and papillose inside, or closed and glabrous. Episeptal
rimae are absent in all Biophytums and in Dapania pentandra from Madagascar.
(1) LF = long-styled form, MF = mid-styled form, SF = short-styled form.
(151)
152 FLORA MALESIANA [ser. Lwoli
Dehiscence of the fruits in Biophytum and Dapania is into a 5-rayed star. In Oxalis the valves remain
united and only longitudinal loculicid slits are formed. Sarcotheca and Averrhoa have indehiscent fleshy
fruits.
Anatomy. CHAUVEL, Rech. sur la famille Oxal. (1903) thesis; Mott & JANss. Mikr. 2 (1911) 9
(Averrhoa); HEtmscu, Lilloa 8 (1942) 97, 191; METCALFE & CHALK, Anat. Dicot. 1 (1950) 299.
Phytochemistry. Few phytochemical investigations were performed with members of the family.
In fact, distinct constituents have been isolated only from a few species of Oxalis. The tendency shown
by many species of Oxalis to accumulate large amounts of oxalic acid in water-soluble form is known
since a long time. Leucocyanidins and leucodelphinidins have been demonstrated to be present in the
leaves of some species of Oxalis and of Averrhoa carambola L. This agrees with the idea that Oxalidaceae
represent the most primitive family of Geraniales. Probably the species described by plant anatomists as
possessing ‘tannin’ cells are the ones which contain leucoanthocyanins in leaves. The bright yellow
flower pigments of Oxalis cernua THUNB. are the aurone glycosides aureusin and cernuoside. A yellow
quinonoid pigment was isolated from the bulbs of one species of Oxalis (O. purpurata Jacq. ?) and later
identified with rapanone, a benzoquinone occurring frequently in myrsinaceous plants. Too little phy-
tochemical information is available at present for a chemotaxonomical appreciation of systematic
relationships of Oxalidaceae. It might be significant, however, that rapanone does also occur in Con-
naraceae. General reference: HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 255. — R. HEGNAUER.
Pollination. Heterostyly is a common feature in the family, heterotristyly is observed in Oxalis,
Biophytum, and Averrhoa bilimbi. This must be assumed to be the primitive condition, as is found in the
allied Connaraceae. From it is derived a heterodistylous condition in Sarcotheca and Dapania, in
certain species of Oxalis and Biophytum, and in Averrhoa carambola. A further derived, homostylous
condition is found in certain species of Oxalis and Biophytum.
DARWIN concluded that the heterotristyly in a Ceylonese Biophytum which he examined, was also
functional in analogy with Oxalis while in cleistogamous flowers } he observed that in some way the
incompatibility factor seemed to be removed (The Different Forms of Flowers, ed. 1877, 181, 323).
SALTER (J. S. Afr. Bot. Suppl. 1, 1944) confirmed by experiments that functional heterotristyly occurs
in Oxalis. This was also found in S. American species of Oxalis. Miss P. MAYURA Devi (J. Genetics 59,
1964, 41) found in Indian Biophytum in her experiments a significant incompatibility in illegitimate
crossings, except for MF selfed with pollen of the long stamens. This means a loss of compulsory he-
terotristyly. Ina later article (ibid. 1966, 245) she described another mid-homostylous form which proved
excellently self-compatible. It is regrettable that she did not conserve voucher specimens bound to her
experiments, as the identification of Indian Biophytum is in distinct confusion and more than one species
is cited as B. sensitivum. The plants used by Devi certainly do not belong to B. sensitivum, as in her
pictures the corolla is much longer than the calyx.
Functional heterotristyly is apparently present in the introduced species of Oxalis, O. deppei and
O. latifolia, which in Malesia occur as SF only and have never been observed with fruit. O. corymbosa,
although present with SF and MF, does not fruit either, but the flowers are often monstruous in this
species. All three reproduce very successfully by bulbils and can become obnoxious weeds.
Biophytum fruticosum, B. adiantoides and B. microphyllum are also heterotristylous. No experiments
have been done.
Biophytum sensitivum and B. reinwardtii sens. str. are both in India and Malesia mid-homostylous.
Reduction to a single stylar form (LF) is found in Oxalis magellanica and O. acetosella ssp. griffithii;
reduction to homostyly (MF) has very far progressed in both O. corniculata where LF occurs rarely (6
out of 105 plants examined; cf. also E1rEN, Am. Mid]. Natur. 69, 1963, 280) and Averrhoa bilimbi,
where LF and SF were only observed once. The introduced Biophytum dendroides is apparently self-
compatible, as only one form (LF) has been found at Bogor and in the glasshouses of the Botanical
Gardens of Leyden and Groningen, where in the apparent absence of pollinators it is fully fertile and weedy.
Heterodistyly (LF and SF) occurs in Biophytum petersianum, Sarcotheca, and Dapania pentandra
from Madagascar. It is plausible that the 3 flowers of the Malesian Dapania originated by an extreme
reduction of the pistil in SF-flowers. Averrhoa carambola is heterodistylous for LF and MF. The latter
type might well correspond with SF, as the shorter stamens are much reduced and without anthers.
More information about heterostyly and literature concerning this matter is given by ORNDUFF and
by Mutcany (Am. J. Bot. 51, 1964, 307 and 1045).
Palynology. See ERDTMAN, Pollen Morphol. & Plant Tax. 1 (1952) 302; HUANG, Taiwania 13
(1967) 70 and Huyun, Bot. Jahrb. 89 (1969) 272.
Uses. Several species of Oxalis are cultivated as ornamentals, Averrhoa for its edible fruit. The wood
of the ligneous Oxalidaceae is useless as timber. In Malaya the fruits of Sarcotheca are sometimes eaten.
See also HEYNE, Nutt. Pl. (1927) 850.
Taxonomy. In Engler’s Syllabus (2, 1964, 248) ScHoLz divided the genera in two groups, A and B,
on the aestivation and the number of ovules per carpel. Through this Averrhoa was joined to the affinity
of Oxalis and Biophytum. However, the aestivation is inconstant. Moreover, the affinity of Averrhoa is
(1) In Biophytum 1 found also in not cleistogamous flowers opened anthers with good pollen already
in bud (protrandry).
1971] OXALIDACEAE (Veldkamp) 153
doubtless with Sarcotheca and Dapania in all other characters. These three genera form a very clear
reticulate affinity.
The family is considered to be the most primitive of the Geraniales by HALLER f. (Arch. Néerl. sér.
3B, 1, 1912, 109) and ENGLER (Nat. Pfl. Fam. ed. 2, 19a, 1931, 10). This primitive position is possibly the
reason for the affinity with the Connaraceae, at which some authors have pointed (HALLIER /. New
Phyt. 4, 1905, 158; Arch. Néerl. sér. 3B, 1, 1912, 109; SCHELLENBERG, Pfl. R. Heft 103, 1938, 127;
TAKHTAJAN, Evol. Angiosp. 1959, 236). The latter family is usually placed near the Leguminosae. As a
matter of fact specimens of Rourea and Sarcotheca were often confused and are very similar, the pistil and
fruit excepted.
The closest allied family is Geraniaceae in which they were merged by BENTHAM & Hooker /. In fact
the American Hypseocharis seems to link both families. The only constant character was said by HALLIER
f. (Beih. Bot. Centralbl. 39, ii, 1921, 172) to be the tenuinucellate ovules in Oxalidaceae and crassinucel-
late ovules in Geraniaceae, but Hypseocharis and many species of other genera have not been checked on
the general validity of this character, as HALLIER f. himself admitted. And one must be very careful in
this respect; for example HUTCHINSON (Fam. FI. Pl. ed. 2, 1, 1959, 494) stated that Oxalidaceae have
albuminous seeds, but some species of Oxalis are exalbuminous (cf. SALTER, J. S. Afr. Bot. Suppl. 1,
1944, 26); he stated also that Oxalidaceae are exstipulate, but stipules are found in all Biophytums and
also in some species of Oxalis (sect. Acetosella, O. corniculata, etc.).
HUTCHINSON’s division of Oxalidaceae into three different families belonging to three different orders
seems not to have a reliable basis as Averrhoa cannot be divorced from Sarcotheca and Dapania and
there is no reason for admitting a preponderant importance to their ligneous habit; besides many species
of Biophytum and Oxalis are dwarf shrubs.
KEY TO THE GENERA
1. Herbs or dwarf shrubs, rarely up to 114 m high. Leaves 3(—4)-foliolate, or paripinnate, herbaceous.
Fruit a dry capsule.
2. Leaflets 3-4. Capsule with the valves remaining attached to the central axis. . . . . . 1. Oxalis
2. Leaflets 6 or more, pinnate. Capsule dehiscing into a 5-rayed star, without leaving a columella.
2. Biophytum
1. Shrubs, trees or lianas, much higher. Leaflets 1 or 3, or leaves imparipinnate, 2-many-jugate. Fruit
fleshy, dehiscent into a 5-rayed star or indehiscent.
3. Leafiets 1 or 3, chartaceous or subcoriaceous. Lateral petiolules articulate, after dropping leaving a
short stalk on the rachis. Ovules 1-2 per cell.
4. Lianas. Inflorescences racemose. Petals inside glabrous. Fruit dehiscent into a 5-rayed star. Seeds
emer ee ils. bs, 2 egoCa ee, (Abs “eeoeel AO LN ee eee 3. Dapania
4. Shrubs or trees. Inflorescences paniculate. Petals minutely papillose ‘inside. Fruit indehiscent
(but episeptal rimae sometimes open!). Seeds exarillate. . . . . . 4. Sarcotheca
3. Leaves imparipinnate, 2-many-jugate, herbaceous to papyraceous. " Lateral petiolules after dropping
net leaving a stalk on the rachis. Ovules 3-7 per cell... 3) sca) Sa. as 5. Averrhoa
1. OXALIS
Linn, Gen. Pl. ed. 5 (1754) 198; Sp. Pl. (1753) 433; Knuth, Pfl. R. Heft 95 (1930).
— Fig. 1.
Annual or perennial herbs (or dwarf shrubs, extra-Mal.), some stemless with
rhizomes or bulbs. Leaves digitately or pinnately 3(—4)-foliolate; leaflets in Mal.
spp. (except O. barrelieri) +- sessile. Stipules when present adnate with the base
of the petiole. Peduncles basal from bulbs or solitary and axillary. /nflorescences
cymose to umbellate, 1-many-flowered. Bracteoles 2-several, sometimes with
apical calli. Pedicels articulate at base and sometimes beneath the calyx. Sepals
shortly connate at base, with or without apical calli. Petals coherent above the
claw, contort, glabrous. Filaments: longer ones sometimes with a dorsal tooth.
Stigma cylindric and minutely bilobed to peltate, sometimes papillose. Ovules |
to c. 10, in 1-2 rows per cell. Capsules loculicid by longitudinal slits, sometimes
with episeptal rimae. Seeds 1-c. 10 per cell, usually few; aril bivalved, ejaculatory;
testa smooth, or with transverse ridges or longitudinal furrows.
[ser. I, vol. 73
$33
Fig. 1. Oxalis magellanica Forst. f. a. Habit, x 1, b. ‘callus’ of leaf, x 12, c. apex sepal, 12, d. bracteoles,
from both sides, x 4. — O. corymbosa DC. e. Leaf, «1. — O. barrelieri L. f. Leaf, nat. size, g. dentate
stamen, X12. — O. acetosella L. ssp. griffithii HARA. h. Leaf, nat. size, i. bracteoles, from both sides,
<4 (a-d BALGooy 252, e BACKER 37150, f—g BOERLAGE s.n. a. 1880, h—-i STEINER 1990).
Distr. Cosmopolitan, at least 700 spp., mainly from S. America and the Cape, in Malesia 3 native
spp.; others introduced and escaping, sometimes becoming weedy. Fig. 2 (§ Acetosellae).
Ecol. O. acetosella ssp. griffithii and O. magellanica are (in Malesia) both characteristic mountain
plants, above 2200 m. O. corniculata is rather indifferent to altitude.
Morph. The longer filaments are frequently thickened in the basal part, as in Biophytum. The apex
of the thickening may emerge as a small tooth, e.g. in O. latifolia, O. deppei, and O. barrelieri. This
tooth does not carry a vascular trace (NARAYANA, J. Jap. Bot. 41, 1966, 321).
For heterostyly see under the family.
An important study of the morphological diversity and taxonomy of the S. African species is provided
by SALTER (J. S. Afr. Bot. Suppl. 1, 1944).
Notes. Pending a revision of the S. American species, Miss LoURTEIG (Paris) advised me to accept the
names of the introduced species in the current sense.
Sizes of petiole and pedicels are given only for the portion above their articulation, as the part below
it is too variable. The length of a 2-lobed leaflet is measured from the base to the apex of the lobe. The
length of the filaments includes that of the basal, fused, annular part.
1971]
OXALIDACEAE (Veldkamp)
155
KEY TO THE SPECIES
1. Leaves along a distinct, supraterranean, creeping to erect stem. No bulbs.
2. Shrublet. Petioles phyllodial, leaflets 0-3, often minute. Peduncles abortive. Flowers 1-9 in a fascicle,
yellow. Cultivated in Java (§ Heterophyllum) (O. rusciformis MIKAN) . .
O. fruticosa RADDI
2. Herbaceous. Petioles not phyllodial, leaflets not reduced. Peduncle distinct.
3. Leaf pinnately 3-foliolate (rachis developed under terminal leaflet). Leaflets elliptic to oblong,
apex not notched. Petals pink with yellow base (§ Thamnoxys). .
3. Leaf digitately 3-foliolate. Leaflets obcordate. Petals yellow (§ Corniculatae).
1. O. barrelieri
2. O. corniculata
1. Stemless herbs with bulbs or subterranean rhizomes; leaves all basal.
4. Rhizome, no bulbs. Inflorescence 1-flowered. Sepals with 0-1 greenish, round apical callus. Petals
white (§ Acetosellae).
5. Stipules conspicuous, much wider than the petiole. Leaflets up to 144 by 144 cm, obcordate;
beneath whitish and glaucous and with a pale callus in the notch. Bracteoles 2, free, not vaginate,
subglabrous. Sepals pubescent, with 1 pale apical callus. . : 3. O. magellanica
. Stipules narrow, slightly broader than petiole, not very conspicuous. Leaflets 1-3 by 14-4 cm,
fishtail-shaped; beneath not whitish and glaucous, ecallose. Bracteole 1, emarginate to bifid, va-
ginate, with an apical brown hair-tuft. Sepals subglabrous, ecallose. 4. ©. acetosella ssp. griffithii
4. Bulbs, no rhizome. Inflorescence 2—many-flowered. Sepals with 2-4 orange apical calli. Petals red
or purple with greenish to yellowish base.
6. Outer tunics many-nerved, fibrous-withering. Leaflets 4(-6), obdeltoid, not notched, not punctate,
at apex sometimes with 2 minute calli. Umbel with many bracts. Filaments crispy-ciliate, the
longer with a dorsal! tooth. Sepals with 4 linear calli. Petals red (bluish when dry) (§ Polyoxalis).
5. O. deppei
6. Outer tunics 3-several-nerved, not fibrous. Leaflets 3, obcordate or fishtail-shaped. Sepals with 2
oblong apical calli. Petals purple (bluish when dry) (§ Jonoxalis).
7. Stolons apically bulbiferous. Outer tunics membranaceous, transparent, several-nerved. Leaflets
fishtail-shaped, not punctate, notch often with 2 minute calli. Bracts of umbellate inflorescence 2,
opposite. All filaments moderately ciliate, the longer with a minute tooth. 6. O. latifolia
. Stolons absent, bulb consisting of many small bulbils; outer tunics papyraceous, brownish,
3-nerved. Leaflets broadly obcordate, minutely punctate all over the surface, especially along the
margins, ecallose. Bracts of the cymose, contracted inflorescence many. Shorter filaments glabrous,
the longer ciliate, without atooth......
1. Oxalis barrelieri LinNE, Sp. Pl. ed. 2 (1763)
624; Rip. Fl. Mal. Pen. 1 (1922) 330; KNuTH,
Pfl. R. Heft 95 (1930) 65; HENDERSON, Mal.
Wild FI. (1959) 47, fig.; BACKER & BAKH. f.
Fl. Java 1 (1963) 245. — O. sepium St. Hi. FI.
Bras. Merid. 1 (1825) 89; ProGEL, FI. Bras. 12, 2
(1877) 505, incl. var. picta PROGEL; KNuTH, Pf.
R. Heft 95 (1930) 64; Herne, Nutt. Pl. (1927)
151. — Fig. 1f-g.
Erect herb, stem up to 1!4 m, without bulbs or
stolons. Stem branched, sparsely patent to ap-
pressed-reflexed pubescent, glabrescent; hairs
simple, straight or bent below the middle, eglan-
dular. Leaves usually more or less opposite,
pinnately 3-foliolate, exstipulate; petiole 2-9 cm,
appressed-puberulous; leaflets elliptic to oblong,
base cuneate to emarginate, apex obtuse to
rounded, not notched, ecallose, glabrous above,
margins strigose, especially at base, beneath pale
and glaucous, sparsely to moderately appressed
pubescent, terminal leaflets largest, 1-34 by
Y4-2' cm. Peduncles 3-5\4 cm, pubescent, once
or twice dichasially forked; branches up to 4144 cm,
4-16-flowered, the bracts opposite the flowers.
Bracteoles minute, ciliate, ecallose. Pedicels 114-3
mm, glabrous. Sepals 2-4 by 14-144 mm, ovate-
Janceolate, acute, glabrous or with a few hairs,
ecallose, 3-nerved. Petals 6-9 by 2-214 mm,
obovate-lanceolate, apex rounded, after anthesis
rolling inwards, pink, lower half greenish with
yellow spots, glabrous. Filaments (MF) %4—1 and
7. O. corymbosa
ered ° elm teery €) berie) thre (s) jie ts
134-214 mm, the shorter glabrous, the longer with
a dorsal tooth, patent-hairy. Ovary 1 by 4 mm,
glabrous; styles (MF) 1-114 mm, ascendingly stri-
gose; stigmas small, capitate; ovules 4 per cell,
in 1 row. Fruit 5-10 by 3-5 mm, slightly ovoid,
5-angular, apex and base 5-lobed, glabrous;
episeptal rimae present. Seeds 3-4 per cell, 144 by
lmm, + flattened-ovoid; testa transversally
ridged.
Distr. Native of tropical S. America, cul-
tivated and established in many places, the oldest
collected specimen from Bogor dates from
BOERLAGE, a. 1888; in Malesia: Sumatra, Banka,
Malaya, Java, Papua (Central Distr.).
Ecol. Around gardens, along roads, in hedges,
fields, and village groves, along rivers, grassy
places with shade, up to 1500 m.
Vern. Tjalingtjing, J, bélimbing tanah, Banka,
kopomani, Tamil.
Uses. The leaves are eaten for their sour taste.
Note. Although only the MF form is found in
Malesia, fertile seed is formed; there is apparently
no functional heterostyly.
2. Oxalis corniculata Linné, Sp. PI. (1753) 435;
Zucc. Abh. Ak. Wiss. Miinch. 1 (1830) 230,
incl. var. repens (THUNB.) Zucc.; PLANCH. in
Houtte, Fl. Serres 12 (1857) 205, incl. var. atro-
purpurea PLANCH.; Mia. FI. Ind. Bat. 1, 2 (1859)
135; Epcew. & Hook. f. Fl. Br. Ind. 1 (1874)
436; F.-VILL. Novis. App. (1880) 32; GUILLAUMIN,
FLORA MALESIANA
[ser.. I. voix
(1911) 170, incl. var. javanica (BL.) BACK.; KNUTH,
Notizbl. Berl.-Dahl. 7 (1911) 300, incl. var. sericea
KNuTH: Rip. FI. Mal. Pen. 1 (1922) 330, incl.
var. villosa Hoox. f. ex RID. [sphalm. ? var.
villosa HOHENACKER, cf. YOUNG, Watsonia 4
(1958) 57]; WiEGAND, Rhodora 27 (1925) 113;
KNuTH, Pfl. R. Heft 95 (1930) 146; MASAMUNE,
Fl. Geogr. Stud. Yakushima (1934) 257, incl.
ssp. repens (THUNB.) MASAM.; KNUTH in Fedde,
Rep. 48 (1940) 3, incl. var. papuana KNUTH;
HarA, Enum. Sperm. Jap. 3 (1954) 8; EITEN,
Taxon 4 (1955) 99; Am. Mid]. Natur 69 (1963)
257; BACK. & BAKH. f. Fl. Java 1 (1963) 246;
VELDK., Fl. Thail. 2 (1970) 17. — O. repens
THuNB. Diss. Oxal. (1781) 16, fig.; Bx. Bijdr.
(1825) 243; B. L. Roprnson, J. Bot. 44 (1906) 311;
Me_rr. En. Born. (1921) 311; En. Philip. 2 (1923)
323. — O. javanica BL. Bijdr. (1825) 243; Mia.
Fl. Ind. Bat. 1, 2 (1859) 135. — O. acetosella
(non L.) BLANco, FI. Filip. (1837) 388. — O.
boridiensis KNUTH in Fedde, Rep. 48 (1940) 3.
Perennial herb, ascending to erect, rooting at the
nodes; main root sometimes much thickened and
woody; stems several from the main root, branch-
ing above the ground, puberulous to pubescent
(hairs mainly 1-celled). Leaves scattered, distant,
or in small tufts. Stipules indistinct to small, some-
times conspicuous, up to 3 by | mm, rectangular.
Petiole 1-514(—10) cm, appressed to patently pu-
berulous. Leaflets broadly to elliptic-obcordate,
4-20(-25) by 5-18(—25) mm, incised up to half-
way, ecallose, lobes rounded, rarely obtuse; upper
surface glabrous to sericeous; beneath paler,
sometimes glaucous, sparsely pubescent to seri-
ceous. Peduncles up to 20cm, usually much
shorter, sparsely puberulous to sericeous. Inflores-
cence cymose to pseudo-umbellate, 1-—S(-8)-
flowered. Flowers usually MF, rarely LF. Bracts
2-several, subopposite to whorled, ovate-lanceo-
late, acute, puberulous, sometimes with septate
hairs. Pedicels up to 20mm, articulate at base
and beneath calyx, in fruit straight to sharply
bent at the articulations, but the fruit always
erect. Sepals lanceolate, obtuse to rarely obliquely
retuse with narrow pale margins, 2-6 by 14-2 mm,
sparsely puberulous to sericeous, sometimes with
septate hairs. Petals spathulate-oblong to -lanceo-
late, 314-10 by 1-7 mm, apex rounded to emarginate,
after anthesis apically crumpled, yellow, with
darker or lighter base. Filaments glabrous, the
longer edentate, in MF 1-4 and 3-6 mm, the
shorter rarely with abortive anthers, in LF
234-314 and 314-414 mm. Ovary 1144-2 by %-1
mm, ellipsoid to cylindric, puberulous; styles in
MF 1-4 mm, in LF 3-4 mm, minutely ciliate,
sometimes mixed with minute septate hairs;
stigmas small, cylindric, sometimes flattened and
minutely bifid, papillose; ovules (1—)5—11 per cell,
in 1 row. Fruit 9-20(-24) by 2-4mm, usually
linear-cylindric, sometimes ellipsoid, pentagonal,
acuminate, minutely puberulous, hairs reflexed or
patent to ascending in upper half, mixed with
patent, septate hairs; episeptal rimae closed, in-
conspicuous; cells inside sparsely to moderately
strigose. Seeds (O—)5—11 per cell, 1 by 34 mm,
flattened-ovoid; testa with c. 3 regular rows of
7-10 transversally connected rows of ridges.
Distr. Cosmopolitan, origin unknown, in Ma-
lesia several forms occur; throughout Malesia,
three times collected in Celebes and Borneo
and scarce in Malaya (Malacca, Perak, Penang;
a common weed in Singapore Bot. Gardens).
Ecol. In many islands a common weed on all
sorts of disturbed soil, in grassfields, gardens
estates, along roads and river-banks, on walls,
etc., in Java up to 2200, in New Guinea to 3000 m.
Vern. Sikap dada, M_ (Sing.), asim-asim,
atim-atim, lela, Atjeh, daun asém (ketjil), (djukut)
tjalintjing, sémanggi, sémanggén, J, tjembitjéna,
Mad., mala mala, Ternate, kuja kawiana, Alor;
Philip.: daraisig; Bik., iayo, kungi, malabalugbug-
dagis, Pamp., kanapa, \g., marasiksik, Uk., pikhik,
Iv., salamdgi, Bon., susokoyili, taingangdaga, Tag.;
New Guinea: keketi, Tumba, puggepagl, Yoowi,
songongom, Wapi, girobi, Musa, Safia, jampijamp,
Enga, Yogos, gagari, Eng, Kapilam, kAwibant,
Maring, kale, Rabaul, tenaquipo, Jimi.
Uses. Cf. HEYNE, Nutt. Pl. (1927) 850. Ac-
cording to VINK (n. 16308) from Uinba, Nona-
Minj Divide, Kubor Range, W. Highlands, New
Guinea (20-8-1963) used 1) ‘“‘In marrying cere-
mony the young woman takes fresh leaves and
makes a gag of it with salt and cold water; the gag
is chewed and the juice is spit on pigmeat, which
is given to the new husband. 2) When the garden
gives a bad production of sweet potato, the woman
looking after the garden puts a bundle of the leaves
in her girdle to get a higher production.”
Note. A most complex and variable species.
Many infraspecific taxa have been described, but
all appear to be linked by intermediate forms. The
following extremes can be recognized;
‘var. repens THUNB.’ Small, decumbent plants.
Leaflets rather dark, small, glabrous to moderately
strigose. Inflorescences few-flowered, flowers
small. Throughout Malesia, a form of exposed,
sunny places.
‘yar. atropurpurea PLANCH.’ Plant brownish to
purplish in vivo, dark green when dry; petals more
or less flushed or blotched with reddish brown in
vivo, fading to whitish or yellowish in drying.
Cultivated and escaping.
‘yar. sericea KNUTH’ (= ‘var. trichocaulon
LEVL.’ according to HARA, 1954; no material of this
seen). Large pubescent plant, foliose apices of
stems erect or ascending; stipules minute, leaflets
strongly pubescent, terminals large, longer than
broad, incised for 1/,—1/, of leaflength, floral parts
relatively large; ovules and seeds 1-5 per cell;
fruit oblong, stout, pubescence ascending at least
in upper half. Eastern New Guinea and New
Britain (Formosa? Japan? Korea?). Shaded river-
banks, open places in forest, edges of paths and
trails, grassfields.
3. Oxalis magellanica Forst. f. Comm. Gott. 9
(1789) 33; Hook. f. Fl. Nov. Zel. 1 (1853) 42,
fig.; Fl. Tasm. 1 (1860) 59; BenTH. Fl. Austr. 1
(1863) 300; Hook. f. Handb. N. Zeal. Fl. (1864)
1971]
OXALIDACEAE (Veldkamp)
38; Reicue, Fl. Chile 1 (1896) 339; KNutTH, Pfi.
R. Heft 95 (1930) 230. — O. lactea Hook. Comp.
Bot. Mag. 1 (1836) 276; Skottss. The Plant World
18 (1915) 129; CHEESEMAN, Man. N. Zeal. Fl. ed. 2
(1925) 536; Curtis, Stud. Fl. Tasm.1 (1956) 96;
ALLAN, FI. N. Zeal. 1 (1961) 238. — Fig. la—d.
Stemless herb with stolons, without bulbs.
Rhizome pink to brownish, glabrous with distinct,
amplexicaul scale-like remains of leafbases.
Stipules much broader than the petiole, con-
spicuous, membranaceous, brown, glabrous. Pet-
ioles 1-814 cm, reddish. Leaflets obcordate,
4-13 by 4-12 mn, incised up to 4 of the length,
lobes rounded; upper surface usually glabrous;
beneath pale, glaucous, appressed-strigose gla-
brescent, with a + prominent, greenish to brown-
ish callus at the notch. Peduncles few, below basal
articulation 14-2 cm, above 1-614 cm, sparsely ap-
pressed- to patently pubescent; bracteoles 2, sub-
opposite, 214-7 by 1-114 mm, lanceolate, acute,
free, very sparsely strigose to glabrous, ecallose,
placed in upper */;—4/,th. Buds erect. Flowers
solitary, only LF. Sepals elliptic to obovate-
oblong, 3—5 by 114-214 mm, acute to truncate and
minutely 3-lobed (apex or middle lobe cucullate,
sometimes inconspicuous by the hairs, seemingly
callose), sparsely to moderately pubescent and
somewhat glaucous, margined, slightly shorter to
equal to the fruit. Peta/s white, 8-1114 by 4-6 mm,
spathulate, somewhat oblique, apex rounded to
emarginate, glabrous to + ciliate. Filaments (LF)
3-444 and 414-5'4 mm long, glabrous, edentate.
Ovary 1-2 by 1-2 mm, glabrous; styles 3-S mm,
157
glabrous; stigma disk-shaped; ovules 2-S per cell,
in 1 row. Fruit 3-6 by 3-5 mm, glabrous, finally
nodding. Seeds 1-3 per cell, 144 by 1mm, +
flattened-ovoid, shiny, brownish, smooth to
slightly lengthwise furrowed.
Distr. Southern Andine S. America, New
Zealand, Tasmania, and the Victorian Alps; in
Malesia: New Guinea, on the high mountains
(Wilhelm, Otto, Kubor Range, Finisterre Range,
Sarawaket Range, Wilhelmina), 2200-3700 m.
Ecol. Subalpine to alpine, on shaded, moist to
wet, humous soil, among grasses and mosses,
sometimes on tree-trunks or rocks. F/. May—Nov.
Vern. Chimbaemagl, Chimbu.
Notes. According to SKOTTSBERG /.c. the S.
American specimens would differ in having less
distinctly obcordate leaflets, peduncles not ex-
ceeding 1 cm, and flowers only c. 5mm across.
The New Zealand botanists accordingly call the
species in New Zealand O. lactea, but ALLAN
added that some specimens from Chile (in K)
approach the New Zealand form.
A comparative study of Chilean, Australasian,
and Papuan specimens showed that SKOTTSBERG’S
differentiation does not hold and that only one
species is concerned.
4. Oxalis acetosella LINNE, Sp. Pl. (1753) 433.
ssp. griffithii (EpGEw. & Hook. f.) Hara, J.
Jap. Bot. 30 (1955) 22; Fl. E. Him. (1966) 168,
638, 661. — O. griffithii EpGew. & Hook. f.
Fl. Br. Ind. 1 (1874) 436; KNuTu, Pfl. R. Heft 95
Fig. 2. Distribution of Oxalis § Acetosellae. 1a. O. acetosella L. ssp. acetosella, \b. ditto, ssp. griffithii
Hara, Ic. ditto, ssp. montana HULTEN, 2. O. obtriangulata MAxiM., 3. O. magellanica Forsv. /., 4. O.
oregana NUTT.
158
FLORA MALESIANA
[ser. I, vol. 72
(1930) 234; Ouwt, FI. Japan (1965) 580. — Fig.
1h-i.
Stemless herb with stolons, without bulbs.
Rhizome dark brown, usually scaley, sparsely
pubescent to glabrous. Stipules shorter and +-
broader than the basal part of the petiole. Petioles
414-22 cm, greenish, red-tinged, pubescent; below
articulation flattened, thickened, persistent, final-
ly + woody, brownish when dry, moderately
to densely pubescent. Leaflets 1-3 by 144-4 cm,
broadly obtriangular, fishtail-shaped, the straight
upper edges forming a very wide angle, incised up
to halfway, lobes rounded to obtuse; upper sur-
face subglabrous to sparsely appressed-strigose;
beneath paler, not glaucous and with denser pu-
bescence, midrib + thickened in notch. Peduncles
1-2, below basal articulation up to 14 cm, above
5-11 cm (shorter in cleistogamic flowers), sub-
glabrous to densely pubescent, especially distally;
bracteoles -- halfway or higher, partly connate-
vaginate, midrib dorsally pubescent, apex with
hairy brown tuft, ecallose. Sepals 3-634 by 114-24
mm, oblong, rounded to emarginate, not cucullate,
ecallose, subglabrous, edge closely ciliate. Petals
8-19 by 4-8 mm, spathulate-oblong, truncate to
emarginate, often oblique, white, often with
lavender veins and yellowish base in vivo. Filaments
(LF) 134-314 and 3-6mm, glabrous, edentate.
Ovary 2-314 by 144mm, glabrous; styles (LF)
3-6 mm, glabrous; stigma minute, more or less
hook-like, entire; ovules 1—S per cell. Fruit 6 by 5
mm, ovoid, acute, glabrous, nodding. Seeds 1-2
per cell, 214 by 114 mm, flattened-ovoid, + smooth
to lengthwise ridged, light brown.
Distr. From India (Sikkim, Bhutan, Khasia)
through China to Japan and Formosa; in Ma-
lesia: Philippines (N. Luzon: Mt Pulog), two col-
lections (STEINER 1990, March 1961; Jacoss 7361,
Jan. 1968).
Ecol. Mossy forest ravine, probably 2500-
2900 m. Fil. Jan-March. Fig. 2.
HarA (1966) stated that it is found more to the
south and at lower altitudes than ssp. acetosella,
both in E. Himalaya and in Japan.
Notes. FERNANDEZ-VILLAR (Novis. App. 1880,
32) recorded true O. acetosella L. from a medical
garden at Manila.
5. Oxalis deppei Lopp. Bot. Cab. 15 (1828) 1500;
Knut, Pfl. R. Heft 95 (1930) 288; Back. &
Baku. f. Fl. Java 1 (1963) 246; MATTHEW,
Rec. Bot. Surv. India 20 (1969) 56. — O. tetra-
phylla (non Cav.) BAcK. & S Loor. Handb.
Thee (1924) 157, t.; HEYNE, Nutt. Pl. (1927) 851.
Stemless herb from bulbous base, no rhizome,
no stolons. Bulb up to 214 by 2 cm, ovoid, acute,
with lateral, sessile bulbils; tunics up to2% by 114
cm, ovate, acute, acuminate or bilobed with
terminal leaf, brown, many-nerved, outer fibrous,
margins bearded, rest glabrous, inner becoming
fleshy. Innovations brownish pilose. Petioles up to
40cm, glabrous or with a few hairs. Leaflets
4(-6), unequal, 214-614 by 134-614 cm, obdeltoid,
entire to slightly retuse and + apiculate (tip
often -++ folded and occasionally with 1—2 minute
calli underneath), glabrous to sparsely pubescent,
usually with dark V-shaped marking in lower third,
epunctate. Peduncles up to 50cm, + glabrous.
Inflorescence pseudo-umbellate, 6-—25-flowered.
Bracts many, up to 214 mm, ovate, acute to acu-
minate, glabrous or with strigose margin, ecallose
or with minute apical calli. Pedicels up to 3 cm,
glabrous. Sepals 5—7(-10) by 114-3 mm, elliptic
to lanceolate, obtuse to emarginate, glabrous,
ciliate on edge, c. 7-nerved, apical calli 2-6, usually
4, linear, inconspicuous, orange. Petals 114-214
by 44-14% cm, spathulate-oblong, rounded to
retuse, often oblique, glabrous, dark red, when
dry often bluish with greenish base. Filaments
(SF) 314-414 and 6-614 mm, (LF) 6 and 10 mm,
the shorter with a few, the longer with many
patent, crisped cilia, dentate. Ovary and staminal
tube c. 4% mm stipitate. Ovary 114-5 by 1-3 mm,
ellipsoid, glabrous; styles (SF) 1 mm long, (LF)
c. 15mm, glabrous, dark when dry; stigmas cy-
lindric, + bilobed, later + peltate, not papillose;
ovules 3—5 per cell, in 1—2 rows.
Distr. Native of Mexico, cultivated and
escaping; in Malesia hitherto only found in Java
(Tjinjiruan); introduced before 1911, now quasi-
spontaneous.
Ecol. Locally naturalized in mountain Cin-
chona estates, c. 1600m, sometimes gregarious,
propagating by bulbils; fruit unknown from Java.
Apparently functionally heterostylous. in Malesia
only the SF has been found so far. Dimensions of
the LF have been taken from MATTHEW, I.c.
Vern. Tjalingtjing badak, tj. gedé, S.
Uses. Ornamental plant and for ground-cover,
but difficult to eradicate; leaves eaten as vegetable
(HEYNE, 1927).
Note. O. tetraphylla Cay., with which this
species has often been confused, has bulbiferous
stolons, emarginate, not apiculate leaflets, and
smaller flowers.
6. Oxalis latifolia H. B. K. Nov. Gen. Sp. 5 (1821)
184, t. 467; KNuTH, Pfl. R. Heft 95 (1930) 273;
HEyYNE, Nutt. Pl. (1927) 851; Symon, Trans. R.
Soc. S. Austr. 84 (1961) 75; YOUNG, Watsonia 4
(1958) 63. — O. intermedia (non A. RICH. ?)
Back. & BAKH. f. FI. Java 1 (1963) 246.
Stemless herb from bulbous base, no rootstock;
bulb ovoid, up to 5 by 2 cm, acute, with numerous,
basal, + erect stolons, with a few small scales,
ending in ovoid, acute bulbils pale brown when
dry; tunics many, outer up to 5 by 2 cm, ovate,
acuminate or with terminal leaf, membranous,
transparent, white, nerves 3 to several, orange;
inner becoming fleshy. Petiole up to 20cm, +
glabrous. Leaflets + equal, 114-714 by 2-814 cm,
broadly obdeltoid, fishtail-shaped, incised up to
halfway, glabrous, beneath subglaucous, often
with 2 orange calli in notch, epunctate. Peduncles
up to 25 cm, slightly hairy. Inflorescence umbellate,
5—13-flowered. Bracts 2, c. 14 mm, ovate, margin
glabrous to strigose, minutely callose or ecallose.
Pedicels up to 2 cm, glabrous, filiform. Sepals
4-414(-6) by 114-2(-3) mm, oblong, obtuse,
glabrous, indistinctly 3—5-nerved; apical calli
1971]
OXALIDACEAE (Veldkamp)
159
2, orange, not confluent, c. 1 mm long, minutely
hastate. Petals 10-20 by 3-6(—8) mm, narrow-
ly obtriangular, truncate, + oblique, red-purplish
with greenish base, crumpled after anthesis.
Filaments (SF) 2'44-3(4) and 4-5(-6) mn,
puberulous, the longer with a minute tooth.
Ovary and staminal tube c. 144 mm stipitate.
Ovary 114 by *% mm, apically sparsely ciliate
on the ribs; styles (SF) c. 1 mm, sparsely
ciliate; stigma peltate, c. 4 mm 9g, not papillose;
ovules 3-6 per cell, in 1-2 rows.
Distr. Native of Central and tropical S. Ameri-
ca; cultivated and escaping, e.g. in Malesia:
Java (Preanger Mts; W. Java: Gedeh, Lembang,
Tjinjiruan).
Ecol. In gardens, fields, and estates, 1250-1550
m. No fruits are recorded from Java, the plant is
apparently functionally heterostylous. It is difficult
to eradicate because of the many bulbils.
Vern. Tjalingtjing, S.
Note. O. intermedia A. Ricn. (Ess. Fl. Cuba,
1842, 315) is said to be more pubescent and to
have edentate, longer filaments. As the teeth are
minute, however, they might easily be overlooked.
7. Oxalis corymbosa DC. Prod. 1 (1824) 696;
Rpt. Fl. Mal. Pen. 1 (1922) 330; Back. &
Stoot. Handb. Thee (1924) 155, t.; HEYNE,
Nutt. Pl. (1927) 851; Symon, Trans. R. Soc. S.
Austr. 84 (1961) 74; BAck. & BAKH. f. Fl. Java 1
(1963) 246. — O. martiana Zucc. Denkschr. K.
Ak. Wiss. Miinch. 9 (1824) 144; Back. Schoolfl.
Java (1911) 170; Merr. En. Philip. 2 (1923) 323;
Knuth, Pfi. R. Heft 95 (1930) 250. — O. violacea
(non L.) Har. f. Med. Rijksherb. Leiden 12
(1912) 19. — Fig. le.
Stemless herb from bulbous base, no rhizome,
no stolons; bulb globose, c. 1 cm 9; bulbils many,
clustered, globular to ovoid, acute; outer tunics
brown, papyraceous, up to 20 by 6 mm, ovate to
oblong, acuminate or with terminal leaf, margins
glabrous to long-bearded, distinctly 3-nerved;
inner pale and fleshy. Petiole up to 30 cm, patently
villose. Leaflets + equal, 114-414 by 134-514 cm,
broadly obcordate, incised for +/,;, lobes rounded,
sometimes + overlapping; upper surface sub-
glabrous, beneath appressed-puberulous; minutely
orange-punctate all over the surface, especially
along the margins. Peduncle up to 35 cm, hairy in
various degree. Inflorescence often up to twice
forked; pseudo-umbels 2-12-flowered. Bracts
many, c. 1 mm long, elliptic, rounded, pale, with
strigose margin and 0-3 orange, linear calli in the
middle. Pedicels up to 214 cm, ascendingly ap-
pressed-strigose. Sepals 314,-S5(-6) by 1-2 mm,
oblong, acute, apex minutely bifid to the 1-3 or-
bicular to oblong, with 2 orange, apical calli, +
puberulous, 3—5-nerved. Petals 11-20 by 4-7 mm,
spathulate-oblong to -lanceolate, obtuse to trun-
cate, often oblique, light reddish purple with
darker veins, yellowish at base. Filaments (MF)
2-3 and 5-6 mm, (SF) 314-4 and 5-6 mm; the
longer dorsally ciliate, edentate. Ovary and
staminal tube c. 34 mm stipitate. Ovary 2 by 1 mm,
abundantly ascendingly ciliate to glabrous; styles
in MF 134-2 mm, in SF 1-114 mm long, abundant-
ly ciliate; stigma bilobed, papillose, c. 1 mm g;
ovules 3-8 per cell, in 1—2 rows. Fruit not seen.
Distr. Native in tropical S. America, natural-
ized in many parts of the World; in Malesia cul-
tivated and escaping in Java (introduced from
Sydney before 1848), W. Sumatra, Malaya, and
Philippines (Luzon).
Ecol. In fields, road-sides, and estates, often as
a gregarious weed, 400-1450 m. No seed is set;
propagation is by the many bulbils, which make it
a difficult plant to eradicate.
Vern. Tjalingtjing beureum, tj. gedé, tj. tégal, S,
kémbang gélas, J, asam-puja, Padang.
Uses. Occasionally cultivated; the leaves are
sometimes used as a substitute for tamarind
(HEYNE).
Terat. No fruits have been recorded, although
two stylar forms occur, possibly in the same
populations, but certainly in the same environ-
ment. The reason for inability of fruit-setting
probably lies in the common occurrence of mon-
struous flowers.
Sometimes the filaments broaden and form
petaloid structures, occasionally still with the
anthers present in reduced state. More often there
is a transition of the anthers towards carpel-like
structures. In the least monstruous forms the con-
nective is flabelliform with patent outer walls of
the anthers. The next step is an apical elongation
and even more developed anther-cells. The elon-
gation may bear a stigma-like structure. Especially
when there is an extra filamentous whorl between
the inner (longer) filaments and the pistil extreme
cases occur. These ‘filaments’ may finally resemble
stipitate, free carpels with a full grown style and
papillose stigma; they are apparently open on the
introrse side. Ovules have not been seen, but may
be present as they are very small and transparent
even in the normal plant. It is possible that these
‘filaments’ fuse with the pistil, as occasionally more
than 5-merous pistils carrying ovules are observed
with the stipitate alongside. Thus there seem to be
transitional stages between the anthers and pistil!
Once a sepal was observed to arise from the cen-
tre of a deformed pistil.
Notes. O. articulata SAviGNy (in Lamarck,
Encycl. Bot. 4, 1798, 686) from Argentina, is often
confused with this species. O. articulata differs
in the presence of a tuberous rhizome, by the deep-
er obcordate leaflets that have large spots, and
the denser pubescence.
O. violacea L. (Sp. Pl. 1753, 434) from N.
America, has emarginate, obdeltoid leaflets,
epunctate with two brown calli at the notch.
2. BLOPHYTUM
DC. Prod. 1 (1824) 689; Epcew. & Hook. f. Fl. Br. Ind. | (1874) 436; KNuTH,
Pfl. R. Heft 95 (1930) 391; Sreen. Bull. Jard. Bot. Btzg III, 18 (1950) 449. —
160 FLORA MALESIANA [ser. I, vol. 71
Oxalis sect. Biophytum ENDL. Gen. Pl. (1839) 1172; Mig. Fl. Ind. Bat. 1, 2 (1859)
134; PRoGEL, FI. Bras. 12, 2 (1877) 482. — Fig. 3-4.
Erect annual herbs or usually sympodially branched dwarf shrubs. Leaves
paripinnate in tufts at the end of the stem or branches, with setaceous stipules;
leaflets opposite, subsessile, terminal pair mostly different from the others, their
acroscopical half of base cuneate, basiscopical half rounded to truncate, rachis
prolonged in a mucro. Flowers terminal in a usually peduncled, bracteate pseudo-
umbel, heterodi-, tri- or homostylous. Pedicels articulate at base. Sepals glabrous
inside, -- free. Petals contort, glabrous, coherent above the claw. Filaments:
shorter ones with a more or less pronounced callus at base, longer ones always
edentate. Styles in LF and MF ciliate; stigmas terminal, subulate and entire or
spoon-shaped to flattened, crenate to bifid; ovules 3-6 per cell, alternating in 2
rows. Capsule finally loculicid to the base, forming a 5-rayed star. Seeds 1-6 per
cell, aril white, thin, at maturity bivalved and ejaculatory.
Distr. Pantropical, possibly some 70 spp., in Malesia 7 spp. among which one escaped and another
introduced.
Ecol. B. dendroides is obviously escaped from the Bogor Botanic Garden and became naturalized in
the immediate vicinity of Bogor. B. reinwardtii and B. sensitivum are weeds in anthropogenic places;
it is doubtful whether the former is indigenous. The three ligneous species, B. adiantoides, B. fruticosum
and B. microphylilum are forest dwellers. B. petersianum is a widely spread annual which prefers places
subject to a pronounced dry season, as also shown by its absence from the great ombrogenous forest belt
in Malaya, Borneo and Sumatra and scarcity in West Java (some local spots N of Bandung).
For heterostyly see under the family.
Germination is insufficiently known and seems variable. SHETTY (Proc. Symp. Recent Adv. Trop.
Ecol. Varanasi, 1968, 213—224) found that in B. sensitivum (a name not checked by me through voucher
specimens) seed collected at Varanasi (Benaras, at c. 25° NL) at the beginning of the (dry) winterseason
failed to germinate ‘under any condition’ for the first 814-9 months and showed hereby to possess a
dormancy period. Germination capacity decreased again considerably after 11-12 months. During the
first month of the germination period it needed thorough washing by which SHETTY concluded to the
presence of a water-soluble inhibitor; besides optimal temperature needed for germination was found
to be 30-40° C. Whether this behaviour holds also for Malesia is unknown; it is common in regions with-
out a dry or cooler season.
I have made germination tests with B. dendroides in the Leyden Hortus where the seeds germinated
in a very short time.
Morph. I have raised seedlings of B. dendroides of which the germination is epigaic. The first leaves,
which bear successively 2, 4, 6, and more leaflets, originate from the plumule which remains between the
cotyledons. Simultaneously the hypocotyl starts to lengthen, forming the initial stem carrying the leaf
tuft at its apex. When this cycadoid first stage is formed, one finds below the tuft usually a collar of
reflexed bristles below the leaf-scars of fallen leaves. In the annual species, characterized by a pithy
(compressible) hypocotylar stem, there is usually one apical tuft, exceptionally there are two, both ses-
sile. In the perennial ligneous species other tufts are formed, either on short-shoots or by genuine lateral
distinctly sympodial branching.
The sympodial branching of the suffruticose species is very characteristic; it is partly by thick short-
shoots on which the cortex is covered by leaf-scars, withering with age, and partly by long-shoots which
carry always under the pseudo-umbel of leaves a reflexed tuft of hairs, the shoot itself showing no leaf-
scars. In the tuft are many, often finally reflexed narrow to needle-like stipules. The peduncle of the in-
florescence resembles the ‘internode’ of the long-shoots, but it carries on top a fascicle of narrow
cuspidate bracts, the inner ones of which bear flowers.
In B. reinwardtii and B. adiantoides it may happen that in the pseudo-umbel some reduced leaves
occur. This can be found also in B. fruticosum (KAUDERN 438); of this species a still more remarkable
specimen has been collected at Sse Mao (PI. Yunnan & Mekong), Prince H. d’ORLEANsS (in P) where the
pseudo-umbel has produced a new umbel with reduced leaves and flowers, exactly similar to a normal
long-shoot. This also occasionally occurs in B. reinwardtii (RANT s.n., 1922, do, 1924, BEUMEE 4848,
KoOoRDERS 29864). A remarkable case of proliferation difficult to account for.
‘Callus’ of the shorter filaments. At the base of the annulus below the shorter (epipetalous) filaments
there is a usually dark coloured tumidity, sometimes cup-shaped, in front of the petals. Similar ones have
already been noticed in Hypseocharis by BAILLON (Adansonia 10, 1873, 363; Hist. Pl. 5, 1874, 26, 41,
f. 56, 57) and in some species of Oxalis by BAILLON (1874) and ProGeEL (FI. Bras. 12, 2, 1877, t. 111
and 113, 6). HALLIER f. discussed their taxonomical value as similar structures occur in Geraniaceae,
1971] OXALIDACEAE (Veldkamp) 161
Limnanthaceae and Linaceae (Beih. Bot. Centralbl. 39, ii, 1921, 165). NARAYANA (in litt.) found they have
no vascular trace in B. candolleanum (called *B. intermedium’, in J. Jap. Bot. 41, 1966, 321). The callosities
are possibly homologous with the scales found in Dapania on the annulus at the base of the shorter
filaments (VELDKAMP, Blumea 15, 1967, 523).
The indument is made up of setaceous, simple hairs. In the inflorescences small, septate, capitate,
glandular hairs and/or minute, red, club-shaped glands may be present.
Syst. I agree with VAN STEENIs that the subdivision of the genus as given by KNUTH is unsatisfactory
and new criteria must be considered. The first provisionally proposed sect. Sensitiva (with the generic
type species B. sensitivum) for annual species and sect. Prolifera (type species B. proliferum) for the suf-
fructicose species. But the situation might be more complicated and a subdivision has to wait for a
complete revision of the genus.
KEY TO THE SPECIES
1. Leaves 3—9-jugate; leaflets orbicular to elliptic; veins -- perpendicular to the midrib, few. Flowers
sessile in centre of tuft, (in Mal.) rarely on a peduncle up to 144 cm. Unbranched annual.
1. B. petersianum
1. Leaves 7—32-jugate; leaflets elliptic to lanceolate; veins oblique to the midrib, many. Flowers in
distinctly peduncled pseudo-umbels.
2. Midrib of leaflets (not terminal pair) at base -- median.
3. Bracts of pseudo-umbel bushy, setaceous, 3-6 mm. Calyx with long, simple hairs. Perennial,
finally (always?) branched; stem woody, not compressible, old parts reflexed. Petals + equalling
the sepals. Sepals 114-2 times as longas thefruit........
3. Bracts of pseudo-umbel ovate-attenuate,
2. B. dendroides
4, ete epi ler ae
1-3 mm. Calyx puberulous and with septate-glandular
hairs. Annual, never branched; stem soft, pithy, compressible, old parts caducous.
4. Sepals + equalling the petals, in fruit 4-7 mm long, longer than the pedicels, 114-2 times as long
as the fri ruit.
oe ». 6 el (ed wearer Bea ee, aceere
Pa ems. te ‘allie woe
3. B. sensitivum
4. Sepals + half as long as the petals, in fruit 2!4—4 mm long, shorter than the pedicels, + as long
as the fruit.
Gp BPNs; 4 ele “s? am, ve, AES
©) 5.5 Oty, O. oS ala 6) Ree ee
4. B. reinwardtii
2. Midrib of leaflets (not terminal pair) at base excentric, in basiscopic corner. Perennials, ligneous,
at least finally branched.
5. Leaves 14~32-jugate. Terminal leaflets 3-614 by 114-3 mm, obovate-oblong, as long as the pre-
ceding. Sepals in fruit 3—7-nerved, 1-2 times as long as the fruit. Fruit puberulous in upper half.
5. B. microphyllum
5. Leaves 18-27-jugate. Terminal leaflets 9-22 by 3-8 mm, oblanceolate, as long as the preceding.
Sepals in fruit 5—12-nerved, 114-2 times as long as the fruit. Fruit glabrous.
6. B. adiantoides
5. Leaves 7—-17-jugate. Terminal leaflets 5-20 by 3-8 mm, obovate to obovate-oblong, and as long as
the preceding (rarely elliptic-oblong and longer than the preceding: var. papuanum). Sepals in
fruit 3-6-nerved, slightly exceeding the fruit or + equalling it (var. papuanum). Fruit apically
RMNUD SS os tah ay Us ee ee
1. Biophytum petersianum KLotTzscu in Peters,
Reise Mossamb. Bot. 1 (1862) 81, t. 15; STEEN.
Bull. Jard. Bot. Btzg III, 18 (1950) 452; Back. &
BAKH. f. Fl. Java 1 (1963) 246; ExeLr, Fl. Zamb.
2 (1863) 158; VeLtpK. Fl. Thail. 2 (1970) 18. —
Oxalis apodiscias Turcz. Bull. Soc. Nat. Moscou
36 (1863) 595. — Oxalis petersianum (KLOTZSCH)
C. MUELL. in Walp. Ann. 7 (1868) 502. — B. apo-
discias (Turcz.) EoGew. & Hook. f. Fl. Br. Ind.
1 (1874) 437; GuILLAUMIN, Bull. Mus. Hist.
Nat. Paris 15 (1909) 124; Fl. Gén. I.-C. 1 (1911)
606. — Oxalis sessilis BuCcH.-HAM. ex WALL.
(Cat. (1831) n. 4344, nom. nud.) ex BALL. Bull.
Soc. Linn. Paris 1 (1886) 598, nom. superfl. illeg.;
BaiLey, Queens]. Fl. 1 (1899) 180. — B. sessile
(Bucu.-HAm. ex BAILL.) KNuTH, Pfl. R. Heft 95
(1930) 406, nom. illeg. — Fig. 3e-f.
“Small annual. Stem simple, rarely up to 15 cm
long. Leaves 3-9-jugate; rachis 14-314 cm, gla-
brous to appressed-pubescent, mainly on the
nodes; leaflets often overlapping, terminals 114-1}
times as long as the preceding, 2-8 by 2—5 mm,
eI Nk eae See ee ee, ee
7. B. fruticosum
obovate, -+- oblique, midrib excentric; other
leaflets triangular to orbicular-elliptic, midrib +
median; apex rounded to obtuse, glabrous or with
sparsely ciliate margin; nerves few, -- perpen-
dicular to midrib, conspicuous. Peduncle (in Mal.)
rarely present up to 1!4 cm, appressed-strigose.
Pedicels 1-3 mm, with some bristly hairs under
calyx. Sepals 3—S by 34-114 mm, ovate-lanceolate,
acute, sparsely hairy to subglabrous, in fruit
5-8-nerved, longer than pedicel, -+- exceeding the
fruit. Petals lanceolate, 5-6 by 1 mm, apex retuse,
yellow in lower half, orange and red in upper, or
orange. Filaments glabrous, (LF) 1-144 and
114-2 mm, (SF) 1 and 2 mm long. Ovary 4-114 by
Y%-1 mm, + glabrous; styles in LF 1-114 mm,
stigma flattened, crenate, in SF 14 mm, stigma
flattened, bifid; ovules 4-5 per cell. Fruit 3-4 by
2-2'4 mm, apically ciliate on the ribs. Seeds
3-4 per cell, c. 4 by 4 mm, with two longitudinal
ridges, in between with transverse rows of small
tubercles.
Distr. Tropical Africa, Madagascar, tropical
162
FLORA MALESIANA
[ser. I, vol. 73
SE. Asia (Ceylon, India, Burma, Thailand, Indo-
China); in Malesia: all islands or island groups,
except the Malay Peninsula, Sumatra, and Borneo.
Ecol. The distribution in Malesia is reflecting
a distinct preference for areas subject to a fairly
well pronounced dry season and besides helio-
philous habitats. This explains the absence from
the Malay Peninsula, Sumatra and Borneo and
its scarcity in West Java (only a few spots N of
Bandung). Ascending to 1500m (Mt Kawi).
Fl. fr. Jan.—Dec.
Fig. 3. Biophytum reinwardtii KLoTzscn. a. Ter-
minal part of leaf, x2, b. flower, x4, c. seeds,
lateral and frontal, x12. — B. microphyllum
VELDK. d. Terminal part of leaf, x2. — B.
petersianum KLotTzscn. e. Terminal part of leaf,
<2, f. seeds, lateral and frontal, x12. — B.
sensitivum DC. g. Flower, X12 (a, c JUNGHUHN
s.n., b GARRETT 243, d CONKLIN 291, e—f HEURN
s.n., g BUNG PHENG 859).
Vern. Kutjinggan, J, babonit, nibuwat-perut,
Andjai, Kebar Valley, W. New Guinea.
Uses. In the Kebar Valley eaten by women and
pigs to increase their fertility (VERSTEEGH BW 738);
a decoction is used in Mozambique as a remedy for
snake-bite (EXELL, Fl. Zamb. 2, 1963, 159) and
in the Congo as a purgative for children (WILCZEK,
Fl]. Congo Belge 7, 1958, 18).
Note. Malesian specimens are always small
with rarely a peduncle. In Africa plants may attain
40 cm with much longer peduncles.
2. Biophytum dendroides (H. B. K.) DC. Prod. 1
(1824) 690; GuILLAUMIN, Bull. Mus. Hist. Nat.
Paris 15 (1909) 125; KNutH, Pfl. R. Heft 95
(1930) 399; STEEN. Bull. Jard. Bot. Btzg III,
18 (1950) 453; Reinwardtia 1 (1952) 477; Back. &
Baku. f. Fl. Java 1 (1963) 247. — Oxalis den-
droides H. B. K. Nov. Gen. Sp. 5 (1821) 194;
PROGEL, Fl. Bras. 12, 2 (1877) 516. — Fig. 4.
Perennial. Stem woody, becoming coarse (not
compressible), up to 15cm, finally (always?)
branched, old parts reflexed. Leaves 9-16-jugate;
rachis 214-7 cm, hirsute mainly at the nodes
leaflets often overlapping, terminals 4-10 by 2-6
mm, asymmetric, falcate, others smaller, truncate
at base, elliptic to oblong, less asymmetric, with
median midrib; apex obtuse, apiculate, upper
surface sparsely hairy, underneath more so;
nerves many. Peduncle 1-31, cm, up to 7-flowered,
appressed-pubescent. Pedicels 114-314 mm, club-
shaped, shorter than the conspicuous, setaceous,
3-6 mm long bracts. Sepals 6-8 by 1-3 mm, ovate,
oblong to lanceolate, apex attenuate, acute, base
hairy, in fruit 5—8-nerved, longer than pedicel,
114-2 times as long as the fruit, equal to + shorter
than the corolla. Petals 7-8 mm, lanceolate,
rounded, pink to lavender. Filaments (LF) 214-3
and 334-5 mm, the longer ones with a few cilia.
Ovary 2 by 1 mm, apically ciliate; styles (LF)
2-234, mm long; stigma subulate, entire; ovules
4-6 per cell. Fruit 3-5 by 24% mm, apically pu-
berulous on the ridges. Seeds 1-6 per cell, 1 by 14
mm, with two longitudinal ridges, in between
with + longitudinal rows of tubercles.
Distr. Native of tropical S. America; in Ma-
lesia: locally naturalized near Bogor (W. Java),
c. 250 m.
Originally grown in the Botanic Gardens, first
collection a. 1893, spontaneous in the Gardens
a. 1905, in 1917 also collected in the vicinity of
Bogor.
Ecol. Shady, grassy places. Fl. fr. Jan.-Dec.
3. Biophytum sensitivum (L.) DC. Prod. 1 (1824)
690; BL. Bijdr. (1825) 242; EpGew. & Hook. f.
Fl. Br. Ind. 1 (1874) 436, incl. var. cumingianum
(Turcz.) EpGEw. & Hook. f.; Kina, J. As. Soc.
Beng. 62, ii (1893) 199; GUILLAUMIN, Bull. Mus.
Hist. Nat. Paris 15 (1909) 126; Fl. Gén. I.-C.
1 (1911) 608; RipL. Fl. Mal. Pen. 1 (1922) 331;
Merr. En. Philip. 2 (1923) 324; Knutn, Pfl.
R. Heft 95 (1930) 393; STEEN. Bull. Jard. Bot.
Btzg III, 18 (1950) 452; Back. & BAKH. f. Fl. Java
1 (1963) 247; VeELDK. FI. Thail. 2 (1970) 19. —
Herba sentiens RumpH. Herb. Amb. 5 (1750) 302.
— Oxalis sensitiva LINNE, Sp. Pl. (1753) 434;
Jaca. Oxal. Monogr. (1794) 42; WILLD. Sp. Pl.
2 (1799) 804; Pers. Syn. 1 (1805) 519; Zucc.
Abh. K. Ak. Wiss. Miinch. 1 (1830) 273; Roxs.
Fl. Ind. ed. Carey 2 (1832) 457; W. & A. Prod. 1
(1834) 142. — Oxalis cumingiana Turcz. Bull.
Soc. Nat. Moscou 31 (1858) 426. — B. cumingii
KLOTZSCH in Peters, Reise Mossamb. Bot. 1
1971]
OXALIDACEAE (Veldkamp)
Fig. 4. Biophytum dendroides DC. as a weed in the Leyden Hortus (Photogr. Miss R. VAN CREVEL).
(1862) 85. — B. cumingianum (TurRcz.) EDGEw.
in Edgew. & Hook. f. Fl. Br. Ind. 1 (1874) 436.
— B. sensitivum var. nervifolia (non EpGew. &
Hook. f.) F.-Vit_. Novis. App. (1880) 33. —
Fig. 3g.
Annual. Stem simple, up to 35 cm, pithy (com-
pressible), smooth. Leaves 7—12(—14)-jugate; rachis
5—10(-16'14) cm, sparsely strigose; leaflets rarely
overlapping; terminals 8-18 by 3-10 mm, + asym-
metric, falcate-obovate, midrib excentric; other
leaflets symmetric, elliptic, margins parallel,
base truncate, not drawn out, midrib median;
apex rounded, apiculate, acroscopic; + gla-
brous; nerves many, little conspicuous. Peduncles
up to 14cm, up to 10-flowered, appressed-
strigose and with patent septate-glandular hairs.
Pedicels 114-3'44mm. Bracts ovate-attenuate,
14%4-3mm. Sepals 4-7 by 14-1144 mm, ovate-
lanceolate, acute, strigose and glandular-hairy,
in fruit 5—9-nerved, longer than pedicels, 114-2
times as long as the fruit, -+- shorter than the corol-
la. Petals 5-7 by 1-2 mm, lanceolate, truncate,
base yellow, limb with purplish and yellow lines.
Filaments (MF) 1-114 and 2-214 mm, the longer
ciliate. Ovary 4-44 by 44-14 mm, apically ciliate;
styles (MF) 14-1 mm, often clasping the anthers
of the longer filaments and tearing them off;
stigmas flattened, crenate to bifid; ovules 2—5 per
cell. Fruit 3-4 by 2 mm, apically puberulous and
minutely septate-glandular hairy on the ribs.
Seeds 0-3 per cell, 34-1 by 4-34 mm, transversely
tubercled and ridged.
Distr. Widely spread in Indo-Malesian tropics,
common and throughout Malesia but not yet
recorded for New Guinea.
Ecol. Shady places, waste land, river-banks,
under damp thickets, efc., up to 250m. Fil. /r.
Jan.—Dee.
164
FLORA MALESIANA
[ser. I, voleg=
Vern. Sumatra: daun kutjingan, si-hirpud, si-
kérpud, Batak, also for Mimosa pudica); krambilan,
kutjingan, turularé, J, indja payong, Djakarta,
kalapaan, ki-payung, S, tindoh-tindoh, Patjira,
Celebes, daun hidoep, kurang-kurang, méniran
utan, Moluccas, bulutu, Kau, Halmahera, gogiolo,
Galela, Halmahera, galofino, ? igo-igo, Ternate,
obat godog, Obi, pagégga, paginga, Sula Is.,
runtili, Talaud; Philip.: damdng-bingkdlat, maka-
hia, makahiang-lalaki, Tag., damon-hiiya, hoya-
hoya, lubi-lubi, Bis., guyankan, Sub., mahihiin,
llk., niug-niug, Sul.
Uses. In the Philippines the powdered seeds
are used as a vulnerary. A decoction of the roots
is used for gonorrhoea and stones in the bladder.
A decoction of the plant is said to cure diabetes
mellitus, in the Sula Is. it is used for pregnancy
diseases (BLOEMBERGEN 4365), on the Karo Pla-
teau of N. Sumatra to diminish the female libido
(GALOENGI 49), in Ternate for chest-complaints,
here the ashes with lime-juice are given for stom-
ach-aches. A reputed medicine for tuberculosis
(HEYNE, 1927, BURKILL, 1935, and QuUISUMBING,
Medic. PI. Philip., 1951).
Note. It has appeared that what in African
Floras has been referred to B. sensitivum is really
a different species, B. helenae Busc. & MUSCHL.,
which has a corolla 114-2 times as long as the
calyx, more pairs of leaflets, is not annual and is
sometimes branched, and has a seed structure as in
B. petersianum (VELDKAMP, Blumea 16 (1968) 137).
4. Biophytum reinwardtii (ZUCc.) KLOTZSCH in
Peters, Reise Mossamb. Bot. 1 (1862) 85; EDGEw.
& Hook. f. Fl. Br. Ind. 1 (1874) 437, incl. var.
metziana EDGEW. & Hook. f. /.c. 438; RIDL. FI.
Mal. Pen. 1 (1922) 331; KNuTH, Pfi. R. Heft 95
(1930) 395; STEEN. Bull. Jard. Bot. Btzg III, 18
(1950) 453; Reinwardtia 1 (1952) 477; Back. &
BAKH. f. Fl. Java 1 (1963) 247; VELDK., Fl. Thail. 2
(1970) 19. — 8B. sensitivum var. reinwardtii
(Zucc.) GUILLAUMIN, Bull. Mus. Hist. Nat. Paris
15 (1909) 127. — Oxalis reinwardtii Zucc. Abh.
K. Ak. Wiss. Miinch. 1 (1830) 274; Mia. FI.
Ind. Bat. 1, 2 (1859) 134. — B. sensitivum (non L.)
F. M. BatLey, Queensl. Agric. J. 23 (1909) 262. —
Toddavaddi RuEEDE, Hort. Mal. 9 (1689) 33. —
Fig. 3a-e.
Annual. Stem simple, up to 35 cm, medullary,
compressible, smooth. Leaves 6—11(—14)-jugate;
rachis 214-6(-914)cm, sparsely puberulous;
leafiets rarely overlapping, terminals largest,
7-18 by 4-7 mm, asymmetric, obovate, midrib
excentric, curved; others elliptic, symmetric, base
truncate, not drawn out, margin + parallel, mid-
rib median, straight to slightly curved; apex
rounded, apiculate, + glabrous, margins sparsely
strigose; nerves many, oblique, little conspicuous.
Peduncles up to 614(-1214) cm, patent to ap-
pressed-strigose and septate-glandular hairy, up
to 8-flowered. Pedicels up to 7 mm; bracts ovate-
attenuate, 1-114 mm. Sepals 244-4 by 14-1 mm,
ovate-lanceolate, acute, in fruit 3-8-nerved, 14-24
as long as pedicel, + as long as fruit, + half as
long as corolla. Petals 6-8 by 1-2 mm, elliptic-
to oblanceolate, apex rounded to emarginate, base
yellow, above with red to purplish veins. Filaments
(MF) 1-144 and 2-3mm, the longer sparsely
ciliate. Ovary \4-1 by '4-!4 mm, glabrous;
styles (MF) 14-34 mm; stigma flattened, crenate
to bifid; ovules 3-4 per cell. Fruit 2-3 by 2-214 mm
apically puberulous and minutely septate-glandu-
lar-hairy on the ribs. Seeds 1-3 per cell, c. 1 by %
mm, transversely ridged and tubercled.
Distr. Tropical SE. Asia, in Malesia: Malay
Peninsula, Java (common), SW. Celebes (Ma-
kassar, a. 1918, one collection), and E. New Guinea
((Boku, BaILey, /.c.).
Ecol. Shady places, waste land, river-banks,
under damp thickets, efc., up to 800 m.
Vern. Biskutjinggan, krambilan, kutjinggan,
piskutjinggan, pis-ngeong, J, inger, Nusa Kam-
bangan, kakalapadn, ki pajong, Md, indja pajung
M.
Uses. Cultivated as medicine, used against
smallpox and rashes (‘sakit inja’) (EDELING s.n.)
and in India against fever (HOHENACKER 144).
Note. The restricted distribution might point to
an early introduction, but it was in BLUME’s and
JUNGHUHN’s time already a common plant in Java.
5. Biophytum microphyllum, sp. nov. — Fig. 3d.
Fruticulus usque ad 30 cm altus, ramosus. Folia
18—32-jugata. Rachis 4-8 cm longa. Foliola minuta,
paria terminalia 3-6 mm longa, 1%-4 mm lata,
obovato-oblonga, precedentibus aequilonga; cetera
lanceolata, basis parte acroscopica truncata,
saepe lobata; apex centralis, rotundatus vel obtusus;
costa basi excentrica. Flores ad pseudo-umbellas
longe pedunculatas digesti. Sepala 4-64 mm longa,
1—2 mm lata, ovato-lanceolata, acuminata, in fructu
3-7-nervosa, pedicellis 1-1.6-plo longiora, fructibus
2-3-plo longiora. Petala sepalis 1% vel duplo
longiora. Fructus distaliter paullo ciliatus. — Typus:
SuLir & CONCKLIN PNH 16905 (L, holo, PNH).
Perennial. Stem woody, up to 30 cm, branched.
Leaves (5—)18—32-jugate; rachis 4-8 cm, slender,
often curved, rusty puberulous, especially at the
nodes; leaflets rarely overlapping, terminals 3-6
by 1144-4mm, obovate-oblong, + as long as the
preceding, midrib -+- median; others lanceolate,
basiscopical half of base rounded, acroscopical
half truncate, often drawn out, midrib excentric,
margins + parallel; apex central, obtuse to
rounded, apiculate; upper surface appressed-
puberulous, beneath + more pubescent; nerves
many. Peduncle 2-534 cm, appressed to patently
pubescent, eglandular. Pedicels 5—-9mm _ long,
appressed- to patently puberulous, eglandular.
Sepals 4-614 by 34-2 mm, lanceolate, acuminate,
the outer three much wider, + half as long as
the corolla, in fruit 3-7-nerved, 3/; to + equalling
the pedicel, c. 14times as long as the fruit
(excl. styles). Petals 7-9 by 1-214 mm, apex
rounded, white to lavender. Filaments (MF)
234-314 and 4-534 mm, (LF) 4 and 144mm,
the longer ciliate. Ovary 144 by 1 mm, glabrous;
styles (MF) 214-3 mm long, (LF) not seen in
anthesis; stigma minutely bifid; ovules 3-4 per cell.
Fruit 2-3 by 2mm, puberulous in upper half.
OXALIDACEAE (Veldkamp)
165
Fig. 5. Distribution of Biophytum fruticosum BL. var. fruticosum (+), ditto, var. papuanum VELDK.
(x), B. adiantoides WiGut (@), B. thorelianum GUILLAUMIN (4), and B. microphyllum VELDK. (@).
Seeds 3-4 per cell (no ripe ones seen).
Distr. Malesia: Philippines (E. Mindoro: Mt
Yagaw), SW. Central Celebes (Pasui), and Lesser
Sunda Is. (W. Flores). Only four collections.
Fig. 5.
Ecol. In crevices of rocks on forest edge, once
on limestone, 400-600 m, apparently /f. and /r.
all year.
Vern. Huya-huya-ili, tagurignik-sa-ili, Mang.,
Philip.
6. Biophytum adiantoides WiGutT ex EpGew. &
Hook. f. Fl. Br. Ind. 1 (1874) 437; Kino, J. As.
Soc. Beng. 62, ii (1893) 200; GuILLAUMIN, Bull.
Mus. Hist. Nat. Paris 15 (1909) 124; Fl. Gén.
I.-C. 1 (1911) 609; Rip. FI. Mal. Pen. 1 (1922)
331; Knutn, Pfl. R. Heft 95 (1930) 397; VELDK.,
Fl. Thail. 2 (1970) 20.
Perennial. Stem woody, usually branched, up to
30 cm high. Leaves 18-27-jugate; rachis 7-17 cm,
yellowish appressed to patently pubescent;
leaflets rarely overlapping, terminals 9-22 by
3-8 mm, oblong to lanceolate, widest at or above
the middle, + as long as the preceding, midrib
+ median; others asymmetric, elliptic to oblong,
basiscopical half of base rounded, acroscopical
half truncate, more or less drawn out, margins
otherwise + parallel, midrib excentric at base;
apex central, rounded, apiculate; above sparsely
strigose, hairs often in three rows, to glabrescent,
beneath more strigose; nerves many. Peduncle
5-1914 cm, up to 9-flowered, puberulous, some-
times with a few septate-glandular hairs. Pedicels
5-17 mm, puberulous and with a few gland hairs.
Sepals 414-6 by 1-1'4 mm, lanceolate, acute,
sparsely strigose to glabrous, sometimes with a
few septate gland hairs, 14-24 as long as the
corolla, in fruit S—12-nerved, half to + equally as
long as the pedicel, 114-2 times as long as the
fruit. Petals 9-10 by 1-214 mm, lanceolate, apex
rounded to truncate, white with yellowish base.
Filaments in SF 114-314 and 414-614 mm long,
in MF 114 and 53% mm long, in LF 1-144 and
2%4-4mm long, the longer ciliate. Ovary 1 by
14-44, mm, glabrous; styles puberulous, in SF
14-114 mm, in MF 2!4 mm, in LF 4-6 mm long;
stigma flattened, crenate to bifid; ovules 2-4 per
cell. Fruit 3-4 by 2-3 mm, glabrous. Seeds 2-3
per cell, 1-14 by %-l1 mm, with transverse
tuberculated ridges.
Distr. S. Vietnam, Cambodia, Thailand,
Tenasserim, and in Malesia: northern half of the
Malay Peninsula (Perlis, Perak, Pahang). Fig. 5.
Ecol. In crevices of (limestone) rocks
166
FLORA MALESIANA
[ser. I, vol, 72
along rivers and in open woods, up to 300 m.
Vern. Daun pajong, maiong, pajong ali, M.
Uses. Given to small children against stomach
trouble in Perak.
7. Biophytum fruticosum BL. Bijdr. (1825) 242;
KnuTH, Pfl. R. Heft 95 (1930) 412 in annot.
sub B. nudum; STEEN. Bull. Jard. Bot. Btzg III,
18 (1950) 454; Reinwardtia 1 (1952) 477, p.p.
— B. esquirolii Lév. in Fedde, Rep. 12 (1913) 181;
Knut, Pfi. R. Heft 95 (1930) 413; Merr.
Lingn. Sc. J. 13 (1934) 31; REHDER, J. Arn. Arb. 18
(1937) 209. — B. thorelianum var. sinensis GUIL-
LAUMIN, Not. Syst. 1 (1909) 25; Bull. Mus. Hist.
Nat. Paris 15 (1909) 128; KNuTH, Pfi. R. Heft 95
(1930) 413. — Oxalis blumei Zucc. Abh. K. Ak.
Wiss. Miinch. 1 (1830) 276; Mia. FI. Ind. Bat. 1,
2 (1859) 134.
var. fruticosum.
Perennial. Stem woody, branched, up to 50 cm.
Leaves 7-17-jugate; rachis 3-9 cm, appressed to
patently strigose, especially on the upper surface;
leaflets rarely overlapping; terminals 5-12 by
3-8 mm, -+ as long as the preceding, obtriangular
to obovate-oblong, -+ symmetric, midrib median;
other leaflets elliptic to lanceolate, basiscopical
half of base rounded, acroscopical half truncate,
more or less drawn out, midvein excentric; apex
rounded, apiculate, central; above densely strigose,
glabrescent, then hairs often in three rows, lower
surface less strigose; nerves many. Peduncle up to
13 cm, strigose, no septate glandular hairs, up to
10-flowered. Pedicels up to 9 mm. Sepals 314-6 by
1-114 mm, ovate-lanceolate, densely to sparsely
strigose, no septate glandular hairs, in fruit
3-7-nerved, */, as long as the pedicel, slightly
exceeding the fruit, c. 24 as long as the corolla.
Petals 514-10 by 1-214 mm, lanceolate, obtuse,
white to pink. Filaments in SF 2-3 and 314-4 mm,
in MF 2-214 and 3 mm, in LF 114-4 and 214-5
mm. long, the longer always ciliate. Ovary 4-1
by 14-1 mm, apically with a few cilia to glabrous;
styles in SF 14mm, stigma bifid, flattened; in
MF 214-4mm, in LF 2-5 mm, stigma subulate
to slightly flattened, entire; ovules 2-4 per cell.
Fruit 214-4 by 214-3 mm, puberulous but eglan-
dular in upper half. Seeds 2-4 per cell, 1 by 34 mm,
transversely tuberculate and ridged.
Distr. SW. China (Yunnan, Hupeh, Kouy-
Tschiou, Canton, Hainan), Indo-China (Mekong),
NE. India (Silhet); in Malesia: Philippines
(Bohol; Sulu Is.: Bengao I.), E. Celebes, and
Lesser Sunda Islands (S. Central Timor; S. Wetar).
Fig. 5.
Ecol. In crevices of rocks, on river-banks, in
thickets and shaded cultivated areas on sand, up
to 800 m.
var. papuanum, var. nov. —Oxalis albiflora F. vy. M.
Vict. Natur. 8 (1892) 164, nom. nud. — B. albi-
florum F. v. M. Vict. Natur. 9 (1893) 112; J. Bot.
31 (1893) 325; KNuTH, Pfl. R. Heft 95 (1930) 397;
STEEN. Bull. Jard. Bot. Btzg III, 18 (1950) 455. —
Oxalis papuana F. v. M. Vict. Natur. 9 (1893)
112, nom. altern.
Terminal leaflets 14-20 by 6-8 mm, elliptic to
oblong, -+ 1.3 times as long as the preceding.
Pedicels 9(—25) mm. Sepals in fruit + ?/; as long
as the pedicel. Fruit + as long as the calyx.
Distr. Malesia: SE. New Guinea (Owen
Stanley Range), two collections (type not seen).
Fig. 5.
Ecol. Stony stream-banks in forests.
Note. In his preliminary revision VAN STEENIS
accepted a much wider specific concept of B.
fruticosum than handled here, in merging B.
adiantoides with B. fruticosum and also referring
the specimens here distinguished as B. micro-
phyllum to it. In fact all three native Malesian
fruticose Biophytums and B. thorelianum GUILLAU-
MIN from Indo-China (fig. 5) differ from allied
continental species (B. nudum, B. proliferum,
B. intermedium, and B. polyphyllum) in having the
midrib obliquely inserted at the base and the
Asian ones having them median. However, the
Malesian material can be sorted into three taxa,
although it must be admitted that the great
scarcity of collections makes it difficult to find the
parameters of their variability.
3. DAPANIA
Kortu. Ned. Kruidk. Arch. 3 (1854) 381; HALL. f. Beih. Bot. Centralbl. 34, ii
(1917) 25; KNutn, Pfl. R. Heft 95 (1930) 419; VELDKAmp, Blumea 15 (1967) 523.
— Fig. 7i.
Glabrous lianas. Leaves estipulate, unifoliolate; petioles articulate with a
constriction (in herb.). Inflorescences racemose, ramiflorous and axillary, solitary
to fascicled. Flowers (in Mal.) androdioecious. Sepals connate in lower half,
margins ciliate, glabrous inside. Petals apotact or paratact, rarely quincuncial,
(in Mal.) free and glabrous inside, minutely clawed, red to white. Filaments (in
Mal.) antheriferous, between the filaments with scales on the annulus, these
sometimes reduced to dark lines. Ovary glabrous, reduced and sterile in ¢ plants.
Ovules in 3 1-2 per cell, in ¢ 0-1. Capsule fleshy, yellowish green (red when dry),
167
1971} OXALIDACEAE (Veldkamp)
loculicid to the base, 1—-6-seeded, valves patent; (in Mal.) episeptal rimae present,
open to base. Seeds up to 2 per cell; testa smooth, hard, yellowish red, terminally
thickened around the micropyle, splitting lengthwise under pressure; aril present,
enveloping the seed, attached to the entire length of the adaxial raphe, fleshy,
bright to whitish yellow, margin irregular crenate, with oily drops; embryo with
the cotyledon 114-2 times as long as the straight, oblique radicle.
Distr. Madagascar (1 sp.), in Malesia (Sumatra, Malaya, Borneo) 2 spp.
Ecol. Lianas in forests, swamps, near rivers, obviously usually on poor soil, at low altitude.
KEY TO THE SPECIES
1. Leaves up to 15cm, base cuneate to rounded. Sepals glabrous, except the ciliate margin.
1. D. racemosa
1. Leaves longer than 15cm, base deeply emarginate. Sepals puberulous outside.
1. Dapania racemosa KorTu. Ned. Kruidk. Arch. 3
(1854) 381; PLANCHON, Ann. Sc. Nat. Bot. IV,
2 (1854) 266; Mia. FI. Ind. Bat. 1, 2 (1859) 134;
Starr in Hook. f. Ic. Pl. III, 10 (1891) t. 1997;
HALL. f. Beih. Bot. Centralbl. 34, ii (1917) 25;
KwnutTH, Pfl. R. Heft 95 (1930) 420; VELDKAMP,
Blumea 15 (1967) 552, f. 1d-g. — D. scandens
Starr in Hook.f. Ic. Pl. III, 10 (1891) t. 1997;
Kina, J. As. Soc. Beng. 62, ii (1893) 201; HALL.
f. Beih. Bot. Centralbl. 34, ii (1917) 25; Rit. FI.
Mal. Pen. 1 (1922) 334; KNuTu, Pfl. R. Heft 95
(1930) 420, f. 28a-e.
Large liana, up to 30m long, 20cm g, rather
profusely branched. Leaves 534-15(-25) by
214-634 cm, oblong to lanceolate, pergamenta-
ceous to subcoriaceous, acute to cuspidate, base
cuneate to rounded; petiole 3-10 by 1-2 mm,
petiolule 2-6 by 1-2mm. Racemes solitary to
fascicled, in g 1-3 together, 3-1314 cm, in ¢
1-15 together, 2—514(—1214) cm; rachis puberulous,
glabrescent. Pedicel (lower joint 0-1 mm, upper
14-14 mm) not covered by the minute, broadly
ovate, acute bract 144-14 by 44 mm. Calyx 1%-2
mm high, glabrous to sparsely puberulous out-
side. Sepals 1-11, by 34-114 mm, broadly ovate
to elliptic, rounded to emarginate. Petals 3-4 by
1-114 mm, obovate-oblong to -lanceolate, often
rolled back at anthesis, obtuse to rounded, darker.
Filaments in g Y%4-¥, and %4-1 mm, in 3 1-3 and
14%4-4mm long. Pistil in Z 114-244 mm, in 3
V4—¥%, mm; styles in ¢ 1-114 mm, in J very short,
acute. Fruit obovoid with cuneate base, 5-11 by
6-10 mm, after dehiscence 9-22mm o. Seeds
4-5 by 1!4mm; radicle 114-134 mm _ long,
cotyledons 2-214 by 34-1 mm.
Distr. Malesia: Sumatra,
Fig. 6.
Ecol. Dense jungle, swamp forest, on poor
soils, up to 750m. Fi. fr. Jan.—Dec.
Vern. Batieg-batieg, dapan, Sumatra; bélimbing
bait, M, jaloai, Brunei, kara-raka, Iban; pau-
kiang, Malacca, sélambun akar, Selangor.
Note. A large-leaved form occurs in North
Sumatra.
2. Dapania grandifolia Vetpkamp, Blumea 15
(1967) 525, f. Ih-p. — Fig. 7i.
Malaya, Borneo.
2. D. grandifolia
a
Fig. 6. Distribution of Dapania racemosa KorTH.
(@) and D. grandifolia VELDK. (A).
Liana, or treelet, up to 27m, 7cm g. Leaves
coriaceous, (1014—)1614-31 by 414-11 cm, oblong
to lanceolate, widest at or below the middle, base
retuse to deeply emarginate, apex acute to at-
tenuate; petiole 2-6 by 2-34 mm, stout, petiolule
1-3 by 2-344 mm. Racemes + densely and +
patently puberulous, not glabrescent, in dg 1-5
fascicled, 3-5 cm long, in g 1-2 together, 4-8 cm
long. Bracts boat-shaped, sometimes recurved,
¥%-1 by 44-1 mm, broadly ovate, acute, covering
the minute (c. 14 mm long) pedicel, but not the
base of the calyx. Calyx 13%4-2'4mm_ high,
pale to brown puberulous. Sepals 1-1'4 by
44-11% mm, suborbicular to ovate, acute to round-
ed. Petals 4-5 by 14-114 mm, obovate-lanceolate,
rounded, base attenuate. Filaments in 3 2-3',4
and 3-4 mm, in d 4%-% and %-1 mm. Pistil in
dé reduced, sterile, 4-4 by %4-\4 mm, ovoid;
styles minute, acute; ¢: ovary 14% by 1mm,
styles 2mm. Fruit before dehiscing 9-11 by 6-8
mm, suborbicular to ovoid, after dehiscence 20-22
mm @. Seeds 444-5 by 114-1344 mm; embryo 4 mm
long, radicle 1144 by 4 mm, cotyledons 2'4 by
14% mm.
Distr. Malesia: NE. to SE. Borneo. Fig. 6.
Ecol. Primary forest, on flat to undulating
land on clay near rivers. Fl. fr. Jan.—Dec.
Vern. Salung kapit, Dusun.
168 FLORA MALESIANA [ser. I, vole?
4. SARCOTHECA
BLume, Mus. Bot. Lugd. Bat. 1 (1850) 241; HALL. f; Med. Rijksherb. Leiden 1
(1911) 1; KNutH, Pfl. R. Heft 95 (1930) 420; VeLDKAmp, Blumea 15 (1967) 527
— Roucheria Mia. FI. Ind. Bat. 1, 2 (1859) 136. — Connaropsis PLANCH. ex HOOK.
f. Trans. Linn. Soc. 23 (1860) 166. — Fig. 7a-h.
Fig. 7. Sarcotheca glauca HALL. f. a. Habit, «24, b. flower, <4, c. petal, <6, d. stamens and ovary
(SF), 10, e. ovary (SF), x10, f. stamens and ovary (LF), x10, g. ovary (LF), x10, A. fruit, x2.
— Dapania grandifolia VELDK. i. Annulus with scales and filaments, 10 (a, h Puasa BNB 5484, b-c,
f-g LAJANGAH SAN 33612, d-e MusIn SAN 33562, i KOSTERMANS 10104).
Shrubs or trees; innovations pubescent. Leaves estipulate, 1- or 3-foliolate;
petioles articulate, with a constriction (in herb). Panicles axillary or pseudoter-
minal, one to few together; flowers in more or less stalked cymes, scattered along a
simple or sparsely branched rachis; cymes subtended by small caducous bracts,
occasionally by a reduced petiole (petioloid), or rarely a small leaf. Flowers
heterodistylous. Sepals unequal, shortly connate at base, inside appressed strigose,
persistent (except in S. diversifolia). Petals contorted, sometimes paratact, inside
with minute papillae in the upper half. Filamental annulus without scales. Ovules
1971] OXALIDACEAE (Veldkamp) 169
2 per cell. Fruit fleshy, red at least when dry, with + distinct episeptal rimae
sometimes lighter and minutely papillose inside (at least when dry). Seeds exaril-
late; testa smooth to transversely rugose, hard, reddish, splitting irregularly by
pressure; embryo with a straight radicle in line with the cotyledons which are 3-5
times as long as the radicle.
Distr. W. Malesia: Sumatra, Malaya, Borneo, Celebes. Fig. 8.
Ecol. Primary and secondary forest on poor soil at low altitude. Fi/. fr. Jan.—Dec.
Uses. The fruit, although acid, is eaten in curry, sajur, and manisan and is said to be a remedy against
coughing. The timber is light and of small dimension, neither very strong nor durable, sometimes used
for roofs.
Notes. The genus has often been confused with Rourea AuBL. (Connaraceae) which differs from
Sarcotheca in having free carpels, 2 collateral ovules, seeds with an aril, and a dry, indehiscent, 1-celled
and 1-seeded fruit.
The species of the genus are closely related and most of them occupy small ranges; in several cases
close allies show replacing areas. With a broader species concept several species would be reduced to sub-
specific rank, notably the triad S. monophylla from Malaya, S. glauca from Borneo, and S. celebica from
Celebes.
KEY TO THE SPECIES
1. Leaves trifoliolate, laterals sometimes caducous, leaving a scar. Mature fruit greenish yellow when
fresh (red when dry!); more than 114 cm long.
2. Calyx 3-5 mm high, outside pale puberulous to glabrous, not persistent in fruit. 1. S. diversifolia
2. Calyx 214-3 mm high, outside brown puberulous, persistent in fruit... . .. . . 2. S. griffithii
1. Leaves unifoliolate; no lateral scars. Mature fruit red when fresh; less than 114 cm long.
3. Mature leaf puberulous to pubescent beneath, also between the nerves.
4. Panicles not or barely exceeding the leaves. Petals 4-7 mm long. Rimae obscure, glabrous inside.
5. Leaves 8-23 by 3-814 cm, margins never paler. Calyx 1144-2 mm high. . . . . . . . 4. S. laxa
5. Leaves 5—1114 by 2-3!4cm, margins usually paler. Calyx 2-334 mm high. . ._ 8. S. ferruginea
4. Panicles exceeding the leaves. Calyx 3-314 mm — Petals 6-8 mm long. Rimae conspicuous,
lighter and minutely papillose inside.. . . | outs... teed OsS. ochracea
3. Mature leaf beneath at most sparsely puberulous | on 1 the nerves.
6. Calyx (and often panicle also) glabrous to finely pale puberulous. Rimae obscure and glabrous
inside.
7. Panicle glabrous, stout. Cymes sessile, flowers in glomerules along the rachis.
3. S. glomerula
7. Panicle puberulous, slender. Cymes stalked. . . . ikaw 14,'Sdaxa
6. Calyx at least at the base (and panicle) rusty puberulous. Rimae conspicuous, inside lighter and
minutely papillose.
8. Leaves 4-1314 cm long. Panicle up to 13 cm, erect, + compact (lax in S. celebica).
9. Cymes not + secund. Basal parts of pedicels all + equal.
10. Leaves when dry brown to reddish brown. Basal parts of pedicels 0-2 mm long. Calyx 134-214
mm high, persistently puberulous outside, reddish brown when dry.
. Leaves 4-10 cm long, acuminate to cuspidate. Panicle + dense, pedicels with shorter and
reduced upper joint (c. 44mm). .. . . . . . . 6. S. monophylla
11. Leaves 6—-1314 cm long, cuneate-acute to ‘faintly acuminate; venation above more prominent
than in the other two species. Panicle slender, lax, joints of pedicel -- equal (up to 114 mm).
7. S. celebica
10. Leaves when dry palish to olive green, 414-11 cm long, acuminate. Basal part of pedicels 2-6
mm long. Calyx 2-3 mm high, outside glabrescent in fruit, except for base and margins,
crimson when dry. . . . 5. S. glauca
9. Cymes + secundly branched. Basal part of one pedicel per cyme usually elongated, up to 5 mm.
Nerves of Jeaf beneath often reddish when dry. .......+4+0+488- 9. S. rubrinervis
8. Leaves 714-2814 cm long. Panicle usually pendulous, slender, ‘usually much longer than 13 cm.
.12. Leaf widest at the middle, apex gradually acute to caudate, base obtuse to rounded. Nerves
often reddish when dry. Petiolules 2~4 by 1-2 mm. Cymes + secundly branched, basal part of
one pedicel elongated, up to 5 mm. Claw of petals 74-1 mm long. . . . . . 9. S. rubrinervis
- 12. Leaf widest at or above the middle, margins + parallel, apex abruptly acuminate to caudate,
base truncate to emarginate. Nerves concolourous with intervenium when dry. Petiolules 3-9
by 1-3 mm. Cymes not + aa ved branched, basal dye of oon + equal. Claw of petal
Yy-Yymmiong. .... . whe ‘ . . 1. S. macrophylla
FLORA MALESIANA
(ser. I, voli 7*
Fig. 8. Distribution of Sarcotheca BL.
1. Sarcotheca diversifolia (M1Q.) HALL. f. Med.
Rijksherb. Leiden 1 (1911) 2; VELDKAmP, Blumea
15 (1967) 529, f. 2a-c. — Rourea diversifolia
Mia. FI. Ind. Bat. Suppl. 1 (1860) 528. — Con-
naropsis diversifolia Kurz, J. As. Soc. Beng. 39,
ii (1870) 69, excl. syn. C. griffithii. — Santalodes
diversifolium O. KUNTZE, Rey. Gen. Pl. 1 (1891)
155. — Connaropsis acuminata PEARSON, Kew
Bull. (1906) 2. — S. acuminata HALL. f. Beih.
Bot. Centralbl. 34, ii (1917) 27. — S. subtripli-
nervis HALL. f. l.c. — Connaropsis grandiflora
Rip. Kew Bull. (1930) 75.
Shrub or tree, up to 27m high, to 90cm @,
often buttressed (90 cm high, 135 cm out, 5 cm 9g).
Leaves trifoliolate, papyraceous to subcoriaceous,
ovate- to elliptic-oblong to -lanceolate, glabrous,
acute to caudate, base acute to truncate, venation
not very conspicuous; nerves 1-5 pairs, usually
-++ prominent, veinlets not as finely reticulate as in
S. griffithii; lateral leaflets 3-914 by 1-334 cm,
terminal ones 514-18 by 2-634 cm; petiole 6-25
by 1-214 mm, rachis (5—)9-27 by >4-2 mm;
petiolules 4-7 by 34-2 mm. Panicles shorter than
subtending leaf, 1-4 together, loosely branched,
1-814(-1314) cm, ferrugineous-puberulous, gla-
brescent; branches flattened, 2—30(-40) mm long;
bract or petioloid ovate, acute, up to 7 mm.
Pedicels: lower joint 214-5 mm, upper 1-3 mm;
Calyx 3-5 mm high, outside pale puberulous to
glabrous, purplish, -- caducous in fruit. Sepals
24%-5 by 114-3 mm, broadly ovate to oblong,
acute to emarginate. Petals 7-11 by 2-5 mm,
obovate-oblong to -lanceolate, rounded to emar-
ginate, 1-2 mm clawed, lilac or pink to scarlet or
red. Filaments in SF 234-344 and 344-414 mm,
in LF 114-24% and 2144-3 mm, the longer oc-
casionally without anthers. Pistil pale puberulous
to glabrous, in SF 114-214 mm, in LF 314-5 mm;
styles in SF 14-1 mm, in LF 214-4; ovary 1-2
by 34-114 mm, ellipsoid. Fruit white to greenish
when fresh (red when dry), ellipsoid, glabrous,
16-31 by 9-20 mm; rimae inconspicuous. Seeds
7144 by 5mm, testa smooth; cotyledons 474 by
4mm, radicle 114-134 by 4/, mm.
Distr. Malesia: N. Sumatra (off Sibolga:
Morsala I.), Borneo.
Ecol. First and second storey of primary and
secondary forest on wet, well-drained, poor soil
(e.g. podsolized sand) on undulating to flat land,
up to 900 m.
Vern. Bélimbing bulat, M, buah piang, \ban,
iba jantan, Suluk, kandis(-daham), Tidong,
kérapa-kérapa, pérapan macas, Tutong Dusun,
kadazan, tabaus, tébarus, Brunei.
2. Sarcotheca griffithii (PLANCH. ex Hoox. f.)
HALL. f. Med. Rijksherb. Leiden 1 (1911) 2;
KnuTH, Pfl. R. Heft 95 (1930) 425; VELDKAmpP,
Blumea 15 (1967) 531, f. 2e-f. — Connaropsis
griffithii PLANCH. ex Hook. f. Trans. Linn. Soc.
23 (1860) 166; EpGew. & Hook. f. Fl. Br. Ind. 1
(1874) 440; Kurz, J. As. Soc. Beng. 39, ii (1870)
69; KING, ibid. 62, ii (1893) 200; Rip. Fl. Mal.
Pen. 1 (1922) 332, fig.; KOCHUMMEN, Res. Pamphl.
(For. Res. Inst. For. Dep. Mal.) 43 (1963) 6, 38. —
Dapania griffithii KNUTH, Bot. Jahrb. 50 (1914) 234.
Tree, up to 42 m, 1 mg (buttresses up to 4m,
1 m out). Leaves trifoliolate, glabrous, pergamen-
taceous, often crisped when dry, venation rather
inconspicuous, veinlets above finely reticulate;
leaflets elliptic-oblong to -lanceolate, occasionally
+ obovate, tapering acuminate to caudate, base
obtuse to acute, lateral ones 2-7 by 34-234 cm,
nerves 1—5 pairs; terminal leaflet 4-11144 by
114-4 cm, nerves 4-6 pairs; petiole 5-12 by
144-2 mm, rachis 3-16 by 34-1 mm; petiolules
3-7 by %-14% mm. FPanicles shorter to +
longer than the subtending leaf, stout, densely
branched, ferrugineous-puberulous, 2-10 cm; bracts
small, ovate or linear, petioloid up to 214 mm.
Flowers pectinately clustered, subtended by
minute bracts. Lower joint of pedicel 1-214 mm,
upper 14-1 mm. Calyx 214-3 mm high, dark red,
outside brown puberulous. Sepals 2-234 by 114-21),
mm, broadly ovate to -obovate, acute to emargi-
nate. Petals + glossy, blackish red, whitish at
base, 414-814 by 2-214 mm, obovate-lanceolate
to -oblong, rounded to obtuse. Filaments in SF
4-414 and 5-514 mm, in LF 114% and 2 mm; the
longer sometimes without anther and club-shaped,
red. Pistil in SF 134 mm, in LF 3-4 mm; pu-
berulous; styles in SF 34 mm, in LF 2-3 mm;
ovary 1 by 34-1 mm, subglobose. Fruit 18-32 by
10-23 mm, ellipsoid, glabrescent, greenish yellow
at maturity, inside yellow, mesocarp glassy;
rimae inconspicuous. Seeds 5—7'4 by 2-4mm;
testa smooth; radicle 114 by 14 mm, cotyledons
5Y% by 3144 mm.
Distr. Malesia: Sumatra, Malay Peninsula.
Ecol. First storey of primary forest on flat to
undulating sandy clay on dry to temporarily
flooded land.
Vern. As(s)am pupy (or pupoii), bélimbing
(hutan), kaju manau, Lampong, kukui, Oesoe,
lain jénis, jinték-jinték, kupoyi, pandija, N. Su-
matra, poké pupoé, pupui, pupoi, pupoy.
Notes. Leaves of seedlings are 1-foliolate.
According to EDGEWoRTH & Hooker f/f. the
leaves are irritable to the touch.
3. Sarcotheca glomerula VELDKAMP, Blumea 15
(1967) 532, f. 2i-j. — Connaropsis macrophylla
Kina, J. As. Soc. Beng. 62, ii (1893) 201, non
S. macrophylla Bui. 1850; Rip. Fl. Mal. Pen.
1971]
1 (1922) 334. — Dapania macrophylla (KiNG)
KnutTH, Bot. Jahrb. 50 (1914) 234. — S. macro-
phylla (non BL.) KNuTH, Pfl. R. Heft 95 (1930)
424, pro specim. malay., excl. specim. born. et syn.
Miq.
Shrub or small tree, up to 9m, 8cm g. Leaves
unifoliolate, (6—)1114-28 by (2144-)4-9 cm, per-
gamentaceous to subcoriaceous, glabrous, oblong
to lanceolate, margins + parallel, contractedly
acute to cuspidate, base broadly truncate to
emarginate, not glaucous; nerves 8-13 pairs,
branching off with wide angle (lower + 65-909),
basals usually recurved; petiole 4-1114 by 114-3
mm, petiolule 3-8 by 2-3 mm, glabrous. Panicles
subterminal or axillary, 1-few together, + erect,
glabrous, stout, 4-19 cm; branches reduced, wart-
like, up to 24mm. Flowers in few-flowered
fascicles. Lower joint of pedicel -+- absent, upper
1-3 by 3% mm. Calyx 114-2 mm high, glabrous
outside except the ciliate margin. Sepals 1144-114
by 144-114 mm, broadly ovate, rounded to obtuse.
Petals 6-84 by 134-214 mm, lanceolate-oblong
to obovate-lanceolate, rounded, red to crimson,
tube lighter. Filaments in LF 1 and 134 mm, in
SF 214-3 and 3144-4mm. Pistil sparsely puber-
ulous, in SF 2mm, in LF 4mm; styles in SF
34-1 mm, in LF 214-3 mm; ovary | by 24-34 mm,
subglobose, pubescence mainly apically. Fruit
8-16 by 5-10 mm, ellipsoid to ovoid, acute to
acuminate, recurved, glabrous, red; rimae in-
conspicuous, in the upper half, neither lighter nor
papillose inside. Seeds 10 by 4mm; testa trans-
versely rugose; cotyledons 714 by 3 mm; radicle
c. 1 mm long, stout.
Distr. Malesia: Malay Peninsula.
Ecol. Primary and secondary forest on flat to
undulating land.
Vern. (Asam) (sé-)tundok, bélimbing bési, b.
hutan, séndok, tétindok.
4. Sarcotheca laxa (RIDL.) KNuTH, Pfl. R. Heft 95
(1930) 422; VELDKAMP, Blumea 15 (1967) 533. —
Connaropsis laxa RipDL. J. Str. Br. R. As. Soc. n. 75
(1917) 9; Fl. Mal. Pen. 1 (1922) 333. — Con-
naropsis glabra Riv. J. Str. Br. R. As. Soc. n. 75
(1917) 9; Fl. Mal. Pen. 1 (1922) 332. — S. glabra
KwnuTH, Pfl. R. Heft 95 (1930) 422. — Connaropsis
sericea Rivv. J. Fed. Mal. St. Mus. 10 (1920) 121;
Fl. Mal. Pen. 1 (1922) 334. — S. sericea KNUTH,
Pfl. R. Heft 95 (1930) 424. — Connaropsis sim-
plicifolia Ripi. Fl. Mal. Pen. 1 (1922) 334. —
S. simplicifolia KNUTH, Pfl. R. Heft 95 (1930) 424.
Shrub or tree, up to 23m high, 45cm @.
Branches glabrous or densely ferrugineous-
tomentose, glabrescent. Leaves unifoliolate, (5—)
8-23 by (2-)3-8!'4cm, oblong to lanceolate,
elliptic to obovate, acuminate to caudate, base
broadly cuneate to truncate, above sparsely silky
pubescent, mainly at base and on the midrib,
beneath sometimes subglaucous, glabrous or pale
to ferrugineous pubescent; nerves 5—11 pairs, basals
sometimes at 90°; petiole 5-25 by 1-314 mm, peti-
olule 3-7 by 1-3 mm, glabrous or pubescent. Panicles
1-3 together, up to 30 cm, erect, pendent in fruit,
glabrous, or finely pale puberulous, or ferrugineous
OXALIDACEAE (Veldkamp)
171
velvety; branches patent to recurved in fruit, up
to 14mm, sometimes once forked near the top,
subtended by a bract, a petioloid, or a small leaf.
Lower and upper joints of pedicel subequal,
%—-1mm. Calyx 1144-2 mm high, outside gla-
brous or very shortly pale puberulous or ferru-
gineous puberulous. Sepals 14%4-2 by 1-2 mm,
broadly ovate to spathulate, obtuse to emarginate.
Petals 4-7 by 1'4%4-2mm, obovate-lanceolate,
rounded to emarginate, whitish to dark red.
Filaments in SF 1-24% and 2-314 mm, in LF
14-1 and 1-134 mm. Pistil glabrous to appressed-
strigose, in SF 1-1!4 mm, in LF 3-4 mm long;
styles in SF 14 mm, in LF 2-3 mm long; ovary
¥%-1 by %-*4mm, ellipsoid. Fruit 6-13 by
4-9 mm, ovoid to ellipsoid, recurved, rounded to
acuminate, glabrescent; rimae inconspicuous,
closed not lighter nor glandular inside. Seeds
5-9 by 3-4 mm; testa smooth to transversely
rugose; cotyledons 4-5 by 2-214 mm, 3-4 times
as long as the radicle measuring 1-114 by 14 mm.
Distr. Malesia: Malay Peninsula, ? NE.
Sumatra.
Ecol. Forest edges and swamps at low altitudes.
Note. A collection from Sumatra (Tenajan
R., Upper Riouw, SOEPADMO 154) of which the
leaves and also the fruit somewhat resemble the
Bornean S. rubrinervis, but of which the panicle
is like the Malayan S. laxa var. sericea, might
be a new variety of S. Jaxa, but better material is
needed to reach a definite conclusion whether this
species occurs also in Sumatra.
a. var. laxa. VELDKAMP, Blumea 15 (1967) 535,
f. 3a—b. — S. laxa (RIDL.) KNuTH. — S. glabra
(RIDL.) KNUTH.
Nearly glabrous throughout. Nerves 5-9
pairs. Branches of panicle distinct, slender, 2-14
mm, near top once forked. Petals retuse to trun-
cate, 444-4% by 1144-144 mm, claw 0-14 mm,
stout. Fruit 6-7 mm long. Seed 5 by 3 mm.
Distr. Malesia: NW. Malay Peninsula.
Vern. Mésékam.
b. var. sericea (RIDL.) VELDKAMP, Blumea 15
(1967) 535, f. 3c—d. — S. sericea (RIDL.) KNUTH.
— 8S. simplicifolia (RipL.) KNUTH.
Leaves glabrous above, sparsely puberulous to
glabrous beneath; nerves 8-11 pairs. Panicles
densely, shortly puberulous; branches reduced,
not forked, 1-6 mm. Petals obtuse to truncate,
5-7 by 12%4-2mm, claw slender, 24-1 mm.
Pistil mainly apically, sparsely strigose. Fruit
8-10 mm long. Seed 714-8 by 3-4 mm.
Distr. Malesia: East coast of the Malay
Peninsula.
Vern. Bélimbing cherchek, b. hutan, gériji,
médang, sétundok.
Uses. Roots for poulticing wounds externally
(BuRKILL & Hanif SFN 17611).
c. var. hirsuta VeLDKAMp, Blumea 15 (1967) 535,
f. 3e-g.
Twigs ferrugineous-tomentose, patchily gla-
brescent. Leaves beneath pale- to rusty-pubescent,
FLORA MALESIANA
[ser. I, vole
above sparsely and silky so, mainly at base and
along the midrib; nerves 5-8 pairs. Panicle rusty-
pubescent, branches reduced, stout, 1-3 mm.
Petals obtuse, sometimes notched; claw 14-1 mm.
Pistil densely appressed-strigose. Fruit 9-13 mm
long. Seed 9 by 4mm.
Distr. Malesia: Malay Peninsula (Johore).
Vern. Bélimbing bési, b. burong, b. hutan, résak
rambai daun.
5. Sarcotheca glauca (Hook. f.) HALL. f. Med.
Rijksherb. Leiden 1 (1911) 2; KNutnH, Pfl. R.
Heft 95 (1930) 421, f. 28f-n.; ANDERSON, Gard.
Bull. Sing. 20 (1963) 162 (ecol.); VELDKAMP,
Blumea 15 (1967) 535, f. 4a—c. — Connaropsis
glauca Hook. f. Trans. Linn. Soc. 23 (1860) 166.
— Fig. 7a-h.
Shrub or tree, up to 21 m, 30cm 9; no buttresses.
Leaves unifoliolate, 444-11 by 134-434 cm (on
sterile twigs larger, up to 1314 by 614 cm), elliptic
to oblong, acute to acuminate, base obtusely cu-
neate to emarginate, glabrous; beneath whitish
green, dull, subglaucous; nerves 5-7 pairs;
petiole 10-27 by 14-1 mm, petiolule 3-6 by 44-114
mm. Panicles 1-2 together, erect, 114-13 cm,
rusty-puberulous; branches up to 10 mm, usually
many-flowered, rather closely placed. Pedicels
pectinately clustered, lower joint 2-6 mm, upper
14-14 mm. Calyx rusty-puberulous outside, gla-
brescent in fruit except for margins and base,
2-3 mm high, crimson when dry. Sepals 134-3 by
%4-14%4 mm, ovate to oblong. Petals red, darker
at apex, 334-6 by 1-2 mm, oblong to lanceolate,
sometimes obovate. Filaments in SF 2-234 and
214-374 mm, in LF 144-1144 and 114-134 mm.
Pistil in SF 1-114 mm, in LF 314-314 mm, ap-
pressed rusty-strigose; styles in SF 14-34 mm, in
LF 214-3 mm; ovary 14-1 by 14-24 mm, ellipsoid.
Fruit bright pink to dark red, 8-12 by 5-11 mm,
subglobose to ellipsoid with obtuse to rounded
apex, glabrescent; rimae open and conspicuous,
inside lighter and papillose. Seeds 6 by 314 mm;
testa shiny, smooth; cotyledons 5 by 214 mm,
elliptic, radicle 1 mm.
Distr. Malesia:
Brunei, Sabah).
Ecol. Rare to very rare in undisturbed forest,
heath forest on well-drained humus podsols,
ground-water podsols, peaty ‘Hochmoors’ and
sand covered clay on terraces and sand stone. Oc-
casionally in secondary forest (BRUNIG).
Vern. Arémajuh, Dajak, asam daham, barus,
bélimbing, b. daham, gitan gizu, kandis daham,
Brunei, médang, piang, Iban, rangkas-rangkas,
Dusun Kinabatangan, ségot, Baju, ftampusi,
temposi(s), Kedayan.
The vernacular epithet daham might point at the
use of this plant against coughing.
Notes. This species is closely allied to S. ru-
brinervis HALL. f. and differs in having a less
puberulous, when dry dark red to crimson calyx,
a short compact panicle with very reduced primary
branches, while the way of branching in the cyme
is indistinct. The pistil in LF is 314-314 mm long,
the filaments in SF 2-234 and 214-324 mm. The
NW. Borneo (Sarawak,
cotyledons are lanceolate.
6. Sarcotheca monophylla (PLANCH. ex Hook. f.)
HALL. f. Med. Rijksherb. Leiden 1 (1911) 2;
KnutTH, Pfl. R. Heft 95 (1930) 422, f. 27; VELD-
KAMP, Blumea 15 (1967) 536, f. 2g—h. — Connarop-
sis monophylla PLANCH. ex Hook. f. Trans. Linn.
Soc. 23 (1860) 166; EpGEw. & Hook. f. FI. Br.
Ind. 1 (1874) 440; Kina, J. As. Soc. Beng. 62, ii
(1893) 200; RipL. Fl. Mal. Pen. 1 (1922) 332, f.
34. — Dapania monophylla KNuTH, Bot. Jahrb.
50 (1914) 648.
Shrub or tree, up to 30m, 38cmg@, bole oc-
casionally buttressed and crooked. Leaves uni-
foliolate, 4-10 by 114-314(-7) cm, elliptic to
oblong, rarely suborbicular, acuminate to cuspi-
date, base rounded to cuneate, glabrous, dark
when dry, subglaucous beneath; nerves 5-7
pairs; petiole 6-31 by 14-34 mm, petiolule 3 by
2-1 mm. Panicles 1-2 together, erect, 1-914 cm,
rusty-puberulous, -+- dense; branches up to 4 mm
long. Pedicels pectinately clustered, upper joint up
to 14 mm, Jower 1-2 mm. Calyx 134-2 mm high,
outside rusty-puberulous, apically less so, red-
dish-brown when dry. Sepals 114-2 by 1-114 mm,
ovate to rectangular, acute to emarginate. Petals
deep red, 3—5 by 34-114 mm, lanceolate, emar-
ginate. Filaments in LF 1-114 and 1144-114 mm,
in SF 114 and 2mm. Pistil densely appressed
rusty-strigose, in LF 3-314, in SF 1 mm;; styles in
LF 2-214 mm, in SF 44 mm long; ovary 14-1 by
14-24 mm, ellipsoid. Fruit up to 13 by 13 mm,
pale to bright red, subglobose to ovoid, rounded
to acute; rimae conspicuous, open, lighter and
papillose inside. Seeds 714 by 3 mm; testa smooth;
cotyledons 4 by 214 mm, radicle 1144 by 14 mm.
Distr. Malesia: W. and Central Malay Pen-
insula.
Ecol. Secondary forest, open jungle on flat to
undulating land on rich soil at low altitude.
Vern. (Asam-)pupoy, bélimbing akar, b.
bésih, b. bulat, b. burong, b. hutan, 6. keris,
b. k(é)ra, b. pénjuru, b. pipit (pépit, pipet), bibit
(Sakai), kilat pivit, sétundok.
Note. The type (CUMING 2324) came from
Malaya, not from the Philippines as KNUTH
supposed.
7. Sarcotheca celebica VELDKAMP, Blumea 15
(1967) 537, f. 4f-m.
Shrub or tree, up to 35m, 17cm g. Leaves
unifoliolate, (314—)6-1314 by 2-5 cm, elliptic to
lanceolate, acute-cuneate, base cuneate to +
truncate, above with prominent, rather coarse
venation, glabrous, subglaucous, sparsely pu-
berulous; nerves 5—6 pairs; petiole 6-21 by 14-1
mm, petiolule 2—5 by 14-114 mm, sparsely strigose.
Panicles 1-2 together, 1-7 cm, erect, slender,
rusty-puberulous; branches up to 2(-7) mm,
distant. Pedicels few, close together with equal
upper and lower joints, up to 144mm. Calyx
2-214 mm high, rusty-puberulous outside, red-
dish-brown when dry. Sepals 114-2 by 1-114 mm,
ovate to rectangular, acute to truncate. Petals
red, 4-414 by 1-114 mm, lanceolate, rounded to
1971]
OXALIDACEAE (Veldkamp)
173
retuse. Filaments in SF 1-2 and 2-214 mm, in
LF 34 and 1mm. Pistil appressed-strigose, in
SF 1 mm, in LF 2!4 mm; styles in SF 14 mn, in
LF 2mm; ovary %4-%4 by %4-34 mm, subglo-
bose to ellipsoid. Fruit 8-13 by 6-9 mm, ellipsoid
rounded to acutish, glabrescent, red; rimae open
from the top to below the middle, + conspicuous,
lighter and papillose inside. Seeds 4-5 by 3 mm;
testa slightly transversely rugose to + smooth;
cotyledons 3 by 234 mm, radicle 1 mm.
Distr. Malesia: Central Celebes (Malili)
and Kabaéna I. (S of SE. Celebes).
Ecol. Rather open country, primary forest on
stony to clayish, flat to undulating land at low
altitude.
Vern. Ko(e)ngilu ((mo-)puté), To Bela, To
Padoé, To Tambeé.
Note. This new species is best distinguished by
its lax, narrow panicles with thin rachis and the
coarser and more prominent venation on the
upper side of the leaf as compared with its nearest
allies S. glauca and S. monophylla.
8. Sarcotheca ferruginea MerRR. Pap. Mich. Ac.
Sc. 19 (1933) 160, fig.; VELDKAMP, Blumea 15
(1967) 538, f. 3h-i.
Tree. Twigs velvety rusty-pubescent, later
patchily glabrescent and dark. Leaves shiny golden
pubescent when young, upper surface soon gla-
brous or with a few appressed, pale hairs at the
basis and along the midrib, beneath densely
ferrugineous-pubescent, not glaucous, (3!4—-)5-
114% by (l-)2-34% cm, oblong to oblanceolate,
acute to caudate, base broadly cuneate to rounded,
margins usually paler; nerves 5—7 pairs; petiole
4-614 by 14-1 mm, petiolule 2-4 by 34-114 mm,
rusty-velvety, glabrescent. Panicles 1—2 together,
short, slender, up to 714 cm, velvety; branches
distant, 14-2 mm. Pedicels with unequal lower
joint, one per cyme elongated, 3-4 mm, others
c. 1mm; upper joint 14-1 mm. Calyx 2-334 mm
high, rusty-puberulous outside. Sepals 134-314
by 14-1 mm, ovate-lanceolate, acute. Petals red,
4-6 by 114-2 mm, obovate-elliptic to -lanceolate,
rounded to obtuse. Filaments in LF 1 and 114
mm. Pistil in LF 3-314 mm, appressed-strigose;
styles in LF 2-214 mm; ovary 1 by 1 mm, sub-
globose. Fruit 7-14 by 6-12 mm, ovoid to sub-
globose, rounded, somewhat shiny, reddish;
rimae inconspicuous, not glandular or ligher inside.
Seeds 7 by 4'4 mm; testa transversely rugose;
cotyledons 5 by 3 mm, radicle 1 mm.
Distr. Malesia: N. Sumatra (West and West
Coast Res.), apparently local and rare.
Vern. Kaju kandis.
Note. I have not seen the short-styled form.
9. Sarcotheca rubrinervis HALL. f. Beih. Bot.
Centralbl. 34, ii (1917) 29; Knuth, Pfl. R. Heft 95
(1930) 425; VetpKamp, Blumea 15 (1967) 539,
.f. 4d-e. — S. oblongifolia Merr. Univ. Cal.
Publ. Bot. 15 (1929) 111; KNuru, Pfl. R. Heft 95
(1930) 421.
Shrub or tree, up to 19m, bole up to 10m,
30 cm @. Leaves unifoliolate, 5-18'4 by 144-7 cm,
oblong to lanceolate, glabrous, acute to caudate,
base obtuse to rounded, beneath not or slightly
glaucous; nerves 4-11 pairs, often with reddish
tinge; petiole 6-20 by 1-2 mm, petiolule 24 by
1-2 mm. Panicles 1-2 together, elongated, lax,
pendulous, rusty-puberulous, 7-38 14 cm; branches
elongated, slender, patent, -+ monochasially
branched, up to 8(-21) mm. Pedicels with un-
equal lower joint, usually one per cyme elongated
to 5 mm, others up to 3 mm; upper joint 14-1 mm.
Calyx 2-3 mm high, rusty-puberulous, at least at
base. Sepals 124-234 by 1-144 mm, ovate to
lanceolate, acute to emarginate. Petals 334-514
by 1-114 mm, lanceolate, rounded to emarginate,
pale red to red, occasionally white, apex usually
darker, claw 24-1 mm. Filaments in SF 124-2
and 214-3 mm, in LF !4—24 and 1-144 mm long.
Pistil in SF1-1%mm, in LF2'4-23/; mm;
styles in SF 44 mm, in LF 134-14/, mm; appressed-
puberulous; ovary \4-4/, by 14-24 mm, sub-
globose, densely appressed-strigose. Fruit 6-10 by
5-10 mm, subglobose, rounded, glabrescent, glau-
cous to glossy, pink to bright red; rimae not al-
ways conspicuous, lighter and papillose inside.
Seeds up to 634 by 5mm; testa smooth to +
transversely rugose; cotyledons elliptic, 434 by
1144 mn, radicle 1 by ?/; mm.
Distr. Malesia: E. Borneo (from Tawao to
Balikpapan).
Ecol. Primary and secondary forest near
rivers on loam, flat to undulating land.
Vern. Asém-asém, Dusun Kinabatangan, iba
talon, Bajau, ira prumpuan, Suluk, kajo badjuk,
Kajam-Dajak, kandis daham, Brunei, l/ampyos,
Dusun Penompang, pinggoh, pinguh, Klabakan,
pingo, Tidon, ténggoh, Kutai.
Note. Sometimes difficult to distinguish from
S. glauca (see there).
10. Sarcotheca ochracea HALL. f. Beih. Bot.
Centralbl. 34, ii (1917) 28; KNuTu, Pfl. R. Heft 95
(1930) 424; VELDKAMP, Blumea 15 (1967) 541,
f. Sc—e.
Treelet, up to 9m, 10cm o. Twigs velvety,
glabrescent. Leaves unifoliolate, 744-2344 by
314-1014 cm, elliptic to oblong, abruptly acu-
minate to cuspidate, base obtuse to truncate,
above glabrous, beneath rusty-pubescent to
-velvety, sometimes subglaucous; nerves 6-11 pairs;
petiole 5-24 by 114-3 mm, petiolule 3-7 by
114-3 mm. Panicles 1-2 together, 7-70 cm, erect
to pendulous, rusty-velvety, compact to elongated.
Branches very short to elongated, often distant,
pectinately branched, many-flowered, patent to
recurved, up to 30 mm long, rusty-velvety. Lower
joint of pedicel 4-8 mm, upper 14-1 mm. Calyx
(24%4-)3-3'44 mm high, rusty-puberulous outside.
Sepals suborbicular to obovate, rounded to retuse,
244-3 by 1%4-3 mm. Petels crimson, apically
darker, (4144—)6-8 by (1-)1'44-144 mm, obovate-
lanceolate to lanceolate, obtuse to rounded. Fila-
ments in SF 214 and 3-344 mm, in LF 114-2
and 2-244 mm. Pistil densely rusty-strigose, in
SF 134-2, in LF 3-4 mm; styles red, in SF 4-%
mm, in LF 2-2!4 mm long; ovary 1-1'4 by | mm,
174
subglobose. Fruit 8-15 by 6-15 mm, subglobose to
oblong, rounded, glabrescent, bright red; rimae
conspicuous, lighter and/or papillose inside. Seed
1-2 per fruit, 544-7 by 314-414 mn; testa trans-
versely rugose. Mature embryo not seen.
Distr. Malesia: Borneo (Sarawak: Bintulu).
Ecol. Along streams in forest on clay.
Vern. [kor mata, Iban, pechi mata.
11. Sarcotheca macrophylla BLUME, Mus. Bot.
Lugd. Bat. 1 (1850) 242; Mia. Fl. Arch. Ind. IIl.
(1870) 70, t. 30; BAmLLon, Adansonia 10 (1873)
364; Hist. Pl. 5 (1874) 26, 47; HALL. f. Beih. Bot.
Centralbl. 34, ii (1917) 29; KNutn, Pfl. R. Heft 95
(1930) 424, pro spec. Born.; VELDKAMP, Blumea 15
(1967) 541, f. S5a—b. — Roucheria macrophylla
Mia. FI. Ind. Bat. 1, 2 (1858) 136; Suppl. 1 (1860)
162.
Shrub or tree, up to 15m, 10cm@. Leaves
unifoliolate, (S5—)16-28 by 6-10cm, oblong to
oblanceolate, margins + parallel, abruptly acu-
minate to cuspidate, base truncate to emarginate,
subcoriaceous, above glabrous, beneath pu-
berulous on the veins, sometimes subglaucous;
nerves 6—13 pairs; petiole 5—12(—25) by 114-3 mm,
petiolule 3-9 by 1-3 mm. Panicles 1-4 together,
slender, lax, 12-85cm, pendulous, brown-pu-
berulous; branches elongated, 6—25(—50) mm,
often flattened. Lower joint of pedicel up to 2 mm,
upper 44-1 mm. Calyx 114-3 mm high, ferru-
gineous-puberulous outside. Sepals 1144-214 by
1-134 mm, obovate to ovate, rounded to obtuse.
Petals dark red, 3144-5 by 1-134 mm, obovate-
oblong to -lanceolate, obtuse to emarginate,
claw 14-14 mm. Filaments in SF 1-2 and 134-3
mm, in LF 34-14, mm and 114-134 mm. Pistil
appressed-strigose, in LF 2-3 mm, in SF 1144-134
mm; styles in SF 4%4-7%4mm, in LF 1-2 mm;
ovary 74-1 by 14-34 mm, subglobose to ellipsoid.
Fruit 6-11 by 5-8 mm, subglobose, rounded, gla-
brescent, shiny, dark red; rimae inside lighter,
papillose, conspicuous. Seeds up to 9 by 4mm;
testa transversely rugose; cotyledons 4 by 214 mm,
radicle 1 mm.
Distr. Malesia: Borneo (Indonesian Borneo;
Sarawak: Marop).
Ecol. Primary and secondary forest on
sand.
Vern. Bélimbing manik, Bakumpai-Dajak, kaju
kim, krumbai mérah, mim, pengu, Dusun, ram(b)-
ajan.
Notes. BLumE cited this species also to occur
in Sumatra, but this seems to be erroneous.
It has been confused with Connaropsis macro-
pPhylla Kinc = S. glomerula.
FLORA MALESIANA
[ser. I, vol: 7?
Dubious
Sarcotheca philippica (F.-ViILL.) HALL. f. Med.
Rijksherb. Leiden 1 (1910) 2. — Connaropsis
Philippica F.-VILL. Novis. App. (1880) 33;
Merr. Sp. Blanc. (1918) 19, 195; KNutn, Pfi.
R. Heft 95 (1930) 417.
Subscandent. Innovations dark purple. Leaves
imparipinnate, 1—4-jugate; leaflets 6-10 by 2-3
cm, oblique-ovate, acute, subcoriaceous, pubes-
cent when young, later glabrous. Panicle axillary
and terminal, shorter than the subtending leaf.
Sepals 2-4 mm long. Petals twice as long, connate
at base when young, later free. Filaments alter-
natingly shorter (‘alterna breviora’), often without
anthers; styles 5, erect, adpressed; stigmas oblong,
apiculate; ovary ovate, pilose, 5-locular, ovules 2
per cell. Fruit 2-3 by 14-1 cm, orange-red, pilose,
5-angular, 5-locular. Seed 1 per cell, ovate, with
white aril (translated from type description).
Distr. Malesia: Philippines (Bugney near
Igbaras, Iloilo).
Vern. Balabangquilin, malabangquilin.
Note. MERRILL noted that he has seen no rep-
resentative of this genus in the Philippines and
none is known to me. Although he is very critical
of F.-VILLAR’s work (cf. I.c. 14-18), he thinks
it to be ‘... apparently a true... Sarcotheca...’.
KNUTH referred it to Averrhoa carambola L., with
which the description agrees for the greater part,
differing, however, by the subscandent habit, the
subcoriaceous leaflets, the 2 ovules per cell, and the
rather small, orange-red, pilose fruit. F.-VILLAR
stated he has seen the plant alive, so presumably
his remarks pertaining to habit and fruit arecorrect;
the more so as in his work this species follows the
description of Averrhoa carambola. As long as no
representative is found the status of this species
will remain in doubt.
Excluded
Connaropsis rubescens RiDL. J. Bot. 62 (1924)
295 = Rourea minor (GAERTN.) LEENHOUTS, Blu-
mea 12 (1963) 20 (Connaraceae).
Sarcotheca paniculata Riv. Trans. Linn. Soc.
III, 2 (1893) 282 = Rourea minor (GAERTN.)
LEENHOuTS, FI. Mal. I, 5 (1958) 515a (Connara-
cede).
Sarcotheca pinnata MERR. J. Str. Br. R. As. Soc.
n. 86 (1922) 314 = Rourea pinnata (MERR.)
VELDKAMP, Blumea 15 (1967) 543 (Connaraceae).
Sarcotheca varians (CRAIB) KNuTH, Pfi. R.
Heft 95 (1930) 425. — Connaropsis varians
CraiB, Kew Bull. (1926) 158 = Rourea minor
(GAERTN.) LEENHOUTS, FI. Mal. I, 5 (1958) 515b
(Connaraceae).
5. AVERRHOA
LINNE, Gen. Pl. ed. 5 (1754) 196; Sp. Pl. (1753) 428; ENDL. Gen. Pl. (1839) 1173,
incl. sect. Bilimbi et sect. Carambola ENDL.; KNUTH, Pfl. R. Heft 95 (1930) 417;
BAck.& BAKH. f. Fl. Java 1 (1963) 247. — Averrhoaceae HUTCHINSON, Fam. FI.
Pl. ed. 2, 1 (1959) 356. — Fig. 9.
1971] OXALIDACEAE (Veldkamp) 175
Evergreen shrubs or trees. Innovations pubescent, glabrescent. Leaves spirally
arranged to terminally clustered, estipulate, imparipinnate, herbaceous; leaflets
entire, subopposite, subsessile, terminal largest. Panicles axillary or cauliflorous.
Flowers in scattered, stalked, loose cymes, heterodi- or -tristylous. Bracts small,
caducous. Sepa/s quincuncial or paratact, shortly connate at base. Petals contort
or paratact, free or coherent above the claw, creamy to dark red with white
markings. Ovary appressed-strigose; ovules (2—)3-—7 per cell, pendulous. Fruit
large, yellowish green and semitransparent in vivo, red when dry; rimae small,
reduced to absent (?). Seeds elliptic, flattened; aril present in one species, attached
to the entire adaxial raphe, bilabiate, enveloping the seed, fleshy, with oily drops;
endosperm present; cotyledons 6-8 times as long as the oblique and stout radicle.
Distr. 2 spp., probably native in Malesia, cultivated pantropically at low altitude for their fruit trees,
often escaping.
It is often assumed that Averrhoa is possibly of American origin. Both species are mentioned in pre-
Linnean literature always from Indo-Malesia (BAUHIN, 1623, being the oldest). The closest relatives are
Malesian-Malagasian. Though I can not forward proof, for or against, an Indo-Malesian origin seems
most probable.
Ecol. Irritability of the leaves has been noted as early as 1785 by Bruce (Phil. Trans. 356), while
DARWIN made a detailed study of it (A. bilimbi L.) in his ‘The Movements of Plants’ (1880) 330, 447.
Uses. Mainly for the fruit; see under the species.
Syst. The two species are remarkably different in important characters which are almost of the same
value as those separating the allied genera Sarcotheca and Dapania as appears from the key to the species.
One is tempted to raise them to the status of sections, as ENDLICHER proposed (Gen. Pl. 1839, 1173).
At the same time this shows the intimate relationship of the three genera. This defeats HUTCHINSON’s
rigorous attempt to assign Averrhoa to a separate family Averrhoaceae, even accommodating it to
another order (Fam. FI. Pl. ed. 2, 1, 1959, 356). The reason for this rests on the supposed presence of a
disk and horizontal ovules; the first I can not find and the ovules are pendulous.
KEY TO THE SPECIES
1. Leaves 3—-6-jugate. Panicles axillary, rarely some ramiflorous. Petals up to 8 mm, minutely puberulous
inside, coherent above the claw. Shorter stamens without anther. Ovules 3-5 per cell. Fruit stellate in
0 ONY rr ee re MEPL Ie ES by op 1. A. carambola
1. Leaves 7—19-jugate. Panicles cauliflorous, rarely some axillary. Petals 10-20 mm, glabrous inside,
not coherent above the claw. All stamens antheriferous. Ovules 4~7 per cell. Fruit terete, slightly
ODER CARITIINIO, ww te eae en ee en ee 2. A. bilimbi
1. Averrhoa carambola LInngé, Sp. Pl. (1753) 428; = stellatum Rumpu. Herb. Amb. 1 (1750) 115, f.
Cav. Diss. 7 (1789) 374, f. 220; DC. Prod. 1
(1824) 689; BL. Bijdr. (1825) 242; W. & A.
Prod. (1834) 141; BLANco, FI. Filip. (1837) 391;
Mia. Fl. Ind. Bat. 1, 1 (1859) 133; EpGew. &
Hook. f. Fl. Br. Ind. 1 (1874) 439; ProGEL,
Fl. Bras. 12, 2 (1877) 520, incl. var. angustisepala;
Trimen, J. Linn. Soc. Bot. 24 (1887) 129 (typi-
fication); NINGRAT, Teysmannia 3 (1892) 754;
K. & VY. Bijdr. 9 (1903) 106; Back., FI. Bat.
(1907) 229; Koorp. Atlas 4 (1918) 603; Merr.
Sp. Blanc. (1918) 194; Ripv. FI. Mal. Pen. 1 (1922)
332; Heyne, Nutt. Pl. (1927) 853; KNutu, Pfl.
R. Heft 95 (1930) 417; Ocuse & BAKH. Vrucht.
(1931) 91, tab. col.; Burk. Dict. (1935) 271;
Quis. Med. Pl. Philip. (1951) 439; Back. &
- BAKH. f. Fl. Java 1 (1963) 247; VeLpK., FI. Thail. 2
(1970) 21. — A. acutangula Stokes, Bot. Mat.
Med. 2 (1812) 543. — A. pentandra BLANCO,
Fl. Filip. (1837) 392. — Tamara tonga RuHEEDE,
Hort. Mal. 3 (1682) 51, f. 43, 44. — Prunum
35. — Fig. 9.
Tree, up to 14 m-by 30cm. Branches often
drooping. Innovations yellowish to reddish pu-
bescent, glabrescent. Leaves 3-6-jugate, not
crowded terminally; rachis up to 20 cm; leaflets
up to 10 by 4 cm, variable in shape, lowest not
reflexed, acute to acuminate, glaucous beneath;
nerves 4-10 pairs. Panicles axillary, rarely rami-
florous, 1-few together, ascending, up to 7cm
long. Flowers heterodistylous (LF, MF), said
to be scentless, melliferous. Pedicels 3-6 mm,
articulated 1-2 mm below the flower. Sepals up
to 4 by 2mm, bright red, yellowish brown with
pale margins when dry, obovate-oblong to trian-
gular, acute to obliquely emarginate, subglabrous.
Petals up to 8 by 2mm, obovate to lanceolate,
coherent, claw c. 144 mm, glabrous, but inside
with minute septate-glandular hairs. Shorter sta-
mens ananthiferous, filaments subulate with often
swollen base, up to 144 mm, longer ones in LF
176
FLORA MALESIANA
[ser. I, vol. 7}
Fig. 9. Averrhoa carambola L. (Courtesy T. B. WORTHINGTON).
up to 3mm, in MF up to 4144 mm. Ovary elliptic,
up to 214 mm long, appressed-puberulous, mainly
on the ribs; styles in LF up to 2 mm, in MF 14 mm;
ovules 3-5 per cell. Fruit up to 1214 by 6cm,
ovoid to ellipsoid, stellate in a, lobed at both ends,
apically with minute rimae on the ribs. Seeds up
to 10, 12 by 5 mm; aril fleshy, bilabiate, enclosing
the seed, lobed at base; cotyledons up to 614 by
314 mm, elliptic.
Distr. Native country unknown, possibly Cen-
tral and East Java, but escaping freely, often a
relict of former cultivation.
Ecol. At low altitude, up to 1000 m, along
rivers, in ravines, primary and secondary forest.
Dispersal. Fruits eaten by bats, mice and Calotes
spp. (Agamidae). Cf. NINGRAT (1892).
Vern. Variations on bélimbing. Sumatra:
asom djorbing, Bat., balimbing manis, b. segi,
b. sajur, b. batu, kémbola, b. bési, b. kéris, Mal.,
kembang bua, kaping, Palemb., b. manih; Java:
balingbing amis, tjalingtjing amis, S, b. légi, b.
lénger, b. lingir, b. manes, Mad., libi melai,
Sawu; Borneo: b. péségi, b. pénjura, Kuching,
b. amamas, Sabah; Celebes: /umpias mamis,
Bentenan, rumpiasa, Bantik, lompiat morominit,
Mongodow, dumpias tariis, lopias emé, Tonsa-
wang, lumpias tombal, Tontembuan, Tonsawang,
leémbétué lombiata, Goronta, lombituko gula,
Buol, takulé, Barée, bainang sulapa, Makass.,
kulirang taning, pulirang taning, N. Salajar,
baknil pasaki; Sepa, S. Ceram, ifél emroro,
Masareta, Buru, malibi totofuo, Weda, S. Halm.,
balibi totofuko, Galelas, Tobelor, totofuko, t. lémo,
Ternate, tufuo, Tidore; Philip.: daligdn, Ilk.,
dalihan, galurdn, \bn., galangan, P. Bis., gardhan,
Bis., malimbin, S. L. Bis., sirinate, Ting.; E. New
Guinea: ibeid, Tehid, painayangara, Tumbunke.
Uses. The fruit is eaten fresh or pickled.
Leaves and fruits are said to be good against fevers,
aphtha, angina, chickenpox, ringworm, headaches,
1971]
OXALIDACEAE (Veldkamp)
177
scurvy, dysentery, piles and affections of skin and
eyes; they excite the appetite, but may cause
vomiting. It is antiphlogistic and a sialogogue. The
fruit removes stains from laundry, hands and
weapons. See NINGRAT (1893), KoorDERS & VaA-
LETON (1903), HEYNE (1927), OcHsE (1931),
BuRKILL (1935), and QUISUMBING (1951).
Phytochem. De PERALTA (Philip. Agr. 17,
1928, 334) has shown the presence of HCN in
leaves, roots and stems; BATE-SMITH (J. Linn.
Soc. Lond. Bot. 58, 1962, 134) the presence of
cyanidin, p-cumaric acid and traces of sinapic
acid, and (ibid. 55, 1957, 687) tannin, leuco-
anthocyanin and leucocyanidin.
Syst. This species is widely cultivated for a long
time and many cultivars are extant (cf. NINGRAT,
1893), differing in taste and size of the fruit. A
f. acida and f. dulcis have been described by Koor-
DERS & WVALETON, and material bearing these
names has been examined, but no characters could
be found to distinguish them. PROGEL’s var.
angustisepala from S. America seems not to war-
rant distinction, as the shape and pubescence of
the sepals in Malesia is very variable. This variety
has possibly been the reason for an assumed Amer-
ican origin of the genus time and again found in
literature.
2. Averrhoa bilimbi LiInNE, Sp. Pl. (1753) 428;
Cav. Diss. 7 (1789) 373, f. 219; DC. Prod. 1
(1824) 689; Roxs. Fl. Ind. ed. Carey 2 (1832)
451; W. & A. Prod. (1834) 142; BLANco, FI.
Filip. (1837) 391; ENpL. Gen. Pl. (1839) 1173;
Mig. FI. Ind. Bat. 1, 1 (1859) 139; EpGew. &
Hook. f. Fl. Br. Ind. 1 (1874) 439; PrRoGEL,
Fl. Bras. 12, 2 (1877) 520; Trimen, J. Linn.
Soc. Bot. 24 (1887) 129 (typification); K. & V.
Bijdr. 9 (1903) 111; Koorp. Atlas 4 (1918) 601,
602; Merr. Sp. Blanc. (1918) 195; En. Born.
(1921) 311; Ripx. FI. Mal. Pen. 1 (1922) 331;
Heyne, Nutt. Pl. (1927) 852; KNutn, Pfl. R.
Heft 95 (1930) 418, incl. f. papuana KNUTH;
Ocuse & BAKH. Ind. Groenten (1931) 546;
BENTHALL, Trees of Calcutta (1933) 70; Burk.
Dict. (1935) 270; SorGDRAGER, Pharm. Tijd. 1
(1941) repr. 4 pp. (anat., uses); Quis. Med.
Pl. Philip. (1951) 438; Back. & BAKu. f. Fl. Java
1 (1963) 247; VeLpk., Fl. Thail. 2 (1970) 22. —
A. obtusangula Stokes, Bot. Mat. Med. 2 (1812)
542. — Bilimbi Rueepe, Hort. Mal. 3 (1682) 55,
f. 45, 46. — Blinbingun teres, Blimbing bula
Rumpu. Herb. Amb. | (1750) 118.
Shrub or tree, up to 15m, 30cmg. Branches
erect. Innovations and green parts long-persisting
pale yellowish to rusty velvety. Leaves 7-19-
jugate, usually terminally tufted; rachis 17—57 cm;
leaflets up to 12 by 4cm, variable in pubescence,
size and shape, lower reflexed, acute to acumi-
nate, not glaucous beneath; nerves 6-14 pairs.
Panicles cauliflorous on tubercles, nearly down to
ground-level, fasciculate and pendulous up to
20 cm, rarely axillary, solitary and erect. Flowers
heterotristylous, usually MF. Pedicels 4-17 mm,
articulated near or below the middle. Sepals 3-8
by 1'4-3 mm, yellowish red to purplish, sparsely
appressed-puberulous outside mainly at base,
inside glabrous or subglabrous, elliptic to lan-
ceolate or spathulate, acute to rounded. Petals
free, 10-20 by 34mm, lanceolate-spathulate,
inside glabrous, claw 3-6 mm. Stamens all fertile,
in SF 314-4 and 7 mm, in MF 2-5 and 9-12 mm,
in LF 5 and 7!4 mm, bases not thickened. Ovary
densely appressed pale strigose and with short,
septate-glandular hairs, in SF 2-214 by 1 mm, in
MF 3-4 by 1-1'44 mm, in LF 4 by 1 mm, elliptic;
styles in SF 14-34 mm, in MF 2-4 mm, in LF
614-9 mm; ovules 4-7 per cell. Fruit terete-
obtusangular, up to 10 by 5 cm, elliptic to obovate,
obtuse, rimae present (?), base tapering. Seeds
up to 14, 6-7 by 4-6 mm, exarillate; cotyledons 4-6
by 3144-5 mm, orbicular, cordate.
Distr. Native country unknown, widely cul-
tivated, escaping freely, often a relict of former
cultivation, for instance in East Malesia.
Ecol. Along rivers, lowland primary and secon-
dary forest.
Vern. Bélimbing and many variations; Sumatra:
liméng, séliméng, thliméng, Atjeh, asom, Batak;
Mal. Pen.: 6. asam, b. bési, b. botol, b. buloh,
b. kéris, b. masam; Borneo: béliwit, Dajak Busang,
iba, Bajan, blimbing puchung, Kedayan; Java:
tjalingtjing, t. wulét, S; limbi, Bima; libi, Sawu;
bélérang, Sangi; Celebes: /Jumpias, Bentenan, Po-
nosakan, Tombulu, Tontembuan, rumpéasa dureng,
Bantik, /ompiat litod, Mongodow, dumpias, d.
tuwama, Tonsawang, wuli(n)dan, Tontembuan,
lémbétué, Gorontalo, lombituko, Buol, sangguléra,
Parigi, tangkurera, Barée, bainang, Makassar,
kulirang, pulirang, N. Salajar, tjaléné, Bugin.;
niniloe daé lok, Roti, kerbol, Timor, baknil, Kai,
ahuréla, Atamano, W. Ceram, Aauréla, Amahai,
S. Ceram, taulela, Nuaulu, takuréla, Ambon,
tahuréla, Sapurua, Ulias, taprera, Kajeli, Buru,
ifél milo, Masarete, Buru, malibi, Weda, S.
Halm., balibi, N. Halm., Ternate; Philip.: iba,
Tag., Sul., P. Bis., ibag, Mbo., kalamids, kala-
nuas, kamids, kolonanas, kolonauas, Tag., kalin-
giua, Bis., kiling-iba, Bik., ibe, Yak., pids, IIk.,
puis, Ig.; New Guinea: miri-miri, Kapur, utéké,
Mimika & Atuka R.., olloloh, Sentani.
Uses. In the Malay Peninsula the leaves are
used as a paste against itches, eaten against syphi-
lis and a decoction is drunk after childbirth. In
Java the leaves are used against mumps, rheu-
matism and pimples, against piles; a decoction of
the flowers is used against cough and thrush. The
fruit juice is generally used against fevers, scurvy,
beri-beri, biliousness, coughs and piles; it also
removes stains from laundry, hands and weapons.
See Koorpers & VALETON (1903), HEYNE (1927),
Ocuse & BAKHUIZEN (1931), BuRKILL (1935),
and QUISUMBING (1951). Fruits and flowers are
edible but more acid than those of A. carambola.
Syst. The species has always been regarded as
possessing only an MF flower. Short-styled or SF
(Ev_mer 15122) and long-styled or LF (HALLIER
Sf. s.n.) forms occur but very rarely so. It is doubtful
whether f. papuana KNUTH merits distinction;
his definition reads: “In omnibus partibus fere
duplo major’’. Material showing these sizes is
178
found on most islands throughout its range in
Malesia.
Excluded
Averrhoa acida LINNE, Sp. Pl. (1753) 428 =
Phyllanthus acida (L.) SKEELS, Bull. U.S. Dep.
Agr. Bur. Pl. Ind. 148 (1909) 17 (Euphorbiaceae).
Averrhoa frondosa SALISBURY, Prod. (1796) 318
= Phyllanthus acida (L.) SKEELS, Bull. U.S. Dep.
Agr. Bur. PI. Ind. 148 (1909) 17 (Euphorbiaceae).
Averrhoa microphylla TARDIEU-BLoT, Not. Syst.
FLORA MALESIANA
[ser. I, vol. 71]
11 (1943) 133; FI. Gén. I.-C. 1, Suppl. (1945) 547,
f. 66, is according to LEENHOUTS = Rourea
harmandiana PiERRE (Connaraceae).
Averrhoa minima PERROTTET, Mém. Soc. Linn.
Paris 3 (1824) 101. ““Espéce nouvelle venue de la
Chine, qui s’éleve 4 1 métre (3 pieds) au plus
(Mascareigne)”. Type-material could not be
found in the Paris Herbarium. Probably it does
not belong to Oxalidaceae, but to Euphorbiaceae?
Averrhoa sinica HANCE in Walp. Ann. 2 (1852)
241 = Rourea minor (GAERTN.) LEENHOUTS, FI.
Mal. I, 5 (1958) 514 (Connaraceae).
VIOLACEAE (M. Jacobs, Leyden; D. M. Moore, Reading)
Shrubs, small trees, or lianas, in Malesia evergreen, or herbs. Stipules present.
Leaves in Malesia spirally arranged, sometimes distichous, simple, the margin
often shallowly incised; generally stalked. Inflorescences axillary variously mo-
dified bundles, or racemes, or panicles, sometimes terminal, or flowers solitary
in the leaf axils; bracts small; pedicels often articulated, whether in the lower or in
the upper part; bracteoles, if present, small and in the lower part of the pedicel.
Flowers bisexual or rarely dioecious, actinomorphic or zygomorphic, particularly
in the corolla; the parts often persistent in fruit. Sepa/s 5, the median one adaxial
(posterior), free or occasionally for a small portion connate, often ciliate. Petals 5,
free, generally sessile, the median one abaxial (anterior), often longer and diffe-
rently shaped, the base then mostly with a sac or spur. Androecium often cylindrical,
stamens 5, episepalous; filaments often more or less connate into a tube, in the
Malesian genera with zygomorphic flowers, those near the odd petal with a re-
curved fleshy appendage; anthers introrse, in Malesia nearly always the connective
at the top produced into an approximately triangular membranous appendage
converging with the others, cells sometimes with a small appendage at the top.
Gynoecium superior, sessile, ovary small, subglobose, one-locular, with generally
3 carpels, the median one adaxial, each carpel with a parietal placenta in the
middle bearing l-many anatropous ovules; style straight or, in the zygomorphic
flowers S-shaped with the stigma curved towards the odd petal and club-shaped
with variations. Fruit in Malesia capsular, the carpels thickened to boat-shaped
leathery or woody valves (in the latter eventually the endocarp separated from the
pericarp) which spread and often compress upon dehiscence. Seeds 1-many,
sessile, one to a few mm in size, often with distinct raphe, sometimes with funicular
outgrowths; rich in endosperm; embryo straight.
Distribution. A pantropical family; only Viola is cold-loving. Hybanthus extends into the subtropics ‘
so does Melicytus (Pacific Plant Areas n. 103, Blumea Suppl. 5, 1966) in Polynesia and New Zealand.
Hymenanthera (congeneric with the former? /.c. n. 104) is temperate in SE. Australia and New Zealand.
Number of genera 16, 8 of them American; the largest are Viola, currently credited with c. 400 spp.,
Rinorea with c. 200, Hybanthus with perhaps 70, and there are about 50 more in the other genera alto-
gether. Total number of species c. 720, in Malesia 31, two of these introduced.
Ecology. Viola is of temperate origin and occurs in Malesia in the mountains. The other Malesian
genera occur in the lowlands; Rinorea and Agatea in everwet forests, Hybanthus in monsoon regions.
Phytochemistry. Accurate chemical information about Violaceae is scanty notwithstanding the
fact that members of this family are used in popular medicine all over the world. The present summary
of chemical characters, therefore, must be considered as a very preliminary one. Different types of
crystals of oxalate of lime occur commonly. Members of the genera Amphirrhox and Allexis accumulate
aluminium according to CHENEREY (Kew Bull. 1948, 173). Leaves and flowers contain rather large amounts
of acidic mucilage in many instances; usually the mucilage is present in epidermal cells. In some taxa
cells with a yellow or reddish ‘resin-like’ content replace mucilage cells; these latter cells also occur in the
mesophyll! and in the cortex, phloem and pith of stems. The chemical nature of the content of these
idioblasts is not known. However, the fact that leucoanthocyanins are rather widespread in Violaceae
suggests that the yellow to reddish cell contents may represent so-called ‘myriophyllin’ or ‘incluses’.
Personal observation on rhizomes of Viola mirabilis L. confirm this supposition. According to PECKOLT
(Ber. Deut. Pharm. Ges. 7, 1897, 97) fresh leaves of Leonia glycycarpa R. & P. are used for the pre-
paration of a bird-lime in Brazil; this suggests that at least some members of the family may have rubber-
containing idioblasts. Roots of several species of Hybanthus (= Ionidium) store inulin-like fructans
instead of starch; other Violaceae store essentially sugars or starch (KRAUS, Sitz. Ber. Naturf. Ges. Halle,
1879, 6).
Preliminary observations about polyphenolic constituents of leaves (LesreTON-BOUCHEZ, Phyto-
(179)
180 FLORA MALESIANA [ser. I, vol. 71
chemistry 6, 1967, 1601) showed a wide range of compounds; leucoanthocyanins, flavono] and flavones
may be present in various combination ; ellagic acid, however, was observed only in trace amount in
three species of Viola. Saponins seem to be rather common in violaceous plants; they have been de-
monstrated to be present in many species of Viola and in some species of Hymenanthera, Hybanthus, and
Melicytus; none of the saponins has been investigated chemically hitherto.
There is much confusion about alkaloids in literature; roots and rhizomes of species of Hybanthus
and of Viola odorata L., and other species of Viola were used formerly as a substitute for ipecacuanha
root. Some authors claimed to have detected emetin or emetin-like compounds (e.g. viola-emetin;
violin) in such crude drugs. Other authors, however, could not find emetin though some of them isolated
minor amounts of alkaloids (e.g. anchietin, ionidin; compare PECKOLT, /.c.; LINDE, Apoth. Z. 34, 1919,
37). Most probably many Violaceae contain small amounts of alkaloids; species of Anchieta, Hybanthus,
Hymenanthera, Rinorea, and Viola are listed in literature as alkaloid-bearing plants; however, the struc-
tures of violaceous alkaloids are totally unknown at present.
Most chemical work has been performed with European species of Viola. Glycosides of delphinidin and
cyanidin occur constantly in blue and purple flowers; violanin is a delphinidin-derived anthocyanin
acylated by p-cumaric acid. Rutin occurs in flowers and in leaves of several species of Viola. From the
herb of Viola tricolor L. HORHAMMER c.s. (Tetrahedron Letters 1965, 1707) isolated apigenin-6,8-di-C-
glucoside, which was called violanthin. The flavone glycoside linarin is the main flavonoid constituent of
leaves of Viola papilionacea PuRSH (V. PLouvirR, C. R. Ac. Sc. Paris 264 D, 1967, 145).
Violatoside is an arabinoglucoside of methyl salicylate; it occurs in species of the section Melanium
but seems to be lacking in other sections of Viola (PICARD, Bull. Soc. Chim. Biol. 8, 1926, 568). From
roots and rhizomes of Viola odorata L. PAILER and Novotny (Naturwissenschaften 45, 1958, 419)
isolated 0.01-0.02 % of nitropropionic acid.
Concluding this survey it must be stated that a thorough chemotaxonomic discussion of Violaceae
is not yet possible. At present no chemical characters are known which contradict the generally accepted
relationships between Flacourtiaceae and Violaceae. — R. HEGNAUER.
Notes. BENTHAM & HOOKER’s subdivision of the family (Gen. Pl. 1, 1862, 115) was extended and
modified by MELCHIOR in E. & P. Nat. Pfi. Fam. ed. 2, 21 (1925) 346; for an account in English of the
latter’s phylogenetic considerations, see EXELL, J. Bot. 63 (1925) 330-333.
Dr. D. M. Moore’s contribution consists of the genus Viola; he also checked the family description.
Indexes to the examined specimens were published in the series ‘Identification Lists of Malaysian
Specimens’, 1. 27 Rinorea (1966) and n. 28 Viola (1967).
KEY TO THE GENERA
1. Flowers actinomorphic, in axillary bundles or racemes. Seeds not winged. Shrubs or trees.
1. Rinorea
1. Flowers zygomorphic.
2. Woody climbers. Flowers in elongate inflorescences. Seeds winged. ......... 2. Agatea
2. Herbs, sometimes with woody base. Flowers solitary in the leaf axils. Seeds not winged.
3. Leaves (sub)sessile. Pedicels jointed. Seeds ribbed sic ie eT Tepe yah cos 3. Hybanthus
3. Leaves stalked. Pedicels not jointed. Seeds smooth. . . dee ec cetcecuss 6°. 5 Ae
1. RINOREA
AUuBL. Hist. Pl. Guian. Fr. 1 (1775) 235; Taus. in E. & P. Nat. Pfl. Fam. 3, 6
(1895) 329; Metcn. ibid. ed. 2, 21 (1925) 349; Jacoss, Blumea 15 (1967) 127;
TATON, Fl. Congo, Viol. (1969) 2. — Pentaloba Lour. Fl. Cochinch. (1790) 154. —
Medusa Lovwr. I.c. 406. — Alsodeia THouARS, Hist. Vég. Isl. Austr. Afr. (1806) 55;
ENDL. Gen. Pl. (1839) p. 911, n. 5047; B. & H. Gen. Pl. 1 (1862) 118; Bogert.
Handl. 1 (1890) 66. — Vareca Roxs. FI. Ind. ed. Wall. 2 (1824) 445. — Jiirgensia
SPRENG. Syst. 3 (1826) 50. — Prosthesia BLUME, Bijdr. (1826) 866. — Dioryktandra
HAssKk. Retzia (1855) 125. — Scyphellandra Tuw. En. Pl. Zeyl. (1858) 21. —
‘Imhofia ZOLL.’ ex TAvuB. (1895) l.c., nomen in synon.
Small shrubs to small trees; innovations mostly laxly pubescent, the hairs some-
times persistent; twigs pithy, generally angular when young, terete when older.
Leaves distichous, rarely in a spiral. Petiole comparatively short, or rarely wanting,
lacking distinct joints at base or top. Stipules caducous to long-persistent, some-
1971] VIOLACEAE (Jacobs & Moore) 181
y
WY
Uy
Y
Za
SER
Fig. 1. Rinorea horneri (KortH.) O.K. a, Habit, x 2, b. venation underneath, «24, c. stipule, 3,
d. flower, *4, e. gynoecium surrounded by stamens, 6, f. stamen from inside, «12, ¢. gynoecium,
«6, h. fruit, subtended by calyx and corolla, «1, i. seed, «1 (a, d-g Curtis 1898, b CLEMENS 9961,
c¢ SAN 42087, A PLeyre 232).
182 FLORA MALESIANA [ser. I, volia®
times quite conspicuous, often striate lengthwise. Leaf blade more or less dis-
tinctly acuminate, domatia occasionally present in the primary vein axils, the margin
shallowly incised to subentire, surfaces generally dull. /nflorescences lateral, the
flowers mostly in bundles, sometimes distichous along a short rachis, sometimes
in a corymb or panicle on a short peduncle, rarely in a raceme or on brachyblasts;
bracts smaller than the stipules; pedicels articulated. Flowers bisexual, rarely di-
oecious by reduction, actinomorphic, calyx quincuncial, covering the bud halfway
to completely, corolla generally contorted in the same direction as the third sepal.
The sepals (almost) equal, fimbriate. Peta/s in Malesia (sub)equal, free, sessile,
thin in texture, to 9 mm long. Androecium nearly as long as the petals, filamental
tube more or less fleshy and sometimes 5-lobed, the filaments inserted on its inner
margin, or seldom the filaments free with only a pair of scales outside at their
base; anthers converging, glabrous or hairy, the connective produced into an entire
dorsal membranous appendage smaller to longer than the thecae, sometimes
bearing each a ventral appendage at the top smaller than the dorsal one. Gynoecium
as longas the androecium, the 3 (—4) placentas with 1-3 ovules each; style straight,
with a terminal more or less distinctly 3-lobed stigma. Fruit capsular, approx-
imately globose before dehiscence, 34-4cm @, generally subtended by the dried up
calyx (which rarely expands), corolla, and androecium, the valves leathery, when
large two-layered with reinforced apex, mostly smooth and glabrous, rarely en-
veloped in a mass of appendages, or hairy. Seeds sessile, 3-6, rarely 1 or 9, ellipsoid,
glabrous, with leathery testa and often clearly defined hilum and raphe.
Distr. Pantropical, richest in Africa with c. 107 spp. on the mainland and 25 spp. in Madagascar;
in the New World with 50-60 spp.; in Indo-Malesia 12-13 spp., altogether extending from S. India/
Ceylon, Assam, Burma, Hainan, to N. Australia, Melanesia, the Carolines; estimated total about
200 spp. Fig. 3.
Ecol. Predominantly in the understorey of primary rainforests, at very low altitude, occasionally up
to 1000 m, on various soils including limestone.
Vern. Common and reliable names seem to be wanting. BURKILL, Dict. (1935) 1912, recorded for the
Malayan species some vernacular names, and some unimportant medicinal uses.
Taxon. A satisfactory subdivision of the entire genus has not yet been made. The one in the Flora
of British India is good for the present area, but is to be reconsidered when the neotropical and African
species will have been comprehensively studied. A/sodeia is there subdivided into 3 sections, viz. I.
Doryctandra (HASSK., genus, actually Dioryktandra) Hook. f. & TH. with stamens “‘exserted”’ (not thus
found by me), filaments slender (not a character), anthers cohering in a cone (notably by peculiar inter-
tangling hairs). Contains R. heteroclita. Il. Unnamed, with stamens included, anthers free, disk cupular
(our filamental tube). Contains most of our species, and the former genera Pentaloba and Prosthesia.
III. Scyphellandra (THw., genus) Hook. f. & Tu., with stamens included, anthers free, disk reduced
to 5 scales, one at the base of each anther (and, we may add, dioecious). Contains our R. virgata.
Kina, J. As. Soc. Beng. 58, ii (1889) gave another subdivision of Alsodeia; on page 400, Sect. I,
Prosthesia (BL.) KING, with 7 spp.; on page 404, Sect. II, Pentaloba (WALL.) KiNG, with 6 spp.; on page
407, Sect. III, unnamed, with 1 sp. Various ‘‘species’’ placed by KiNG in different sections have in the
present revision been merged.
Notes. The taxa in Rinorea east of the Indus have all been worked up in the same way, but the datz
have been sorted out for publication between the Flora Malesiana and its precursor in Blumea 15 (1967)
127-138. The precursor accounts for all names of all taxa, with their first references and their type
specimens, but gives descriptions, distribution and ecology only of non-Malesian taxa. The Flora Ma-
lesiana accounts for Malesia as far as names and literature are concerned, later references included, but
the descriptions cover the taxa found in Malesia over their full range, and the same holds for distribution
and ecology. The key in the Flora Malesiana is also complete for all taxa.
Six species belonging to other families have been ascribed to Rinorea; see under Excluded.
KEY TO THE SPECIES
1. Stipules caducous. Seeds 6 or fewer.
2. Primary veins curving approximately parallel towards the margin at distances of 1-2 cm; secondary
venation scalariform.
1971]
VIOLACEAE (Jacobs & Moore)
183
3. Inflorescences fasciculate (rarely stalked in R. anguifera).
4. Dorsal appendage of the anthers small. Style hairy. Fruit enveloped in moss-like appendages.
1. R. anguifera
4. Dorsal appendage of the anthers distinct. Style glabrous. Fruit smooth.
5. Anthers with small ventral appendages. Ovules 3. Fruit subglobose, with 3 seeds.
2. R. bengalensis
5. Anthers with ventral appendages about as long as the dorsal one. Ovules 6. Fruit conical, sub-
tended by the expanded calyx, with 1 seed.
3. R. horneri
3. Inflorescences on a peduncle (if racemose, see R. ‘longitticembsays
6. Gynoecium glabrous. Petals to 4 mm long. Anthers with small dorsal appendage. Fruit triangular
with rounded corners, 1—3 cm long, valves thick. . .
6. Gynoecium hairy. Petals to 9 mm long.
. 4. R. sclerocarpa
7. Pedicels jointed near the base. Anthers with small dorsal appendage. Fruit subglobose to ovoid,
8-11 mm long, hairy, with a hairy style remnant.
5. R. lanceolata
7. Pedicels jointed about the middle. Anthers with a ‘dorsal appendage 12 2 times as long as the cells.
(Fruit unknown.) .
6. R. macrantha
Primary veins few, independently | curv ed towards ‘the margin, secondary venation irregular.
re Inflorescences more or less elongate. Anthers with distinct ventral appendages. Fruit 114-4 cm o.
9. Stipules 1-314 mm long. Inflorescence axis 14—-2'4 cm long. Leaves often obovate.
7. R. javanica
9. Stipules (2—)4—21 mm long. Inflorescence axis to c. 10 cm long. Anthers often bearded at the base.
8. R. longiracemosa
8. Inflorescences fasciculate, or flowers densely set on a short rachis. Anthers with small or no ventral
appendages. Fruit 14-114 cmg.
10. Leaves petiolate, more or less concolorous. Anthers with a dorsal appendage about as long as the
cells or longer.
. Leaves 1—7(—14) cm long. Petiole 1-5 mm. Plant dioecious; in the ¢ flowers only a style, in the
2 flowers the anther cells vestigial, ovules 6. :
Leaves 6-36 cm long. Petiole 5-35 mm. Flowers bisexual. Ovules SP
9. R. virgata
ial Stipules | 14-6 mm, scarcely striate. Leaves dark-coloured in the dried state. Fruit sparsely hairy.
10. R. macrophylla
12. Stipules 4-16 mm, distinctly striate. Leaves greenish in the dried state. Fruit glabrous.
2. R. bengalensis
10. Leaves subsessile, discolorous, pale underneath, 314-6 cm long. Anthers with a small dorsal
appendage. Style hairy in the middle only. Ovary glabrous. Andamans, India?
1. Stipules long-persistent. Ovules and seeds 9. . .
1. Rinorea anguifera (Lour.) O.K. Rev. Gen. PI. |
(1891) 42; Crais, FI. Siam. En. 1 (1925) 88;
Merr. Comm. Lour. (1935) 270; GAGN. FI. Gén.
I.-C. Suppl. 1 (1939) 187; Jacoss, Blumea 15
(1967) 127. — Medusa anguifera Lour. FI.
Cochinch. (1790) 406. — Alsodeia echinocarpa
KortuH. Ned. Kruidk. Arch. | (1848) 360; Mita.
Fl. Ind. Bat. 1, 2 (1859) 116; OupeM. in Miq.
Ann. 3 (1867) 68; Hook. f. & Tu. in Hook. /.
Fl. Br. Ind. 1 (1872) 188; KING, J. As. Soc. Beng.
58, ii (1889) 406; Borssieu, FI. Gén. I.-C. 1
(1909) 214, f. 20: 6; RipL. Fl. Mal. Pen. 1 (1922)
134, f. 12. — Pentaloba corylifolia Turcz. Bull.
Soc. Nat. Moscou 27, ii (1854) 341. — Alsodeia
corylifolia (Turcz.) Turez. Bull. Soc. Nat.
Moscou 36 (1863) i 559. — Alsodeia capillata
KING, J. As. Soc. Beng. 58, ii (1889) 407; KING,
Ann. R. Bot. Gard. Calc. 5 (1896) 127, pl. 146A;
Rip. Fl. Mal. Pen. | (1922) 133. — Alsodeia
comosa KING, J. As. Soc. Beng. 58, ii (1889) 407;
KING, Ann. R. Bot. Gard. Calc. 5 (1896) 127,
_pl. 146B; Boissieu, FI. Gén. I.-C. 1 (1909) 213;
Ripi. Fl. Mal. Pen. 1 (1922) 133. — Alsodeia
echinocarpa Kortu. var. nervosa Capit. Bull. Soc.
Bot. Fr. 57 (1910) 394, et var. zollingeri BOERL. ex
Capit. Lc. — R. anguifera var. nervosa (CAPIT.)
R. heteroclita
11. R. iliaspaiei
@ en Se ee epee” OPE ae eee
Me_rr. Philip. J. Sc. 11 (1916) Bot. 100. — R.
comosa (KING) Merr. En. Born. (1921) 410. — R.
echinocarpa Burk. Gard. Bull. S. S. 6 (1930)172.
Young twigs terete, generally straight, densely
covered with a persistent brown tomentum;
pith narrow. Leaves distichous (spirally arranged
on terminal or crooked branches). Stipules (4—)
5-9(-il) mm long, triangular, not appressed,
with a distinct keel sometimes protracted into a
slender tip, more or less striate, in Borneo per-
sistent, otherwise the scars not conspicuous.
Petiole (1—)2—8(-10) mm. Blade thin-coriaceous,
to (8—-)14-28(-—37) by (3—)4-8(-10'4) cm, widest
at the middle to above, index 2.5—3.7(—4.4);
base acute to (mainly in Borneo) narrowly cor-
date, top acuminate; midrib and veins flat above,
prominent beneath, sometimes scarcely different
in colour, sometimes lighter; no domatia; primary
veins fairly regular and parallel, with distances in
the order of 1 cm between, no smaller intermediate
veins, secondary venation mostly distinct, scalari-
form, reticulation obscure; margin dentate, with
often larger and smaller teeth alternating, to
subentire; surfaces dull, brownish, often with a
livid tinge, somewhat paler beneath, midrib and
veins hairy beneath, to a smaller extent also
184
FLORA MALESIANA
[ser. I, voln7
above, seldom nearly glabrous. Flowers with up to
c. 20 in dense axillary (sub)sessile clusters on
twigs and sometimes also on branchlets; bracts
acute, resembling the sepals, pedicels very short.
Sepals c. 4-5 mm long, fleshy, hairy, the inner ones
narrower than the outer and slightly exceeding
them. Petals c. 5-7 mm long, strap-shaped but
wider in the middle, top recurved, often the whole
petal S-shaped, outside with a median band of
appressed silvery hairs. Stamens with distinct fi-
lamental tube, slender sometimes hairy filaments
and small glabrous anthers; dorsal appendage 14
of the anther’s length or shorter, ventral appenda-
ges none or vestigial. Ovary with its long appressed
hairs obconical, ovules 6, style with patent hairs,
glabrous towards the top. Fruit subtended by the
persistent sepals, before dehiscence globose, hidden
in a (mostly) dense, moss-recalling mass of more or
less branched, hairy appendages on the valves, in
all 114-3144 cm go. After dehiscence the seeds
become visible, about 3 in number, ellipsoid, 4~7
mm long, evenly straw-coloured, glossy.
Distr. In Indo-China scattered S of the 17th
parallel. In Malesia common in the Malay Pe-
ninsula S of 7°35’N in Thailand; throughout
Sumatra where very common in the East Coast
Residency; scattered in Borneo N of the Kapuas in
Kalimantan and Sarawak (all divisions) to Brunei;
also at 2°50’ N, 117°40’ E in NE. Kalimantan.
Ecol. Primary forest, in humid places, some-
times recorded from heath-forest, swamp forest,
light forest or forest edges, or secondary forest,
mostly on sandy or loamy, rarely from calcareous
soil; altitudes low, seldom up to 600 m.
Notes. A shrub 1m to a treelet 10m tall.
Bark smooth, mottled. Flowers mostly white to
sometimes yellow, facing upwards from the
horizontal branches, anthers light brown. Fruit
greenish at maturity. KING’s collector’s notes to
10084 from Perak: “in dense bamboo forest,
flowers deep-red with pale reddish green, waxy
tinged’’, do not fit in with any other data.
In RIDLEY 4824 from Singapore, the pedicels are
3 mm.
In TEIJSMANN fi. 30.VIII.1872 from Lingga, Bt.
Sipindjang off Sumatra, the branchlets are slender,
somewhat crooked, the leaves almost glabrous,
concolorous, and not distichous, 2-3 mm petioled,
c. 4-6 by 1144-2 cm. TEISMANN fl. 14.TX.1872
from Lingga, Sg. Tanda, is similar but the leaves
are c. 7-9 by 114 cm. Habit unknown.
In RAHMAT 1674 from Sumatra East Coast, the
globose, cluster-like inflorescences are 5mm pe-
dunculate.
2. Rinorea bengalensis (WALL.) O.K. Rev. Gen.
Pl. 1 (1891) 42; Gacn. FI. Gén. I.-C. Suppl. 1
(1939) 190; Merr. & CHUN, Sunyatsenia 5 (1940)
127, f. 12; Jacoss, Blumea 15 (1967) 128. —
Alsodeia bengalensis WALL. Trans. Med. Phys.
Soc. Calc. 7 (1835) 224; Hook. f. & Tu. in Hook.
f. Fl. Br. Ind. 1 (1872) 186; Kurz, For. Fl. Burma
1 (1877) 70; Kans. & Das, Fi. Assam 1 (1934)
82. — Alsodeia obtusa KorTH. Ned. Kruidk.
Arch. 1 (1848) 359; Mia. Fl. Ind. Bat. 1, 2 (1859)
115; OupEem. in Mig. Ann. 4 (1869) 215. —
Pentaloba fasciculata Turcz. Bull. Soc. Nat.
Moscou 27, ii (1854) 341. — Pentaloba semigyrata
Turcz. l.c. 342. — Alsodeia fasciculata (TURCZ.)
TurRcz. Bull. Soc. Nat. Moscou 36 (1863) i 559. —
Alsodeia paradoxa BL. ex OUDEM. in Miq. Ann. 3
(1867) 71. — Alsodeia wallichiana Hook. f. &
Tu. in Hook. f. Fl. Br. Ind. 1 (1872) 187; Kina,
J. As. Soc. Beng. 58, ii (1889) 400; Rip. FI.
Mal. Pen. 1 (1922) 128. — R. wallichiana (HooK.
f. & Tu.) O.K. Rev. Gen. Pl. 1 (1891) 42; HENbD.
J. Mal. Br. R. As. Soc. 17 (1939) 35. — R. obtusa
(KoRTH.) O.K. Rev. Gen. Pl. 1 (1891) 42. —
Gelonium glandulosum Em. Leafl. Philip. Bot.
3 (1910) 917. — Gelonium trifidum Exo. I.c. 918.
— Alsodeia lankawiense Rivv. J. Str. Br. R. As.
Soc. 59 (1911) 70; Fl. Mal. Pen. 1 (1922) 129. —
Alsodeia pruinosa PULLE, Nova Guinea 8 (1912)
669. — Alsodeia salomonensis RECH. in Fedde,
Rep. 11 (1912) 184; Denkschr. K. Ak. Wiss.
M.-N. KI. Wien 89 (1913) 137, t. 6, f. 11b. —
R. paradoxa (BL. ex OUDEM.) J. J. SMITH in K. &
V. Bijdr. 13 (1914) 67. — Alsodeia semigyrata
(Turcz.) J. J. Smitu, l.c. 72. — R. semigyrata
(Turcz.) J. J. Smiru, /.c. 73; MerR. J. Arn. Arb.
35 (1954) 145; BAcK. & BAKH. f. Fl. Java 1 (1964)
194. — R. fasciculata var. minor Em. Leafl.
Philip. Bot. 8 (1915) 2877. — R. fasciculata
(Turcz.) MErRR. Philip. J. Sc. 12 (1917) Bot. 286;
En. Philip. 3 (1923) 104; J. Arn. Arb. 32 (1951)
80. — R. glandulosa (ELM.) MeErR. Philip. J. Sc.
12 (1917) Bot. 286; En. Philip. 3 (1923) 104;
J. Arn. Arb. 32 (1951) 80. — R. lankawiensis
(RIDL.) CRAIB, Fl. Siam. En. 1 (1925) 90. — R.
pruinosa (PULLE) MELCH. in E. & P. Nat. Pfi.
Fam. ed. 2, 21 (1925) 352; Bot. Jahrb. 62 (1929)
369. — R. salomonensis (RECH.) MELCH. in E. &
P. Nat. Pfl. Fam. ed. 2, 21 (1925) 352. — Suregada
glandulosa (ELM.) CROIZAT, Bull. Jard. Bot Btzg
III, 17 (1942) 216. — Suregada trifida (ELM.)
CroizatT, /.c.
Growth apparently in flushes, but internodes of
equal length. Young parts with lax to rarely dense
yellow-brown pubescence, glabrescent. Twigs
greenish, furrowed when young, often zig-zag,
nodes distinct, with stipular scars taking 14 of
the twig’s circumference. Leaves often disti-
chous. Stipules appressed, triangular, slender,
(4-)8-16 mm long, brown and distinctly striate.
Petiole 14-214 (-3'4) cm, may differ in length from
the stipules. Blade coriaceous, 6—22(—36) by
214-9(-1214) cm, widest at the middle or slightly
above, index (1.9—)2-3(—3.3); base acute, rarely
to subcordate, top gradually acuminate; midrib
and veins distinct, yellowish to sometimes brownish,
often with hairy pit-domatia in the primary
axils, major veins about 7-13 on either side, the
basal one always thin; reticulation on both sides
distinct, not very regular; margins shallowly
crenate, rarely entire; surfaces distinctly greenish
with variants, darker, glossy, glabrous above,
beneath sometimes with a few hairs on the veins.
Inflorescences on the young twigs; from an axil-
lary rosette of caducous hypsophylls 1-2 mm long
and similar in kind as the stipules, a few (seldom
1971]
one) partial inflorescences crop out like curving
fingers 44-1 cm long, which grow at the top
while producing flowers in two rows, the pedicels
leaving distinct scars, eventually 10-30. Flower
buds initially protected by 2 keeled brownish
bracteoles (?) c. 144 mm long, the outer ones
inserted on the dorsal side of the “‘finger’? where
they are to leave a broad pale scar. Sometimes the
pattern is less clear, there being a cluster of flowers
that may be dense and large in Philippine plants,
seldom there is (in New Guinea) a stout common
peduncle a few mm long. Pedicel 3—10(—20) mm,
jointed at or near the base. Sepals 13—-!4 the
length of the petals, rarely as long; fleshy, ciliate.
Petals strap-shaped and often S-curved, (214-—)3-4
(-5)mm long. Androecium glabrous (occasion-
ally a few hairs at the base of the thecae), filamental
tube 0.1-0.5 times as long as the stamens, the
filaments 0.1-0.4, the anthers 0.4—0.8; dorsal
appendage varying in size and shape, mostly
broad-triangular; ventral appendages mostly
small. Ovary sometimes hairy, with 3 ovules,
style glabrous. Fruit at the base with the dried
striate sepals and petals, before maturity globose,
(A4-)1-144 cm @, with a rest of the style, dull,
glabrous. Seeds 3, subglobose, 3-71 mm, yellow-
brown with specks of purple to dull purple all over.
Distr. Ceylon, S. India, Assam, Burma,
Indochinese Peninsula and Hainan, Andamans,
scattered through Malesia (not in S. Sumatra and
Java); Northern Queensland, Melanesia, Ca-
rolines.
Ecol. An understorey plant of mixed rain-
forest at low elevation; as an exception on Mt
Kinabalu up to 1500 m. On various soils but often
on steep limestone hills; once recorded from
swampy forest; no apparent periodicity.
Notes. A shrub which may already be fertile
when 2 feet tall, to a tree of c. 20m by 28 cm.
Wood yellow, flowers green to white, fruit orange
with red seeds, or cream-brown, or pale green.
Several records mention the rich green leaves.
A very polymorphic species.
As the plant has never been found again in
Java after BLUMe’s collection from Mts Salak and
Burangrang, which was described as Alsodeia
paradoxa, 1 think the record is apocryphous.
ZOLLINGER 2979, the type of Pentaloba semigyrata,
was from a plant in the Bogor Gardens.
3. Rinorea horneri (KorTH.) O.K. Rev. Gen. PI.
1 (1891) 42. — Alsodeia horneri Kortu. Ned.
Kruidk. Arch. 1 (1848) 360; Mia. FI. Ind. Bat.
1, 2 (1859) 116; Oupem. in Miq. Ann. 3 (1867) 70;
Mio. Ann. 4 (1869) 214; Merr. En. Born. (1921)
410; Jacops, Blumea 15 (1967) 130. — Alsodeia
browni Kortu. Ned. Kruidk. Arch. 1 (1848) 361;
Hassk. Retzia ed. nov. (1858) 33 (‘A. Brownei’),
name only, the description being of R. lanceolata;
Mia. FI. Ind. Bat. 1, 2 (1859) 116 (‘A. Brownii’);
Oupem. in Mig. Ann. 3 (1867) 70. — Alsodeia
‘kunstleriana KinG, J. As. Soc. Beng. 58, ii (1889)
401; Kinc, Ann. R. Bot. Gard. Calc. 5 (1896)
122, pl. 141; Ript. Fl. Mal. Pen. 1 (1922) 128, —
R. brownii (KortH.) O.K. Rev. Gen. PI. 1 (1891)
VIOLACEAE (Jacobs & Moore)
185
42. — Alsodeia astrolabes K. ScH. & Laut. FI.
Schutzgeb. (1901) 451. — R. copelandii MERR.
Philip. Gov. Lab. Bur. Bull. 35 (1906) 45;
En. Philip. 3 (1923) 103. — R. acuminata MERR.
Philip. J. Sc. 5 (1910) Bot. 201; En. Philip. 3
(1923) 103. — Alsodeia hirtella Rip. Bull. Misc.
Inf. Kew (1914) 377. — R. amboinensis MERR.
Philip. J. Sc. 11 (1916) Bot. 292. — R. hirtella
(RIDL.) MerRR. En. Born. (1921) 410; Philip. J.
Sc. 30 (1926) 413. — R. castilloi MERR. Philip.
J. Sc. 21 (1922) 530. — R. astrolabes (K. Scu. &
LauT.) MELCH. in E. & P. Nat. Pfl. Fam. ed. 2, 21
(1925) 352; Bot. Jahrb. 62 (1929) 371, incl. var.
hirta MELCH. I.c. 372. — R. ledermannii MELCH.
lic. 370. — R. elmeri MERR. Un. Calif. Publ.
Bot. 15 (1929) 206. — R. celebica MELCH. Notizbl.
Berl.-Dahl. 12 (1934) 205. — R. hamelii MERR.
Pap. Mich. Ac. Sc. 19 (1934) 171. — Fig. 1, 2.
An undershrub (35—)60 cm to a treelet 5 m tall.
Twigs brownish yellow, furrowed when young,
pithy, laxly to densely set with a greyish to fulvous
indumentum puberulous to velvety. Terminal bud
not completely enveloped by the stipules, the
latter (3—)6—-11(—13) mm long, rather wide at the
base, brown and striate, more or less hairy, leaving
inconspicuous scars. Leaves spirally arranged.
Peticle (3—)8-25(—32) mm, hairy like the twig.
Blade thin in texture, to 11-34 by 4-914(-11) cm,
widest above the middle, sometimes about, index
(2.2—)2.7—3.4(-3.8); base tapering, top acuminate;
midrib and veins distinct, flat above, prominent
beneath, yellowish brown, no domatia; primary
veins fairly regular and parallel towards the margin
at distances of c. 1(—2) cm, nearly always without
smaller intermediate veins; secondary venation
scalariform, often crossing the primary veins;
reticulation distinct beneath; margins dentate to
sometimes entire; surfaces fairly dull, paler be-
neath, some hairs on the midrib and major veins
beneath often remaining, otherwise glabrous,
brown-green in the dried state (yellowish when
Schweinfurthed). Jnflorescences originally sub-
tended by a few stipule-like hypophylls in the
axils of the (Ist-)2nd-6th leaves of twigs, or some-
times the flowering part of a twig leafless, rarely
flowering on an older branchlet, consisting of a
more or less contracted (sub)sessile panicle with
unequal branches, the main axis to 1'4 (excep-
tionally to 7) cm long; flowers few-20 (rarely more
spirally arranged; bracts and bracteoles c. | mm;
all pedicels jointed at 44-1 mm above their in-
sertion, the distal part 3-10(-15) mm, thickened
towards the top, puberulous. Buds and often
flowers ovoid, but sometimes the outer sepals
spreading in the dried state; calyx somewhat
decurrent into the pedicel; petals converging,
corolla with a narrow opening at the top. Sepals
2-4(—-5) by 1-—5(-6) mm, more or less fleshy,
variously hairy, exceptionally the margins of the
outer sepals recurved. Petals (2—)3—6(—8) mm long,
sometimes S-shaped or with recurved top, mostly
thin but sometimes fleshy. Androecium glabrous
but exceptionally a few hairs at the base of the
anthers or also the filaments hairy; filamental tube
0.1-0.3 part of the length of the stamens, the fila-
186
FLORA MALESIANA
[ser. I, vol. 7}
ments 0.1—0.3(—0.5) part, the anthers 0.4-0.7 part;
dorsal and ventral appendages always well
developed. Ovary mostly hairy, sometimes gla-
brous, style glabrous, rarely hairy; ovules 6(—7).
Calyx enlarging with the fruit, eventually leathery,
funnel-shaped, 8-12mmg, on a firm pedicel,
about 3-5 per inflorescence. Fruit ovoid, 10-15 mm
long, glabrous, turbinately splitting, the valves
thin but firm; seed one, ellipsoid to ovoid, c. 8-11
mm long, glossy even light brown.
Fig. 2. Rinorea horneri (KORTH.) O.K., deviating
specimens. a. An elongated inflorescence, x 2,
b. flower with sepal margins recurved, c. such a
sepal in cross-section (a KOSTERMANS 21471,
b-c JAcogs 5431).
Distr. Peninsular Siam; scattered throughout
Malesia except Java and the Lesser Sunda Is.;
Solomons.
Ecol. In various kinds of primary forest,
sometimes depleted, rarely in secondary forest, up
to 600 m (to 1500 m on Mt Kinabalu); often on
limestone, but also on sandstone or clay.
Notes. An undershrub (35—)60 cm to a treelet
5m tall. Leaves dark green above, pale beneath;
flowers weakly fragrant, white to pale green-
yellow, the anthers pale brown. Fruit apparently
green and hard at maturity.
For KING’s remark of 1896 on the occurrence
of both bisexual and 2 flowers, I found no con-
firmation in the material. That “the anthers
have no pollen” may be due to their very early
loosing it.
The name R. strobobas is sometimes found as a
corruption of astrolabes.
4. Rinorea sclerocarpa (BURGERSD.) JAcoss,
Blumea 15 (1967) 131. — Alsodeia sclerocarpa
BURGERSD. in Miq. Pl. Jungh. (1852) 122. —
Alsodeia rugosa MiQ. Sumatra (1860) 390; Ann.
4 (1869) 215. — Alsodeia condensa KiNG, J. As.
Soc. Beng. 58, ii (1889) 405; Ann. R. Bot. Gard.
Calc. 5 (1896) 126, pl. 145; Rip. FI. Mal. Pen. 1
(1922) 132. — Alsodeia pachycarpa KiNG, J. As.
Soc. Beng. 58, ii (1889) 407; Ript. FI. Mal.
Pen. 1 (1922) 131. — Alsodeia scortechinii KiNG,
J. As. Soc. Beng. 58, ii (1889) 405; Ann. R. Bot.
Gard. Calc. 5 (1896) 126, pl. 144; Rrpv. FI. Mal.
Pen. 1 (1922) 131. — R. pachycarpa (KING)
CraisB, Fl. Siam. En. 1 (1925) 91. — R. scorte-
chinii (KING) Crats, l.c. — R. sumatrana MERR.
Pap. Mich. Ac. Sc. 19 (1934) 172. — R. condensa
(KING) Merr. l.c. 173.
Growth apparently in flushes; internodes of
varying length. Twigs + terete, dull greyish or
sometimes purplish tinged, initially with greyish
pubescence but soon glabrescent; branchlets
slightly crooked, bark with faint longitudinal ribs,
pith narrow. Leaves mostly distichous. Stipules
(3—)4-7(-10) mm Jong, not appressed, triangular,
with a pale-hairy keel, more or less striate, their
scars not conspicuous. Petiole 5-12 mm, dull
greyish, often wrinkled. Blade thin-coriaceous, to
1314-25(—38) by 414-9(-11) cm, widest about the
middle to sometimes above, index (2.1—)2.6-3.5
(-3.9); base acute and slightly decurrent, top
acuminate; midrib and veins often in a similar
colour as the surface to somewhat lighter; no
domatia; primary veins fairly regular and conver-
ging within the margin, with distances between in
the order of 114 cm at the midrib, no smaller
intermediate veins, secondary venation thin,
scalariform, reticulation indistinct; margin shal-
lowly dentate, entire near the base; surfaces dull,
above grey-green-brown, glabrous, beneath with
a characteristic chocolate-brownish colour, gla-
brous or seldom sparsely hairy on the main nerves.
Inflorescences in the axil of the 2-4 latest leaves,
2-5 cm long, often rich-flowered and sometimes
quite densely, peduncle mostly near the base
divided into 2-4 main axes, which after anthesis
may long persist as partly empty stalks; rather ir-
regularly dichasially branched, axes laxly greyish
puberulent; bracts late caducous, triangular, 1-2
mm; pedicels jointed below the middle, i.e. at
1-3 mm above their insertion. Flowers pear-
shaped, the corolla converging. Sepals subor-
bicular, (2—)2!4-3(-4) mm, fleshy, sparsely hairy.
Petals 314-4 mm long, sometimes the top slightly
recurved, outside sparsely hairy towards the top.
Androecium glabrous, filamental tube 0.3 part of
the length of the stamens, filaments 0.4—0.5 part,
anthers 0.2—0.3 part; dorsal appendage pale, short
and narrow, no ventral appendages. Pistil gla-
brous; ovules 6. Fruits seldom more than 2 per
inflorescence, subtended by the persistent sepals,
before maturity mostly tapering to both ends,
(0.9-—)2.3-3 cm long, bluntly triangular with
hollow sides (0.8—)1.5-2cm wide, dull dark-
purplish, the valves eventually spreading, com-
pressed and separating into two layers, the inner
one cartilaginous and straw-coloured, the outer
one leathery. Seeds 3-6, c. (5—)6-8 mm long,
evenly dull brownish.
Distr. Malesia: Malay peninsula (scattered
between 6°17’ N in S. Siam and Johore), Sumatra
(scattered in Atjeh and East Coast, also near
Palembang).
1
i
1971]
Ecol. Primary forest, on moist slopes or by
small rivers, lowlands to 1000 m. Fi. March, /r.
June—Sept.
Notes. Tree, more or less crooked, sometimes
shrubby, to 10(—20)m tall, bole 7-l3cm a,
fluted or buttressed at the base, bark mostly
smooth, fawn or grey; slash inner bark granular,
yellow or dull orange, wood cream or white.
Branches brittle. Leaves above dark green, be-
neath light green with darker veins. Flowers
white, or pale green. Ripe fruits triangular, dark
green, sometimes flushed with orange, 314 cm 9;
unripe fruits may be globose.
MIQUEL, FI. Ind. Bat. 1, 2 (1859) 115, had it
under Alsodeia obtusa, our Rinorea bengalensis;
BoERLAGE, Cat. Hort. Bog. 1 (1899) 48, under
Alsodeia javanica.
Alsodeia rugosa was described with a hairy
ovary; in the buds available this was glabrous.
A Cur. SMITH specimen (BM), stated to be from
Ceram, fi. Il. 1802, must have been wrongly la-
belled.
5. Rinorea lanceolata (WALL.) O.K. Rev. Gen. PI. 1
(1891) 42; Merr. J. Arn. Arb. 35 (1954) 145;
Jacogs, Blumea 15 (1967) 133. — Vareca lan-
ceolata WALL. in Roxb. Fl. Ind. ed. Wall. 2
(1824) 445; ed. Carey 1 (1832) 648. — Pentaloba
lanceolata (WALL.) ARN. Mag. Zool. Bot. 2
(1848) 544. — Alsodeia brownei (non KorTH.)
Hassk. Retzia ed. noy. (1858) 22. — Alsodeia
cymulosa Miq. Fl. Ind. Bat. 1, 2 (1859) 116,
non WELW. 1868; Back. Schoolfl. (1911) 67. —
Alsodeia dasypyxis Mia. Sum. (1860) 391; Ann.
4 (1869) 215 (‘dasyprysis’ in Kew Index). —
Alsodeia lanceolata OupeM. in Miq. Ann. 3
(1867) 68; Arch. Néerl. Sc. Ex. Nat. 2 (1867) 196;
Hook. f. & Tu. in Hook. f. Fl. Br. Ind. 1 (1872)
188; Kuna, J. As. Soc. Beng. 58, ii (1889) 404;
Ripi. FI. Mal. Pen. 1 (1922) 131. — Alsodeia
maingayi Hook. f. & Tu. in Hook. f. Fl. Br. Ind.
1 (1872) 188; KiNG, J. As. Soc. Beng. 58, ii (1889)
402; Rip-. Fl. Mal. Pen. 1 (1922) 130. — Alsodeia
mollis Hook. f. & Tx. in Hook. f. Fl. Br. Ind. 1
(1872) 188; Kurz, For. Fl. Burma 1 (1877) 70. —
Alsodeia floribunda KiNG, J. As. Soc. Beng. 58,
ii (1889) 406; Ann. R. Bot. Gard. Calc. 5 (1896)
125, pl. 143B; Rip. FI. Mal. Pen. 1 (1922) 132.
— R. cymulosa (Miq.) O.K. Rev. Gen. Pl. 1
(1891) 42; Koorp. Exk. Fl. Java 2 (1912) 65;
J. J. Smiru in K. & V. Bijdr. 13 (1914) 70; Back.
& Baku. f. Fl. Java 1 (1964) 193. — R. maingayi
(Hook. f. & Tu.) O.K. Rev. Gen. Pl. 1 (1891) 42;
Taus. in E. & P. Nat. Pfl. Fam. 3, 6 (1895) 329.
— R. mollis (Hook. f. & Tu.) O.K. Rev. Gen. PI. 1
(1891) 42; Crais, Fl. Siam. En. 1 (1925) 90. —
Alsodeia grandiflora Rivi. Bull. Misc. Inf. Kew
(1914) 377. — R. floribunda (KiNG) MerR.
Pap. Mich. Ac. Sc. 19 (1934) 171.
Growth mostly in flushes; the first internodes
shorter. Twigs slender, + straight, generally
purplish, terete when young, lengthwise striate,
pith narrow; sometimes laxly fulvous velvety but
mostly glabrous from an early stage. Terminal
bud with some tips as the stipules do not com-
VIOLACEAE (Jacobs & Moore)
187
pletely envelop it. Leaves mostly spirally arranged,
sometimes distichous. Stipules 4-11 mm long,
rather wide at the base, brown and striate, leaving
rather distinct scars. Petiole 2—7(-10) mm, ge-
nerally dark-coloured. Blade thin-coriaceous in
texture, to 10-29(-42) by (2.3—)3.3-9(-20) cm,
widest about the middle to sometimes above,
index (2—)2.5-4.8(—6.1); base tapering to obtuse,
top acuminate; midrib and veins flat above, pro-
minent beneath, mostly in a similar colour as the
leaf surface; no domatia except in Java; primary
veins fairly regular and converging within the
margin, with distances between them in the order
of 34-1(-114) cm at the midrib, smaller inter-
mediate veins rare, secondary venation indistinct,
scalariform, reticulation obscure; margin (often
minutely) dentate; surfaces generally dull, and
concolorous, green-brownish, midrib and veins
on both sides hairy like the twigs, if so, sparsely
hairy between the veins underneath but mostly
glabrous. Inflorescences on the young twigs rarely
also on branchlets, laxly globose to compound
corymbose in shape, 1-9 cm long in all; peduncle
a few mm to 3(-4!4) cm long, hairy like the twig,
coloured like the petioles, mostly divided into two
subequal axes of about 1; its length; bracts c. 2-3
(if on a main axis to 10) mm long, lanceolate;
number of flowers few to several dozens; pedicels
to 4(-8) mm, jointed in their basal half. Sepals
214-5 by 134-414 mm, suborbicular, elliptic or
ovate, more or less fleshy, with thin margin, the
outer sepals somewhat shorter and wider than the
inner ones and sometimes with a cucullate top,
somewhat hairy particulary in the centre. Petals
314-7(-9) by 114-2 mm, widest about the middle,
thin, top sometimes recurved, outside with a pale
median band of hairs. Stamens consisting of a
filamental tube of 0.2—0.3 part of the total length,
a filamental of 0.4—0.5 part, and an anther of
(0.2—)0.3-0.4 part; upper part of the filament and
back of the connective between the thecae hairy,
exceptionally glabrous; dorsal appendage tri-
angular, 14-14 the length of the thecae, ventral
appendages up to half the length of the dorsal one,
or wanting. Pistil hairy, ovules 6, once 3. Fruits
subtended by the persistent sepals and petals,
before dehiscence subglobose to slightly ovoid,
the hairy style always persistent, 8-11 mm long,
the valves leathery, brownish, more or less hairy,
splitting straightly; seeds c. 3, c. 3-6 by 2-4 mm,
dull evenly light brown.
Distr. Burma: S. Tenasserim; Peninsular Siam;
in Malesia: common in Malaya, scattered in
Sumatra (also Lingga Is.) and in the very SW of
Java (also Peutjang I. in Sunda Straits.)
Ecol. Primary forest and (open) jungle, also
forest edges, in fertile volcanic loam, also on
coral limestone, below 750m.
Notes. A shrub 14-3 m to a tree 6—-13(-21) m
by 4-30 cm @, bark grey smooth thin, the twigs
may be drooping at the top. Leaves light or yel-
lowish green, once reported darker above, and
once glossy. Flowers with green calyx and (waxy)
white-yellow corolla. Fruits (pale) green when ripe.
In one specimen the fruits are 4-merous.
188
The species is quite polymorphic. All specimens
from Penang (the type locality) are glabrous,
with narrow leaves: index 3.3—6.1, and small in-
florescence: peduncle 144-5 mm. All specimens
from Perak are hairy, with rather wide obovate
leaves: index 2.6—3.3, with broad base, and large
inflorescences: peduncle 214-3 cm. All specimens
from Peninsular Burma and Siam (‘R. mollis’)
are hairy, with still wider leaves: index 2.4~—2.8,
and very wide sepals. All specimens from West
Java and Peutjang are glabrous, with 1 cm stalked
large leaves with domatia, and lax inflorescences
5-8 cm long in all. KoorpDERS 4498 and 4499 are
placed here on account of their domatia. Except
for the domatia, the above forms are not restricted
to the districts mentioned, and in other provinces
various forms have been found.
Fig. 3. Rinorea, number of species in Malesia and
adjacent regions, the number of endemics, if any,
below the hyphen.
6. Rinorea macrantha JAcoss, Blumea 15 (1967)
133 5th als
Twigs angular, glossy light brown, glabrous;
branchlets terete, zig-zag. Stipules 7mm long,
brown, striate. Leaves probably spirally arranged,
subsessile to 34 cm petiolate; blade (sub)coria-
ceous, 15-33 by 514-10 cm, widest about the
middle; base blunt to obtuse and subcordate, top
long-acuminate; midrib strong and prominent,
veins regularly curved with c. 114 cm between,
reticulation distinct beneath, more or less scalari-
form; margin crenate; surfaces dull, and glabrous
from the beginning. Flowers with several in lax la-
teral racemes 1-114 cm long, bracts 2-3 mm long,
pedicels 6-7 mm, jointed just above the middle.
Sepals thin, 3144-5 by 214-414 mm, glabrous or
nearly so. Petals thin, 6-9 by 2-314 mm, glabrous.
Androecium glabrous, filamental tube fleshy and
distinctly 5-lobed, the filaments as long or shorter,
anthers with a triangular dorsal appendage 1-2
times as long as the cells; ventral appendages none.
Pistil club-shaped, brown-puberulous in the basal
4-24, part; ovules 6. Fruit unknown.
Distr. Malesia: W. Central Borneo.
Notes. For the slight differences between the
two collections on which the above description has
been based, see JAcoss, /.c.
A sterile collection from Singkep in the Lingga
FLORA MALESIANA
(ser. I, voly 7?
Archipelago, BUNNEMEVER 7384 (BO) may also
belong here.
7. Rinorea javanica (BL.) O.K. Rev. Gen. Pl. 1
(1891) 42; Koorp. Exk. Fl. Java 2 (1912) 625;
J. J. SMitH in K. & V. Bijdr. 13 (1914) 64; BAcK.
& BAKH. f. Fl. Java 1 (1964) 194; Jacoss, Blumea
15 (1967) 134. — Prosthesia javanica BL. Bijdr.
(1826) 867. — Alsodeia haplobotrys WHassk.
Vers]. Kon. Ak. Wet. 4 (1856) 137; Retzia ed.
nov. (1858) 19; Mia. Fl. Ind. Bat. 1, 2 (1859) 116.
— Alsodeia javanica (BL.) Mig. Fl. Ind. Bat. 1,
2 (1859) 117; OupDEM. in Miq. Ann. 3 (1867) 71;
Mia. Ann. 4 (1869) 216; BAck. Schoolfl. (1911) 68.
— Alsodeia hookeriana KinG, J. As. Soc. Beng.
58, ii (1889) 402; Ann. R. Bot. Gard. Calc. 5
(1896) 123, pl..142A; Ripv. Fl. Mal. Pen. 1 (1922)
130. — Alsodeia wrayi KinG, J. As. Soc. Beng.
58, ii (1889) 403; Ann. R. Bot. Gard. Calc. 5
(1896) 124, pl. 142B; Rip. Fl. Mal. Pen. 1 (1922)
129. — R. haplobotrys (Hassk.) O.K. Rey. Gen.
Pl. 1 (1891) 42; Koorp. Exk. Fl. Java 2 (1912)
624. — R. wrayi TAUB. in E. & P. Nat. Pfl. Fam. 3,
6 (1895) 329. — Alsodeia minahassae Koorp.
Minah. (1898) 645. — Alsodeia formicaria ELM.
Leafl. Philip. Bot. 5 (1913) 1850. — R. formicaria
(Ecm.) MErRR. Philip. J. Sc. 12 (1917) Bot. 286;
En. Philip. 3 (1923) 104. — R. hookeriana (KING)
Burk. Bull. Misc. Inf. Kew (1935) 319; Dict.
(1935) 1913.
Growth in flushes, internodes rather variable in
length. Young twigs grooved, purplish to light-
coloured but soon with grey bark; pith narrow;
glabrous almost from the beginning to sometimes
rather densely short-pubescent. Leaves largely
distichous. Stipules 1-3(-314)mm long, with
broad base and narrow top, keeled, sometimes
late caducous. Petiole 3-15 mm, leaving a con-
spicuous scar. Blade more or less thinly coria-
ceous, 10-17(—-20) by 234-514(-7) cm, widest at
the middle or often well above, index (2.5-)
2.8-3.3(-4); base cuneate and slightly decurrent
to seldom rounded, top acuminate; midrib flat
to subprominent above, veins 5—9(—14) on either
side, converging within the margin, comparatively
slender, no domatia; reticulation distinct (less so
in Indo-China), without much regularity; margins
more or less distinctly dentate, rarely (sub)entire;
surfaces dark green-brown with variants, mostly
lighter underneath, dull but sometimes very glossy
above, glabrous or with some hairs on the midrib
beneath. Inflorescences mostly on the branchlets,
axillary (sometimes a whole row of them on a
leafless part of a twig), generally consisting of a
bracteate pubescent axis 0.5—2.6 cm long, some-
times cluster-like or rarely simple to 3 cm long and
then the flowers distichous, with 3—7 lateral cymes
c. % cm long with a few to several flowers each;
bracts resembling the stipules; pedicels 3-5 mm,
jointed about the middle. Sepals more or less
fleshy, triangular to elliptic, 1-2 by 34-1 mm,
outside with some hairs. Petals thin, 3 by 1-114
mm, sometimes in the apical half the margins
like pinched together, outside often with some
hairs. Filamental tube about 14 part the total
1971]
length of the stamens, glabrous; filaments 0.2-0.3
part, sometimes hairy; anthers 0.3-0.2 part, gla-
brous; dorsal appendage broad, triangular, as
long as or somewhat longer than the cells, ventral
appendages about half as long as the dorsal one.
Ovary sometimes hairy, with 3-9 ovules; style
glabrous. Fruit subtended by the dried diverging
perianth, on a very short stipe, globose to elliptic
before dehiscence and c. 114-2'4 cm long; valves
glabrous or short-hairy, dark-coloured, with some
veins like a man’s skin, leathery with woody back
and top, the latter folded inwards upon dehiscence.
Seeds 3-6, c. 8-9 by 6-7 mm, with irregular pattern
of dark and light colour, sometimes evenly dark-
coloured, hilus distinct.
Distr. Indo-China (only in Annam _ near
Quang-tri); Southwestern and Peninsular Siam;
in Malesia frequent in the everwet parts of Ma-
laya; in Sumatra very rare and not in the northern
part; in W. Java in several places; in the Philippines
once found in Palawan; in Celebes once found in
the NE. Peninsula.
Ecol. Mostly in primary, also in light forest,
sometimes in swamp or peat forest, at low ele-
vations and on hillsides up to 1000 m. No records
from calcareous soil.
Notes. Tree 5-10m with spreading hanging
branches. Leaves bright and sometimes glossy
green, paler beneath. Flower white to pale green.
Capsules dark greenish brown or black, c. 154
by 1 cm, dehiscing explosively.
In BLUME’s type specimen and some others the
outer layer of the fruit is 4-5 mm thick and
corky — a malformation?
Alsodeia formicaria owes its name to the small
black ants which Ecmer found inhabiting the hollow
and sometimes bladdery capsules, “‘passing in
and out through the distinctly circular aperture
at the top.”
As the only collection from Borneo is one by
KoORTHALS, whose labelling was often inaccurate,
this record is considered apocryphous; it may be
from Sumatra.
8. Rinorea longiracemosa (KURZ) Craip, Fl. Siam.
En. 1 (1925) 90; Jacoss, Blumea 15 (1967) 134. —
Alsodeia longiracemosa Kurz, J. As. Soc. Beng.
39, ii (1870) 63; For. Fl. Burma 1 (1877) 70. —
Alsodeia cinerea KING, J. As. Soc. Beng. 58, ii
(1889) 403; Ann. R. Bot. Gard. Calc. 5 (1896) 124,
pl. 143A; Rip. Fl. Mal. Pen. 1 (1922) 130. —
Alsodeia cinerea var. hirsutiflora KiNG, J. As.
Soc. Beng. 58, ii (1889) 404; Ann. R. Bot. Gard.
Calc. 5 (1896) 125; Ripv. Fl. Mal. Pen. 1 (1922) 130.
— Alsodeia macropyxis Capit. Bull. Soc. Bot. Fr.
57 (1910) 395. — R. macropyxis (CapiT.) MERR.
En. Born. (1921) 411. — R. gaultheriiflora J. J.
Smitu, Bull. Jard. Bot. Btzg III, 4 (1922) 231,
t. 5; Back. & Baku. f. Fl. Java 1 (1964) 193. —
R. borneensis Mere. Un. Calif. Publ. Bot. 15
(1929) 206.
Growth apparently in flushes, internodes of
largely similar length. Twigs when young furrowed
and more or less puberulous; branchlets somewhat
crooked, the bark brownish or greyish, sometimes
VIOLACEAE (Jacobs & Moore)
189
dull purplish, corky with faint longitudinal ribs,
pith narrow. Leaves mostly distichous. Stipules
with broad base, (2—)4-14(—21) mm long, striate,
more or less closely appressed, often late caducous.
Petiole 2-10 mm. Blade firmly thin-coriaceous, to
(35%4-)8-18(-22) by (13%4-)24%4-5144--74%4) cm,
widest about the middle, index 2.3—-3.4(-4); base
blunt to acute, top acuminate; midrib narrow,
prominent on both sides, major veins about 6-11
(—14) on either side, converging within the margins,
their axils mostly with hairy domatia (not in
North Borneo), secondary veins and reticulation
lax and rather irregular; margin with small
roundish teeth; surfaces dull, olive-green to
brownish, with a greyish aspect, mostly some-
what lighter beneath, glabrous but sometimes
the midrib above sparsely pubescent, rarely pilose
on the veins beneath, too. Inflorescences axillary
(sometimes the subtending leaf gone), on the
mainland of Asia and in Sabang generally a
simple, rather densely-flowered raceme to 9cm
long 114 cm wide, initially resembling a cone due
to the conspicuous bracts which are persistent,
in Borneo and Java a lax-flowered panicle to 10 cm
long, 314 cm wide, with an occasional leaf or only
stipules subtending the first branches, the bracts
less conspicuous, and caducous; axes sometimes
puberulous, bracts 2-3 mm, striate, pedicels 2-4
(-6) mm, jointed in the basal or apical part.
Flower when young conical, the calyx covering
the basal half. Sepals thin, (sub)equal, more or
less triangular, surface glabrous or pubescent,
margin ciliate. Petals thin, straight, (214-)314-4
(-5'4) by 1-3 mm, glabrous, sometimes sparsely
ciliate. Androecium consisting of a filamental
tube 0.1-0.3 part of the total length, filaments
0.1-0.4 part, mostly glabrous, anthers 0.5-0.6
part, bearded at the base, the dorsal appendage
broad, triangular, nearly as long as the thecae or
somewhat longer, ventral appendages nearly
reaching the top of the dorsal one. Ovary mostly
glabrous, sometimes sparsely hairy, style glabrous,
ovules 6. Fruit in Asia (few known) globose before
dehiscence, 114-114 cm @ with seeds subglobose
4mm g, in Borneo (well-known) on woody stalk,
glabrous, before dehiscence 3-sided with blunt
corners, 234-4 cm Jong, 214 cm wide, the base not
or scarcely narrowed, valves leathery, veined,
mostly 2-layered after dehiscence, with woody
back; the seeds obconical, c. 8 mm long, 7 min @.
Seeds to 6 in number, with a circular depression
at the place of insertion, and evenly straw-co-
loured.
Distr. In SE. and Peninsular Burma; Indo-
China (Laos: near Vientiane, Cambodia, and Blao
NE of Saigon); Hainan (one coll.);in Siam at 15° N
in the West, and further S in the SW and the Pe-
ninsula, also in SE. Siam. In Malesia in Sabang I.
(N off Sumatra), Malay Peninsula (Perak and
Trengganu), W. Java, Borneo (northern central
Kalimantan, Sarawak, Brunei, and very common
in Sabah); scattered in all other regions mentioned.
The sole record from India, Assam, is uncertain,
KANJILAL & Das having “not found it any-
where in this Province”’.
190
Ecol. Mostly primary forest, but also from
secondary, bamboo and swampy forest and forest
edges, on clayey or loamy soil, also from lime-
stone and basalt, often by streams mostly at
low altitude but in Hainan from 1000 m.
In the monsoon area generally at least partly
deciduous, the fresh leaves then cropping out at
flowering time.
Notes. A shrub, sometimes climbing or spraw-
ling, ((44—-)1144—4 m, to a treelet or small unbuttres-
sed tree 3-12 m. Leaves light green; calyx (green-
ish) white, corolla (yellow) white, anthers yellow,
the dorsal appendage white with brown base.
Fruit green when more or less ripe, to light brown
when fully ripe.
The species is very polymorphic and looks
different in various parts of its area, but although
in Sabah it is both strikingly common and uni-
form (‘horneensis’), there is not enough reason for
the recognition of subspecies. The material from
Java (‘gaultheriiflora’) resembles that of Sabah,
although the leaves in the former are smaller. The
material of R. macropyxis (two out of the four
collections outside Sabah) with its domatia, hairy
leaves 7-9 cm long, and hairy twigs is unique for
Borneo, but agrees quite well with CHEW 63
from Malaya. The Sabah type of inflorescence is
also found in ScHmip anno 1961 from Blao
in S. Vietnam. Moreover, throughout the area,
all the examined flowers have a great mutual re-
semblance.
9. Rinorea virgata (THw.) O.K. Rev. Gen. Pl. 1
(1891) 42; Jacoss, Blumea 15 (1967) 135. —
Scyphellandra virgata THw. En. Pl. Zeyl. (1858)
21. — Alsodeia minutiflora Rip. J. Str. Br. R.
As. Soc. 59 (1911) 71; Fl. Mal. Pen. 1 (1922) 131.
Growth apparently in flushes. Twigs mostly long,
slender, but rarely crooked; when young angular,
greenish, often puberulous with curved hairs.
Branchlets terete, with longitudinal ridges of
brown-grey bark; the very short axillary brachy-
blasts sometimes producing an extra, smaller
leaf. Leaves generally distichous, on the same plant
variable. Stipules triangular to subulate, 1-2 mm
long, brownish. Petiole 1-5 mm, hairy as the twig.
Blade firmly herbaceous to thinly coriaceous,
more or less rhombic, 1—7(—1314) by 34-3(-6) cm,
index 1.7-4.7; base cuneate to rounded, top
acute to obtuse or sometimes acuminate, oc-
casionally mucronate; midrib slender, above
mostly prominent, major veins 2-6 on either side,
thin; no domatia; reticulation lax and irregular;
margins dentate; surfaces sometimes with a few
hairs on the midrib above, otherwise glabrous,
dull, concolorous, greenish. Dioecious; flowers
with a few apparently spirally arranged on the
knob-like brachyblasts; bracts persistent, tri-
angular, about 4% mm long, brownish; pedicels
slender, 3-6 mm, jointed near the base, puberulous.
Sepals more or less equal, thin, often triangular,
1-2 by 34-1 mm, sometimes hairy. Petals thin,
134-3144 by 34-144 mm, glabrous, sometimes the
top recurved, the margin sometimes undulate.
Androecium about as long as the petals, glabrous,
FLORA MALESIANA
(ser. I, Valea
in the 3 flowers the anthers sessile, about as wide as
the petals, with outside at the base an elongate
scale 14-14 mm long; thecae 0.40.5 part of the
total length of the anther, dorsal appendage ta-
pering towards the top. ventral appendages none
or vestigial. In the 2 flowers the staminodes sub-
sessile, sometimes narrower than the petals, the
basal gland at the outside minute, the thecae abor-
tive. Gynoecium glabrous, as long as the petals;
in the 2 flowers the ovary globose with thick style
and (5—)6 ovules; in the g flowers ovary none,
only a slender style rising from a flat receptacle.
Fruit before dehiscence (sub)globose, c. 1 cm g,
after dehiscence valves spreading, olive-green,
leathery, veined. Seeds subglobose, 214-3 by 2-214
mm, even straw-coloured, with distinct raphe.
Distr. Ceylon; Burma, NE. part: Lace
5462; Siam (scattered, not in the E. part); Laos
(near Pak Lay W of Vientiane: KINGDON WARD
9026); S. Vietnam (scattered S of 12°30’); Hainan.
In Malesia: Perlis in the very NW of the Malay
Peninsula.
Ecol. In more or less open evergreen forest, in
thickets, in teak plantations, on dry slopes and in
ravines, mostly on rocky, sometimes on sandy soil,
up to 600 m.
Note. A shrub (0.2—)0.5—1.5(-4) m, rarely
a tree 2m. Leaves paler green beneath. Flowers
white.
10. Rinorea macrophylla (DECNE) O.K. Rey. Gen.
Pl. 1 (1891) 42; Jacoss, Blumea 15 (1967) 135. —
Alsodeia macrophylla DECNE, Nouv. Ann. Mus.
Hist. Nat. Paris 3 (1834) 468, t. 19 = Herb. Timor.
Descr. (1835) 100, t. 19; Mig. Fl. Ind. Bat. 1,
2 (1859) 115; OUDEM. in Mig. Ann. 3 (1867) 69. —
Alsodeia horsfieldii Mia. Fl. Ind. Bat. 1, 2 (1859)
117; Mig. Ann. 4 (1869) 216; Back. Schoolfl.
(1911) 216. — Alsodeia dasycaula Miq. Sumatra
(1860) 391; Ann. 4 (1869) 216; Rip. FI. Mal.
Pen. 1 (1922) 129. — Alsodeia horsfieldii TuRCZ.
Bull. Soc. Nat. Moscou 36 (1863) i 559. —
Alsodeia membranacea KING, J. As. Soc. Beng.
58, ii (1889) 402; Ann. R. Bot. Gard. Cale.
5 (1896) 122, pl. 140B. — R. horsfieldii (MtQ.)
O.K. Rev. Gen. Pl. 1 (1891) 42; Koorp. Exk.
Fl. Java 2 (1912) 625; J. J. SmirH in K. & V.
Bijdr. 13 (1914) 68; Back. & Baku. f. Fl. Java
1 (1964) 194. — R. palawanensis MeErRR. Philip.
J. Sc. 1 (1906) Suppl. 210; En. Philip. 3 (1923) 104,
incl. var. mollis. — Alsodeia hispida Riv. J. Str.
Br. R. As. Soc. 6 (1912) 50. — R. dasycaula
(Miq.) Cras, Fl. Siam. En. 1 (1925) 88; GAGN.
Fl. Gén. I.-C. Suppl. 1 (1939) 194. — R. yatesii
Merr. Pap. Mich. Ac. Sc. 19 (1934) 173.
Growth apparently gradually; internodes of
similar length. Young parts velvety with persistent
brownish hairs to sparsely puberulous. Twigs
initially dark purplish tinged, later greyish, length-
wise striate, terete, comparatively thick with wide
pith, often slightly zig-zag, stipular scars indistinct.
Leaves mostly distichous. Stipules not always ad-
pressed, 114-5(-6) mm, subulate and not visibly
striate, often late caducous. Petiole 34-2 cm. Blade
before unfolding curved inward like a bow, of
1971]
papery texture, 1014-25 by 4-814 cm, widest at
the middle to well above, index (2.2—)2.6—-3.9
(4.2); base more or less tapering and slightly
decurrent, top acuminate; midrib and particularly
the veins thin, flat above, no domatia, about
7-10(-12) major veins on either side converging
within the margin, secondary veins and reti-
culation Jax and rather irregular, dark-tinged;
margins from base to top more or less distinctly
dentate-serrate; surfaces dull and mostly con-
colorous in variants of brown and green, glabrous
to sparsely hairy on the midrib and veins mainly
beneath. Inflorescences mostly densely hairy, in
structure like those of R. bengalensis but hypso-
phylls more scale-like, the axes no more than 4
in number, with c. 5—25 flowers each and these
but seldom in a clearly distichous arrangement,
the pedicel scars mostly obscure; bracts minute,
triangular; pedicels 3-10 mm, jointed near the
base. Sepals + equal, fleshy, 114-3 mm long,
bluntly triangular to elliptic, dark-coloured, hairy
outside. Petals fleshy, 244-514 mm long, widest
about the middle, more or less diverging, top
sometimes recurved, more or less hairy outside.
Androecium often glabrous, consisting of a fila-
mental tube 0.2-0.4 part of the total length, some-
times with a few hairs, filaments up to 0.2 part,
rarely hairy, and anthers 0.5—0.8 part, the dorsal
appendage broad, triangular, 1-2 times as long
as the thecae, ventral appendages tiny to half
as long as the dorsal one, sometimes wanting;
sometimes a few hairs on the base of the thecae
and on the back of the connective between.
Ovary hairy, each carpel with 1 ovule; style
glabrous. Fruit subtended by the dry spreading or
reflexed sepals and petals, before dehiscence
(sub)globose, c. 114 cm @, valves leathery, dark
purplish brown, sparsely hairy; seeds 3, c. 6-7
by 5mm, glossy brown, with distinct raphe.
Distr. Asia and Malesia; in the very east of
Siam and in Indo-China at + 16-17° N, SE. Siam,
and Peninsular Siam southwards to Pahang in the
Malay Peninsula (also Langkawi Is.); Sumatra
(East Coast Res. and Lampongs, ? Palembang);
Java (frequent on Nusa Kembangan, rare in
Central Java, also Kangean Is.); Lesser Sunda Is.
(Timor); Philippines (on the smaller islands S of
the 12th parallel; Palawan, Culion, Guimaras,
Cebu, Jolo). Scattered everywhere.
Ecol. Light forest, secondary forest, teak forest,
jungle, forest edges, mostly on limestone, some-
times on other rich soils, below 600 m.
Notes. A shrub lax!ly branched often crooked
(although in the herbarium many twigs are long
and simple) 1-3 m tall, to a tree 6 m tall. Leaves
bright green. Flowers with pale green calyx and
yellow-green-white corolla. Fruits first pale
green, greyish brown when ripe.
Occasionally the fruit 4-merous with 4 seeds.
AI. Rinorea iliaspaiei Jacons, nov. sp.
Stipulae longe-triangulares, persistentes. Folia
majora angusta minute serrata, basi obtusa apice
acuminata, Flores majores, staminibus glabris
anthera appendice dorsale magna late triangulare
VIOLACEAE (Jacobs & Moore)
191
appendicibusque ventralibus distinctis, ovario cy-
lindrico dense pubescente ovulis 9. Fructus maior
apicem versus pubescens seminibus 9. — Typus:
S 28099 ILiAs Pate (E, K, L holo, SAR, SING),
Borneo, Sarawak, Serian subdistr., Bk. Emperan
Ra’a, fl. 30-IX-1968.
Branchlets straight, with short internodes and
distinct leaf scars, bark greyish. Stipules rather
adpressed, long-triangular, dull dark purple-
brown, to 12-15 mm long (shorter on the smaller
branchlets), base half-amplexicaul, more or less
distinctly keeled and/or striate, initially with
minute hairs, long persistent. Petiole 3—7 mm.
Leaf blade coriaceous, to (10—)15—27 by (2-)3-614
cm, widest somewhat above the middle, index
(3.5—)3.8-5; base blunt and slightly unequal, top
gradually and long-acuminate; midrib narrow,
prominent on both sides, major veins c. 13-17 on
either side, with often in between a secondary vein
half as long, the major ones converging and con-
nected to an intramarginal vein, reticulation rather
irregular; margin with minute dark coloured teeth
c. Y%-\4 cm apart; surfaces dull, above olive-
green to brownish, beneath somewhat paler,
glabrous (youngest leaves unknown); no domatia.
Inflorescences axillary (sometimes the subtending
leaf gone), axis single or branched, 2—3 cm long;
bracts like the stipules but smaller; pedicel c. 4
mm, initially with minute hairs, jointed in the
apical part. Sepals rather unequal, the outer
ones fleshy, 4mm @, outside with some adpressed
hairs, the inner ones membranous, 6 by 5mm,
glabrous but ciliate at the top. Peta/s not very
fleshy, rather concave, glabrous, 8 by 3 mm, top
slightly cucullate. Androecium 3 mm high, gla-
brous, filamental tube c. 0.3 part of the total length,
free portion of the filaments c. 0.2-0.3 part,
anthers 0.4-0.5 part, the dorsal appendage broad,
triangular, somewhat longer than the thecae,
ventral appendages nearly reaching the top of the
dorsal one. Ovary cylindrical, densely soft white
hairy; style glabrous; ovules 9. Fruit (only known
after dehiscence) with thick leathery valves
214-3% cm long, wrinkled and, especially towards
the top, with short soft patent hairs. Seeds 9 in
number, obconical, 8 by 7 by 6mm, with a
circular depression at the place of insertion, evenly
straw-coloured.
Distr. Malesia: Borneo (Sarawak, 4 collections).
Ecol. Mixed dipterocarp and kerangas forests
at 100-600 m, on sandstone or clayey soil or shale.
Fl. Sept., fr. March, Aug., Nov.
Notes. Tree 2-4 m tall. Flower buds white,
when open yellow with reddish white stamens.
Ripe fruit green.
Close to R. longiracemosa (to which S 18905
and SF 35738 were reckoned in the Identification
List; actually they belong here, like S 20935),
and to R. macrantha; the differences have been
pointed out in the Latin diagnosis.
Excluded
Alsodeia chrysodasys Mi1Q. Sumatra (1861) 390 =
Paropsia vareciformis (Grire.) Mast, (Flacoartia-
ceae),
192
FLORA MALESIANA
[ser. I, vol. 71
Alsodeia dubia Em. Leafi. Philip. Bot. 8 (1915)
2875 = Trigonopleura malayanum Hook. f. (Eu-
Phorbiaceae).
Alsodeia glabra BURGERSD. in Migq. Pl. Jungh.
(1852) 122. — Rinorea glabra (BURGERSD.) O.K.
Rev. Gen. Pl. 1 (1891) 42 = Celastrus pani-
culatus WILLD. (Celastraceae).
Eriobotrya bengalensis (ROxB.) HOOK. f. (Rosaceae).
Rinorea pulgarensis ELm. Leafil. Philip. Bot. 5
(1913) 1849 = Diospyros cauliflora BL. (Ebena-
cede).
Vareca moluccana Roxs. FI. Ind. ed. 2, 1 (1832)
647 = Pittosporum moluccanum (LAMK) Mia.
(Pittosporaceae). In Fl. Mal. I, 5 (1954) 33 it was
Alsodeia grandis Mig. Sumatra (1861) 391 = but tentatively assigned to Rinorea.
2 AGATEA
A. GRAY, Proc. Am. Ac. Arts Sc. 2 (1852) 323; Bot. U.S. Expl. Exp. 1 (1854) 89;
BRONGN. Bull. Soc. Bot. Fr. 8 (1861) 80 (‘Agation’); B. & H. Gen. Pl. 1 (1862) 118;
BRONGN. & Gris, Ann. Sc. Nat. V, 1 (1864) 346; GumL. Ann. Mus. Col. Mars.
II, 9 (1911) 98; Bull. Mus. Hist. Nat. Paris 17 (1911) 350; ibid. 26 (1920) 362;
F. SARASIN & Roux, Nova Caled. B I, 3 (1921) 184; BAKER f. J. Linn. Soc. Bot.
45 (1921) 263; MeLcu. in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 360; GuILL.
Bull. Soc. Bot. Fr. 89 (1942) 20; A. C. Smitu, Sargentia 1 (1942) 57.
Scandent shrubs to lianas, innovations mostly laxly pubescent, indumentum
sometimes persistent. Twigs slender, with dull dark violet bark finely longitudinally
striate. Leaves in a spiral, petiolate. Stipules deltoid, c. 1mm long, persistent,
often inconspicuous. Leaf blade thinly to thickly coriaceous, c. 9-13 by 4-9 cm,
index c. 1.6—-2.2(—2.6); widest about the middle to somewhat lower; base acute
to rounded, top acute to abruptly acuminate with a short tip; veins (3—)5—7(-10)
on either side, reticulation more or less distinct; margin entire to crenate; surfaces
dull, glabrous. Inflorescences racemes but mostly panicles, few to 30cm long,
axillary and terminal mostly combined, pubescent; bracts small, triangular; pe-
dicels with 2 bracteoles above the base, articulated higher up. Flowers bisexual,
zygomorphic. Sepals subequal, 1-2 mm, covering a minor part of the bud, not
persistent. Petals unequal, sessile, the posterior also outer pair c. 3-4mm, sym-
metrical, the middle pair somewhat larger, asymmetrical, the inner, anterior one
c. 7-10 mm, the base gibbose, the apical part wide, thus making a lip, in bud
longitudinally rolled up, surface sometimes woolly above. Androecium equalling
the smallest petals, stamens subequal, (2—)4 connate at the base and converging
at the top but the posterior one apart; the two anterior ones at the base with a
fleshy outgrowth into the sac of the odd petal; filaments broad, flat; inner locules
longer than the outer ones, their thecal appendage small, single or bifid or double
or vestigial; connective appendage about twice as long as the anther, triangular,
brown-membranous. Gynoecium slightly exceeding the androecium, ovary sub-
globose, sometimes hairy, with 3 placentas and many ovules; style S-curved, gla-
brous, stigma anterior, clavate. Fruit (see note) a capsule, elongate, 144-5 cm,
with 3 leathery to woody valves. Seeds many, imbricately arranged, flat, with an
irregularly elliptic wing; testa with a hard black inner layer and membranous
yellowish outer layer.
Distr. New Guinea, Solomons, New Caledonia, Fiji, Tonga. Number of species probably 1; see below.
Ecol. Primary rain-forest in the lowland.
Notes. The above generic description covers the whole diversity of the genus; the specific description
applies to the New Guinea materials only. For the insignificant differences with the conspecific A. salo-
monensis MERR. & PERRY, J. Arn. Arb. 24 (1943) 209, see the description of the only collection, KAJEWSKI
2309 (A! BO! BRI! P!), from Bougainville.
1971] VIOLACEAE (Jacobs & Moore) 193
A cursory inspection of several dozens of Fiji and New Caledonian plants revealed that no two spe-
cimens match in all details — in climbers not an unusual feature. But while all plants from Fiji were
placed under A. violaris, in New Caledonia, where there is no more diversity than in Fiji, 7 species
were described, on trifling characters and mostly without fruits; see GUILLAUMIN, Bull. Soc. Bot. Fr. 89
(1942) 20. The differences can be summarized: in Fiji generally the young parts are sometimes glabrous,
the stipules sometimes wanting, the leaves acute and tending to be ovate, with (sub)entire margin, the in-
Fig. 4. Agatea violaris A. Gray. a. Habit, x %/, b. flower, «4, c. the odd petal, from above, 6, d. an-
droecium and stigma, from the right, x6, e. stamens, from inside, «8, /. gynoecium, from the right,
x6, g. fruit, immature, x 2/4, A. another fruit, mature, with i. seeds, both x %, j. leaf axil, x6 (a, g, /
Brass 14057 from New Guinea, b-f A. C. Smiru 66, h-i A. C. SmirH 6287, both from Fiji).
194 FLORA MALESIANA (ser. I, vol. 72
florescence up to 7(—14) cm, the lip 5-7 mm; in New Caledonia the young parts are mostly hairy, the sti-
pules always present, the leaves acuminate and widest at the middle, with often an incised margin, the in-
florescences up to 6-20 cm, the lip 6-914 mm. The few fruits collected in New Caledonia are 12-34 mm
long, the valves thick-coriaceous and covered with a grey felt, the seeds are c. 11 by 6-7 mm. The dozen
or so fruiting numbers from Fiji have valves 23-50 mm long, thin-woody and mostly glabrous but in
SEEMANN 12 (unripe) and A. C. SMITH 346 (ripe) they are more or less felty, too; the seeds are 13-22 by
9-10 mm, depending on the size of the fruit. As the fruit characters seem to overlap, and regional dif-
ferences are so slight, a comprehensive study of this genus may well result in the retention of only one
polymorphic species, A. violaris. MELCHIOR’s sections Macrobotrys and Euagatea, based on subtle dif-
ferences in the filamental gland, are to be reduced anyway.
BRONGNIART’S alteration of the name to Agation for fear of confusion with Agathaea (Compositae)
seems unnecessary.
A fine description of the ripening and dehiscence of the fruit was given by BRONGNIART, Bull. Soc.
Bot. Fr. 8 (1861) 78; the same text occurs in BRONGN. & Gris, Ann. Sc. Nat. V, 1 (1864) 346-350.
The ‘‘wing”’ of the seed is in fact a flattened and enlarged part of its funicle, which grows out mainly
towards base and top, its irregularities in shape due to cramming in the fruit. The approximately trian-
gular seed is pressed against the outside of its wing, dull grey-brownish as is the rest of the wing outside;
the wing otherwise glossy dark purple-blackish. When ripe, the valves, firm in texture and boat-shaped,
split while the strap-shaped basal part of the funicles remain attached to the receptacle, tearing loose
gradually towards the top.
1. Agatea violaris A. Gray, Bot. U.S. Expl. Exp. 1
(1854) 89, t. 7. — A. macrobotrys K. ScuH. & LAuT.
Fl. Schutzgeb. (1901) 453, t. 14; MERR. & PERRY,
J. Arn. Arb. 24 (1943) 208. — A. salomonensis
Merr. & Perry, J. Arn. Arb. 24 (1943) 209. —
Fig. 4.
Large climber. Twigs laxly pubescent, gla-
brescent, pith narrow. Stipules deltoid, 1 mm,
brown-purple, fimbriate. Petiole 1144-2144 cm,
glabrescent. Leaf blade thin-coriaceous, 9-1214
by 514-634 cm; base rounded, top gradually
acuminate, the tip c. 1 cm; midrib narrow, sub-
prominent, veins 5—7 on either side, rather parallel
and arcuating, reticulation fairly distinct on both
sides; margin subserrate; surfaces dull brown-
greenish, glabrous. Inflorescences axillary, sparsely
branched racemes to 8(—20) cm long, also terminal
to 17(—-20) cm long, fulvous pubescent; pedicels
Sepals 1-2 by 1 mm. Petals: the smallest 314-4
by 114 mm, the middle ones 4 by 2 mm, the lip
7(-9) by 5mm, hairy, with two extra hair tufts
distally near the basal sac. Stamens 2-3 mm;
filaments narrower than the anthers; fleshy
appendages about as long as the anther cells;
thecal appendage small, sometimes bifid. Ovary
sparsely pilose, size c. 1mm. Fruit (immature)
344 by 134 cm, ellipsoid, brown, glabrous.
Distr. Pacific: Fiji, Tonga, Solomons; in
Malesia: northern New Guinea (West: Rouffaer
and Idenburg R.; East: Bismarck Mts).
Ecol. Rain-forest, also marginal growth of
flooded rain-forest, at low elevation; in Fiji up
to 1000 m.
Note. Besides BrAss 14057 and DocTERS VAN
LEEUWEN 9708, I had the description of the type,
Ropatz & KLINK 236, probably lost in B, from
at some mm distance, c. (3-)5-8 mm long. which the figures in brackets are derived.
3. HYBANTHUS
Jaca. En. Pl. Carib. (1760) 2; BAILL. Bot. Médic. 2 (1884) 841; MELCH. in E. & P.
Nat. Pfl. Fam. ed. 2, 21 (1925) 357; Doin, Bibl. Bot. 89 (1928) 982-985; SCHULZE,
Bot. Jahrb. 67 (1936) 437-489; Morton, Contr. U.S. Nat. Herb. 29 (1944) 74-82;
SPARRE, Lilloa 23 (1950) 522-574, 4 pl.; RoBson, Bol. Soc. Brot. 32, ii (1958)
164-170; H. Perr. in Humb. FI. Madag. fam. 139 (1959) 3-10, f. 1: 2-11. — Solea
SPRENG. in Schrad. J. Bot. 4 (1800) 192. — Ionidium VENT. Jard. Malmais. (1803)
sub t. 27; B. & H. Gen. Pl. 1 (1862) 117; Bogert. Hand]. 1 (1890) 65.
Small gnarly shrubs or half-shrubs or herbs, rarely trees up to 8m, in a few
species the twigs thorny and microphyllous. Innovations sprouting from axillary
or terminal buds, simple themselves, more or less densely covered with simple,
rarely stellate hairs. Stipules generally small and mostly caducous. Leaves in a
spiral or rarely (sub)opposite, often herbaceous, sometimes leathery, more or less
sessile with a tapering base, to c. 15 by 4cm, but mostly smaller to sometimes
almost needle-shaped, margin entire to incised, rarely thickened underneath, the
1971] VIOLACEAE (Jacobs & Moore) 195
Fig. 5. Hybanthus enneaspermus (L.) F.v.M. in Sumbawa, Lesser Sunda Islands (photogr. DE Voocp).
teeth sometimes glandular. Flowers generally solitary in the leaf axils, rarely in more
or less reduced axillary cymes or dichasia up to several cm long or in raceme-like
elongated monochasia, rarely in a terminal leafy panicle; bracts mostly present;
pedicel slender, with 2 bracteoles beneath an articulation, the top curved down-
wards to give the flower an horizontal position. Flower 0.3—2 cm, bisexual,
but rarely cleistogamous, zygomorphic. Sepals subequal, small, more or less
triangular, persistent, rarely the margin with deep incisions. Pefa/s unequal, per-
sistent, posterior ones small and straight, middle ones longer and falcate, anterior
one extended to a lip, with a claw more or less deeply saccate to shortly spurred
and distinct blade often hairy above. Filaments free or seldom partly connate,
sometimes short; the anterior pair with a patent gland at the base outside; anthers
with rounded to elongate thecae (which rarely have an appendage at the top)
and a distinct membranous connective appendage. Ovary with 3 placentas bearing
196 FLORA MALESIANA
[ser. I, vol. 7?
Fig. 6. Hybanthus enneaspermus (Ie)
x6, e. filaments and pistil, also fro
fruit surrounded by floral parts, x
m the right, x8, f. stamen from inside,
3, i. seed, X8, in j. from above (a, d-g
Ramos, h-j JENSEN 313).
F.v.M. a., b. Two different plants, x 7%, ¢. stipules, x8, d. flower,
12, g. pistil, x8, 4. open
Brass 8434, b, c BS 27307
1971] VIOLACEAE (Jacobs & Moore) 197
3-24 ovules; style S-shaped with anterior stigma. Fruit subtended by the dried up
flower, c. | cm in size, subglobose before maturity, capsular with 3 leathery valves.
Seeds a few mm, ellipsoid, mostly with a small caruncula.
Distr. Tropical and subtropical regions of the world, preferring a dry period. The number of species
that will stand in a critical monograph is hard to estimate; there may be about 50 in the New World,
some in Africa and Madagascar, some in Australia, a few woody ones in New Caledonia. The one
Malesian species is palaeotropical (seems not to occur in New Caledonia) and possibly pantropical.
Notes. The literature cited under the genus refers chiefly to the most recent revisions in all parts of the
world. The name was in 1905 conserved over Calceolaria LoEFL. 1758. The above description applies to
the entire genus; it was largely based on SCHULZE. The specific description below was made on material
from Malesia, where only part of the diversity of the species is represented.
1. Hybanthus enneaspermus (L.) F. v. M. Fragm.
Phyt. Austr. 10 (1876) 81; BAck. Onkr. Suiker.
(1930) 454, pl. 429; Rosson, Fl. Zambes. 1
(1960) 254, t. 40; TENNANT, Kew Bull. 16 (1963)
431. — Viola enneasperma L. Sp. Pl. (1753) 937.
— lonidium enneaspermum (L.) VENT. Jard.
Malmais. | (1803) sub t. 27; GiNG. in DC. Prod. 1
(1824) 308; DaLz. & Giss. Bombay FI. 1 (1861)
12; OuDEM. in Miq. Ann. 3 (1867) 72; Mia. Ann. 4
(1869) 217; Merr. En. Philip. 3 (1923) 106. —
Tonidium heterophyllum VENT. Jard. Malmais. 1
(1803) sub t. 27; Back. Schoolfl. (1911) 66. —
Tonidium frutescens Bui. Bijdr. (1825) 58. —
Tonidium thymifolium PrResL, Rel. Haenk. 2 (1835)
97, non Bot. Bemerk. (1844) 11, nec MERR. Philip.
J. Sc. 10 (1915) Bot. 191. — Jonidium molle
Turcz. Bull. Soc. Nat. Moscou 27, ii (1854) 338.
— fonidium zippelii OupemM. in Miq. Ann. 3
(1867) 73. — H. suffruticosus (L.) BAIL. Bot.
Médic. 2 (1884) 841; Back. & BAKH. f. Fl. Java
1 (1963) 194. — Fig. 5, 6.
Herb 15-60cm tall, more or less profusely
branched, often woody at the base; young stems
angular, more or less pubescent. Stipules tri-
angular to subulate, 1-2 mm long, thin in texture
and light in colour, sometimes fimbriate towards
the base, the tip sometimes glandular. Leaves
spirally arranged, herbaceous, in one plant va-
rying somewhat in size; in different plants 2-7
cm long, 2-17 mm wide, index c. 2.4-20, widest
sometimes above the middle becoming spathu-
late; base decurrent sometimes into a petiole 5-7
mm, top acutish or blunt to gradually acuminate,
often with a mucro; margin entire to serrate or
remotely crenate. Flowers solitary; pedicels fili-
form, 34—2(—4) cm; bracteoles at the joint in the
upper part, their tip sometimes glandular. Sepals
subequal, triangular, membranous, 2-4 by 4-1
mm, glabrous or ciliate. Petals: posterior pair
244-344 by %4-1 mm, middle pair 244-4 by
1-114 mm, falcate; odd one c. 544-19 mm long,
the lip 4-10 mm wide, the base 1-2 mm saccate.
Stamens c. 2-3 mm long, the filaments and the
dorsal appendage each c. 0.3 times that long, the
anther c. 0.4 times; anterior stamens with a small
recurved fleshy appendage, glabrous or hairy,
‘sometimes their connective woolly outside. Pistil
glabrous, ovary subglobose, 4-11. mm @, the
placentas each bearing 2-5 ovules attached in
the middle; style 1'4-2mm long. Fruit sub-
globose, 4-5 mm 9, valves boat-shaped, eventually
Fig. 7. Distribution of Hybanthus enneaspermus(L)
F.v.M. in Malesia.
compressed, light green, each with 2—3 seeds which
are ellipsoid c. 2 by 1mm, _ straw-coloured,
lengthwise ribbed, at the base obliquely truncate,
with distinct raphe, the top with a shallow
crater.
Distr. Widely in Africa and Madagascar,
scattered in India and Ceylon, in SE. China and
Hainan, a few localities in the Indochinese
Peninsula, but not in Peninsular Thailand. In
Malesia: E. Java (Sumbarwaru and Baluran in the
very northeast), also Madura J.; Lesser Sunda Is.;
Borneo (Kudat in NW. Sabah); the Philippines
(locos Norte in Luzon, Cotabato in Mindanao,
Golo I.; Moluccas (Ternate, Kai Is.); New Guinea
(SW. Papua) and Thursday I. in Torres Straits;
tropical Australia. Fig. 7.
Ecol. With irregular frequency in monsoon
areas, at roadsides, savannahs, grasslands, pas-
tures, in the open or sometimes in shade, on sandy
or calcareous or volcanic soils; generally at low
altitudes, but in Timor to 750 m, in Flores to 1200
m.
Notes. After study of the type and other ma-
terials at the British Museum and Kew, I am
inclined to agree with Mr J. R. TENNANT’s
conception of the species. With his excellent paper
on African, Indian, and Chinese Hybanthus
handy, only the names recorded for Malesia need
to be evaluated here and the number of references
can be restricted to a minimum.
As Mr TENNANT found out, DaALzett &
198 FLORA MALESIANA fser. I, vol
Gipson, /.c., were probably the first to make a 195, t. 85, or on Jonidium frutescens ROEM. &
choice between the two Linnean epithets ennea- SCHULT. Syst. Veg. 5 (1819) 394; GinG. in DC.
sperma and suffruticosa, as early as 1861. Prod. 1 (1824) 311 (which in turn may have been
It seems that Jonidium frutescens BL. was based based on BURMAN’s species), from Ceylon and
on Polygala frutescens BURM. Thes. Zeyl. (1737) India, but BLUME refers to none of them.
4. VIOLA
LINNE, Gen. Pl. ed. 5 (1754) 402; Sp. Pl. (1753) 933; BL. Cat. Gew. Buitenzorg
(1823) 57; Bijdr. (1825) 57; KoRTHALS, Ned. Kruidk. Arch. | (1848) 357; BURGERS-
DUK in Mig. Pl. Jungh. 1 (1852) 118; OUDEMANs in Mig. Ann. Mus. Bot. Lugd.-
Bat. 3 (1867) 73; Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 217; Botsstieu & Ca-
PITAINE, Bull. Soc. Bot. Fr. 57 (1910) 337; BECKER, Beih. Bot. Centralbl. 34, i
(1916) 208; ibid. (1917) 373; Bot. Jahrb. 54 (1917) Beibl. 120, 156; Beih. Bot.
Centralbl. 36, ii (1918) 15; ibid. 40, ii (1923) 20, 69, 119; BECKER & MELCHIOR
in E. & P. Pfl. Fam. ed. 2, 21 (1925) 363; MELCHIOR, Bot. Jahrb. 62 (1929) 368;
Notizbl. Berl.-Dahl. 12 (1934) 205; STEEN. Bull. Jard. Bot. Btzg III, 13 (1934)
258; Riv. J. Bot. 73 (1935) 13; Lin, Taiwania 1 (1950) 269; D. M. Moore in
Fedde, Rep. 68 (1963) 81-86; Nova Guinea, Bot. n. 11 (1963) 177.
Perennial, rarely annual herbs (in Mal.). Leaves alternate, petioled. Stipules
free or adnate to petiole, persistent, often conspicuous, usually serrate or fimbriate.
Leaves suborbicular to linear-lanceolate, the margin serrate to crenate or subentire,
often glandular in indentations. Flowers bisexual, solitary, axillary, with a pair of
bracteoles usually in the upper half of the peduncles. Sepa/s equal, entire to den-
ticulate or fimbrio-dentate, prolonged into appendages below the point of their
insertion, persistent. Corolla zygomorphic. Peta/s unequal, the lower saccate or
spurred and usually broader than the others, the lateral pair smaller than the
upper pair, the lateral petals often, the others more rarely, bearded inside. Androe-
cium shorter than the petals; filaments connivent around the gynoecium; anthers
2-celled, subsessile, the 2 lower with appendages projecting into the spur; connecti-
ve produced into an apical appendage. Gynoecium as long as the androecium or
longer, the ovary glabrous or pubescent, with 3 placentas and ~ ovules; style
straight, curved or geniculate, filiform to conspicuously clavate, often lobed at
apex; with the stigma terminal or anterior and subterminal. Fruit a 3-valved lo-
culicidal capsule, subtended by dried-up calyx, globose to cylindrical or ellipsoidal,
4-16 mm long; valves boat-shaped, usually with thick rigid keels and thin sides
so that on drying they contract and forcibly discharge the seeds, glabrous or
pubescent. Seeds ~, usually ellipsoidal, glabrous, with leathery testa, usually with
terminal elaiosome.
Distr. About 400 spp. occurring in temperate regions throughout the world.
Ecol. Terrestrial, generally in open or lightly shaded places in the mountains above 1000 m, excep-
tionally descending to c. 250 m.
The elaiosome may contain oils attractive to dispersal of seeds by ants.
Notes. With the exception of V. biflora (sect. Dischidium G1NG.) and the introduced V. tricolor (sect.
Melan um Gina.), all species occurring in Malesia may be included in sect. Viola (sect. Nomimium GING.).
The separation of V. hederacea and its allies into sect. Erpetion (SwEET) BECKER (in E. & P. Nat. Pfl. Fam.
ed. 2, 21, 1925, 376) does not appear to be justified in view of the intermediates between this and sect.
Viola. The various subsectional groupings within sect. Viola (see BECKER in E. & P. /.c.) prove impos-
sible to apply effectively and have been omitted in this account. Any useful subsectional treatment must
await a monographic revision of the whole genus.
Chromosomes. There has been little cytological work on Malesian violets, and chromosome numbers
1971] VIOLACEAE (Jacobs & Moore) 199
1mm
14
Fig. 8. Style of all Malesian Viola species. The number sequence is the same as in the text, | referring to
1. V. tricolor, ete.
206 FLORA MALESIANA [ser. I, vole
are available for only’5 of the 16 native species. The counts on Malesian material were all made on plants
from New Guinea and were published by Moore in Fedde, Rep. 68 (1963) 81 and BORGMANN, Z. Bot. 52
(1964) 118. Chromosome numbers based on extra-Malesian material are indicated in the text.
Cleistogamous flowers. Except for V. tricolor, the species in Malesia produce during some, usually
the later, part of the growing season or under abnormal environmental conditions so-called cleistogamous
flowers. These have a shorter peduncle, a reduced corolla not expanded beyond the calyx; up to 3 anthers
may be aborted and the amount of pollen produced is greatly reduced, while the style is usually much
shorter than in chasmogamous flowers and may be much contorted. Such flowers produce plenty of seed,
from self-pollination. The variation within species resulting from this sort of reproduction often does not
conform to a pattern amenable to formal taxonomic recognition, and may partly explain the multi-
plicity of names applied to some taxa.
Chasmogamous (i.e. normal, expanding) flowers are essential for the reliable determination of most
violets. In the absence of suites of flowering specimens for comparison, fruiting or cleistogamous material
can rarely be confidently identified, particularly since, in such material, related species often show con-
vergence in leaf characters.
KEY TO THE SPECIES
(see fig. 8 for the shape of the styles)
1. Stipules, at least the upper ones, pinnatifid or palmatifid. Style expanded above into a globose head.
1. VY. tricolor
1. Stipules entire to long-fimbriate. Style not as above.
. Style hooked at apex to give stigmatic beak about as long as the diameter of the style. Capsule on
decumbent peduncle, not explosive... . . . 3. V. odorata
. Style not markedly hooked, stigmatic beak much shorter than the diameter ‘of the style. Capsule on
erect peduncle, explosive.
3. Flowers yellow. Leaves reniform to broadly ovate, apex rounded or obtuse. Stipules entire or
sparsely denticulate. Style with apex lobes fused behind, no stigmatic beak. . . . . 2. V. biflora
3. Flowers purple or white, rarely yellow (sp. 13).
4. Stems or stolons absent.
5. Spur 214-6 mm, 2-4 times as long as wide, eae e downwards curving, 2—5 times as long as
calycine appendages. . . . . 7. V. philippica
5. Spur up to 314 mm, up to 1, times as long as s wide, straight o or slightly upcurved, up to 3 times as
long as calycine appendages.
6. Calycine appendages (1—)2—4.8 mm, incised, often conspicuous, 14-34(-1) as long as sepal.
6. V. inconspicua
6. Calycine appendages 0—2(—214) mm, rounded or subdentate, not conspicuous, up to 1% (rarely
14) as long as sepal.
7. Leaves at least 16 mm, long-decurrent on petiole, glabrous... . . . . 5. V. betonicifolia
7. Leaves up to 16 mm, not or scarcely decurrent on Boule, with clusters of hairs in marginal
dentations on upper side. . . : ee ere AY ETT
4. Well-developed decumbent or ascending ‘stems ¢ or ‘stolons.
8. Style filiform.
I, SYMP aE Reith Ree Mele aye ais acetate ae en Poe ee aos eee 18. V. papuana
9. Spur up to 244 mm
10. Stipules long- fimbriate. Calycine pada at least 0.8 mm, pointed or denticulate, usually
HITSWtes Rigaca hey oe 12. V. pilosa
10. Stipules sparsely fimbrio- dentate. Calycine appendages up to 1mm, rounded, glabrous.
11. Leaves and flowers grouped in distinct rosettes at intervals along stolons. Stipules free.
17. V. hederacea
11. Leaves and flowers not grouped in distinct rosettes. Stipules half-adnate to petiole.
16. V. kjelibergii
8. Style clavate.
12. Leaves usually broader than long, arcuate or ovate, usually with prominent upturned basal
lobes, crenulate or dentate.
13. Style with separate rounded marginal lobes at apex. pens usually entire or shortly fimbrio-
dentate... mais 10. V. arcuata
13. Style with apex having flattened marginal lobes which 2 are 2 fused ‘posteriorally. Stipules long-
fimbriate. . . oop ce: ae ee javanica
12. Leaves usually longer than broad, if broader than long ‘then with rounded apex and promi-
nently round-crenate (see V. diffusa).
14. Leaves with hairs on upperside confined to zone towards margins, glabrous in central portion.
8. V. merrilliana
14. Leaves glabrous or, if hairy, hairs not distributed as above.
1971]
VIOLACEAE (Jacobs & Moore)
201
15. Leaves + twice or more as long as broad, elliptical to lanceolate-ovate. 14. V. curvistylis
15. Leaves up to 134 times as long as broad, suborbicular to ovate or elliptical.
16. Petiole conspicuously winged. Leaves orbicular to orbicular-ovate, rarely elliptical when
base cuneate to subcordate.
4. V. diffusa
16. Petiole not or scarcely winged. Leaves ovate to triangular-ovate, rarely elliptical when base
with narrow deep sinus.
17. Style not or slightly marginate at apex. .
17. Style distinctly marginate at apex.
18. Leaves white-hirsute above and beneath, elliptical to ovate.
12. V. pilosa
. 15. V. rupicola
18. Leaves glabrous or sparsely hirsute in marginal indentations or on veins, especially below,
ovate to triangular-ovate.
19. Lateral petals not bearded. .
19. Lateral petals bearded.
. 13. V. sumatrana
20. Sepals at least 4 by 1 mm, margins entire, appendages rounded or denticulate. Leaves
subcoriaceous, glabrous or sparsely hirsute on veins towards base.
11. V. javanica
20. Sepals c. 3 by 1 mm, margins denticulate, appendages dentate. Leaves herbaceous,
sparsely hirsute in crenations.
1. Viola tricolor LINNE, Sp. Pl. (1753) 935;
Back. & BAKH. f. Fl. Java 1 (1963) 195. — Fig. 8.
Annual, biennial or perennial; rhizome short
or none. Stems 10-40 cm, ascending or erect,
usually branched. Leaves ovate-lanceolate to
-elliptical, cuneate or subcordate at base, obtuse,
crenate. Stipules deeply and pinnately lobed,
terminal segment larger than the others, lanceolate,
entire or crenate, leaf-like. Flowers c. 2-3 cm,
violet, yellow or parti-coloured, very variable in
colour and size; peduncle 3-10 cm, exceeding
leaves. Sepals 7-14 by 2—5 mm, linear-lanceolate,
acute, glabrous; appendage 3-5 mm, prominent.
Petals obovate, 1-2 times as long as broad; spur
4—6 mm, variable up to twice as long as calycine
appendages. Style 2-214 mm, geniculate at the
base, clavate distally with globose apex having
prominent broad, anterior stigmatic beak. Cap-
sule 8-14 mm, ellipsoid, glabrous.
Distr. Widespread throughout Europe ex-
tending to Asia Minor and Himalayas, in Malesia:
garden-ornamental in mountain regions of Java
and the Philippines (Luzon), not known to be
naturalized. Fi. Jan.—Dec.
Note. These garden forms probably represent
derivatives [V.wittrockiana (= V. hortensis
auct.)| from hybridization with V. lutea Hups.
and V. altaica KER-GAWLER.
2. Viola biflora LINNE, Sp. PI. (1753) 936; BECKER
Bot. Centralbl. 36, ii (1918) 39; Lin, Taiwania 1
(1950) 280. — Fig. 8.
Perennial; rhizome horizontal or oblique, more
or less stout, roots on lower surface, leaves and
stems at apex. Stems up to 8 cm, slender, ascending
to erect, with leaves along their length and flowers
distally. Leaves 34-114 by 1-2'4 cm, 14-2 times
as broad as long, reniform to broadly ovate,
more or less deeply cordate, rounded to broadly
obtuse, subcrenate to repand-dentate, especially
on lobes, subcoriaceous, glabrous beneath,
sparsely hirsute towards margins and on veins
above, dark green; petiole '4—-Scm, slender.
Stipules 2-4 by 2-3 mm, ovate, acute, entire to
sparsely denticulate, glabrous, fuscous, free.
Flowers 8-12 mm, yellow with brownish-purple
9. V. mearnsii
veins; peduncle 1.4-8 cm, much exceeding leaves,
slender. Sepals 3-4 by c. 1 mm, linear to linear-
oblong, acute, glabrous or shortly ciliate on margin,
fuscous, with margin scarious; appendage up to
14mm, rounded, glabrous. Petals 2-3 times as
long as broad, obovate to almost oblong, basal
exceeding the others, laterals not bearded; spur
1-114 mn, cylindrical, obtuse. Style 1.2-1144 mm,
slightly geniculate at base, clavate distally, apex
with 2 lateral lobes fused behind, no stigmatic
beak. Capsule 4-6 mm, oblong to ellipsoid, gla-
brous.
Distr. Circumboreal, south to 40° N in North
America, to c. 38° N in Europe. Through Hi-
malayas to Japan, China, Korea, and Formosa, in
Malesia: N. Sumatra (Mts Losir and Kemiri).
Fig. 9.
Ecol. Alpine grassland, among rocks and grass
tussocks, 3000-3500 m. F/. Oct.—March.
3. Viola odorata LINNE, Sp. PI. (1753) 934; BACK.
& SLtoot. Handb. Theeonkr. (1924) 178, t. 178;
Back. & BAKH. f. Fl. Java 1 (1963) 195. — Fig. 8.
Perennial; rhizome vertical, stout, bearing ro-
sette of leaves and usually long, procumbent
rooting stems at apex. Leaves 21-6 by 214-61,
cm, orbicular-reniform to -ovate, deeply cordate,
rounded to obtuse, shallowly crenate to crenate-
serrate, glabrous or sparsely pubescent on veins
and margins; petiole up to 20cm. Stipules 8-12
by 3-5mm, ovate to ovate-lanceolate, usually
glandular-fimbriate, glabrous, free. Flowers 10-15
mm, purple or white; peduncles 5—14 cm, slender.
Sepals c. 5 by c. 2 mm, ovate, obtuse, entire, with
ciliate margins; appendage I—2 mm, dentate.
Petals broadly obovate, laterals bearded or not;
spur c. 4mm, straight or slightly upcurved. Style
c. 2mm, uncinate at apex. Capsule globose, pu-
bescent.
Distr. Throughout Europe except the extreme
north, Asia Minor, Caucasus, in Malesia: garden-
ornamental in mountain regions of Java and the
Philippines (Luzon); not known to be naturalized
but Backer & BAKHUIzEN/. Stated that “itis plant-
ed to prevent erosion by rain-wash’’. F/. May-Oct.,
perhaps all the year.
202
FLORA MALESIANA
[ser. I, vol. 7}
4. Viola diffusa GinGins in DC. Prod. 1 (1824)
298. — Fig. 8.
See for synonyms under the subspecies.
Perennial or annual; rhizome vertical, rather
thin, fibrillose, bearing rosette of leaves and flowers
and procumbent rooting stems. Stems up to 12 cm,
rooting at ends and producing dense rosettes of
leaves and flowers. Leaves 14-314 by 1-2 cm,
smaller on stolons, suborbicular to ovate or
elliptic, cordate to cuneate at base, subacute to
obtuse or rounded, crenate to serrate-crenate on
margin, long-decurrent on petiole; hirsute or
rarely glabrous; petiole 1-7 cm. Stipules 5-9 by
c. 1mm, lanceolate or ovate-lanceolate, long-
acute, dentate to fimbriate, green or pale brown,
free. Flowers up to 9mm, pale violet to almost
white; peduncle 114-6cm. Sepals 3-6 by 1-114
mm, lanceolate to ovate, acute, sparsely hairy,
fimbrio-ciliate, green; appendage c. ‘14mm,
rounded, sparsely fimbrio-ciliate. Petals 114-2
times as long as broad, obovate, the basal smaller
than the others, the laterals not bearded; spur
0.4 to c. 1 mm, obtuse. Style c. 114 mm, slightly
geniculate at base, clavate distally, apex with 2
lateral lobes, with small anterior beak between
lobes. Capsule 4-6 mm, ellipsoid, glabrous.
Distr. Himalayas, China, Indo-China, Japan,
Formosa, in Malesia: Philippines, New Guinea.
Fig. 9.
Fig. 9. Distribution in Malesia of Viola diffusa
GING. ssp. diffusa (circles), ssp. apoensis (ELM.)
D. M. Moore (dots), and V. biflora L. (+).
Ecol. Open places in montane forest and grassy
clearings, often along tracks and other disturbed
places, a weed of cultivation, 1400-2500 m. Fi.
Aug.—March, probably later.
This species comprises two subspecies, as
follows:
ssp. diffusa. — V. diffusa GING.; MERR. Philip.
J. Sc. 5 (1910) Bot. 201; Becker, Philip. J. Sc.
19(1921) 713; Beih. Bot. Centralbl. 40, ii (1923) 114;
Lin, Taiwania 1 (1950) 271. — V. tenuis
BENTH. in Hook. Lond. J. Bot. 1 (1842) 482. —
V. diffusa GING. ssp. tenuis (BENTH.) BECKER,
Philip. J. Sc. 19 (1921) 714, pro parte.
Leaves (114-)134-3'4 by (0.9-)1-2 cm, ovate
to ovate-elliptic, cuneate to shallowly cordate at
base, obtuse to acute, rarely rounded, with
crenate-serrate to serrate margins; petiole 1-114
times as long as the blade. Chromosome number:
2n = 26 (Japan, Miyas1, Cytologia 1, 1929, 28).
Distr. Himalayas, China, Indo-China, Japan,
Formosa, in Malesia: Philippines (Luzon: Mts
Data and Pukis).
ssp. apoensis (ELtM.) D. M. Moore, comb. nov. —
V. apoense E_o. Leafi. Philip. Bot. 8 (1915) 2878.
— V. diffusa GING. ssp. tenuis (BENTH.) BECKER,
Philip. J. Sc. 19 (1921) 714, pro parte; D. M.
Moore, Nova Guinea, Bot. n. 11 (1963) 181.
Leaves 34-—114(-2) by 1-114 cm, suborbicular to
broadly ovate, deeply, rarely shallowly, cordate
at base, rounded at apex, with round-crenate
margins; petiole 1—3.2 times as long as the blade.
Distr. Malesia: Philippines (Luzon: Mt Data;
Mindanao: Mt Apo), New Guinea.
Note. It is not known whether the two sub-
species occupy different habitats on Mt Data,
but some hybridization is likely since these po-
pulations, although readily distinguished, are
closer morphologically than other populations of
the two taxa. It is possible that they have come into
contact by recent introduction of one of the sub-
species, in view of the weedy habitats favoured by
the species.
5. Viola betonicifolia J. E. SMITH in Rees, Cyclop.
37, I (1817) n. 7. — Fig. 8.
See for synonyms under the subspecies.
Perennial; rhizome short, oblique or vertical,
rather stout, bearing rosette of leaves and flowers
at apex, acaulescent. Leaves 114-714 by 14-3 cm,
1-19 times as long as broad, linear-lanceolate to
triangular-hastate or triangular-ovate, cuneate,
truncate or widely and shallowly cordate with
basal lobes often laterally prominent, acute or
sometimes roundish obtuse, shallowly and distant-
ly crenate, sometimes dentate on basal lobes,
usually long-decurrent on petiole, glabrous; pe-
tiole 1-1014 cm. Stipules 2-7 by 14-1 mm, ovate-
lanceolate, acuminate, sparsely short-fimbriate,
glabrous, fuscous, adnate to petiole to 34 length.
Flowers 6-144 mm, white to purple with darker
veins; peduncle equalling or exceeding leaves,
glabrous. Sepals 3-614 by 0.6—-2.5 mm, ovate to
ovate-lanceolate, acute or acuminate, glabrous or
ciliate, green with scarious margins; appendage
0.2-214 mm, or absent, (144-)!4-+/, as long as
sepa], rounded to squarish or somewhat sub-
dentate, glabrous. Petals 114-3.4 times as long as
broad, obovate, laterals usually bearded; spur
0.6-314 mm, half to twice as long as broad, cy-
lindrical, obtuse, straight or slightly upcurved.
Style 144-244 mm, geniculate at base, clavate
distally, apex with prominent marginal lobes fused
behind, anterior stigmatic beak. Capsule 7-10 mm,
ellipsoid to oblong, glabrous.
Distr. Himalayas, India, Ceylon, China, Japan,
Indo-China, Australia, in Malesia: Sumatra,
E. Java (Mts Tengger and Idjen), Lesser Sunda
Fig. 10. Distribution in Malesia of Viola betonici-
folia J. E. SmitH ssp. betonicifolia (dots), an
intermediate in N. Queensland, and ssp. nova-
guineensis D. M. Moore (circles).
Islands (Alor, Bali, Timor), Celebes, Philippines
(Mindanao, Luzon), and New Guinea. Fig. 10.
Ecol. Usually in grasslands, sometimes in open
montane woodland of Eucalyptus, (650—)1100—
2100(—2700) m. Apparently sometimes a ruderal
and occurring along path margins and in cul-
tivated ground. F/. Jan.—Dec.
Notes. This species shows considerable va-
riation in leaf-shape, on the basis of which two
subspecies have been recorded.
V. betonicifolia differs from V. inconspicua in
having the calycine appendages less than half as
long as the sepals, in leaf-shape and in the lamina
generally being long-decurrent on the petiole.
It differs from V. philippica by its generally shorter
spur which is less than twice as broad as long, by
the heavy bearding of the lateral petals, and by its
leaf-shape.
The widespread references to V. patrinii in
Malesian literature concern V. betonicifolia and
V. inconspicua. In an analysis of this group
Becker (Bot. Jahrb. 54, Beibl. 120, 1917, 156)
has shown that the name V. patrinii DC. should be
restricted to the plant from northern E. Asia.
This species appears to be diploid (Mryay1, Cy-
tologia 1, 1929, 28) and some of the taxonomic
difficulties may be due to its genome being present
in the species further south, of which those known
cytologically are tetraploid or hexaploid.
ssp. betonicifolia. — V. betonicifolia J. E. SM. —
V. betonicifolia J. E. SM. ssp. australis BECKER,
Bot. Jahrb. 54 (1917) Beibl. 120, 166. — V.
betonicifolia J. E. SM. ssp. nepalensis BECKER,
Lc.; Philip. J. Sc. 19 (1921) 716; Merr. En.
Philip. 3 (1923) 105; Oosrstr. & H. J. LAM, Blu-
mea 5 (1945) 594. — V. patrinii DC. var. na-
paulensis DC. Prod. 1 (1824) 293. — V. caespi-
tosa D. Don, Prod. FI. Nepal. (1825) 205. — V.
patrinii DC. var. caespitosa Rivi. J. Bot. 73
(1935) 17. — V. patrinii (non DC.) Merr. Philip.
J. Sc. 5 (1910) Bot. 200, pro parte; Boissieu &
CAPITAINE, Bull. Soc. Bot. Fr. 57 (1910) 337;
Koorp. Exk. Fl. Java 2 (1912) 627, pro parte.
VIOLACEAE (Jacobs & Moore)
203
Leaves linear-lanceolate to triangular-ovate.
Chromosome number: 2n = 48, 72 (Australia,
Moore in Fedde, Rep. 68, 1963, 84).
Distr. As the species but in New Guinea only
in extreme west.
Notes. The northernmost (including the type
of ssp. nepalensis) and southernmost (including the
type of ssp. betonicifolia) populations cannot be
differentiated and intergrade continuously with
the various leaf-forms present throughout much
of Malesia, though there is a tendency for more
definitely triangular cordate forms in the centre
of the range. Confusion between this and the next
subspecies may result partly from hybridization
with V. inconspicua. Populations from N. Su-
matra have a conspicuously narrow cuneate leaf
which is 4-19 times as long as broad.
ssp. nova-guineensis D. M. Moore in Fedde,
Rep. 68 (1963) 82.
Leaves triangular-hastate with basal lobes pro-
minent laterally, even deltoid. Chromosome
number: 2n = 72.
Distr. Malesia: Sumatra, Timor, Philippines,
Celebes, New Guinea (except extreme west).
Note. This subspecies, which is centred in New
Guinea, intergrades with ssp. betonicifolia in
N. Queensland, and it is possible that many of the
triangular-leaved forms north from New Guinea
represent intermediates between the two subspecies.
6. Viola inconspicua BLUME, Cat. Gew. Buitenzorg
(1823) 57; Bijdr. 1 (1825) 58; KortTH. Ned.
Kruidk. Arch. 1 (1848) 357; Mia. Fl. Ind. Bat.
1, 2 (1858) 112; Becker, Bot. Jahrb. 54(1917) 167;
Philip. J. Sc. 19 (1921) 718; Mere. En. Philip. 3
(1923) 105; Ripx. J. Bot. 73 (1935) 14; BAcK. &
BAKH. f. Fl. Java 1 (1963) 195. — V. trinervis
Kortu. Ned. Kruidk. Arch. 1 (1848) 357;
Mia. Fl. Ind. Bat. 1, 2 (1858) 112. — V. confusa
CHAMP. ex BENTH. in Hook. J. Bot. Kew Misc. 3
(1851) 260, pro parte; Riv. J. Bot. 73 (1935) 14.
— V. patrinii (non DC.) Boissteu & CAPITAINE,
Bull. Soc. Bot. Fr. 57 (1910) 340; Merr. Philip.
J. Sc. 5 (1910) Bot. 200, pro parte; Koorp. Exk.
Fl. Java 2 (1912) 627, pro parte; BACK. & SLooT.
Handb. Theeonkr. (1924) 179, t. 179. — V.
serpens (non WALL.) RiDL. Fl. Mal. Pen. 1 (1922)
126, pro parte. — Fig. 8.
Perennial; rhizome vertical or oblique, rather
short and stout, with rosette of leaves and flowers
at apex, acaulescent. Leaves 134-9 by 1-744 cm,
1-2.2 times as long as broad, broadest at base,
triangular-ovate, cordate at base with usually
prominent, rounded basal lobes, acute, crenate to
subdentate, sometimes dentate on basal lobes,
decurrent to 4 length of petiole, glabrous or
sparsely pubescent in sinus; petiole 1-10 cm. Sti-
pules 3-10 by 1-2 mm, ovate-lanceolate, acute,
sparsely short-fimbriate, glabrous, fuscous, adnate
to petiole to % of length. Flowers 9-11 mm, pale
purple with darker veins; peduncle shorter than
to somewhat exceeding leaves, glabrous or pu-
bescent distally. Sepals 24-7 by 1-1.8 mm, ovate-
lanceolate, acute, glabrous or sometimes ciliate,
Fig. 11. Distribution of Viola inconspicua BL.
green with scarious margin; appendage (1!4—)2—
4.9mm, at least half as long as sepal, incised.
Petals 2-3 times as long as broad, obovate, the
laterals bearded; spur 1.8-2.6 mm, 1.1—1.3 times
as long as broad, #/; to 14 as long as calycine
appendages, cylindrical, straight, obtuse. Style c.
2mm, geniculate at base, clavate distally, apex
with large marginal lobes fused behind, anterior
stigmatic beak. Capsule 9-16mm, oblong, glabrous.
Distr. Himalayas, Burma, China, in Malesia:
Sumatra, Malay Peninsula (Penang), Java,
Philippines (Luzon). Fig. 11.
Ecol. Grasslands, fallow lands, path margins;
apparently normally ruderal, 500-1700m. FI.
Nov.—Febr.
Notes. Closely related to V. betonicifolia but
differing by its shortly decurrent lamina and by the
conspicuous leaf-like calycine appendages which
are more than half as long as the sepals. Inter-
mediates between the two species occur in several
places throughout the range of V. inconspicua.
These are considered to be hybrids rather than
clinal variants, because no geographical pattern
can be discerned, because the weedy habitats pre-
ferred by both species are conducive to inter-
specific crossing, and both species occur sym-
patrically in some Philippine and Sumatran lo-
calities. Only occasional collections, from Su-
matra and the Philippines, have chasmogamous
flowers and this, together with the apparently
weedy nature of V. inconspicua, supports the
suggestion of BECKER (1917) and of BACKER &
BAKHUIZEN/. (1963) that the species isa relatively
recent immigrant from continental Asia.
7. Viola philippica CAVANILLES, Icon. Descr. 6
(1801) 19; Mia. Fl. Ind. Bat. 1, 2 (1858) 113;
Becker, Bot. Jahrb. 54 (1917) 174, incl. ssp.;
Philip. J. Sc. 19 (1921) 719; Merr. En. Philip. 3
(1923) 105; BAck. & Baku. f. FI. Java 1 (1963)
195. — V. confusa CHAMP. ex BENTH. in Hook.
J. Bot. Kew Misc. 3 (1851) 260, pro parte; BACK.
Schoolfl. (1911) 66. — V. selkirkii (non PursH)
BoulissIEU & CAPITAINE, Bull. Soc. Bot. Fr. 57
(1910) 337. — V. edanoii BECKER, Philip. J. Sc.
19 (1921) 722; Merr. En. Philip. 3 (1923) 105. —
Fig. 8.
Perennial;
rhizome vertical or suboblique,
FLORA MALESIANA
(ser. I, volyg?
rather slender, bearing rosette of leaves and flowers
at apex, acaulescent. Leaves 1-4 by 14-3 cm, 1-2
times as long as broad, ovate-elliptic to ovate or
triangular-hastate, deeply to shallowly cordate
to subtruncate at base, with the basal lobes usually
somewhat converging, obtuse to rounded, round-
crenate to crenate-serrate, usually long-decurrent
on petiole, glabrous or puberulent beneath on
veins and margins; petiole 1-6 cm. Stipules 2-8
by 0.8-3mm, ovate-lanceolate, acute, shortly
fimbrio-dentate, glabrous or sparsely pubescent,
green to rather fuscous, adnate to petiole to 34
length. Flowers 914-1414mm, pale to dark
violet; peduncle usually much exceeding leaves,
glabrous or puberulent distally. Sepals 3—5.2 by
0.7-1.8 mm, ovate to ovate-lanceolate, acute or
rounded entire, glabrous or ciliolate, green with
scarious margin; appendage 14-2 mm, (14-)14-’%
as long as sepal, rounded to denticulate, glabrous
or sparsely pubescent. Petals 1.2—3 14 times as long
as broad, obovate, the laterals not or lightly beard-
ed; spur 214-6 mm, 2-4 times as long as broad,
2-5 times as long as calycine appendages, usually
slightly tapering and downwards curving. Style
114-2.4mm, geniculate at base, clavate distally,
apex with prominent lateral lobes partially fused
behind, anterior stigmatic beak. Capsule 5-8 mm,
ellipsoid, glabrous.
Fig. 12. Distribution of Viola philippica Cav.
Distr. Himalayas, India, Mongolia, China,
Burma, Korea, Japan, in Malesia: Sumatra, Java,
Lesser Sunda Islands (Flores), Celebes (Loha),
Philippines (Luzon, Mindanao). Fig. 12.
Ecol. Grasslands; path margins; sometimes
ruderal, 250-1700 m. F/. Oct.—July, probably all
year.
Notes. BECKER (1917, lc.) recognized two
subspecies distinguished mainly on leaf-shape,
viz ssp. munda BECKER, lamina lanceolate or oblong,
cuneate to subcordate at base, and ssp. malesica
BECKER, lamina broader and shorter with cordate
base. The type material contains leaves referable to
both these entities, as do collections from a variety
of areas (cf. STEWARD, Man. Vasc. Pl. Lower
Yangtze Valley, 1958, Corvallis), and no sub-
specific differentiation seems justified on present
knowledge. V. philippica is only likely to be con-
1971]
VIOLACEAE (Jacobs & Moore)
205
fused with V. betonicifolia and the distinguishing
characters are summarized under that species.
8. Viola merrilliana BECKER, Beih. Bot. Centralbl.
34, ii (1916) 234; Philip. J. Sc. 19 (1921) 712;
Merk. En. Philip. 3 (1923) 105. — Fig. 8.
Perennial; rhizome vertical to oblique, rarely
horizontal, bearing leaves and stems at apex.
Stems up to 15 cm, creeping to ascending, rooted
at some nodes, with leaves and flowers at some
nodes. Leaves ¥,—-2% by 1-134 cm, (1-)1 14 times
as long as broad, ovate, with cordate base, and
rounded basal lobes, acute, crenulate or crenate-
serrate, especially on basal lobes, upper surface
with hairy zone towards margin and glabrous in
central portion, glabrous or sparsely pubescent on
veins beneath; petiole 1-8 cm, slender, glabrous.
Stipules 3-6 by 1-114 mm, the lower ovate-
lanceolate, fimbrio-dentate, the upper lanceolate,
dentate near base, all glabrous or sparsely pu-
bescent, fuscous or greenish, adnate to petiole
to 4 length or more. Flowers 10-15 mm, white or
pale purple, with darker veins; peduncle 4-9
cm, usually exceeding the leaves. Sepals 3-5 by
0.6-1.7 mm, lanceolate, acute, entire, glabrous;
appendage 0.4-0.8 mm, rounded, glabrous.
Petals 2-3 times as long as broad; the basal
obovate; the laterals oblanceolate, bearded; spur
1144-2 mm, exceeding calycine appendages, cy-
lindrical, obtuse. Style 1.2-1.6 mm, geniculate
at base, clavate distally, apex with lateral margins
fused behind, with stigmatic beak projecting an-
teriorally and somewhat upcurved. Capsule 5-7
mm, oblong, glabrous.
Distr. Malesia: Philippines (Luzon). Fig. 13.
Eco!. Damp places along streams and in mossy
forest, c. 2100 m. Fi. March—June.
Notes. Distinguished from V. mearnsii by the
distribution of hairs on its upper surface of the
leaf, the narrower and deeper basal leaf sinus, the
larger flowers, the entire sepals and the small
rounded calycine appendages. V. merrilliana
appears to be sympatric with V. rupicola on Mt
Pulog but the latter species differs in its hairy
petioles, completely hairy uppersides of the leaves,
ciliate sepals and free or slightly adnate stipules.
Fig. 13. Distribution of Viola pilosa Bi. (dots)
and V. merrilliana Becker (triangles).
9. Viola mearnsii MERRILL, Philip. J. Sc. 5 (1910)
Bot. 200; Becker, Beih. Bot. Centralbl. 34, i
(1916) 411; Philip. J. Sc. 19 (1921) 715; MeErrR.
En. Philip. 3 (1923) 105; MELCHIOR, Notizbl.
Berl.-Dahl. 12 (1934) 206. — Fig. 8.
Perennial; rhizome vertical or oblique, slender,
bearing rosette of leaves, flowers and sometimes
stems at apex. Stems, when present, up to 3 cm,
rooted at terminal node. Leaves 4-114 by
34-114 cm, about as long as broad, triangular-
ovate, broadly and shallowly cordate at base,
obtuse, rarely acute, crenulate or serrate-crenate,
pubescent in crenations above, glabrous beneath,
herbaceous; petiole 1-4cm, slender, glabrous.
Stipules 3-5 by c. 1 mm, linear-lanceolate, acute,
fimbrio-dentate, fuscous or greenish, slightly
adnate to petiole. Flowers 7—-1114 mm, whitish
with purple veins; peduncle 3-9cm, slender,
glabrous. Sepals c. 3 by c. 1mm, lanceolate,
acute, glabrous or ciliate near base, sparingly
denticulate on margin; appendage c. 1mm,
dentate. Petals 2-214 times as long as broad;
basal obovate, shorter than others; laterals ob-
lanceolate, bearded; spur 114-214 mm, exceeding
calycine appendages, cylindrical, obtuse. Style c.
114 mn, geniculate at base, clavate distally, apex
with margins fused behind and stigmatic beak
projecting anteriorally. Capsule c. 6 mm, globose,
glabrous.
Distr. Malesia: Philippines (Luzon, Negros,
Mindanao), Celebes. Fig. 18.
Ecol. Along streams and in montane forest,
c. 2200 m. Fi. April-May.
Notes. Differs from V. merrilliana in its more
triangular, shallowly cordate leaves which have
hairs only in the marginal crenations on the
upper side, smaller flowers, denticulate sepals and
dentate calycine appendages. The occurrence in
Celebes is based on a single collection (KJELL-
BERG 2658) which differs from Philippine ma-
terial in having glabrous leaves, and entire gla-
brous sepals and calycine appendages. The ter-
minal portion of the style is absent. Further ma-
terial is required to confirm this determination.
10. Viola arcuata Briume, Bijdr. (1825) 58;
Kortu. Ned. Kruidk. Arch. 1 (1848) 357; Mia.
Fl. Ind. Bat. 1, 2 (1858) 114; Koorp. Exk. FI.
Java 2 (1912) 629; Becker, Beih. Bot. Centralbl.
34, ii (1916) 226; Philip. J. Sc. 19 (1921) 710;
Merr. En. Philip. 3 (1923) 104; Back. & SLoor.
Handb. Theeonkr. (1924) 177, t. 177; RipL. J.
Bot. 73 (1935) 14; Back. & Baku. f. Fl. Java 1
(1963) 195; D. M. Moore, Nova Guinea, Bot. n.
11 (1963) 178. — V. distans Wai. [Cat. (1831)
n. 4022] Trans. Med. Soc. Calc. 7 (1835) 227;
Back. Schoolfl. (1911) 66. — V. japonica (non
LANGsD.) Kortu. Ned. Kruidk. Arch. 1 (1848)
357. — V. alata BURGeRSDUK in Mig. Pl. Jungh. 1
(1852) 121; Mio. Fl. Ind. Bat. 1, 2 (1858) 114;
Back. Schoolfl. (1911) 65; Koorp. Exk. Fl. Java
2 (1912) 629; Becker, Beih. Bot. Centralbl. 34,
ii (1916) 227; Ript. J. Bot. 73 (1935) 13. —
V. toppingii Em. Leafl. Philip. Bot. 2 (1908) 504.
— V. semilunaris (MAxiM.) Becker var. philip-
206
FLORA MALESIANA
[ser. I, vol. 7}
( £ ¢
Fig. 14. Viola arcuata BL. a. Habit, x 2, b. stipule, x4, c. & d. stamens, 6, e. fruit, x3, f. seeds, x6
(a VAN STEENIS 7446, b, e BUNNEMEIJER 9595, c-d VAN STEENIS 4306, f BACKER 12848).
pinarum BECKER, Beih. Bot. Centralbl. 34, ii
(1916) 231; Philip. J. Sc. 19 (1921) 712; MeErr.
En. Philip. 3 (1923) 106. — V. lunata RIDL.
Trans. Linn. Soc., Bot. 9 (1916) 18. — V. herbi-
vaga Ripi. J. Bot. 73 (1935) 17. — Fig. 8, 14.
Perennial; rhizome vertical to oblique, slender
to rather stout, bearing stems and leaves at apex.
Stems up to 80cm, slender, procumbent to as-
cending or erect, often rooted at lower nodes, with
leaves and flowers along their length. Leaves
1-334 by %-4144 cm, (1.1—)1.3-2.3 times as
broad as long, broadly hastate, sometimes ovate,
broadly and deeply to shallowly cordate, with pro-
minent basal lobes which are (3—)6-—21 mm long,
typically curved upwards and usually rounded,
cuspidate to obtuse at apex, crenulate, rarely
dentate on basal lobes, glabrous, rarely sparsely
ciliate ‘or pubescent on veins beneath, usually
herbaceous, sometimes thicker and darker green;
petiole 1-14 cm, slender. Stipules 4-14 by 1—3(-6)
mm, generally prominent and sometimes leaf-like,
lanceolate to oblong-lanceolate, acute to mu-
cronate, dentate to shortly fimbrio-dentate, gla-
brous or rarely white to pale purple, with darker
veins; peduncle 2-15 cm, slender. Sepals 214-5
by 1.2—2 mm, ovate-lanceolate to lanceolate, acute,
entire, glabrous, green or fuscous with scarious
margins; appendage c. 1 mm, rounded, glabrous.
Petals (144-)2-4(-414) times as long as broad,
oblong to oblanceolate; the basal shorter than
the others; laterals not or lightly bearded; spur
14-2 mm, equalling or slightly exceeding calycine
appendages, cylindrical, obtuse. Style 1-2 mm,
geniculate at base, clavate distally, with rounded
marginal lobes at apex not joined behind, stig-
matic beak projecting anteriorally. Capsule 4-10
mm, oblong, glabrous. Chromosome number:
22a.
Distr. India, N. Burma, China, Indo-China, in
Malesia: Sumatra, Java, Philippines (Luzon),
Moluccas (Buru), New Guinea. Fig. 15.
Ecol. Typically a plant of wet meadows,
marshes, stream and lake margins, ditches and
grassy clearings, 1000-3000m. Fl. Jan.—Dec.
Notes. In its typical, widespread form V. ar-
cuata is readily recognised by its glabrous, arcuate
leaves and long prostrate stems. It varies in the
size and marginal dissection of the stipules and
in the presence or absence of bearding on the
lateral petals, while some forms have thicker,
more pubescent and deeply cordate leaves than
is usual. Variation in these characters cannot be
correlated with each other, or with any distri-
butional pattern, and intermediates occur. No
formal recognition of such variation is possible
and much of it may be due to plants which occupy
drier habitats than usual.
11. Viola javanica BECKER, Beih. Bot. Centralbl.
34, ii (1916) 260; Back. & Baku. f. FI. Java 1
(1963) 195. — V. arcuata (non BL.) Mia. FI. Ind.
Bat. 1, 2 (1859) 686; RipL. J. Bot. 73 (1935) 14. —
Fig. 8.
Perennial; rhizome vertical, rather stout,
bearing leaves and stems at apex. Stems up to 26
cm, rather slender, creeping or somewhat as-
cending, rooted at some nodes, bearing leaves
along their length and flowers usually distally.
Leaves 14-2'4 by 34-3 cm, 1-1.6 times as broad
as long, triangular-ovate, rarely subhastate,
shallowly and rather broadly cordate at base,
acute to obtuse or rounded, crenate or subden-
tate, glabrous or sparsely pubescent on veins
towards sinus, subcoriaceous, dark green. Sti-
pules 5-10 by 1-2 mm, lanceolate, acute, fimbriate,
glabrous, fuscous, free. Flowers 6-12 mm, purple
with darker veins; peduncle 2-9 cm, exceeding
leaves, sparsely hirsute distally. Sepals 4-514 by
114-1.8 mm, ovate-lanceolate, acute, entire, gla-
brous or sparsely pubescent near base, green with
scarious margin; appendage 14-1 mm, rounded or
slightly dentate. Petals about 3 times as long as
broad, oblong to obovate-oblanceolate; laterals
heavily, rarely lightly bearded; spur 14-114 mm,
shorter than to slightly exceeding calycine ap-
pendages, cylindrical, obtuse. Style 114-2 mm,
VIOLACEAE (Jacobs & Moore)
207
rs = : ; ; = = aI
Fig. 15. Distribution of Viola arcuata BL. in
Malesia (dots) and V. javanica BECKER (triangles).
geniculate at base, clavate distally, apex with
flattish marginal lobes fused behind and anterior
stigmatic beak. Capsule 5-9 mm, oblong, glabrous.
Distr. Malesia: E. Java (Mts Ardjuno, Teng-
ger & Jang). Fig. 15.
Ecol. Grasslands, 2000-3000 m. Fi. Jan.—Nov.,
probably all year.
Notes. Closely related to V. arcuata but
readily distinguished from typical forms of that
species by its subcoriaceous, strongly crenate,
ovate leaves and its long-fimbriate stipules. Some
forms of V. arcuata, particularly in drier habitats,
approach V. javanica in these characters but the
stigma has the rounded separate lobes of V.
arcuata and not the posteriorally fused, flatter
margin of V. javanica. There is a superficial re-
semblance to some forms of V. pilosa but in that
species the style is scarcely clavate and the stigma
is emarginate.
12. Viola pilosa BLUME, Cat. Gew. Buitenzorg
(1823) 57; Bijdr. (1825) 57; Kortu. Ned. Kruidk.
Arch. 1 (1848) 357; Mia. FI. Ind. Bat. 1, 2 (1858)
113; Back. & SLoor. Handb. Theeonkr. (1924)
180, t. 180; DocTERS VAN LEEUWEN, Trop.
Natuur 16 (1927) 187; Verh. Kon. Ak. Wet.
A’dam II, 31 (1933) 184. — V. serpens WALL.
ex GING. in DC. Prod. 1 (1824) 296; Mia. FI. Ind.
Bat. 1, 2 (1858) 113; Back. Schoolfl. (1911) 66;
Koorpb. Exk. Fl. Java 2 (1912) 628, pro parte;
BECKER, Beih. Bot. Centralbl. 40, ii (1923) 103;
Ooststr. & H. J. LAM, Blumea 5 (1945) 594;
BAcK. & BAKH. f. Fl. Java 1 (1963) 195. — V.
palmaris BUCH.-HAM. ex GING. in DC. Prod. 1
(1824) 298; BURGERSDIJK in Miq. PI. Jungh. 1
Fig. 16.
Viola pilosa BL. on Mt Gedeh in W. Java (photogr. VAN WOERDEN).
208
(1852) 120; Mia. Fl. Ind. Bat. 1, 2 (1858) 113.
— V.hamiltoniana D. Don, Prod. FI. Nepal. (1825)
206; BoissteU & CAPITAINE, Bull. Soc. Bot. Fr.
57 (1910) 340. — V. sarmentosa BURGERSDIJK
in Miq. Pl. Jungh. 1 (1852) 120; Mia. FI. Ind.
Bat. 1, 2 (1858) 114. — V. burgersdijkii OUDEMANS
in Miq. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 77;
BorssteU & CAPITAINE, Bull. Soc. Bot. Fr. 57 (1910)
337, incl. f. nodosa Boiss. & Capit. l.c. 338;
Becker, Beih. Bot. Centralbl. 40, ii (1923) 106;
Rip. J. Bot. 73 (1935) 16, incl. var. timorensis
Rip_. — V. glaucescens OUDEMANS in Miq. Ann.
Mus. Bot. Lugd.-Bat. 3 (1867) 74; BECKER, Beih.
Bot. Centralbl. 40, ii (1923) 109. — V. confusa
(non CHAMP.) BortssIEU & CAPITAINE, Bull. Soc.
Bot. Fr. 57 1910) 339. — V. canescens (non
WALL.) BotsstEU & CAPITAINE, /.c. 338; BECKER,
Beih. Bot. Centralbl. 34, ii (1916) 256. — ?
V. celebica BECKER in Fedde, Rep. 14 (1916)
321; Beih. Bot. Centralbl. 34, ii (1916) 416. —
Fig. 8, 16, 17.
Perennial; rhizome vertical to oblique, 114-4
mm @, bearing leaves and stems at apex. Stems
up to 1m, slender, rooted at some nodes, leafy,
with flowers at some nodes. Leaves 1-10 by
FLORA MALESIANA
[ser. I, vol. 71
1-614.cm, 1-2times as long as broad, ovate,
deeply cordate at base, acute to acuminate, ser-
rate or serrate-crenate, pubescent to hirsute above
and beneath, especially on veins, rarely glabrous,
usually pale green; petiole 1-17 cm, pilose especi-
ally distally, rarely glabrous. Stipules 7-30 by
114-5 mm, lanceolate, Jong-acute, long-fimbriate,
pubescent, fuscous or green. Flowers (5—)8-14 mm
long, purple to white with darker veins; peduncle
4-12 cm, pilose especially distally, rarely glabrous.
Sepals 314-9 by 1-2mm, _linear-lanceolate,
acute, entire or denticulate, usually pilose especial-
ly near base, ciliate; appendage 0.8—214 (-3) mm,
pointed or denticulate, rarely rounded, pilose or
rarely glabrous. Petals 114-4times as long as
broad; basal obovate, sometimes slightly bearded;
laterals oblanceolate, bearded; upper pair usually
lightly bearded; spur 1-214 mm, slightly shorter
than to just exceeding calycine appendages, cy-
lindrical, obtuse. Style 114-3 mm, geniculate at
base, somewhat thickened distally, apex gently or
strongly curved to appear flattened or sometimes
hooked, with simple emarginate stigma. Capsule
5-10 mm, ellipsoid, glabrous or pubescent in
centre of valves.
Fig. 17. Viola pilosa Bu. a. Habit, x24, b. stipule, x2, c. flower, x2, d. & e. stamens, x6, f. pistil,
<6, g. young fruit, «3, h. ripe fruit, «3, i. & j. seed, * 6 (a—~f KOORDERS 43653, g—j KUHL 5.n.).
en
1971]
Distr. Himalayas, India, Burma, Thailand,
China, in Malesia: Central Sumatra (Mt Kerintji),
Java, Lesser Sunda Islands (Bali, Timor), SW.
Celebes (Mt Bonthain), Moluccas (Buru, Ce-
ram). Fig. 13.
Ecol. Grasslands, alpine woods,
1100-3300 m. F/. Jan.—Dec.
Notes. This is a rather variable species within
which no satisfactory formal subdivision seems
possible at present. The variation is principally
in such characters as size, leaf-shape and in-
dumentum, which are notoriously subject to
environmental modification in this genus and no
correlation can be detected between such vari-
ation and distribution. Conspicuously large and
robust specimens, undoubtedly the result of parti-
cularly sheltered habitats, have been separated as
V. burgersdijkii and V. sarmentosa. The most
distinctive populations within V. pilosa are those
from the Moluccas. Two collections from Buru
(Toxoreus on | and 3 III 1922), at different
elevations on Mt Fakal, superficially resemble
V. sumatrana in leaf-colour, and in having gla-
brous, more or less entire calycine appendages;
they are undoubtedly, however, subglabrous forms
of V. pilosa. Material from Mt Pinaia in Ceram
(STRESEMANN 302, EyMA 2259) is superficially
like V. papuana in its coriaceous leaves and like
V. kjellbergii in its rounded calycine appendages;
it differs from both in having free stipules and from
each in other characters. It is possible that these
populations are worthy of subspecific status, but
further material would be desirable since some of
these characters are undoubtedly the result of
the exposed alpine habitat which the species oc-
cupies in Ceram.
V. celebica BECKER was described from a
SARASIN collection on Mt Bonthain in SW. Ce-
lebes. The undistributed collection was destroyed
at Berlin, but the subsequent extensive collec-
tions by BUNNEMEVER from Mt Bonthain leave
little doubt that it is referable to V. pilosa.
It may be that a fragment of the type is present
in the diary of the SARAsINs which is preserved at
the Botanical Institute at Basel, but it has not
been possible to examine this.
pathsides,
13. Viola sumatrana MiQueL, Sum. (1860) 389;
Becker, Beih. Bot. Centralbl. 40, ii (1923) 108;
Ript. J. Bot. 73 (1935) 18. — V. sumatrana Mia.
var. caerulescens BoissteU & CAPITAINE, Bull.
Soc. Bot. Fr. 57 (1910) 341. — V. hossei BECKER,
Beih. Bot. Centralbl. 34, ii (1916) 257, pro parte;
Ript. J. Bot. 73 (1935) 18. — V. sikkimensis
Becker var. acuminatifolia Becker, Beih. Bot.
Centralbl. 34, ii (1916) 260. — V. serpens (non
WALL.) Ript. Fl. Mal. Pen. 1 (1922) 126. —
V. korinchensis Rivv. J. Bot. 73 (1935) 15. —
V. robinsonii Rao. Lc. 16. — V. jugalis Rio. Le.
18. — Fig. 8.
* Perennial; rhizome vertical to oblique, 114-6
mm @, bearing leaves and stems at apex. Stems up
to 60 cm, slender, rooted at some nodes, with leaves
and flowers at many nodes. Leaves %-7 by “4-4
cm, 1-15 times as long as broad, ovate, mod-
VIOLACEAE (Jacobs & Moore)
209
erately to deeply cordate at base, acute, serrate,
glabrous or rarely with scattered hairs below,
especially on veins, dark green or purplish above,
glaucous below; petiole 1-15 cm, glabrous. Sti-
pules 6-14 by 1-3 mm, linear-lanceolate, long-
acute, long-fimbriate, fuscous. Flowers 10-1414
mm, purple to creamy white, with darker veins;
peduncle 5-l3cm, glabrous or rarely hispid
distally. Sepals 5—7 by c. 1 mm, linear-lanceolate,
acute, entire, glabrous or sparsely pubescent,
rarely ciliate; appendage 0.4—1 (—1.4) mm, rounded
or slightly denticulate, glabrous or rarely hispi-
dulous. Petals 2-4 times as long as broad; basal
rather shorter than the others, obovate; laterals
oblanceolate, not bearded; spur 1—2 mm, usually
twice as long as calycine appendages, cylindrical,
obtuse. Stvle 1.7—-2 mm, geniculate at base, cla-
vate distally, apex with two lateral lobes and
anterior stigmatic beak. Capsule 7-11 mm,
ellipsoid, glabrous.
Fig. 18. Distribution of Viola sumatrana Mia.
(dots) and V. mearnsii MERR. (triangles).
Distr. China, Burma, Thailand, Indo-China,
in Malesia: Sumatra, Malay Peninsula, Borneo
(Sarawak: Batu Tibang; North Borneo: Mt Ki-
nabalu). Fig. 18.
Ecol. River banks, pathsides, openings in
montane woodland, 900-2800 m. F/. Jan.—Dec.
Notes. This species has close affinities with
V. pilosa and V. curvistylis. It differs from V. pi-
Josa in the marginate stigma, unbearded lateral
petals, short rounded calycine appendages, entire
and usually glabrous sepals, and the generally
glabrous leaves which are usually much darker on
the upper side and which have glandular-serrate
margins. V. curvistylis differs in its leaf-shape,
bearded lateral petals, frequently hirsute sepals
and its pale green, usually pilose leaves. V. suma-
trana is apparently sympatric with V. curvistylis
in the Malay Peninsula (Pahang), North Borneo
(Mt Kinabalu) and Sumatra (Mt Kerintji), and
with V. pilosa in Sumatra (Mt Kerintji).
The type description gives flower colour as
yellowish, hence the recognition of var. caerules-
cens by Boussieu & Capriratne to account for their
purple-flowered specimens. Plants with yellowish
210
FLORA MALESIANA
[ser. I, vol. 7}
or whitish flowers have been encountered spo-
radically throughout the range of the species,
sometimes with the typical purple-flowered form.
These specimens cannot otherwise be distinguished
from the rest of the material examined and are
undoubtedly colour mutants which do not merit
formal taxonomic recognition.
14. Viola curvistylis BoissiEU & CAPITAINE, Bull
Soc. Bot. Fr. 57 (1910) 339; BECKER, Beih. Bot.
Centralbl. 40, ii (1923) 168. — V. ovalifolia
BECKER, Beih. Bot. Centralbl. 34, ii (1916) 256;
RIDL. J. Bot. 73 (1935) 15. — V. malvina RIDL.
J. Fed. Mal. St. Mus. 15 (1917) 135. — V. serpens
(non WALL.) RIDL. FI. Mal. Pen. 1 (1922) 126,
pro parte. — V. lecomteana BECKER ex GAGNEP.
Suppl. Fl. Gén. I.-C. 1 (1939) 181, descr. gall.,
illeg. — Fig. 8.
Perennial; rhizome vertical to oblique, slender
to rather stout (4mm @), bearing leaves and stems
at apex. Stems up to 35cm, slender, creeping,
bearing rooted rosettes of leaves and flowers at
some nodes. Leaves 1-9 by '4-5cm, about
twice as long as broad, elliptic to lanceolate-
ovate, shallowly cordate to cuneate at base,
acute, serrate or crenate-serrate, hirsute on both
sides, rarely pubescent or glabrous, pale green;
petiole 0.6-10 cm, pubescent or scabrid, especially
distally, rarely glabrous. Stipules 7-11 by 1-5 mm,
linear-lanceolate, long-acute, long-fimbriate, gla-
brous, green or fuscous, free. Flowers 8-11 mm,
violet or white with darker veins; peduncle 2-13
cm, slender, pubescent or scabrid, especially
distally, sometimes glabrous. Sepals 4-6 by 0.6—1
mm, triangular- to linear-lanceolate, acuminate.
entire, sometimes pubescent near base, usually
ciliate; appendage 14-1 mm, rounded or slightly
denticulate, glabrous or scabrid. Petals 2-3 times
as long as broad; basal 24 as long as others,
obovate; laterals oblanceolate, bearded; spur 1-1 %
mm, exceeding calycine appendages, cylindrical,
obtuse. Style 114-2 mm, geniculate at base, cla-
vate distally, with two large marginal lobes and
anterior stigmatic beak at apex. Capsule 6-10 mm,
globose to oblong, glabrous or pubescent.
Fig. 19. Distribution of Viola curvistylis Boiss. &
Capir. in Malesia (dots), V. rupicola ELM.
(circles), and V. kjellbergii MELCH. (triangles).
Distr. Indo-China, Burma, in Malesia: Su-
matra, Malay Peninsula, North Borneo (Mt Ki-
nabalu). Fig. 19.
Ecol. Openings in montane woodlands, river
banks, (600—)1000-—3000 m. Fi. Jan.—Dec.
Notes. Differs from V. sumatrana in having
bearded lateral petals and usually pilose leaves,
and from V. pilosa in the prominent lateral
margin of the style apex and in the usually rounded
short calycine appendages, and from both species
in leaf-shape. It is apparently sympatric with
V. sumatrana in the Malay Peninsula (Pahang)
and North Borneo (Mt Kinabalu), and with both
V. sumatrana and V. pilosa on Mt Kerintji,
Central Sumatra. The specific name derives from
the curved, short style in the cleistogamous flowers
of the type specimen; the species would be much
better designated by BECKER’s epithet ovalifolia.
15. Viola rupicola ELmer, Leafl. Philip. Bot. 1
(1908) 324; Becker, Beih. Bot. Centralbl. 34,
ii (1916) 415; Philip. J. Sc. 19 (1921) 712; Beih.
Bot. Centralbl. 40, ii (1923) 171; Merr. En. Philip.
3 (1923) 106; Lin, Taiwania 1 (1950) 273. —
V. adenothrix HAYATA, Ic. Pl. Formos. 3 (1913) 23.
— V. effusa BECKER, Beih. Bot. Centralbl. 34, ii
(1916) 424. — Fig. 8.
Perennial; rhizome vertical to oblique, rather
stout, bearing leaves and usually stems at apex.
Stems up to 25 cm, creeping to ascending, some-
times rooted at lower nodes. Leaves 34-214 by
34-134 cm, (1-)114-114 times as long as broad,
elliptical to elliptic-ovate, cordate at base with
narrow sinus and basal lobes rounded and some-
what converging, acute to subobtuse, shallowly
crenate, subdentate on lobes, pubescent, often
densely so; petiole 34-1214 cm, slender, hirsute.
Stipules 3-9 by 0.6 to c. 1 mm, lanceolate, acute,
fimbriate to dentate, glabrous, fuscous. Flowers
9-12 mm, pale purple with darker veins; peduncle
4-9 cm, exceeding leaves, slender. Sepals 2-4 by
c. 1mm, ovate-lanceolate, acute, entire, sparsely
pubescent on back and margin; appendage c.
14 mm, rounded, ciliate. Petals 2-3 times as long
as broad, obovate to oblanceolate; basal usually
shorter than the others; laterals and usually
upper bearded; spur 0.8—2 mm, exceeding calycine
appendages, cylindrical, obtuse. Style 1.2-1.4
mm, geniculate at base, clavate distally, with apex
distinctly lobed on both sides and anterior stig-
matic beak. Capsule 4-5 mm, globose to oblong,
glabrous.
Distr. Formosa,
(Luzon). Fig. 19.
Ecol. Damp ravines, rocky ledges and stream
margins, c. 2000-2500 m. Fi. Febr.—May.
Note. Sympatric with V. merrilliana on Mt
Pulog.
in Malesia: Philippines
16. Viola kjellbergii MELCHIOR, Notizbl. Berl.-Dahl-
12 (1934) 206. — V. arcuata (non BL.) OOSTSTR.
& H.J. LAM, Blumea 5 (1945) 594. — V. lagaipensis
D. M. Moore, Nova Guinea, Bot. n. 11 (1963)
182. — Fig. 8.
Perennial; rhizome more or less vertical, up to
1971]
214 cm, bearing leaves, flowers and stems at apex.
Stems up to 25 cm, slender, glabrous, leafy, with
flowers and rooted at some nodes. Leaves 0.4-3
by 0.6-2.8cm, ovate to somewhat reniform,
shallowly to deeply cordate at base, obtuse to
acute, crenate to distantly dentate, glabrous,
sometimes sparsely pilose near sinus; petiole
0.2-8 cm, glabrous or pubescent, especially dis-
tally. Stipules 2-3 mm, ovate-lanceolate to lan-
ceolate, acute, sparsely dentate or dentate-fim-
briate, glanduliferous on margin and apex, gla-
brous or slightly ciliate, fuscous, adnate to petiole
for 14 length. Flowers 4-10 mm, pale purple to
white, with darker veins; peduncle 2!4-25 cm,
glabrous or sparsely pubescent distally. Sepals
24%4-644 by %-1'%4 mm, lanceolate to ovate-
lanceolate, acute, entire or sparsely fimbrio-
dentate, glabrous, green or fuscous, with scarious
margin; appendage up to | mm, rounded. Petals
114-2 times as long as broad; basal obovate;
others oblanceolate; laterals bearded; spur
0.2-2.4 mm, shorter than to slightly exceeding
calycine appendages, rounded. Style 1.8-214 mm,
geniculate at base, somewhat curved, filiform;
stigma simple, somewhat cupuliform at maturity.
Capsule 5-7 mm, ellipsoid, glabrous, purple to
green. Chromosome number: 2n = 48.
Distr. Malesia: SW. Central Celebes (Lati-
modjong Range), E. New Guinea. Fig. 19.
Ecol. Alpine heath and grasslands, occasion-
ally wood margins, c. 2000-3540 m. F/. June-
March.
Notes. Superficially similar to the populations
of V. pilosa from Ceram but differing in its ad-
nate, shorter fimbriate stipules and in stigma shape.
This species is related to V. papuana from which
it is distinguished by the short spur, smaller
flowers, shorter style and bearded lateral petals
(occasionally so in papuana), and to V. hederacea
from which it differs in the adnate stipules, the
absence of leaves grouped in distinct rosettes,
and in the much less marked bearding of the la-
teral petals.
17. Viola hederacea Lapitt. Pl. Nov. Holl. 1
(1804) 66, t. 91; Becker, Beih. Bot. Centralbl. 34,
ii (1916) 428; D. M. Moore, Blumea 11 (1963)
535, map. — Fig. 8.
Perennial; rhizome horizontal to oblique or
vertical, rather stout, bearing leaves, flowers and
stems at apex. Stems up to 25 cm, slender, pros-
trate, with rosettes of leaves and flowers at some
nodes, rooted at some nodes. Leaves 0.9-2 by
0.9-3 cm, (1—)1.2-114 times as broad as long,
reniform to suborbicular, cordate, broadly obtuse,
distinctly repand-crenate, sparsely pubescent
beneath, more heavily above, subcoriaceous,
dark green; petiole 0.8—9 cm, glabrous or sparsely
pilose. Stipules 2-3 by 1-2 mm, ovate-lanceolate,
acute, fimbrio-dentate to fimbriate, glabrous,
green to fuscous, free. Flowers 7-9 mm, white with
purple centre; peduncle 2-19cm, much ex-
ceeding leaves, slender. Sepals 4-6 by c. 144 mm,
linear-lanceolate, acute, entire, glabrous, green
with scarious margin; appendage up to 4 mm,
VIOLACEAE (Jacobs & Moore)
211
rounded, glabrous. Petals about twice as long as
broad, obovate-oblong: laterals bearded; spur
absent or rudimentary. Style c. 244 mm, genicu-
late at base, slightly curved, filiform; stigma
terminal, simple. Capsule c. 8 mm, ellipsoid, gla-
brous.
Distr. Tasmania, Australia (S. Australia, Vic-
toria, New South Wales, Queensland), in Ma-
lesia: Malay Peninsula (Cameron Highlands,
Pahang). Fig. 20.
Fig. 20. Distribution of Viola hederacea LABILL.
in Malesia (triangles) and V. papuana BECKER &
PULLE (dots).
Ecol. Montane grassland, 2030 m. Fi. Sept.-
Nov.
Note. The isolated occurrence in Malaya of
this Australian species is surprising, and falls
outside the accepted migration track of cold-
loving plants from the southern hemisphere into
Malesia. However, in view of its relatively recent
discovery in the Cameron Highlands, an area
which has been well-collected in the past, in a
locality close to the radio station on Batu Brin-
chang, it is possible that this species has been in-
troduced by man.
18. Viola papuana BECKER & PULLE, Nova Guinea
8 (1912) 670; Becker, Beih. Bot. Centralbl. 34, ii
(1916) 431; D. M. Moore, Nova Guinea, Bot. 7.
11 (1963) 184. — V. klossii Rip. Trans. Linn.
Soc., Bot. 9 (1916) 18. — Fig. 8.
Perennial; rhizome more or less vertical, stout,
bearing leaves and stems at apex. Stems up to
24 cm, rather slender, rooting at nodes with leaves
and flowers along their length. Leaves 0.4-1.4 by
0.6-1.8 cm, ovate, rather deeply cordate at base
with rounded basal lobes, acute to subobtuse,
more or less repandate to undulate-serrate, gla-
brous or sparsely pubescent on veins above and
sparsely ciliate, rather coriaceous, dark green and
often purple-tinged; petiole 44-4 cm. Stipules 3-4
by c. 1 mm, lanceolate to ovate-lanceolate, acu-
minate, subentire to sparsely fimbriate, sometimes
glanduliferous, glabrous, fuscous. Flowers (ex-
cluding spur) 8-13 mm, pale violet to white with
violet centre; peduncle 4-14 cm, glabrous. Sepals
3-6 by c. 1 mm, lanceolate to ovate-lanceolate,
212
FLORA MALESIANA
[ser. I, vol. 77]
acute, sparsely fimbrio-dentate; appendage c.
4mm, rounded, glabrous. Petals 2-3 times as
long as broad; basal obovate, slightly shorter than
others; laterals oblanceolate to obovate, usually
not bearded; spur (3-)4-9mm, _ cylindrical,
straight or slightly upcurved, obtuse. Style 214-3
mm, geniculate at base, slightly curved, filiform;
stigma simple, more or less indistinctly cupuliform.
Capsule 7-9 mm, oblong-ellipsoid, glabrous. Chro-
mosome number 2n = 48.
Distr. Malesia: New Guinea. Fig. 20.
Ecol. Pathsides and open places in montane
forest and among rocks of alpine areas, 2100-3350
m. F/. Oct.—Febr.
Doubtful
Viola ramosiana BECKER, Philip. J. Sc. 19 (1921) 716.
Based on a fruiting specimen from “‘c.
1350 m,
along streams, Mt. Masingit, Lubuagan, Kalinga
Subprovince, Luzon, Philippines, on ii. 1920, by
Ramos & EDANO, Bur. Sci. 37548’. The holotype
was destroyed in the Manila herbarium. An iso-
type (US) does not agree with the description in
that it is not stoloniferous, the stipules are
partially adnate to the petiole, and some of the
leaves are sparsely pubescent. This material may
be referable to V. philippica, which is known from
the same locality, but it differs from that
species in the well-developed glands on the
marginal crenations of some _ leaves. This,
together with the absence of flowers and the firm
reference to stolons in the destroyed holotype,
precludes any definite application of this specific
name.
Excluded
Tribe Sauvagesieae, including the genera Neckia KorTH., Sauvagesia L., Schuurmansia Bu., and Indovethia
BOERL. were sometimes arranged in Violaceae, but are now unanimously classified in Ochnaceae.
Gestroa Becc. Malesia 1 (1877) 184 is, according to SLEUMER, FI. Mal. I, 5 (1954) 6, a synonym of Ery-
throspermum (Flacourtiaceae).
Perissandra GAGNEP. Bull. Soc. Bot. Fr. 95 (1948) 27, from Indo-China is, according to JAcoss, Blumea
15 (1967) 138, a synonym of Vatica (Dipterocarpaceae).
APONOGETONACEAE (H. W. E. van Bruggen, Heemskerk)
1. APONOGETON
LInnE f. Suppl. (1781) 32; ENGLER & Krause, Pfl. R. Heft 24 (1906); STEEN.
Fl. Mal. I, 4 (1948) 11; BRUGGEN, Blumea 18 (1970) 457-486. — Fig. 1-5.
Perennial waterplants with a tuberous, elongate or cylindrical and often
branched rootstock or rhizome which produces a tuft of leaves and the inflores-
cences. Leaves submerged and/or floating (very seldom emerged), with a mostly
distinct midrib and one or more pairs of parallel main nerves, connected by
numerous cross-veins. Inflorescence long-peduncled, emerging above the water
surface, in bud enveloped by a caducous or rarely persistent spathe, composed of 1
(in Mal.) or 2-11 spikes. Flowers (in Mal.) bisexual, spirally arranged, turned
towards all directions. Tepal/s 2, mostly persistent, rarely caducous. Stamens 6,
in 2 whorls. Ovaries 3(-—4—5), free, sessile, narrowed into the style with a stigmatic
ridge on the inner side; ovules 2-8 per carpel. Fruits with a mostly distinct, lateral
or terminal, often curved beak. Seeds without endosperm; testa mostly a single
envelope, sometimes, however, split into two envelopes, the inner one, brown and
closely fitting the embryo, the outer loose, transparent and reticulately veined;
embryo with the plumule fitting in a groove or not, or without plumule (the
embryos of all species with a double testa seem to have no plumule).
Distr. About 40 spp. described, from Africa (Ethiopia to the Cape), Madagascar, India & Ceylon,
through SE. Asia (to c. 30° NL) and Malesia to SW., N. and E. Australia (to 34° SL), centering in Africa
and Madagascar.
Ecol. In stagnant and running, shallow water, mostly in the lowlands, ascending to c. 1000 m;
A. crispus THUNB. in Ceylon 1000-2300 m, also in Africa and Madagascar some species to c. 2500 m.
Phytochem. Very little is known about the chemistry of this highly interesting family. Since no
recent chemical investigations were published, the reader is referred to the discussion of Aponogetonaceae
in HEGNAUER, Chemotax. d. Pfl. 2 (1963) 70-73. — R. HEGNAUER.
Uses. In Malesia none. The starchy tuberous rootstock of most species is edible and seems to be in
some areas an important food source in times of famine. The leaves and inflorescences can serve as a
vegetable. Many species are in cultivation with aquarists.
KEY TO THE SPECIES
1. Tepals longer than 5 mm, caducous. Ovules 2. Plant stoloniferous. 1. A. undulatus
1. Tepals shorter than c. 3 mm, persistent. Ovules 4-8. Plant not stoloniferous.
Dee nDIS, Lepais yellow. . . . +» + «+ < + « s inuk ie. lel? gees eee 2. A. lakhonensis
2. Testa simple.
3. Cross-veins at a -+- 50° angle. Tepals yellow. Filaments only slightly widening towards the base.
memerauntir wv cermimal beak, . . 0 « «8 deine > wa Ws. Opa) 5) dee 3. A. loriae
3. Cross-veins at a +. 70° angle. Tepals white or cream. Filaments strongly widened and applanate.
Fruit with a short lateral beak. 4. A. womersleyi
re ee) ee a oe ee oe i ee) he ee ee ee er
1. Aponogeton undulatus Roxs. [Hort. Beng. Med. Landb. Hogeschool Wagen. 2 (1958) 96,
(1814) 26, nomen] FI. Ind. ed. Carey 2 (1832) 211;
ENGLER & Krause, Pfl. R. Heft 24 (1906) 11,
pro nomen, excl. descr.; BRUGGEN, Blumea 18
(1970) 465, f. 2%, t. I-Il. — Spathium undulatum
Epcew. Calc. J. 3 (1843) 534, f. 15. — Ouvirandra
undulata Evcew. in Hook. Lond. J. Bot. 3
41844) 404, f. 18. — A. microphyllum Rox. FI.
Ind. ed. Carey 2 (1832) 211; Hook. f. Fl. Br.
Ind. 6 (1894) 565. — A. stachyosporus De Wit,
f. 1-3. — A. crispus (non TuunB.) Hook. f/f. Fl.
Br. Ind. 6 (1898) 564, pro parte; TRIM. Fl. Ceyl. 4
(1898) 372; GamBLe, FI. Pres. Madras 3 (1931)
1597, — A. monostachyon (non L. f.) ANDR. Bot.
Rep. 6 (1797) t. 406. — Fig. 1', 2.
Tuber globular, obovoid, or elongate, 6-25 mm
@, smooth. Submerged leaves 10-25 by 4-44 cm,
alternately transparent or opaque in an irregular
pattern; base (narrowly) cuneate or rounded,
(213)
FLORA MALESIANA
[ser. I, vol. 74
ae ? t fe.
aos) eA a> i P ei
R71
Fig. 1. Floral details of Aponogeton. — 1. A. undulatus Roxs. — 2. A. lakhonensis A. CAMuS. — 3. A.
loriae MARTELLI. — 4. A. womersleyi BRUGGEN.
In all figures a = tepal(s), b = stamen, c = ovaries, d = fruit(s), e = seed, f= embryo. All x7
(Courtesy Blumea).
apex (narrowly) cuneate or rounded, seldom
emarginate, tip blunt; margin undulate, midrib
wide with (2—)3(-4) main nerves on either side;
petiole 10-35 cm. Floating leaves rather rare, up
to 20 by 314 cm; base rounded or cordate, apex
cuneate with a blunt tip; main nerves 5 or 7;
petiole up to 70 cm. Peduncle up to 55 cm, thick-
ening towards the inflorescence. Spathe up to
17mm, persistent or caducous. Spike up to 10
cm, laxly flowered. Tepals (1—)2(—3), caducous,
spathulate or obovate, 1(—2-3)-nerved, white or
pinkish, (2—)314-6(-12) by 1-214(-4) mm. Fila-
ments slightly widening towards the base, white
or pinkish; anthers light yellow, pollen yellow.
Ovaries 3(-4), light pink, 114-2 by 14-1 mm;
ovules 2. Infructescence up to 16cm. Fruit 5-7
by 4mm with a short, terminal, curved beak.
Seeds with a simple testa; embryo up to 5 by 2
mm; plumule attached near the base of the
embryo and lying in a very wide groove.
Runners resembling peduncles, ascending, up
to 35cm, not or slightly thickened towards the
tip. The plantlet is developed at the tip of the run-
ner, in an early stage it is enveloped by a persistent
spathe of up to 20 mm. The young plant itself may
also put forth one or two short runners, and by
repeating this up to 7 plantlets may be produced.
Distr. NW., N. and NE. India, E. Pakistan,
Burma, ?Thailand; in Malesia: ?7Malay Peninsula
(Johore: Sg. Sedili at Mawai).
Ecol. Ponds and ditches, 300 m. F/. July—Aug.
and Nov.
Notes. There are no specimens which specimens
proved to come from Malaya. The only have been
I have seen were collected in the Van Cleef Aqua-
rium at Singapore. They are said to have been
collected in the Sg. Sedili at Mawai, but I am
not convinced that this record is reliable. These
specimens have been described as A. stachyo-
sporus. Also the occurrence in Thailand has not
yet been proved. Yearly many thousands of
corms are exported from Bangkok for aquarium
purposes. They are claimed to have been collected
near Chanthaburi and Haadyai (close by the Ma-
APONOGETONACEAE (van Bruggen) 215
1971]
Ze
__—
SS
ee
a
i
ae
men in the Van Kleef Aqua-
Johore, by Ernest Tort, x 4.
er a living speci
Sedili,
have been collected originally in the Sg
. Aponogeton undulatus Roxs. Drawn at Singapore aft
rium, said to
Fig. 2
216
layan frontier), but up till now I have not seen
herbarium specimens collected in Thailand either.
It is remarkable that A. undulatus flowers very
seldom in cultivation. I have been cultivating
specimens from different localities for more than
twelve years, but up till now only once an in-
florescence has developed.
It seems that in the eastern part of its range the
leaves are more narrow than in the western part;
moreover, the eastern populations seem not to
tend to develop floating leaves.
2. Aponogeton lakhonensis A. CAMus, Not. Syst. 1
(1909) 273, f. 18; Fl. Gén. L-C. 6 (1942) 1226;
BRUGGEN, Blumea 18 (1970) 479, f. 212, 3a. —
A. pygmaeus KRAUSE, Bot. Jahrb. 44 (1910)
Beibl. 101: 8; A. Camus, Fl. Gén. I.-C. 6 (1942)
1227.— A. luteus A. Camus, Not. Syst. 2 (1911) 204;
FI. Gén. I.-C. 6 (1942) 1227. — A. monostachyon
(non L. f.) A. Camus, Fl. Gén. I.-C. 6 (1942)
1225; LARSEN, Dansk Bot. Ark. 20, 2 (1962) 134.
— A. loriae (non MARTELLI) STEEN. Fl. Mal. I,
4 (1948) 11, pro descr. — Fig. 1°, 3.
Tuber elongate or obovoid, up to 2 cm @.
Submerged leaves very variable in shape and size,
narrowly oval to linear, up to 25 by 6cm (e.g.
25 by 6cm, 15 by 0.9 cm); base (very) narrowly
© 70
Fig. 3. Aponogeton lakhonensis A. Camus. Habit,
with submerged leaves (HARMAND s.7.), X14
(Courtesy Blumea).
FLORA MALESIANA
(ser. I, vole?
cuneate, apex narrowly cuneate with a blunt tip or
rounded; main nerves 7 or 9; petiole 7-35 cm.
Floating leaves (not seen in Mal.) up to 17 by
4 cm, base cordate or (seldom) rounded; apex
cuneate with a blunt tip or rounded; main nerves
7 or 9. Peduncle only slightly thickening towards the
inflorescence. Spathe c. 17 mm, caducous, rarely
persistent. Spike up to 8 cm, rather laxly, some-
times densely flowered. Tepals 2, obovate, 1-2
by 34-1144 mm, yellow, 1-nerved. Stamens 6,
114-3 mm, filaments widened towards the base.
Ovaries 3—4(—5), 1-134 by 34-1 mm; ovules 4-8.
Infructescence up to 17 cm. Fruits up to 3 by 2 mm,
with a terminal or lateral beak. Seeds 2-3 by 1 mm;
testa double; embryo 114-214 by 14 mm; plumule
absent.
Distr. SE. Asia (India: Assam, Thailand,
Cambodia, Vietnam, China); in Malesia: SW.
Celebes (Maros; Pangkadjene), two collections.
Ecol. Slow running streams, flooded rice fields
and ponds, in Celebes in a stream in limestone
country, not frequent but rather abundant lo-
cally, 200-800 m. F/. March, May, Sept. and Oct.
Note. The Malesian specimens are very frag-
mentary with regard to their generative parts,
and have only some almost ripe fruits. Therefore,
I can not be completely certain that these speci-
mens belong indeed to the continental A. lakhonen-
sis. They share with this species the shape of the
tepals and the structure of the seed, which has a
double testa. This last characteristic distinguishes
them from A. loriae from Papua, which has a
single testa and with which they were confounded
before. It is highly desirable to have fresh material
from Celebes for further study.
3. Aponogeton loriae MARTELLI, Nuova Giorn.
Bot. Ital. 2, 3 (1897) 472, t. 8; ENGLER & KRAUSE,
Pfi. R. Heft 24 (1906) 12; RENDLE, J. Bot. (1923)
Suppl. 58; STEEN. Fl. Mal. I, 4 (1948) 11, excl.
descr.; BRUGGEN, Blumea 18 (1970) 473, f. 29, Sb.
— A. crispus (non THUNB.) F. v. M. Descr. Not.
Pap. Pl. 8 (1886) 51; Ripv. J. Bot. 24 (1886) 359.
— A. monostachyon (non L. f.) HEMSL. Kew Bull.
(1899) 113. — Fig. 1%, 4b, 5.
Tuber up to 214 cm g. Leaves submerged, very
hard in texture, greenish red, 15—65(—80) by 1-334
cm; base narrowly cuneate, apex (narrowly)
cuneate with a blunt tip; main nerves 7-9, con-
nected by numerous cross-veins at a 50° angle;
margin flat, waved or slightly crisped; petiole
2-15(—30) cm. Peduncle up to 60 cm, c. 3144 mm @,
not or slightly thickening towards the inflores-
cence. Spathe up to 22 mm, persistent, seldom
caducous; spike up to 12cm, densely or rather
laxly flowered, scentless. Tepals 2, obovate,
(greenish) yellow, 1-2 by 1-134 mm, 1-nerved.
Stamens 6, 114-2 mm, filaments not or slightly
widened towards the base; filaments, anthers, and
pollen yellow. Ovaries 3, 144-144 by 34-1 mm,
yellow; ovules 4-8. Infructescence cylindrical,
very dense. Fruits up to 6 by 3144 mm, with a
terminal beak, greenish brown. Seeds with a
simple testa; embryo 214-4 by 34-1 mm, greenish
brown; plumule very small, completely covered
APONOGETONACEAE (van Bruggen) 2ty
1971]
MMT LL
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‘Fig. 4.4
‘, Habit. — A. loriae MARTELLI. b. Fragment of leaf showing
, b Brass 5567) (Courtesy Blumea).
a’ NGF 17717
nogeton womersleyi BRUGGEN. a~a
venation. All = 4 (a Brass 8671
po:
218
FLORA MALESIANA
[ser. I, vol. 77]
yi
we)
SSS
To
AO
=
=
~
ns
Fig. 5. Aponogeton loriae MARTELLI. From the
type, after MARTELLI, X c. 4.
by the margin of the cotyledon, and therefore
nearly invisible.
Distr. Malesia: East New Guinea (Papua,
Central District: Laloki R., c. 3 miles E of Sogeri
patrol station, 147°25’E, 9°30’S; Sogeri: Si-
rinumu area, c. 3 miles S of Sogeri; Kubuna;
Koitaki; Magibiri; affluent of Laragi; stream near
Ower’s Corner).
Ecol. Common in swiftly flowing, shallow,
stony streams, 100-600 m. F/. June—Jan.
Note. The description is partly based on cul-
tivated specimens from Sirinumu, | received thanks
to the co-operation of Dr. RosppBins and Mrs.
PuULSFORD of the University at Pt. Moresby.
4. Aponogeton womersleyi BRUGGEN, Blumea 18
(1970) 477, f. 21° Sa. — Fig. 14, 4a.
Tuber up to 214 cm @. Leaves submerged, 20-50
by 114-314 cm; base very narrowly cuneate, apex
narrowly cuneate with a blunt tip; main nerves
7 or 9, connected by numerous cross-veins at a
+ 70° angle; margin flat or waved; petiole 5-35
cm. Peduncle up to 114m, strongly thickened
towards the inflorescence. Spathe unknown,
caducous. Spike up to 15 cm, very laxly flowered.
Flowers very small. Tepa/ls 2, white to cream,
broadly obovate or wedge-shaped, 34-1 by 1-1%
mm, 1l-nerved. Stamens 6, 74-14% mm, filaments
strongly widened towards the base and applanate.
Ovaries 3, up to 1 by 34 mm, ovules 4-6. In-
fructescence very dense. Fruits greenish brown,
7-8 by 2-3 mm, laterally beaked. Seeds with a
simple testa; embryo 4-5 by 14 mm; plumule
c. 4mm, attached at + 1/, of the length of the
embryo and partly covered by the margins of the
cotyledon.
Distr. Malesia: East New Guinea (Papua,
Western District: Oriomo R., mouth of Yakup
Creek, c. 64km from sea, 143° E, 8°50’ S; Pen-
zara, between Morehead and Wassi Kussa R.),
2 collections.
Ecol. Usually rooting on muddy bottoms in the
shallower parts of lowland rivers in savannah
forests. Fl. fr. Sept. and Dec.
Vern. Zo-inge, Penzara.
Note. A. womersleyi is clearly different from
A. loriae from the Central District and even in a
sterile state they cannot be confused, as its cross-
veins are at a 70° angle (in A. Joriae at a + 50°
angle).
LEMNACEAE (F. van der Plas, Leyden)
Monoecious, very rarely dioecious, small to minute aquatic annuals, floating
at the surface of the water, or floating just below the surface whereby only a very
small part of the frond is exposed to the air, or completely submerged and then
rising to the surface in the flowering period. Fronds either solitary or connected in
small groups by short to very short hyaline or rarely elongate green stipes (fig. 1
Aa, 1 Ba), symmetric or asymmetric, with reniform, round, linear-lanceolate or
angular dorsal outline, green, with or without red or brown pigment cells, some-
times with both types of pigment; base symmetric or asymmetric, obtuse, emar-
ginate or narrowing into the stipe; apex symmetric or asymmetric, round, obtuse
or acute; margin entire or slightly dentate; dorsal side flat to slightly convex,
smooth or with one or more small papillae; ventral side flat to strongly inflated;
somewhere in the median provided with a ‘node’ (fig. 1 Ac). There the roots,
nerves, new fronds, and flowers emerge. Nerves 0-l-~, running towards the
apex. New fronds attached to the node of the mother frond by means of a
‘stipe’ which is sometimes hardly visible and is connate with their ventral side.
Daughter frond sometimes (in Spirodela) provided with 2 basally connate, roundish
scales inserted at the base of the stipe (fig. 1 Ad, 2 e), unequal, one connate with
the yentral side as far as its node. Roots several, one or none, unbranched, growing
downward from the node; in root-producing species the root(s) closely enveloped
by a sheath, which during growth is circumscissile-dehiscing, leaving a basal
sheath (in some species soon hardly visible) and a ‘calyptra’ on top. Budding
pouches 2 (fig. | Ab) or | (fig. 1 Cb); if there is one budding pouch this is basal,
median, dorso-ventrally flattened or funnel-shaped and it produces only new
fronds; when there are 2 budding pouches these are lateral, one on either side of
the axis, dorso-ventrally flattened and produce new fronds, one pouch may
give rise to an inflorescence. In taxa with only | budding pouch (fig. 1 Cj; subfam.
Wol ffioideae) the inflorescence is borne in a median or lateral dorsal flowering
cavity (an exception is extra-Mal. Wolffiopsis which has 2 dorsal flowering ca-
vities ), without a spathe and consisting of | female and | male flower. In taxa with
2 budding pouches (fig. 3 k, 6 e-f) the inflorescence is surrounded by a spathe and
consists of 1 female and 2 male flowers (female flower rarely absent). Perianth
none. — Male flower consisting of | stamen, anther uni- (fig. 8 d) or bilocular
(fig. 2 f), apically or transversely dehiscent; filament short or long and slender;
pollen grains 17-21 «2, spinose. — Female flower consisting of 1 globular ovary
with a short persistent style (fig. 3 k, 6 f), and containing 1-4 ovules. Ovules
orthotropous, amphitropous or anatropous. Fruit symmetric (fig. 5 d—e) or asym-
metric, 1—4-seeded, globose or laterally compressed, winged or without wings.
Seeds smooth or ribbed (fig. 4 1), with little or no endosperm; operculum and
chalaza prominent.
Distribution. There are 6 genera with c. 30 spp. all over the world, obviously introduced in oceanic
islands (see under dispersal). The genera Spirodela, Lemna, and Wolffia are widely distributed in the
temperate and tropical zones; the other genera have a more restricted range. Wolffiella occurs in the
subtropical and tropical parts of America and in South Africa, Pseudowolffia is restricted to tropical
Africa, and Wolffiopsis has been found in the tropics of Africa and America. See Den HARTOG & VAN DER
PLas (Blumea 18, 1970, 355-368).
(219)
220 FLORA MALESIANA (ser. | vole
Fig. 1. Diagrams showing the external morphology of Lemnaceae.
A-B. Spirodela polyrhiza (L.) SCHLEID. A. Dorsal side of the frond, B. ventral side of the frond. —
C. Wolffia sp.
In all figures: a = stipe, b = budding pouch, c = node, d = young frond showing the dorsal scale,
e = ventral scale, f = primary root, g = secondary root, h = root sheath, i = root cap, j = flowering
cavity with 3 and 92 flower.
In Malesia the first three mentioned genera occur with together 6 spp. As Lemnaceous plants are dis-
tinctly under-collected, their precise distribution is very incompletely known, and consequently range
extensions may be expected. For this reason 3 spp. are added here which have not yet been found in Ma-
lesia, but which may be found in future.
In oceanic islands Lemnaceae are very scarce and possibly all introduced according to Guppy. From the
Pacific islands the following are known to me from collections: Hawaiian Is. Spirodela polyrhiza and
Lemna perpusilla (HELLER 1895, LE Roy TopPpinG and DEGENER 1927). L. perpusilla is, besides in Hawaii,
also collected in Polynesia already during the Wilkes’ Exploring Exp. from Fiji (non vidi), and further
known from New Caiedonia (last century), Samoa (1893) and Tonga (1926). Spirodela punctata is also
known from Fiji and New Caledonia. This is also recently collected on Mahé Atoll (Maldives, Indian
Ocean). —
Ecology. Lemnaceae are found in fresh water, or sometimes in brackish water or on wet mud; in
stagnant waters of canals, ditches and small ponds, sometimes also in small, sluggish streams. They
usually indicate eutrophic conditions, and are generally favoured by organic pollution. In Malesia they
occur up to 2100 m, outside Malesia they have been found much higher, e.g. Lemna trisulca up to 3000 m.
They occur in monospecific growths as well as mixed with other Lemnaceous species; they associate also
with other small aquatics, e.g. with water ferns of the genera Salvinia and Azolla or with the liverwort
Ricciocarpus natans. Submerged Lemnaceae sometimes associate with liverworts of the Riccia fluitans
complex. They mostly form a thick layer at the surface of the water or just below it, and form a nuisance
when they choke drainage ditches or cover fish ponds.
The most common species in Malesia are Spirodela polyrhiza, S. punctata (Java!), Lemna perpusilla,
and Wolffia globosa (Malay Peninsula).
The ecological demands of the various species are generally similar, although not exactly the same, and
the tolerance to various environmental conditions differs from species to species. In India MAHESHWARI &
Kapiti (Amer. J. Bot. 50, 1963, 679) only found Wolffia microscopica in association with Lemna perpusilla,
while Wolffia arrhiza (no doubt W. globosa was meant) was only found together with Spirodela polyrhiza.
In the tropics of East Asia and Australia, however, mixed populations of Wolffia globosa and Lemna
1971] LEMNACEAE (van der Plas) 221
perpusilla, with or without Spirodela polyrhiza, are not rare. Thus the combinations of these species ap-
pear to be of local interest.
Between the roots and in the root sheaths of the Lemnaceae blue-green Algae of the genera Nostoc
and Anabaena have often been found.
Lemnaceae are attacked especially by non-aquatic insects; MCCANN (J. Bomb. Nat. Hist. Soc. 43,
1942, 152) found beetles (Curculionidae) and the larvae of a moth (Nymphula responsalis WALKER)
feeding on the upper parts of the fronds of Lemna and Spirodela. Eggs, larvae and cocoons were found
within the fronds. Many small aquatic animals choose the fronds as a substratum but generally do not
feed on them.
Vegetative and sexual reproduction. Multiplication takes place mainly by budding. Some species,
e.g. Spirodela polyrhiza, produce specialized resting buds, i.e. modified fronds consisting of compact tissue
filled with reserve food, for surviving less favourable periods (drought, low temperatures). In Spirodela
and Lemna, the most primitive members of Lemnaceae, the budding pouches serve for sexual as well as
for vegetative reproduction. In the more evolved genera, Wolffia, Wolffiella, Pseudowolffia and Wolffiopsis
the budding pouch serves only for vegetative reproduction, while the inflorescence is borne in 1 or 2
special dorsal flowering cavities, which are only formed when the frond starts flowering.
For a number of species flowering is a rare feature, and of a few species the flowers are still unknown.
However, as the flowers are small, inconspicuous and ephemeral, it is possible that flowering has often
remained unobserved; only when a whole population is flowering this phenomenon is easily detected.
Gregarious flowering is possibly restricted to very specific ecological conditions. In herbarium material
one single flowering or fruiting plant is often to be found among many sterile ones.
Monoecious plants are most common; plants carrying flowers of one sex only have rarely been found in
Lemna gibba (DEN HARTOG, Gorteria 4, 1968, 90-92). In Spirodela polyrhiza SCHLEIDEN (Beitr. Bot. 1844,
230) found “male” inflorescences, in which the male flowers were normally developed but the female
flower was abortive.
The male and female flowers of one inflorescence are usually not mature at the same time; proterandry
or proterogyny are more or less a rule. In Lemna species even the two male flowers do not grow out of
the spathe at the same time (fig. 3 k, 4i). The various types of flowering sequence may be exhibited in a
single species, although the various populations are always uniform in this respect. As a consequence
of this, self-pollination is excluded. Cross-pollination is possible by contact between the stamens of one
plant with the stigma of another one. This contact pollination can easily be brought about by water
movements (TRELEASE, Proc. Boston Soc. Nat. Hist. 21, 1882, 410-415; MCCANN, J. Bomb. Nat. Hist.
Soc. 43, 1943, 151; DEN Hartoc, Gorteria 2, 1964, 68-72). MCCANN /.c. reports pollination by small
insects. How pollination takes place in the species with a dorsal flowering cavity is not known.
In Lemna the seed is usually liberated from the fruit by decay or, more seldom, by splitting of the
pericarp. Sometimes the fruit becomes detached from the frond and the seed remains enclosed by the
pericarp. In autumn the fruits often remain attached to the dying fronds and sink with them to the
bottom. When the seed germinates the pericarp is often still present (DE SLOOvER, Naturalistes Belges 47,
1966, 449). Germination (fig. 4 n) can take place immediately and sometimes starts while the fruit is
still attached to the frond (Lupwic in Kirchner, Loew & Schréter, Lebensgesch. Bliitenpfl. Mittel-
europas I, 3, 1934, 59). In general the seeds germinate in the next favourable period, i.e. after the winter
or after a period of drought. After having lain at the bottom during the resting period, the seed rises to
the surface when germination starts. First the operculum is pushed off by the expanding plumule (fig. 4n);
the latter is green, rootless and does not resemble the normal frond. It gives rise to a new root-bearing
frond, which in its turn gives off new fronds, and decays after some time. The first frond is slightly curved,
the side that is more developed has vascular tissue and a budding pouch, both lacking on the other side.
The second frond is normal, but is curved in the same way as the first frond.
All fronds arising from one seed are homodromous, i.e. all daughter fronds are similar to the mother
frond. It appears namely that the 2 budding pouches of a frond are not exactly identical; one of them is
somewhat larger and gives off more daughter fronds than the other. This better developed budding
pouch may be the left one in the one population and the right in another one (De SLoover, Naturalistes
Belges 47, 1966, 443-456; ENGLER in E. & P. Nat. Pfl. Fam. 2, 3, 1889, 156).
Dispersal. Dispersal of the whole plant as well as the seeds over short distances may be effectuated
by birds; Wolffia brasiliensis was found by WeppeLv on the feathers of a shot bird (Ann. Sc. Nat. 3,
Bot. 1849, 155). Over very short distances transport by amphibians is possible (RipLey, Disp. of PI.
1930, 542-543). Water currents transport the plants and seeds as well. Transport over long distances is
restricted by the fact that desiccation of the plants is a distinctly limiting factor. Seeds sink after coming
free. Several islands or groups of islands have populations which are characterized by a particular pattern
of pigmentation (Spirodela punctata in Fiji), a large number of roots (S. punctata of Java), or a particular
shape (Lemna perpusilla of New Guinea). This ‘raciation’ implies that there is no regular exchange between
‘the various populations, and that the island populations have obviously largely developed from isolated
clones. This, and the fact of an endemic species in India (Wolffia microscopica), pleads against long-
distance dispersal. In the oceanic islands of the Pacific (New Caledonia and Fiji) and the Indian Ocean
(Maldives) Spirodela punctata occurs, but this species is notorious for being dispersed by man, e.g. in the
rice fields of northern Italy and is said by Dauss to be introduced in America, Gurry concluded (Nat.
222 FLORA MALESIANA {ser. I, vol. 7}
Pac. 2, 1906, 407-408) to its introduction into the Pacific islands and excluded dispersal of Lemnaceae
over wide seas.
Taxonomy. Two subfamilies can be recognized within the Lemnaceae, the Lemnoideae and the Wolffi-
oideae. The Lemnoideae are characterized by the presence of roots, 2 budding pouches, an inflorescence
with 2 male flowers and a membranous spathe. The Wolffioideae have only 1 budding pouch and an in-
florescence with 1 male flower; roots and spathe are absent. See for our new system of Lemnaceae
C. DEN HARTOG & F. VAN DER PLAS (Blumea 18, 1970, 355-368).
Affinity within Lemnaceae. In comparing the genera, it is clear that there is an increase in reduction
from Spirodela, via Lemna, to the Wolffioideae, Lemnoideae having 2 male flowers with bilocular stamens,
roots, and a spathe, against Wolffioideae with 1 male flower with unilocular stamens but with neither roots
nor spathe. The latter appear further derived because of the differentiation of the reproductive structures,
viz the (vegetative) budding pouch and dorsal flowering cavity.
Compared with Spirodela, Lemna seems the more reduced or specialized genus, each frond producing
only one root, or none at all; the scales are connate with the root sheath (only visible as two lateral wings
on the root sheath in L. perpusilla), the stipe is entirely merged with the daughter frond, while the budding
pouches are more marginal.
As to the position of the flowers, the gynoecium is placed at the dorsal side of the pouch, the 2 anthers
at the ventral side, their insertions forming a sort of triangle.
Affinity of Lemnaceae with other families. The interpretation of the flower structure has been various.
EICHLER (Bliithendiagramme 1, 1875, 73-80), and recently LAWALREE (Bull. Soc. R. Bot. Belg. 77, 2, 1945,
27-38) homologize the spathe of Lemnoideae with the membranous floral envelope in Najadaceae and
accept the floral structure of each frond to represent a single bisexual flower.
Most authors, however, homologize the floral envelope as a true spathe comparable with that found in
Araceae (in the aquatic Pistia it is also membranous) and the gynoecium and stamens as representing each
a reduced unisexual flower. Furthermore, the seed structure of Lemnaceae closely resembles that of
Araceae according to S. C. MAHESWARI & R. N. Kapri (J. Ind. Bot. Soc. 43, 1964, 270-277); in both
groups there are 2 integuments. The same authors (Am. J. Bot. 50, 1963, 907-914) could show earlier
that the endosperm in Lemnaceae is cellular, and not helobial as stated by LAWALREE, which is also in
favour of affinity with Araceae and not with Helobiae. Furthermore, seeds of Lemnaceae have an oper-
culum as in Araceae according to JoHRI (Rec. Adv. Embr. Angiosp. 1963, 398) which does not occur in
Helobiae.
Among Araceae, the aquatic genus Pistia is possibly most related to Lemnaceae, in having in each spathe
a single female flower and a clump-like whorl of 2-8 stamens separate from it. Furthermore, ENGLER
showed (in E. & P. Nat. Pfl. Fam. 2, 3, 1889, 154-164) that the germination of the seeds in Pistia and
Lemna proceeds in a rather similar way. Besides, both Pistia and Lemna have in common the occurrence
of a similar vegetative reproduction. In Pistia stolons are developing from leaf-axils, producing at the
end a new rosette. In Lemnoideae new fronds are produced on a stipe emitted from the budding pouch
of the mother frond at the node and partly connate with it. In Pistia the young plant is originally sur-
rounded by a membranous early disrupted sheath. A very comparable structure is found in Spirodela,
where the daughter frond is basally enveloped by the partially connate dorsal and ventral scale.
It seems almost impossible to escape from the impression that these scales are homologous with the
envelope in Pistia. In passing, it may be remarked that in Aponogeton undulatus Roxs., which also pro-
duces runners with apical plantlets, the latter are surrounded by a membranous spathe. The morphologic
interpretation of the spathe cg. scales is unclear.
Whereas these arguments indubitably show the affinity of Lemnaceae with Araceae, the homology of
the ‘frond’ of Lemnaceae is less evident. VELENOVSKI (Vergl. Morph. Pfl. 2, 1907, 339-345) mentioned
the occasional reduction of the leaves in Pistia to a few or even one. In both the frond of Lemnoideae
and Pistia veins are curvinerved. I envisage to elaborate the idea of the homology of the Lemnaceous
frond and the leaf rosette of Pistia in another paper.
Palynology. The palynological data give no indication about the systematic relationship of the
Lemnaceae with other families. ERDTMAN (Pollen Morph. Pl. Tax. Angiosp. 1952, 232) found that the
pollen of some Lemnaceae have some characters in common with the pollen of some Araceae, while
the pollen of the Najadaceae seems to be less similar. THANIKAIMONI (Trav. Sect. Sc. & Tech. 5, 5, 1969,
19-26) found no affinity between Pistia and the Lemnaceae. The pollen type of the Lemnaceae has not
been found in the genera of the Araceae so far studied by THANIKAIMONI.
Palaeobotany. Fossil Lemnaceae are very scarce; most fossils described as Lemna are not Lemna-
ceae at all. HENsLow (J. Linn. Soc. Bot. 29, 1893, 486-487) reported a Protolemna described by SAPORTA
from strata above the Later Jura. This plant resembles the present Lemnaceae in some respects, but it
possesses an axis bearing leaves with a dicotyledon-like venation. BEATSON (New Phytol. 54, 1955,
208) reported subfossil Lemna-like pollen in Quarternary deposits. DARRAH (Textbook of Paleobotany,
1939, 300-302) reported a Spirodela from the Miocene and questioned the identity of some species re-
ported from the Cretaceous and the Tertiary.
Phytochemistry. Oxalate of lime (raphides; clustered crystals) is present in many, but by no means
in all, members of the family. So-called myriophyllin cells (see sub Haloragaceae) occur in species of
Spirodela, Wolffia and Wolffiella. The flavonoid constituents were investigated thoroughly. Four main
1971] LEMNACEAE (van der Plas) 223
types of flavonoid compounds occur in the family and each species and genus is said to be characterized
by a distinct pattern of flavonoids. According to J. W. McCLure & R. E. Alston (Am. J. Bot. 53, 1966,
849-860) and B. L. TURNER (Chemistry Nat. Prod., 4th Int. Symp. Stockh. 1966, 1967, 201-205)
Spirodela shows the most complex pattern of flavonoid compounds; anthocyanins, flavonols, flavones
and C-glycoflavones are present. In Lemna the property to produce flavonols is lost. The only flavonoid
compounds observed in Wolffiella are flavonols. Three species of Wolffia were found to contain only
flavonols and two species to produce only flavones and glycoflavones. Two lines of reductive biochemical
evolution, both starting with Spirodela are evident: Spirodela — Lemna — Wolffia p.p. and Spirodela >
Wolffiella — Wolffia p.p. The American authors assume that the evolution of taxa most probably fol-
lowed the same pathway and that Wolffia, as defined by morphology, represents a biphyletic group.
Regrettably, members of the genera Pseudowolffia and Wolffiopsis were not included in these studies.
The pectic substances of cell walls of Lemnaceae contain galacturonic acid, apiose and xylose (E. BECK,
Z. Pflanzenphys. 57, 1967, 444-461). Such apiose-containing pectines occur in all members of the family
investigated hitherto. This striking chemical feature, however, seems to be linked with ecology rather than
with systematics (C. F. VAN BEUSEKOM, Phytochemistry 6, 1967, 573-576).
Most authors assume intimate relationships between Araceae and Lemnaceae. The chemical characters
so far known from both families (HEGNAUER, Chemotax. d. Pfl. 2, 1963, 73-99, 267-269, 483) are in
perfect agreement with such a hypothesis. The fact, however, that chemical information about these
families is still scanty implies that such a statement indicates lack of negative evidence, rather, than strong
positive evidence. — R. HEGNAUER.
Uses. Because of their high rate of vegetative reproduction and the ease with which they can be cul-
tured, Lemnaceae are often used in plant physiological experiments (HILLMAN, Bot. Rev. 27, 1961,
221-287).
Collecting & preservation. For the identification of Wolffia species it is necessary to know the
exact shape of the fronds. Unfortunately the usual procedure of boiling dried specimens for a while is
inadequate as the pressed fronds generally remain flat. Those which do not are mostly resting buds
which usually have a somewhat aberrant shape. Therefore, it is recommended that when collecting the
plants should be put in fluid at once. Formalin is not advisable since the plants become very fragile in it,
although they keep their original form. They remain more plastic and cannot be damaged in any way
when collected in FAPA (ingredients for | liter: 50 cc formalin 40 %, 25 ce acetic acid, 25 cc propionic
acid and 900 cc spirit 50-70 %).
KEY TO THE GENERA
1. Roots present. Budding pouches 2, basal, lateral, one on either side of the axis. Inflorescence de-
veloping from one of the budding pouches, consisting of 1 2 and 2 g flowers enclosed by a membranous
spathe. Anther bilocular, transversely dehiscent. Stipe present. Raphides present.
2. Fronds with a dorsal and a ventral scale, one to many roots and 3-15 nerves. Stipe ventrally at-
tached. Brown pigment cells and druses in the parenchyma. .......... . 1. Spirodela
2. Fronds without dorsal and ventral scales, with one root (rarely none) and 1-3, often indistinct
nerves. Stipe marginally attached. Brown pigment cells and druses absent... . .. . 2. Lemna
1. Roots none. Budding pouch 1, median, never giving rise to an inflorescence. Inflorescence 1, dorsal,
consisting of a cavity containing 1 2 and | 3 flower, without a spathe. Anther unilocular, apically
dehiscent. Stipe not visible to the naked eye. Raphides absent... .......4... 3. Wolffia
1. SPIRODELA
ScHLeEID. Linnaea 13 (1839) 391, nom. gen. cons. prop.; HEGELM. Lemnac. (1868)
147; Bot. Jahrb. 21 (1895) 283; DAauss, Monogr. Lemnac. (1965) 8; HARTOG,
Taxon 19 (1970) 647-648; HARTOG & PLAS, Blumea 18 (1970) 358. — Lenticularia
Sfécuter, Pl. Veron. 3 (1754) 129, nom. gen. rejic. prop. — Lemna sect. Spirodela
Coss. & GERM. FI. Env. Paris 2 (1845) 577. — Lemna subg. Spirodela PETERM.
Deutschl. Fl. (1849) 540; Rouy, FI. Fr. 13 (1912) 285, — Fig. 1 A-B, 2-3.
Small water plants, floating at the surface. Fronds either solitary or connected
in groups of 2~5 (sometimes even more), symmetric or asymmetric, reniform to
obovate, flat or distinctly inflated; dorsal side flat or slightly convex, smooth
or with some small median papillae; underside often red due to pigment cells
in the epidermis; brown pigment cells, raphides and druses in the parenchyma;
stomata on the dorsal side; margin entire; nerves 3-15. Stipe hyaline, fugacious,
224 FLORA MALESIANA (ser. I, voli #4
short or relatively long, attached to the underside of the frond. Dorsal scale
present, fugacious in older plants. Ventral scale broad, often pigmented. Roots
1-18, more or less vascularized; root sheaths short, clustered together and co-
vered by the ventral scale with which some of them are adnate; one or more
roots perforating this scale (primary roots); root cap straight or slightly curved,
acute or rarely obtuse. Budding pouches 2, basal, lateral, one on either side of the
axis, dorso-ventrally flattened, more or less triangular in outline, opening by a
transverse slit, which is ventral to the margin of the frond. Jnflorescence 1, lateral,
developing from one of the budding pouches, consisting of 1 2 and 2 3 flowers,
enclosed by a membranous spathe. Anther bilocular, transversely dehiscent.
2 Flower superior and lateral to the 2 3 flowers, consisting of a globular ovary
with a short, partly persistent style and 1 amphitropous or 2-4 anatropous ovula.
Fruit asymmetric, slightly or distinctly winged. Seed smooth or longitudinally
ribbed.
Distr. Worldwide, the Arctic regions excepted, consisting of 4 spp.
KEY TO THE SPECIES
1. Fronds 5-12 mm long. Roots 7-16, one perforating the ventral scale. Nerves 7-12. Seeds smooth.
1. S. polyrhiza
1. Fronds 2-5 mm long. Roots 1—9(—12), all perforating the ventral scale. Nerves 3—5. Seeds ribbed.
1. Spirodela polyrhiza (L.) ScHLEID. Linnaea 13
(1839) 392; ZoLt_. Nat. Geneesk. Arch. N. I. 2
(1845) 212; Hassx. Flora 30 (1847) 469; Mia.
Fl. Ind. Bat. 3 (1855) 222; Kurz, Nat. Tijd. N. I.
27 (1864) 220; HEGELM. Lemnac. (1868) 151-152;
Bot. Zeit. 29 (1871) 621, 645; FRANCH. & SAV.
En. Pl. Jap. 2 (1879) 13; HEGELM. Bot. Jahrb. 21
(1895) 284; THomPpson, Rep. Mo. Bot. Gard. 9
(1897) 27; UsTeri, Vierteljahrschr. Naturf. Ges.
Ziirich 50 (1905) 453; Merr. Fl. Manila (1912)
134; En. Philip. 1 (1923) 190; Back. Handb. FI.
Java 3 (1924) 1; Onkruidfl. Jav. Suikerr. 1 (1928)
175; SteEN. Arch. f. Hydrobiol. Suppl. 11 (1932)
292; HAND.-MAzz. Symb. Sin. 7 (1936) 1368;
JUMELLE in Humbert, Fl. Madag. fam. 32 (1937)
3-4; McCann, J. Bomb. Nat. Hist. Soc. 43 (1942)
156-157; MASAMUNE, FI. Kainantensis (1943) 394;
Sc. Rep. Kanazawa Univ. 5 (1957) 92-93;
Dauss, Monogr. Lemnac. (1965) 10-13, incl. var.
masonii DAuBS; Hess, LANDOLT & HIRZEL, FI.
Schweiz 1 (1967) 497; Back. & BAKH. f. FI. Java
3 ( 1968) 127; CLARK & THIERET, Mich. Botanist
7, 2 (1968) 69; HARTOG & PLAs, Blumea 18 (1970)
360. — Lemna polyrhiza LINNE, Sp. Pl. (1753)
970; Kurz, J. Linn. Soc. Lond. 9 (1866) 267-268;
BENTH. FI. Austr. 7 (1878) 164; NaAves, Novis.
App. (1882) 296; Hook. f. Fl. Br. Ind. 6 (1893)
557; Trim. Handb. FI. Ceyl. 4 (1898) 367; RIDL.
J. Str. Br. R. As. Soc. n. 33 (1900) 179; BRowN,
FI... Trop. Afe:: 8, 2) (1901) 2013 t@orters,, Fl:
Simlensis (1902) 545; Wricut, J. Linn. Soc. Bot.
36 (1903) 188; PRAIN, Bengal Pl. 2 (1903) 841;
RIDL. Mat. Fl. Mal. Pen. 3 (1907) 53; F. M. BAILEy,
Queensl. Agric. J. 2, 1 (1914) 78; Ripi. J. Mal.
Br. R. As. Soc. 1 (1923) 106; FI. Mal. Pen. 5
2. S. punctata
(1925) 132; HEND. Gard. Bull. S. S. 4 (1928) 352;
FISCHER in Gamble, FI. Madras 3, 9 (1931) 1593;
GAGNEP. FI. Gén. I.-C. 6 (1942) 1198; Mirra,
Fl. Pl. E. India 1 (1958) 86; HUNDLEY & U CHIT
Ko Ko, List trees etc. Burma ed. 3 (1961) 292;
LarsEN, Dansk Bot. Ark. 20 (1962) 136; Onw1,
Fl. Japan (1965) 264. — Lenticula polyrhiza (L.)
LAMK, FI. Fr. 2 (1778) 189. — Lemna orbicularis
Kit. ex SCHULTES, Oesterr. FI. ed. 2, 1 (1814)
64, nomen seminudum. — Lemna_ thermalis
BEAvuV. J. Phys. Chim. Hist. Nat. 82 (1816) 102,
113, f. 23; NUTTALL, Gen. Amer. 1 (1818) 19. —
Lemna orbiculata Roxs. [Hort. Beng. (1814) 66,
nomen] Fl. Ind. ed. Carey 3 (1832) 565. — Tel-
matophace polyrhiza (L.) Gopr. Fl. Lorr. ed. 1,
3 (1843) 18. — Lemna major [C. A. Meyer, Ind.
Cauc. (1831) 11, nomen] GrirF. Not. 3 (1851)
216; Ic. Pl. As. 3 (1851) t. 264. — Telmatophace
orbicularis ScHuR, En. Pl. Transs. (1866) 635. —
S. atropurpurea MONTAND. Guide Bot. (1868)
309. — Lemna maxima BLATTER & HALLB. J.
Ind. Bot. Soc. 2 (1921) 49. — S. maxima (BLAT-
TER & HALLB.) MCCANN, J. Bomb. Nat. Hist.
Soc. 43 (1942) 158. — Fig. 1 A-B, 2.
Fronds solitary or cohering in groups of 2-5,
symmetric or asymmetric, reniform to round or
obovate, 3-12 by 214-8 mm; base obtuse; apex
obtuse or round; dorsal side flat, smooth; ventral
side flat to strongly inflated; green, ventral side
and margin often red-purple; nerves 7-12; stipe
hyaline, often long, prominent; dorsal scale mem-
branous, orbicular to reniform, attached to the
base, disappearing in older plants; ventral scale
membranous, broad, with much brown pigment,
almost centrally attached near the node, covering
|
.
1971]
LEMNACEAE (van der Plas)
Fig. 2. Spirodela polyrhiza (L.) SCHLEID. a. Habit, mother frond with adhering offspring, dorsal view, x 6,
b. habit, ventral view, the sheaths of the secundary roots covered by the ventral scale, x6, c. detail from
ventral side, the primary root piercing the ventral scale, «12, d. root cap, x12, e. turion, dorsal view
(left) showing dorsal scale and ventral view (right) showing ventral scale, primordium of primary root,
stipe a minute protuberance, 18, f. detail of flowering frond, with style and 2 stamens enveloped
by the spathe, « 18 (a—d VAN DER PLAS 23-9-69, e HALL Sept. 1867, f STALIN 451).
the bases of the roots. Roots 7-16, one (very
rarely 2) perforating the ventral scale (primary
roots), the others passing between the scale and
the frond (secondary roots); primary roots ap-
pearing earlier than the secondary ones. Root
cap acute. Ovary with 1 amphitropous ovule or
2 anatropous ones. Fruit a 1-2 seeded utricle,
slightly winged. Seed smooth.
Distr. Almost cosmopolitan, common in the
northern hemisphere, rather rare in Africa and
Australia, in South America replaced by two
closely related species; in Malesia common:
Sumatra, Malay Peninsula, W.-E. Java, Flores
Sea (Tanah Djampea, S of Celebes), N. Celebes,
Philippines (Luzon to Mindanao), New Guinea.
Also in the Hawaiian Is. (1895 Heiter; 1927
Le Roy Torrina), together with Lemna perpusilla.
Ecol. Stagnant fresh water habitats such as
sawahs, ponds and ditches, rarely in slightly
brackish water, from the lowland to c. 2100 m.
Vern. Kakaréwoan, 8, mata leéle, J.
Notes. Like many other Lemnaceae this species
often forms resting-buds to survive less favourable
times. In S. polyrhiza these resting-buds are
easier to distinguish from the normal fronds than
in the other Lemnaceous species and are often
called ‘turions’. They are produced in the normal
way in the budding pouch. The turion is reniform,
dorso-ventrally flattened, variable in size, con-
sisting of dense tissue. Its roots are still present
but when dormant only the primary roots are
visible. On both sides it bears a membranous
scale as does a normal young frond.
2. Spirodela punctata (G. F. W. Meyer) THomp-
SON, Rep. Mo. Bot. Gard, 9 (1897) 28; Dauss,
Monogr. Lemnac. (1965) 15; Clark & THieret,
Mich. Bot. 7, 2 (1968) 69; Harroc & PLAs,
Blumea 18 (1970) 360. — Lemna punctata G. PF.
W. Meyer, Prim. Fl. Esseq. (1818) 262. —
Lemna gibba (non L.) Bianco, FI. Filip. ed. 1
(1837) 672; ed. 2 (1845) 468; ed. 3, 3 (1879) 78;
226
Naves, Novis. App. (1882) 296. — Lemna oli-
gorrhiza Kurz, J. Linn. Soc. Lond. 9 (1866) 267,
t. 5, f. 1-6; BENTH. FI. Austr. 7 (1878) 163;
Naves, Novis. App. (1882) 296; Hook. f. FI. Br.
Ind. 6 (1893) 557; PRAIN, Bengal PI. 2 (1903) 841;
Guppy, Obsery. Nat. Pac. 2 (1906) 407-408;
SCHWARZ in Fedde, Rep. 24 (1928) 80; GREENWOOD
Proc. Linn. Soc. 154, 2 (1943) 104; GUILLAUMIN,
Fl. Nouv.-Caléd. (1948) 48; Mitra, Fl. Pl. E.
India 1 (1958) 86; GuILLAUMIN, Mém. Mus.
Hist. Nat. Paris n.s. sér. B, Bot. 8 (1959) 189;
ibid. (1962) 213; PARHAM, PI. Fiji Is. (1964) 268.
— Lemna melanorrhiza F. y. M. ex KurRzZ, J. Bot.
5 (1867) 115. — Lemna pleiorrhiza F. vy. M. ex
Kurz, lc. — S. oligorrhiza HEGELM. Lemnac.
(1868) 147-150, incl. var. B melanorrhiza (F. vy. M.
ex KURZ) HEGELM., var. y pusilla HEGELM., var. 6
pleiorrhiza (F. vy. M. ex KURZ) HEGELM., var. €
javanica HEGELM.; ENGL. Bot. Jahrb. 7 (1886) 448;
HEGELM. Bot. Jahrb. 21 (1895) 287; F. M. BAILEY,
Queens]. Fl. 5 (1902) 1700; Miki, Bot. Mag.
Tokyo 48 (1934) 333; McCann, J. Bomb. Nat.
Hist. Soc. 43 (1942) 157; BLAcK, FI. S. Austr. ed.
2 (1948) 172; Dauss, Monogr. Lemnac. (1965)
14-15; Onwt, FI. Japan (1965) 264; McComsB &
McComs, J. R. Soc. W. Austr. 50, 4 (1967) 107;
Back. & BAKH. f. Fl. Java 3 (1968) 127. — S.
melanorrhiza HEGELM. Bot. Jahrb. 21 (1895) 287.
— §. pusilla HEGELM. I.c. — S. pleiorrhiza HE-
GELM. /.c. 288. — S. javanica HEGELM. I.c. — Fig 3.
Fronds solitary or cohering in groups of 2-6,
asymmetric, elliptic to obovate, 2.8—4.5 by 1.4-2.6
mm, base and apex asymmetric, obtuse; mostly
thick, dorsal side flat to slightly convex, smooth
or with a median row of small papillae; ventral
side flat to convex, sometimes slightly inflated and
with relatively few large air spaces; green, some-
times red; nerves 3—5; stipe hyaline, often long and
distinct; dorsal scale strongly reduced, fugacious;
ventral scale membranous, often pigmented,
perforated by the short sheaths of all (1-9,
sometimes 12) roots; root cap straight or slightly
curved, acute, with or without pigment. Ovary
with 1 amphitropous ovule. Fruit asymmetric,
winged. Seed longitudinally ribbed.
Distr. Apparently rare in Africa and South
FLORA MALESIANA
[ser. I, volews
America, common in Asia and Australia, in the
Indian Ocean in the Maldives (Mahé Atoll),
in the Pacific Ocean in New Caledonia and Fiji;
in Malesia: Malay Peninsula (Pahang) and Java
(west to east).
According to Dauss (1965) introduced in
America, but this must then have been before 1814
when G. W. F. Meyer recorded it from there
already. According to Guppy (1906) introduced in
the Pacific islands. Dispersal by man has cer-
tainly given this species a wider distribution in
recent time; it is e.g. not rare in the rice-fields in
Italy. Also its obviously local occurrence in Ma-
lesia might point to introduction.
Ecol. From sea-level to c. 2100m, both in
everwet areas and areas subject to a severe dry
season.
Notes. S. punctata is a very variable species;
several of its characters show considerable va-
riations in different localities. As a result of this
there are some populations which are characterized
by a certain peculiarity not occurring in other
populations. In some collections from Java the
number of roots is extremely large (up to 12!),
in other areas the number of roots rarely exceeds
5. The plants from New Caledonia and Fiji are
characterized by a much heavier pigmentation
than in other areas. These characters vary in-
dependently of each other, and do not seem to be
constant even within one collection.
It is thus understandable that F. VON MULLER
(ex KuRZ, 1867) on the grounds of a small number
of collections came to recognize 3 species in this
complex, and that HEGELMAIER (1868) distinguish-
ed 5 varieties within S. oligorrhiza (raised to the
rank of species in his paper of 1895). My in-
vestigation of much more material than these
authors had at their disposal leads, however, to
the conclusion that these taxa all belong to a single
species, S. punctata, in which they are linked by
all possible transitions.
From BLANCO’s description of Lemna gibba it is
obvious that the plant in question was in fact
S. punctata. MERRILL (Sp. Blanc. 1918, 93) er-
roneously interpreted L. gibba ‘BLANCO’ as L.
paucicostata (= L.. perpusilla).
2. LEMNA
LINNE, Gen. PI. ed. 5 (1754) 417; Sp. Pl. (1753) 970; HeGeLM. Lemnac. (1868) 134;
Bot. Jahrb. 21 (1895) 288; Dauss, Monogr. Lemnac. (1965) 16; HARTOG & PLAs,
Blumea 18 (1970) 360. — Lenticula [HILL. Brit. Herb. (1757) 530] BOEHMER in
Ludwig, Def. Gen. Pl. (1760) 499. — Hydrophace HALL. Hist. Stirp. Indig. Helv.
Fig. 3. Spirodela punctata (G. F. W. MEYER) THOMPSON. a. Habit, mother frond with fruit and daughter
frond, x 12, b. ditto, x 12, c. ventral view roots piercing ventral scale, «12, d. detail of c, «24, e. ventral
view, l-rooted frond, x12, f. ditto, detail from e, x24, g. habit, dorsal view, 12, h. lateral view, x 12,
i. ventral view of a plant from Java with many roots, x12, j. ventral view of a frond with 2 roots,
< 12, k, detail of the inflorescence, x 24 (a DEN HARTOG 779, b-d DEN HARTOG 5.n., e-f, j-k DEN HARTOG
1161, g-h DEN Hartoc 190, i Koorpers 21531).
1971] LEMNACEAE (van der Plas) 227
a
> ttt le Lid | \
iv. Bee], aT
228 FLORA MALESIANA (ser. I, vole Ye
3 (1768) 68. — Telmatophace SCHLEID. Linnaea 13 (1839) 391. — Staurogeton
Reus. [Consp. (1828) 288, nomen] Nom. (1841) 33. — Lenticularia MONTANDON,
Guide Bot. (1868) 308, non SEGuIER, Pl. Veron. 3 (1754) 129. — Fig. 4-7.
Small aquatics, floating at the surface or sometimes completely submerged in
which case they come to the surface only in the flowering period. Fronds either
solitary or connected in groups of 2-10 (sometimes even more), symmetric to
slightly asymmetric, round, elliptic, oblong, obovate or lanceolate, flat or slightly
swollen, sometimes considerably inflated at the ventral side, often with a median
row of papillae on the flat or slightly convex dorsal side; with or without red
pigment cells in the epidermis; raphides in the parenchyma; no brown pigment
cells; druses none; stomata on the dorsal side of floating plants, but absent in the
submerged plants; margin entire, rarely denticulate; nerves 1-3(—5). Stipe hyaline
and fugacious or green and persistent, attached to the margin. Dorsal as well as
ventral scale absent. Root | (rarely absent), slightly vascularized, with or without
a distinct root sheath; root cap straight or curved, obtuse, mucronate or acute.
Budding pouches 2, basal, lateral, one on either side of the axis, dorso-ventrally
flattened, more or less triangular in outline, opening by a transverse slit; slit
coinciding with the margin of the frond, rarely ventral or dorsal to the margin.
Inflorescence 1, lateral, developing from one of the budding pouches, consisting
of 1 2 and 2 ¢ flowers enclosed by a membranous spathe. Anther bilocular, trans-
versely dehiscent. 9 Flower superior and lateral to the 2 3 flowers, consisting of a
globular ovary with a short, persistent style. Ovary with | orthotropous or amphi-
tropous ovule or with 2-4 anatropous ovules. Fruit symmetric or asymmetric,
more or less compressed, with or without wings. Seeds longitudinally ribbed,
rarely smooth.
Distr. 9 spp., worldwide with the exception of the Arctic regions.
Taxon. The genus can be subdivided into 2 subgenera:
1. Subg. Lemna. Floating water plants with round, elliptic, oblong or obovate fronds. Root 1, with a
distinct root sheath. Slit of budding pouch coinciding with the margin of the frond, rarely ventral to
the margin. Ovary with 1-4 ovules.
2. Subg. Staurogeton Rcus. Fl. Germ. 1 (1830) 10. Submerged water plants, only rising to the surface
in the flowering period. Fronds oblong to linear-lanceolate, with or without a root. Root sheath in-
distinct. Slit of budding pouch just dorsal to the margin of the frond.
KEY TO THE SPECIES
1. Floating water plants. Fronds round to oblong or obovate, with 1 root. Slit of budding pouch coin-
ciding with the margin of the frond, rarely ventral to the margin. Subg. Lemna.
. Root sheath with lateral wings. Fruit asymmetric; ovule 1, orthotropous. . . . . 1. L. perpusilla
. Root sheath without wings. Fruit symmetric; ovules amphitropous or anatropous.
oe)
Fig. 4. Lemna perpusilla Torrey. a. Habit, mother frond with adhering offspring, dorsal view, x 18,
b. root cap, x36, c. lateral view, frond with apical and nodal papillae, x 18, d. ditto, only with apical
papilla, x18, e. ditto, without papillae, 18, f. habit, plant from New Guinea, dorsal view, elongated
fronds, x 18, g. ventral view, showing root emitted, x 18, A. detail of root sheath provided with 2 lateral
wings, x 36, i. mother frond and daughter frond, both with inflorescence, x 18, j. detail of a flowering
frond in which inflorescence and daughter frond develop simultaneously in the same reproductive
pouch, x 36, k. frond with fruit and daughter frond, dorsal view, x 18, /. seed, x 40, m. ditto, longitudinal
section, X 80, 7. germinating seed, showing plumule and first frond, dorsal (left) and ventral view (right),
x18 (a-c, i-n DEN HARTOG 632, d BACKER 19104, e BACKER 25650, f BRASS 8112, g LUTJEHARMS 5387,
h DEN HaArRTOG 188).
LEMNACEAE (van der Plas)
2
=
=
&
:
E
Ee
$
=
=
230
FLORA MALESIANA
iser. I, volo?
3. Frond mostly symmetric, ventral side flat to slightly convex, never inflated. Cells on ventral side
not visible to the naked eye. Fruit without wings. Ovule 1, pane Slit of budding pouch
coinciding with the margin of the frond
L. minor
3. Frond mostly asymmetric, ventral side flat to strongly inflated. Cells on ‘ventral side clearly visible
to the naked eye.
Slit budding pouch ventral to the margin of the frond .
Fruit with lateral wings. Ovule 1,
amphitropous or ovules 2-4, anatropous.
L. gibba
. Submerged water plants, only floating at the surface when flowering. Fronds oblong to linear-lan-
ceolate, often without a root. Slit of budding pouch just dorsal to the margin of the frond. Subg.
Staurogeton.
4. Margin denticulate towards the apex. Stipe usually strongly elongate, green, persistent.
4. Margin entire, Stipe short, hyaline, fugacious. .
1. Lemna perpusilla Torrey, Fl. N. Y. 2 (1843)
245; AusTIN in A. Gray, Man. Bot. ed. 5 (1867)
479, incl. var. trinervis AUSTIN; HEGELM. Lemnac.
(1868) 139: Bot. Jahrb. 21 (1895) 294; THOMPSON,
Rep. Mo. Bot. Gard. 9 (1897) 33; DauBs, Mo-
nogr. Lemnac. (1965) 25-27; BAck. & BAKH. f.
Fl. Java 3 (1968) 127. — L. minor (non L.)
Lour. Fl. Cochinch. 2 (1790) 550; Hassk. Cat.
Hort. Bog. (1844) 53, 302; JUNGH. Reisen (1845)
42: Grirr. Not. 3 (1851) 216; Mia. Pl. Jungh.
(1852) 169; JUNGH. Java ed. 2, 1 (1853) 285-287;
Mio. FI. Ind. Bat. 3 (1855) 221; Kurz, Nat.
Tijd. N. I. 27 (1864) 220; J. Linn. Soc. Lond.
9 (1866) 266; J. Bot. 5 (1867) 115; BENTH. FI.
Austr. 7 (1878) 163, p.p.; NAves, Novis. App.
(1882) 296; Hook. f. Fl. Br. Ind. 6 (1893)
556, p.p.; MASSART, Bull. Soc. R. Bot. Belg.
34 (1895) 184; Koorp. Med. Lands Plan-
tentuin 19 (1898) 305; F. M. BAILEY, Queensl.
Fl. 5 (1902) 1700, p.p.; Guppy, Observ. Nat.
Pac. 2 (1906) 407-408; Koorp. Exk. Fl. Java 1
(1911) 268; Boxtp. Zakfl. Landbouwstr. Java
(1916) 125; Burk. Dict. (1935) 1328; GAGNEP.
FI. Gén. I.-C. 6 (1942) 1197; GuUILLAUMIN,
FI. Nouv.-Caléd. (1948) 48; LaArseEN, Dansk
Bot. Ark. 20 (1962) 136; GUILLAUMIN, Mém.
Mus. Hist. Nat. Paris n.s. sér. B, Bot. 8 (1962)
213; PARHAM, PI. Fiji Is. (1964) 268. — L. aequi-
noctialis WELW. Apontam. Phytogeogr. FI. Prov.
Angola, Ann. Conselho Ultram. Dec. 1858, n. 55.
(1859) 578; Brown, FI. Trop. Afr. 8, 2 (1901) 203;
GIARDELLI, Darwinia 11 (1959) 584-590. — L.
angolensis WELW. ex HEGELM. J. Bot. 3 (1865)
112; H&GeLM. Lemnac. (1868) 141-142; Bot.
Jahrb. 21 (1895) 296. — L. paucicostata HEGELM.
Lemnac. (1868) 139, incl. var. membranacea
HEGELM. /.c. 141; Kurz, J. As. Soc. Beng. 45, ii
(1876) 153; FRANCH. & Sav. En. Pl. Jap. 2
(1879) 12; Hook. f. Fl. Br. Ind. 6 (1893) 556;
HEGELM. Bot. Jahrb. 21 (1895) 294-296; TRIM.
Handb. Fl. Ceyl. 4 (1898) 366; Ripv. J. Str. Br.
R. As. Soc. n. 33 (1900) 179; BRown, FI. Trop.
Afr. 8, 2 (1901) 202-203; Pratin, Bengal Pl. 2
(1903) 841; Wricnt, J. Linn. Soc. Bot. 36 (1903)
188; Usteri, Vierteljahrschr. Naturf. Ges. Ziirich
50 (1905) 453; Guppy, Observ. Nat. Pac. 2 (1906)
407-408; RipL. Mat. Fl. Mal. Pen. 3 (1907) 52;
Me_rr. FI. Manila (1912) 134; Ripu. J. Mal. Br.
R. As. Soc. 87 (1923) 106; Merr. En. Philip. 1
(1923) 190; Back. Handb. FI. Java 3 (1924) 2;
RIDL. FI. Mal. Pen. 5 (1925) 132; HEND. Gard.
2. L. trisulca
3. L. tenera
Bull. S. S. 4 (1928) 352; Back. Onkruidfi. Jav.
Suikerr. 1 (1928) 175; FiscHER in Gamble, FI.
Madras 3 (1931) 1593; STEEN. Arch. f. Hydrobiol.
Suppl. 11 (1932) 291; Merr. Trans. Am. Phil.
Soc. 24, 2 (1935) 100; CuristopH. B. P. Bish.
Mus. Bull. 128 (1935) 44; JUMELLE in Humbert, FI.
Madag. fam. 32 (1937) 2; MCCANN, J. Bomb.
Nat. Hist. Soc. 43 (1942) 153; MASAMUNE, FI.
Kainantensis (1943) 394; Back. Bekn. Fl. Java
(em. ed.) 10 (1949) fam. 226, p. 2; HEND. Mal.
Wild FI. Monocot. (1959) 210; MASAMUNE,
Sc. Rep. Kanazawa Univ. 5 (1957) 94; MitTrRA,
Fl. Pl. E. India 1 (1958) 86; HuNDLEY & U
Cuit Ko Ko, List trees etc. Burma ed. 3 (1961)
292; LARSEN, Dansk Bot. Ark. 20 (1962) 135;
Ouwl, FI. Japan (1965) 265. — L. trinervis
(AUSTIN) SMALL, FI. SE. U.S. (1903) 230; DAuss,
Monogr. Lemnac. (1965) 27. — Hydrophace
perpusilla (TORREY) LUNELL, Am. Midl. Nat. 4
(1915) 237. — L. minima Buiatr. & HALLB. J.
Ind. Bot. Soc. 2 (1921) 50. — L. blatteri MCCANN
J. Bomb. Nat. Hist. Soc. 43 (1942) 153. — L.
eleanorae MCCANN, I. c. 154. — Fig. 4.
Small floating aquatic. Fronds solitary or co-
hering in groups of 2—5, asymmetric, ovate to
obovate or sometimes obovate-oblong, 1.2-4.8
by 0.7-2.8 mm, thin, membranous to thickened,
green, no red pigment cells; base strongly asym-
metric, obtuse or slightly acute; apex nearly sym-
metric or asymmetric, obtuse to slightly acute;
dorsal side slightly convex, with or without a row
of median papillae, but often with a distinct,
hook-shaped papilla near the apex; ventral side
flat to convex, not inflated; nerves 3, sometimes in-
distinct; margin entire. Stipe hyaline, fugacious,
sometimes persistent. Root sheath cylindrical with
2 lateral wings; root cap acute. Slit of budding
pouch coinciding with the margin of the frond.
Ovary with 1 orthotropous ovulum, style sub-
apical, persistent. Fruit asymmetric, ellipsoid,
laterally slightly compressed. Seed with 12-18
distinct longitudinal ribs, laterally slightly com-
pressed.
Distr. Worldwide in the tropics and sub-
tropics. In the Pacific islands in several Ha-
waiian Is. (first 1895), Samoa (1893), Tonga
(1926), Fiji (1842), New Caledonia; common in
Malesia: Sumatra (also Enggano), Malay Penin-
sula, W.-E. Java (also Madura & Kangean Is.),
Lesser Sunda Is. (Timor), Flores Sea (Tanah
Djampea), North Borneo, Philippines (Luzon to
1971]
LEMNACEAE (van der Plas)
231
Mindanao), Moluccas (Ambon; Aru Is.: Maikoor),
and W.-E. New Guinea.
Ecol. The most common duckweed in Malesia.
It occurs mostly in fresh water, but is sometimes
found in slightly brackish water. In sawahs,
ditches and small streams up to 2100m, often
together with other Lemnaceae (Spirodela). Flower-
ing and fruiting occur commonly in this species.
Vern. Kiambang, M (Sum.), ganggéng, S;
Philippines: inalia, lia, liya, Tag.
Notes. BACKER (1924) was the first to point
out that L. minor does not occur in Malesia;
he referred this material to L. paucicostata.
L. perpusilla is an extremely variable species.
HEGELMAIER (1868) distinguished in his mono-
graph 3 species, viz L. perpusilla (seed with 30-60
ribs, restricted to the eastern part of the U.S.A.),
L. paucicostata (seed with 12-20 ribs, occurring
in the tropics of the Old and the New Worlds;
also in the S. and SE. parts of the U.S.A.) and
L. angolensis (seed with 12-15 ribs, and moreover
a conspicuous apical papilla on the frond, restric-
ted to tropical Africa). L. angolensis, already
earlier described as L. aequinoctialis by WELWITSCH,
is with respect to the seed character within the
range of variation of L. paucicostata. The de-
velopment of the apical papilla on the frond is
subject to so much variation, even within one
sample, that it is not useful as a specific character.
THOMPSON (Ann. Rep. Mo. Bot. Gard. 9, 1897, 33)
who investigated L. perpusilla and L. paucicostata
in the U.S.A. found the number of ribs on the
seeds of these species so variable that he could not
distinguish the species with certainty, and for that
reason he regarded them as forms of one species.
Some flat, membranous forms with a distinct
venation, but otherwise not different from L.
perpusilla, have been distinguished as L. per-
pusilla var. trinervis, L. paucicostata var. mem-
branacea or as L. trinervis. They have been re-
corded from America and India. In my opinion they
represent only forms of L. perpusilla, induced by
poor environmental circumstances. MCCLURE
& At.ston (Am. J. Bot. 53, 1966, 849-860),
however, found L. trinervis to be quite distinct
from L. perpusilla with respect to the flavonoid
chemistry. Therefore, membranous forms require
re-investigation.
The fronds from New Guinea are often twice
as long as wide, in contrast to those from the rest
of Malesia.
Merrixt (Sp. Blanc. 1918, 93) referred L. gibba
(non L.) Bianco to L. paucicostata (= L.
perpusilla). As BLANCO recorded the occurrence
of 2 roots, his plants must have belonged to
Spirodela punctata. MERRILL’s representative speci-
men Sp. BI. 131 belongs to L. perpusilla.
Lemna minor Linné, Sp. PI. (1753) 970; He-
Gem. Lemnac. (1868) 142-144; Hook. f. FI. Br.
Ind. 6 (1893) 556, pro parte; HeGcetm. Bot.
Jahrb. 21 (1895) 291-292; Dauss, Monogr.
Lemnac. (1965) 21-25. — Fig. 5 f-h.
Small floating aquatic. Fronds solitary or co-
hering in groups of 2-5 (sometimes many more),
symmetric to slightly asymmetric, round-elliptic
to elliptic or ovate; 1.5-4.6 by 1-2.9 mm); base
++ symmetric, obtuse; apex symmetric or slightly
asymmetric; margin entire; dorsal side flat to
slightly convex, often with a median row of
papillae; ventral side flat to slightly convex;
cells on ventral side not visible to the naked eye,
green; nerves 3. Stipe often persistent and pro-
minent; root sheath cylindrical, without wings,
root cap obtuse. S/it of budding pouch coinciding
with the margin of the frond. Ovary with 1
amphitropous ovule. Fruit ellipsoid, symmetric,
laterally compressed, without wings. Seed smooth.
Distr. Widely distributed in the temperate zone
of the northern hemisphere, reaching as far north
as Alaska and northern Scandinavia. All records
of L. minor from tropical Asia, Malesia and
tropical Australia relate to L. perpusilla.
Lemna gibba LINNE, Sp. Pl. (1753) 970; HEGELM.
Lemnac. (1868) 145-147; Hook. f. Fl. Br. Ind. 6
(1893) 556-557; HEGELM. Bot. Jahrb. 21 (1895)
289-290; Koorpb. Exk. FI. Java 1 (1911) 268-269;
Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam.
226, p. 2; Dauss, Monogr. Lemnac. (1965)
17-19; Back. & BAKu. f. Fl. Java 3 (1968) 127.
— Telmatophace gibba (L.) SCHLEID. Linnaea 13
(1839) 391; Mig. Fl. Ind. Bat. 3 (1855) 222;
Kurz, J. Linn. Soc. Lond. 9 (1866) Bot. 266. —
Fig. 5 a-e.
Small floating aquatic. Fronds solitary or co-
hering in groups of 2-4, mostly asymmetric, some-
times almost symmetric; round, ovate or obovate,
2.1-4.9 by 1.7-3.7 mm; base almost symmetric,
obtuse to emarginate; apex mostly asymmetric,
sometimes symmetric, obtuse to round; margin
entire; dorsal side flat or convex, smooth or with
a median row of papillae; ventral side flat to
strongly inflated and globular, with a relatively
small number of large air cavities; green or red
on both sides; nerves 3—5, sometimes indistinct.
Stipe often prominent and persistent. Root
sheath cylindrical, short, without wings; root cap
obtuse or sometimes acute. Slit of budding pouch
ventral to the margin of the frond. Ovary with
1 amphitropous ovule or 2-4 anatropous ones.
Fruit symmetric, ellipsoid, with lateral wings,
dorso-ventrally compressed, with 1-4 seeds.
Seeds longitudinally ribbed.
Distr. Reported from the temperate and tropical
zones of the whole world, but doubtless many
records relate to other species.
The records from Malesia are doubtful;
among the material studied not one specimen of
L. gibba was found. The record from the Philip-
pines, listed by Dauss, no doubt refers to Mer-
RILL’s Sp. Bl. 131, which is L. perpusilla, The
Australian records of L. gibba probably must be
referred to L. disperma HEGELM.
2. Lemna trisulea Linné, Sp. Pl. (1753) 970;
Scucem. Linnaea 13 (1839) 391; Mig. FI. Ind.
Bat. 3 (1855) 222; Grises. Fl. Br. W. Ind. Is.
(1864) 512; Kurz, J. Linn. Soc. Lond. 9 (1866)
268; J. Bot. 5 (1867) 166; Heoetm. Lemnac.
252 FLORA MALESIANA [ser. I, vol7?
Fig. 5. Lemna gibba L. a. Habit, dorsal view, x 8, b. ventral view showing subepidermal air cavities, 8,
c. lateral view, 8, d. winged fruit with 2 seeds, x18, e. ditto, with 1 seed, x 18. — L. minor L. f. Habit
dorsal view, x8, g. ditto ventral view, <8, h. lateral view, x8 (a-e HELLER 5571, f-h VAN DER PLAS
29-4-69).
1971]
(1868) 134-136; BenTH. FI. Austr. 7 (1878) 162;
FRANCH. & SAv. En. PI. Jap. 2 (1879) 12: Hook.
f. Fl. Br. Ind. 6 (1893) 557: HEGELM. Bot. Jahrb.
21 (1895) 293; Massart, Bull. Soc. R. Bot. Belg.
34 (1895) 184; F. M. Battey, Queensl. Fl. 5 (1902)
1700; Pratn, Bengal Pl. 2 (1903) 841; K. Scu. &
Laut. Nachtr. FI. Schutzgeb. (1905) 62; Koorp.
Exk. Fl. Java 1 (1911) 268; MeRR. En. Philip. 1
(1923) 190; Back. Handb. FI. Java 3 (1924) 2:
STEEN. Arch. f. Hydrobiol. Suppl. 11 (1932) 291;
HAND.-Mazz. Symb. Sin. 7 (1932) 1368; GAGNepP.
Fl. Gén. I.-C. 6 (1942) 1198; McCann, J. Bomb.
Nat. Hist. Soc. 43 (1942) 154; Black, FI. S. Austr.
LEMNACEAE (van der Plas)
233
ed. 1, 2 (1948) 172; Back. Bekn. Fl. Java (em.
ed.) 10 (1949) fam. 226, p. 2; HERTER, Rev.
Sudam. Bot. 9 (1954) 185: Mitra, FI. Pl. E. India
1 (1958) 86; HUNDLEY & U Cuit Ko Ko, List
trees etc. Burma ed. 3 (1961) 292; WILLIS, Handb.
Pl. Vict. 1 (1962) 271; ZutsHr & KAuL, Trop.
Ecol. 4 (1963) 95-96: Dauss, Monogr. Lemnac.
(1965) 32-34; Ouw!, Fl. Japan (1965) 265;
Hess, LANDOLT & HirZEL, Fl. Schweiz 1 (1967)
496; CLARK & THIERET, Mich. Bot. 7, 2 (1968) 73;
Back. & Baku. f. Fl. Java 3 (1968) 127.5 —
Lenticula trisulca Scop. Fl. Carn. ed. 2 (1772) 213;
MOENCH, Meth. (1794) 319; St. Hi. Exp. fam.
Fig. 6. Lemna trisulca L. a. Habit, submerged plant, x 6, b. floating plant with inflorescence, dorsal view,
*6, c. frond with fruit, ventral view, 6, d. ditto. *6, e. part of a flowering frond, dorsal view, x 12,
f. detail of inflorescence surrounded by spathe, x 24, g. stamen, x 24 (a Den HARTOG 1162, b-¢ Hotm-
BERG 45
3).
234
FLORA MALESIANA
[ser. I, vol? 7?
nat. germ. pl. 1 (1805) 53. — Lenticula ramosa
LAMK, FI. Fr. 2 (1778) 189. — L. cruciata ROxB.
[Hort. Beng. (1814) 66, nomen] FI. Ind. ed. Carey
3 (1832) 566. — L. intermedia RUTHE, Fl. Mark
Brandenb. Niederlausitz, ed. 2 (1834) 277. —
L. bisulca C. A. Meyer, Beitr. Pfl. Russ. Reich
9 (1854) 104. — Staurogeton trisulcus (L.) SCHUR,
En. Pl. Transs. (1866) 636. — Hydrophace
trisulca (L.) BUBANI, Fl. Pyren. 4 (1897) 23;
LUNELL, Am. Midl. Natur. 4 (1915) 237. — Fig. 6.
Submerged aquatic, only floating at the surface
when flowering; when sterile many fronds con-
nected in large groups, when fertile only 2 fronds
cohering. Fronds symmetric, or almost so, oblong
to linear-lanceolate, 2.5-6.9 by 1.1-2.9 mm;
base slightly asymmetric or symmetric, narrowing
into the stipe; apex symmetric or almost so,
acute to obtuse; margin distinctly or indistinctly
denticulate towards the apex; frond thin or
slightly thickened when flowering; dorsal side
flat, smooth; ventral side flat to slightly convex;
green, without red pigment; nerves 1-3, sometimes
indistinct. Stipe strongly prolonged, flat, up to 11
by 0.2-0.3 mm, green, persistent; when flowering
@reo
less distinct or not developed at all. Root often
absent in submerged plants; root sheath very
short, most indistinct; root cap acute. Slit of
budding pouch just dorsal to the margin of the
frond. Ovary with 1 amphitropous ovulum. Fruit
broad, more or less symmetric. Seed ribbed.
Distr. Worldwide, but curiously absent from
South America, extremely scarce in Malesia:
North Sumatra (Toba Lake, not collected, cf.
STEEN. 1932), Philippines (Luzon, Mindanao),
S. Moluccas (Aru Is.: Maikoor I. leg. BECCARI,
26-6-73), and New Guinea.
For Java recorded from Bogor by MIQUEL
(1855, l.c.) and again by MAssarT (Un botaniste
en Malaisie, 1895, 184) from inundated rice-
fields near Bandung and Garut. No other botan-
ists or collectors have ever found it in Java, so
that we may safely assume that these two records
rest on errors.
Ecol. Stagnant waters up to 1800 m, outside
Malesia even recorded from 2800-3000 m.
Mostly submerged, forming a dense layer below
the surface of the water, sometimes together with
Riccia spp.
Fig. 7. Lemna tenera Kurz. a. Habit, dorsal view, x12, b. ventral view of floating frond with inflores-
cence, X12, c. ventral view, of submerged frond, x12, d. habit, dorsal view of submerged plant, x12
(a, c-d RiDLEY 10778, b MOLLER 30).
1971]
LEMNACEAE (van der Plas)
23)
3. Lemna tenera Kurz, J. As. Soc. Beng. 40, ii
(1871) 78; HEGELM. Bot. Jahrb. 21 (1895) 293-
294; Ripi. Mat. Fl. Mal. Pen. 3 (1907) 52:
Fl. Mal. Pen. 5 (1925) 132. — Lemna sp. near
L. trisulca, Ripv. J. Str. Br. R. As. Soc. n. 33
(1900) 179. — Fig. 7.
Submerged aquatic, only floating at the surface
when flowering. Fronds solitary or connected in
groups of 2, seldom 3, slightly or strongly asym-
metric, often curved; oblong to linear-lanceolate;
3-7.8 by 1-2.3 mm; frond when floating slightly
convex; base slightly to strongly asymmetric, atte-
nuate; apex symmetric, acute; margin entire:
dorsal side smooth or with an indistinct apical
papilla; ventral side flat, or convex when floating;
green without red pigment; nerves 3, sometimes
indistinct. Stipe short, hyaline, sometimes per-
sistent. Root 1, slender, very long; root sheath ab-
sent or most indistinct; root cap strongly curved,
sometimes straight, acute. Slit of budding pouch
just dorsal to the margin of the frond. Fruit
unknown.
Distr. Burma (Pegu, type), in Malesia: Malay
Peninsula (Singapore; Wellesley; Dindings), Ga-
lang (possibly Galang I. in Riouw Arch.,
RIDLEY 10374, Sept. 1899, SING).
Ecol. In ditches and ponds, in rice-lands often
together with L. perpusilla, at low altitude, in
swamp forest (KURZ), rare.
Note. Only one collection of flowering L. tenera
is known (MG6LLER 30 from Singapore); although
only a few plants have flowers, all the plants are
thickened and it is, therefore, probable that they
were floating at the surface; they have stomata at
the dorsal side. Some other collections also
consist of completely or partly thickened plants.
They are very similar to L. perpusilla in their shape
and in bearing an apical papilla, however, they
can be recognized by their more acute apex and
in the absence of a distinct root sheath. L. tenera
can easily be distinguished from the related L.
trisulca by its entire margin, the absence of a long
stipe, and by the fact that only 2-3 fronds remain
connected.
3. WOLFFIA
HoRKEL ex SCHLEID. Beitr. Bot. 1 (1844) 233, nom. gen. cons. prop.; DAUBS,
Monogr. Lemnac. (1965) 41, pro parte; HARTOG, Taxon 18 (1969) 591-592:
Hartoc & Pas, Blumea 18 (1970) 366. — Grantia Grirr. ex Voict, Hort. Suburb.
Calc. (1845) 692; Not. 3 (1851) 223. — Bruniera FRANCH. Billotia 1 (1864) 25,
30. — Wolffia sect. Uniflorae subsect. Estipitatae HEGELM. Lemnac. (1868) 122;
Bot. Jahrb. 21 (1895) 301. — Fig. 1 C, 8.
Minute aquatics, floating at the surface. Fronds either solitary or connected in
groups of 2, symmetric, thick, more or less globular, ellipsoid or ovoid, sometimes
dorsally flattened, with fleshy parenchyma without air spaces, green with or without
brown pigment cells in the epidermis; raphides none; stomata at the dorsal part of
the frond; margin entire or with a few more or less prominent papillae; nerves and
roots absent. Vascular tissue absent, except for an occasional trace in the stamen.
Budding pouch basal, median, funnel-shaped with circular opening and with a
rudimentary stipe (not visible to the naked eye). Inflorescence 1, dorsal, consisting
of a cavity containing | 3 and | ° flower, without a spathe, proterogynic. Anther
unilocular, dehiscent along an apical, pigmented line. 2 Flower situated nearer
to the budding pouch than the ¢ flower, consisting of a globular ovary with a
short persistent style. Ovule 1, orthotropous. Fruit globular. Seed globose or
slightly compressed, smooth.
Distr. At least 7 spp. in the tropical, subtropical and temperate regions of the world.
KEY TO THE SPECIES
1. Dorsal outline of the frond round or elliptic. Ventral side inflated, globular.
2. Dorsal outline of the frond elliptic. Dorsal side flat to convex. Width greatest below the dorsal! plane,
TPO, |... oes e whe eheue © kee meets 1. W. globosa
2. Dorsal outline of the frond round to ovate. Dorsal side flat. Width greatest in the dorsal plane.
W. arrhiza
1. Dorsal outline of the frond angular. Ventral side with a central conical protuberance.
W. microscopica
erp. (8)\mas) 6
236
1. Wolffia globosa (Roxs.) HArToG & PLAS,
Blumea 18 (1970) 367. — Lemna globosa Roxs.
[Hort. Beng. (1814) 66, nomen] FI. Ind. ed.
Carey 3 (1832) 565; GRAHAM, Cat. Bomb. PI.
(1839) 252; Daz. & Giss. Bomb. FI. (1861) 281.
— Grantia globosa (ROxB.) GRIFF. ex VOIGT,
Hort. Suburb. Calc. (1845) 692; Not. 3 (1851)
229. — W. schleideni Mig. Ned. Kruidk. Arch.
3 (1855) 428; Fl. Ind. Bat. 3 (1855) 221; Nat.
Tijd. N. I. 10 (1856) 399-402; Kurz, ibid. 27
(1864) 220; Naves, Novis. App. (1882) 296;
Back. Handb. FI. Java 3 (1924) 2; ibid. 2 (1928)
290; Bekn. Fl. Java (em. ed.) 10 (1949) fam.
226, p. 3; BACK. & BAKH. f. Fl. Java 3 (1968) 128.
— W. delilii var. schleideni (M1Q.) KURZ, J. Linn.
Soc. Bot. 9 (1866) 265. — W. arrhiza (non Win-
MER) MARTENS, Preuss. Exp. O.-Asien, Bot.
Teil, Tange (1866) 143; HEGELM. Lemnac. (1868)
124-126, p.p.; Hoox. f. Fl. Br. Ind. 6 (1893)
557-558, p.p.; HEGELM. Bot. Jahrb. 21 (1895)
301-302, p.p.; Trim. Handb. FI. Ceyl. 4 (1898)
367; RipL. J. Str. Br. R. As. Soc. n. 33 (1900)
79; CoLLeTT, FI. Simlensis (1902) 545; Prain,
Bengal Pl. 2 (1903) 841; RipL. Mat. FI. Mal. Pen.
3 (1907) 53; Koorpb. Exk. Fl. Java 1 (1911) 269;
Me_err. Philip. J. Sc. 12 (1917) Bot. 101; En.
Philip. 1 (1923) 190; BAck. Handb. Fl. Java 3
(1924) 2; Riv. Fl. Mal. Pen. 5 (1925) 132;
HEND. Gard. Bull. S. S. 4 (1928) 352; FiscHER in
Gamble, Fl. Madras 9 (1931) 1593; Burk. Dict.
(1935) 1328; GAGNeEP. FI. Gén. I.-C. 6 (1942)
1198; McCann, J. Bomb. Nat. Hist. Soc. 43
(1942) 159-160; Back. Bekn. Fl. Java (em. ed.)
10 (1949) fam. 226, p. 3; HEND. Mal. Wild FI.
Monocot. (1954) 210; MASsAMUNE, Sc. Rep.
Kanazawa Univ. 5 (1957) 94; Mitra, FI. Pl. E.
India 1 (1958) 86; LarsEN, Dansk Bot. Ark. 20
(1962) 136; Onw1, FI. Japan (1965) 265; SHRI &
Naik, Fl. Osmanabad (1968) 682; BAcK. &
BAKH. f. Fl. Java 3 (1968) 128. — W. cylindracea
HEGELM. Lemnac. (1868) 123; Bot. Jahrb. 21
(1895) 302; Brown, FI. Trop. Afr. 8, 2 (1901)
205. — W. delilii (non SCHLEID.) NAVES, Novis.
App. (1882) 296. — W. microscopica (non KURZ)
MATSAMURA & HAYATA, En. Pl. Form. (1906)
463; Miki, Bot. Mag. Tokyo 48 (1934) 334.
— Lemna arrhiza (non L.) Back. Bull. Jard. Bot.
Btzg II, 12 (1913) 21. — Fig. 8 a-a?.
Fronds with elliptic to oblong dorsal outline,
c. 144 times as long as wide, 0.30-0.73 by 0.20-0.33
mm, 0.23-0.55 mm high; base and apex obtuse;
margin with a few papillose cells; dorsal side flat
near the apex and convex near the base, with
a few papillose cells; ventral side strongly inflated,
globular, with large cells; width greatest below the
dorsal plane, at about half the height; green,
without brown pigment cells in the epidermis.
Budding pouch often with a distinct collar of elon-
gate cells. Fruit unknown.
Distr. E. Asia (Japan; China: Fukien; Ryukyu
Is.), SE. Asia (Tonkin; India: Sibpur, Dehra Dun,
Peshawar, etc.; Ceylon), also in Africa and
Australia; in Malesia: Malay Peninsula (Singa-
pore; Selangor; Kuala Lumpur; Malacca),
E. Java (near Surabaja and near Redjasa E of
FLORA MALESIANA
(ser. I, volad=
Pasuruan), Kangean Is. (Sepandjang), Lesser
Sunda Is. (Flores: Rana Mesé), Philippines
(Central Luzon: Bulacan, Rizal).
Pampanga,
bi
a
a a2
Fig. 8. Wolffia globosa (RoxB.) HARTOG & PLAS.
a. Habit with a daughter frond, lateral, x18,
a. ditto, X18, a®. dorsal view, x18. — W.
arrhiza var. australiana BENTH. b. Habit, lateral
view, X18, b1. ditto, x18, b®. ditto, dorsal view,
x18. — W. arrhiza (L.) HORKEL ex WIMMER.
c. Habit with a daughter frond, x18, c1. ditto,
dorsal view, x18. — W. microscopica (GRIFF.)
Kurz, d. Habit of a flowering plant with a
daughter frond, showing elongated ventral side,
and dorsal inflorescence, x18, d'. ditto, frond
without protuberance, 18, d*. dorsal view,
x18, d%. ditto, more angular outline, 18,
d‘. detail of the inflorescence (1 pistil, 1 stamen),
X36 (a-a? JAAG 1956, b-b? DEN Hartoc 1186,
c—c! VAN DER PLAS 28-9-67, d-d* DESHPANDE 1727)
— aS? lee ee
1971]
BACKER & BAKHUIZEN f. note this also from
Mts Jang & Idjen in E. Java at 2200m, but I
believe this to be an error.
Ecol. Stagnant water and ditches, once in cal-
ciferous water, in Malaya in everwet climate, in
E. Java under severe monsoon conditions, all
localities in Malesia at low altitude. Near Redjasa
under a thick cover of Azolla pinnata; at 1000m
in Rana Mesé.
Vern. Philippines: dugmdn, Bis., lid, Tag.
Notes. The specimens of W. globosa of Ma-
lesia are c. 1144 times as long as wide, while the
Australian specimens are usually twice as long as
wide. The size of both the Malesian and Australian
plants of W. globosa is smaller than that of W.
arrhiza. In Australia still another Wolffia is found
with an elliptic dorsal outline and which is
much higher than all other species in the genus.
It is known as W. arrhiza var. australiana
BENTH. but no doubt represents a good species
(fig. 8 b-b?).
Wolffia arrhiza (L.) HorRKEL ex Wimmer, FI.
Schles. ed. 3 (1857) 140; HEGELM. Lemnac.
(1868) 124-126, p.p.; Hoox. f. Fl. Br. Ind. 6
(1893) 557-558, p.p.; HEGELM. Bot. Jahrb. 21
(1895) 301-302, p.p.; DAuBs, Monogr. Lemnac.
(1965) 48-49. — Lemna arrhiza LINNE, Mant.
(1771) 294. — W. michelii SCHLEID. Beitr. Bot.
(1844) 233. — Fig. 8 c-c!.
Fronds with round to ovate dorsal outline,
usually as long as wide, rarely somewhat longer,
0.8-1.3 by 0.6—0.8 mm; 0.6—-0.8 mm high; base and
apex obtuse; margin entire or with papillae;
dorsal side flat to slightly convex; ventral side
strongly inflated, globular, consisting of large cells;
width greatest in the dorsal plane; green, without
brown pigment cells in the epidermis.
Distr. Temperate zone of the Old World, in
Africa also in the tropics, not yet found in Ma-
lesia and probably absent from the whole of
East Asia and Australia. Records of W. arrhiza
from these areas seem to relate to the closely
allied W. globosa.
LEMNACEAE (van der Plas)
237
Ecol. Stagnant waters, often mixed with other
floating aquatics.
Wolffia microscopica (GRIFF.) KuRz, J. Linn.
Soc. Bot. 9 (1866) 265; HEGELM. Lemnac. (1868)
127-128; Hoox. f. Fl. Br. Ind. 6 (1893) 558;
HEGELM. Bot. Jahrb. 21 (1895) 301; Pram,
Bengal PI. 2 (1903) 841; Mitra, FI. Pl. E. India 1
(1958) 86; DAuBs, Monogr. Lemnac. (1965) 44. —
Grantia microscopica GRIFF. ex WVoiGcT, Hort.
Suburb. Calc. (1845) 692; Not. 3 (1851) 226. —
Fig. 8 d-d*.
Fronds with pentagonal or hexagonal dorsal
outline, 0.4—0.6 mm o; margin entire except for the
papillose corner cells; dorsal side flat; ventral side
elongated into a central, conical, parenchymatic
protuberance up to 1.8 mm long; green, without
brown pigment cells in the epidermis.
Distr. So far this species is only known from
India. It is probable that this minute plant has
been often overlooked; therefore, it is expected to
occur also elsewhere in East Asia and even in
Malesia.
Ecol. Occurs usually mixed with other floating
aquatics, especially with Lemna perpusilla. Ac-
cording to MAHESWARI and KapiL (Am. J. Bot.
50, 1963, 679) it never co-exists with Spirodela
polyrhiza.
Note. Records of W. microscopica from For-
mosa and Japan relate to W. globosa.
Doubtful
Mique. (Ned. Kruidk. Arch. 3, 1855, 429; FI.
Ind. Bat. 3, 1855, 221) reported that a second con-
siderably larger Wolffia species was found to-
gether with W. schleideni near Surabaia. He
thought that this species was identical or related
to W. delilii SCHLEID., but unfortunately he restrict-
ed the description to a few remarks, which are
repeated here:
‘Frond complanate, slightly lenticular, 2-244
mm long; epidermis cells gyrose.’
I have not seen this material.
Dubious
Lemna obcordata WINBERG in Thunberg, FI. Jav. 1 (1825) 2, nomen.
HALORAGACEAE (R. van der Meijden & N. Caspers, Leyden)
Perennial (rarely annual) herbs, or undershrubs, terrestrial or aquatic, sometimes
stoloniferous (Gunnera). Leaves opposite, spiral, or verticillate, in the terrestrial
species nearly always simple, in the aquatic ones always partly pinnately divided,
pinnately nerved or (in Gunnera) palmately nerved. Stipules 0, but the leaves often
flanked by small, subulate and caducous enations. Flowers mostly in spike-like
inflorescences, sometimes in a compound panicle, mostly solitary or (sometimes )
in clusters of up to a dozen flowers in the axil of a bract or reduced leaf, °, mono-
ecious, dioecious or polygamous, perigynous, actinomorphous, mostly 4-merous,
or 2-, or (not in Mal.) 3-merous. Sepals 4 or 2, rarely (not in Mal.) 3, in ¥ flowers
sometimes much reduced to 0, free or little connate, mostly persistent. Petals
alternisepalous, 4, 2 or 0, rarely 3 (not in Mal.), free, in 2 flowers absent or strongly
reduced, often soon caducous, mostly more or less unguiculate and cochleari-
form, longer than the sepals. Stamens as many as sepals and then epi- or alterni-
sepalous, or twice as many, 8, 4 or 2, rarely (not in Mal.) 3 fertile and 3 sterile,
or 1, in 2 flowers completely reduced; filaments mostly filiform, long and very
thin, rarely (not in Mal.) short and thick; anthers 2-celled, basifixed, latrorse,
mostly oblong to linear, ay + elliptic. Disk 0. Ovary 1- or 4-, rarely 2- or
(not in Mal.) 3-celled, in the 3 flowers 0 or reduced; style alternisepalous, free,
mostly short, grading into the preter or subulate stigmas which spread in fruit,
the stigmatic, more-celled papillae hair-like elongating towards the end of the
anthesis (except in Gunnera). Ovules as many as styles, or (in Gunnera) single,
apical, pendulous, anatropous and apotropous. Fruit nut-like or (in Gunnera) a
drupe, variously sculptured, indehiscent l-seeded or breaking up into 4(-2) I-
seeded mericarps. Seed witha thin testa; embryo cylindrical, surrounded by a thick,
white, oily albumen, or (in Gunnera) obcordate and in top of a very copious and
oily albumen.
Distribution. Genera 7, with c. 150 spp., nearly all over the world, but rather rare in the tropics.
Ecology. Malesian Haloragaceae have a varied ecology; in general they prefer damp, often poor
soils. Myriophyllum consists mostly of aquatics. Myriophyllum and Haloragis species grow in the lowland
ascending into the hills; other representatives are decidedly restricted to the montane zone and the
lower part of the alpine zone, e.g. Gunnera macrophylla, Haloragis halconensis, H. micrantha, H. philip-
pinensis, Laurembergia coccinea, and Myriophyllum pedunculatum. There is even one restricted to the
alpine zone, viz. Myriophyllum pygmaeum. A few species are bound to a seasonal climate, viz Myrio-
phyllum dicoccum, M. coronatum, and Haloragis acanthocarpa. Gunnera shows a remarkable symbiosis
with algae.
Flower biology of Haloragis, Laurembergia and Myriophyllum. It seems that at least the Malesian
species are protandrous; in the initial stage of anthesis the stigmatic papillae are very short; at this stage
the anthers grow to proportionally large size, protrude from the flower on elongating filaments, and shed
their pollen; they soon wither and are caducous. In the meantime the ovary has already distinctly en-
larged. Then the stigmatic papillae elongate and obviously reach the receptive stage. The structure seems
to point to cross-pollination by wind, as already pointed out by Miss Grers (1917). Consequently, sizes
of flower parts can only be compared when full-grown. The petals also reach their full size only in just
opened flowers.
Taxonomy. Haloragaceae are generally agreed to be classified with the Myrtales. The circumscription
of the family has differed and has included the genera Callitriche and Hippuris. These are now generally
regarded as two separate ae though HUTCHINSON still includes Hippuris (Fam. Fl. Pl. ed. 2, 2,
1959, 448; Fvol. Phyl. Fl. Pl. 1, 1969, 546). They were sometimes placed near Haloragaceae, but for
_example PULLE Pep ale ed "regarded both as reduced sympetalous families and assumed the re-
ductions to be convergent coinciding with the aquatic habitat, not expressing systematic affinity. This
view is sustained by HeGNaver (Chemotax. 4, 1966, 238; Pharm. Act. Helv. 41, 1966, 585) and WIEFFERING
(239)
240 FLORA MALESIANA [ser. I, volar
(Phytochem. 5, 1966, 1064) on chemotaxonomical arguments. MELCHIOR (Engl. Syll. ed. 12, 2,1964,
438) and TAKHTAJAN (Evol. Angiosp. 1959, 228; Fl. Pl. 1969, 225) place only Callitrichaceae near or in
Lamiales. HUTCHINSON /.c. places Callitriche in Onagrales.
Within the family Gunnera occupies an isolated position, in which it is mostly regarded as a separate
subfamily (SCHINDLER, MELCHIOR); also HUTCHINSON includes it in the family. Others regard it to re-
present a separate family, placing it next to Haloragaceae, e.g. VON WETTSTEIN, PULLE, and TAKHTAJAN.
The differences between the two groups are the following:
Haloragoideae: Vessels not polystelic. Ligule-like structures absent. Hairs many-celled, rarely reduced
few-celled. Bracteoles present, rarely fully absent. Stigma short, capitate or shortly ligulate, on a more or
less distinct style. Ovules 2-4, as many as the styles, crassinucellate, with 2 integuments. Fruits without a
stone. Endosperm nuclear. Embryo large, cylindrical, with a long radicle.
Subfam. Gunneroideae: Vessels polystelic. Ligule-like structure distinct, sometimes ochrea-like.
Hairs one-celled. Bracteoles absent. Stigma long, subulate, sessile. Ovule 1, tenuinucellate, with 1 in-
tegument. Fruits with a stone. Endosperm cellular. Embryo small, obcordate, with a short radicle.
However, in general the floral structure is in good agreement to include both in one family. The
Australian genus Glischrocaryon (Loudonia) combines characters of both Gunnera and Haloragis, being
macromorphologically distinctly allied to Haloragis, but possessing pollen which resembles that of
Gunnera (similar type of apertures and exine), thus giving an additional strong argument for keeping
Gunneroideae as a subfamily of Haloragaceae.
Subfam. Haloragoideae is divided into tribe Halorageae (with a 1-seeded fruit) and tribe Myrio-
phylleae (with the fruit breaking up into 4(—2) 1-seeded mericarps).
Chemotaxonomy. As a consequence of the small economic importance of this family chemical
information is scanty. Flavonols, leucoanthocyanins, ellagic acid and rather large amounts of tannins
have definitely been demonstrated to be present in members of Haloragaceae. The tannins are most
probably mixtures of hydrolysable and condensed tannins; this is indicated by the presence of ellagic acid
and leucoanthocyanins. The terms “myriophyllin cells’ and “‘myriophyllin” often used in anatomical
literature are derived from Myriophyllum. Myriophyllin cells are idioblasts which are coloured purple by
vanillin and hydrochloric acid and myriophyllin is their tannin-like content which gives this reaction
(indicating leucoanthocyanins or catechins). Species of Mpyriovhyllum bear trichomes giving the “my-
riophyllin”’ test. Two species of Haloragis have been found to produce prussic acid; the cyanogenic com-
pounds, however, have not been investigated. Oxalate of lime occurs frequently, especially in the form of
small clusters.
Accumulation of tannins containing ellagic acid and of condensed leucoanthocyanins fits well with
the inclusion of Haloragaceae in Myrtales in the wider sense. The genera Callitriche and Hippuris have
by some authors been included in or thought affiliated with Haloragaceae, but they are biochemically so
different that such a relationship seems highly improbable; their chemical characters rather indicate affi-
nities with sympetalous taxa. Reference: HEGNAUER, Chemotaxonomie der Pflanzen 4 (1966) 235-238.
— R. HEGNAUER.
Morphology. Though the enations or trichomes occurring near the leaf-base have been sometimes
interpreted as stipules, they are according to SCHINDLER (Pfl. R. Heft 23, 1905, 4) enatia te which he
ascribes a gland function. However this may be, they are also found on the leaves and in other not
prescribed places on the stem, showing that they are not organs in the proper sense. Recently PRAG-
Lowsky (Grana 10, 1970, 159-239) revised the palynology of the family.
Uses. The only useful plant in Malesia is Myriophyllum brasilse ienwhich is cultivated in fish-ponds.
Note. The treatment of Haloragis was done by Mr. N. CAspPERS several years ago; while working on
Gunnera and Laurembergia, he fell ill and could not finish this work.
KEY TO THE GENERA
1. Leaves radical, long-petioled, palmately nerved. . . . . - - - - e+ ee eee eee 4. Gunnera
1. Leaves not radical, sessile or shortly petioled, pinnately nerved.
2. At least the lower (submersed) leaves pinnately divided, verticillate.. . .... 3. Myriophyllum
2. All leaves entire to crenate-serrate, opposite, alternate or sometimes in whorls of 3-4.
3. Fruit breaking up into 2-4 mericarps. Leaves sessile. .........-.-.- 3. Mpyriophyllum
3. Fruit a one-seeded nutlet. Leaves mostly shortly petioled.
4. Flowers in axillary clusters, the middle one ¢ or sometimes § and (in Asiatic spp.) very long-
Sta led the ther caeast Gk peaks bin Wea icat is’ aa) “ies meth eae wl leony See AOS 2. Laurembergia
4. Flowers solitary, all $, not long-stalked.. . . «© - 6 6 ee eee ee ees 1. Haloragis
1. HALORAGIS
J. R. & G. Forst. Char. Gen. Pl. (1776) 31, t. 31; ScHINDL. Pfl. R. Heft 23 (1905)
1-133, 36 fig. (‘Halorrhagis’); WENT f. Nova Guinea 14 (1924) 105-109, t. XI. —
Gonocarpus THUNB. Nov. Gen. PI. 3 (1783) 55; Fl. Jap. (1784) 69, t. 15; SCHREB.
1971] HALORAGACEAE (van der Meijden & Caspers) 241
Gen. Pl. 1 (1789) 86 (‘Gonatocarpus’); KON. & Sims, Ann. Bot. 1 (1805) 546
(‘Goniocarpus’). — Gaura Lour. Fl. Coch. (1790) 225. — Fig. 1.
Herbs or small shrubs, prostrate or erect; stems mostly angular by the decurrent
edges of the petiole. Leaves (in Mal.) simple, decussate or in whorls of 3-4, rarely
alternate, pinnately nerved, the margin almost always dentate, serrate, or crenate.
Inflorescences terminal. Flowers (in Mal.) solitary in the axil of a bract, bisexual,
(in Mal.) 4-merous. Sepals (in Mal.) 4, valvate in bud, mostly triangular, persistent.
Petals (in Mal.) 4, imbricate in bud, boat-shaped, caducous. Stamens (in Mal.)
8, rarely 4, in the latter case episepalous (in one extra-Mal. sp. epipetalous).
Ovary (in Mal.) often 4-gonous, 8-ribbed, the mid-sepaline ribs less distinctly
raised than the others, 4-celled; pericarp enlarging to fruit-size long before the
seed is set; styles (in Mal.) 4, cylindric, often incurved or the stigmas sessile.
Fruit nut-like, pericarp hard.
Distr. About 60 spp., almost all in Australia (51), also in Chatham I., Carolines (Yap, Palau), a
few endemic in Tasmania and New Zealand, in the Pacific eastwards in Rapa I. and Juan Fernandez,
in continental SE. & E. Asia in the Himalayas, Japan, and Korea; 3 extra-Australian species in Asia
and Malesia, and 1 in New Caledonia; in Malesia 5 spp., some in the lowland, others on the
mountains.
According to SKOTTSBERG (Nat. Hist. Juan Fern. 2, 1921, 151) the opinion that H. erecta would have
been introduced in Chile rests upon a mislocalized specimen. SKOTTSBERG described three new species
from the Juan Fernandez group all of the affinity of H. erecta.
Ecol. Three species are montane to subalpine and prefer generally damp or marshy habitats (H. mi-
crantha, H. halconensis, H. philippinensis). H. chinensis is usually found on poor soils from sea-level up to
2200 m; H. acanthocarpa is bound to a lowland seasonal steppe and savannah climate on badly drained
soils.
Taxon. SCHINDLER divided the genus into two subgenera by the number of stamens, viz 8 (or 6 in 3-
merous flowers) in subg. Haloragis, or 4 in one W. Australian species forming a monotypic subg. Pseudo-
haloragis in which the 4 episepalous stamens are aborted.
However, in H. philippinensis, and some other extra-Malesian species, there are also only 4 stamens,
but here by abortion of the epipetalous stamens. Furthermore, PRAGLOWSKI (Svensk Bot. Tidskr. 63, 1969,
486) observed in two Australian species sterility of anthers, 4 out of 8 stamens or 2 out of 4 stamens,
hence, a sort of intermediate stage towards full abortion.
Consequently, it tends to be doubtful to distinguish subgenera on the basis of the number of stamens.
KEY TO THE SPECIES
1. Small, caespitose herb, glabrous in all parts. Stems rooting at the nodes, at least at the base, apex
often ascending. Sepals with a thickened and shallowly cordate base. Anthers 0.3-0.7 mm long.
3. H. micrantha
1. Plant hairy, at least in the younger parts. Stems not rooting at the nodes, generally erect. Base of the
sepals not thickened and subcordate. Anthers 0.8—2.3 mm long.
2. Fruit with 8 longitudinal ridges alternating with 8 rows of (3-4) blunt-spiny tubercles, glabrous
or appressedly hairy only at the apex. . «© 6 6 eee ee ee ee es 4. H. acanthocarpa
2. Fruit without rows of intercostal tubercles, or occasionally with 1(—3) blunt, low tubercles between
the ribs, and then the fruit covered with appressed, curved hairs.
3. Robust plants with semi-woody, stiff stems, mostly very much branched. Leaves decussate or,
especially in the middle of a branch, in whorls of 3-4; if decussate, then the index of the leaves of the
OE SS SP en eee Mare et alte 5. H. halconensis
3. Perennial, slender herbs, the stem mostly branched only at the base, rather thin. Leaves always
decussate; Jeaf-index (2—-)2.8-4(—5.5).
4. Anthers 8. Ripe fruit erecto-patent to patent, mostly not thickened below the sepals, 0.7-1 by
0.8-1 mm (excluding the sepals), with a granular surface, the ribs often =: tuberculate, sometimes
with 1(—3) intercostal tubercle(s). Inflorescence mostly much branched. . . . . 2. H. chinensis
4. Anthers 4. Ripe fruit nodding, with a distinct, + triangular thickening at the base of the sepals,
1.3-1.6 by 11.3 mm (excluding the sepals), the ribs smooth, the surface in between smooth or
granular. Inflorescence little or not branched. . «6 6 6 we ee es 1. H. philippinensis
242
FLORA MALESIANA
[ser.-I, volvy=
Fig. 1. Haloragis micrantha (THUNB.) S. & Z. a-b. Habit, a in exposed, b in shaded situation, nat. size,
c. leaf, 7, d. flower, 15, e. fruit, x15 (W. Java, Papandajan, VAN STEENIS 4246, 4296).
1. Haloragis philippinensis Merr. Philip. J. Sc. 1
(1906) Suppl. 216; En. Philip. 3 (1923) 221. —
H. tetragyna (non Hook. f.) CLARKE, FI. Br. Ind. 2
(1878) 430, pro specim. khasyan. — H. scabra
var. abbreviata SCHINDL. Pfl. R. Heft 23 (1905) 29;
? GUILLAUMIN, FI. Gén. I.-C. 2 (1920) 715, var.
‘attenuata’ (sic). — H. isomera PARKER in Fedde,
Rep. 29 (1931) 104; HUNDLEY & U Cuit Ko Ko,
List Trees Burma (1961) 98; TARDIEU-BLOT,
Fl. Laos, Camb. & Vietn. 4 (1965) 119, t. 1
f. 11-13.
Perennial herb, 9-30 cm, not or little branched,
with a number of stems from the base, each ap-
pressed-hairy and with ascending apex. Leaves
decussate, ovate- to obovate-oblong, or lanceolate,
4-25 by 214-7mm, often appressed-hairy on
both sides or the upper surface glabrous, margin on
both sides with 2-10 small teeth; petiole 0-2 mm.
Racemes not or hardly branched, 214-12 cm.
Bracts ovate-elliptic to lanceolate, the lower ones
almost as large as the upper leaves, gradually
diminishing in size upwards, 12-114 mm, with 1-6
teeth on either side. Bracteoles of various shape,
14-1 mm, widened at the base. Pedicel 0-34 mm,
1971]
HALORAGACEAE (van der Meijden & Caspers)
glabrous or hairy. Sepals 34-1 by 1 mm, at the
base often somewhat thickened, glabrous. Petals
14%4,-2mm, the midrib dorsally mostly hairy.
Stamens 4, episepalous; filaments c. 0.2—0.3 mm;
anthers c. 1-1'4mm. Styles 4, 0.2-0.5 mm.
Fruit nodding, ovoid to broad-ellipsoid, rarely
globular, c. 1144-114 by 1-1'4 mm, with a distinct,
+ triangular thickening at the base of the sepals,
glabrous or with straight, appressed hairs, surface
smooth or granular.
Distr. Assam (Khasya Mts), Burma (Tenasse-
rim, Mergui), Thailand (Penins.: Ranong), S.
Vietnam; in Malesia: N. Sumatra (Gajo Lands;
Tapanuli: Sidikalang; Toba-Batak area) and
Philippines (N. Luzon: Benguet Prov.). Fig. 2.
Fig. 2. Distribution of Haloragis philippinensis
Merr. (except the Thailand locality)
Ecol. Mountain heathland and blangs, plateaus
and ridges, sedge turf, Sphagnum bogs, and open
pine forest, 1200-3400m. Fi. fr. Dec.—May.
Notes. In addition to the morphological
characters H. philippinensis has another ecology
than H. chinensis; it is a true mountain species,
although in the highest localities of H. chinensis
the two species are found almost together (Toba-
Batak Lands, Benguet).
In his original description MERRILL mentioned
three syntypes of which I have examined MERRILL
4357 and 4444 (both in US, the latter also in L).
Only the first fits the description and I have chosen
this as lectotype; the second is wrongly identified
and belongs to H. chinensis.
Until 1931 the Khasya specimens have always
been confused with H. scabra by British botanists,
and have been cited under H. fetragyna var.
micrantha by those whose contention it was to
merge these two species.
2. Haloragis chinensis (Lour.) Merr. Trans. Am.
Phil. Soc. n.s. 24, 2 (1935) 290, 39; Tuvama, J.
Jap. Bot. 16 (1940) 284, f. Sb, 6, incl. var. yapensis ex
descr.; Merr. Sunyatsenia 5 (1940) 150; Merr. &
Perry, J. Arn. Arb. 23 (1942) 407; ibid. 29 (1948)
161; Sincram, Gard. Bull. Sing. 15 (1956)
24; Tarpieu-BLort, FI. Laos, Camb. & Vietn. 4
243
(1965) 120; KENG, Mal. Nat. J. 23 (1970) 123,
t. 13. — Gaura chinensis Lour. Fl. Coch. (1790)
225; ed. Willd. 1 (1793) 276. — Goniocarpus
scaber KON. & Sims, Ann. Bot. 1 (1805) 547, t. 12
f. 6. — H. scabra Bru. Fl. Hongk. (1861) 139,
excl. specim. khasyan.; SCHINDL. Pfl. R. Heft 23
(1905) 28, f. 9A, incl. var. elongata; MERR.
Philip. J. Sc. 1 (1906) Suppl. 216; BRITTEN, J. Bot.
45 (1907) 135; VALET. Bull. Dép. Agr. Ind. Néerl.
n. 10 (1907) 41, incl. var. novaguineensis VALET.;
Me_rr. J. Str. Br. R. As. Soc. nm. 76 (1917) 102;
GUILLAUMIN, FI]. Gén. I.-C. 2 (1920) 715, f.
74-14; Merr. En. Born. (1921) 455; En. Philip. 3
(1923) 221; Went f. Nova Guinea 14 (1924) 108;
KANEH. En. Micr. (1935) 383; H. J. LAM, Blumea 5
(1945) 581. — AH. tetragyna [non (LABILL.)
Hook. f.] Bru. FI. Austr. 2 (1864) 484, incl.
var. micrantha, quoad specim. asiat. et mal.,
khasyan. except.; CLARKE, Fl. Br. Ind. 2 (1878)
430; Fors. & HEMsL. J. Linn. Soc. Bot. 23 (1887)
292.
Erect or suberect, obviously perennial, mostly
branched herb, 10-60 cm; stems appressed-hairy,
mostly distinctly 4-angular with straw-coloured
edges. Leaves decussate, ovate to narrow lan-
ceolate, 2-26 by 1-9 mm, hairy, especially the
underside of the midrib, margin on either side
with 3-10 teeth; petiole 0-14, mm. Racemes
mostly more or less strongly branched, 2-22 by
1-7 cm. Bracts ovate to elliptic, 44-114 by 4-Y%
mm, pubescent, sometimes with 1 tooth on either
side. Bracteoles 0.2-0.7 mm long, widened at
base. Pedicel 0.2-0.6 mm, glabrous. Flowers
erecto-patent to patent. Sepals 0.5-0.9 by 0.4-0.6
mm, glabrous, often incurved. Petals 1-134 mm,
hairy on the midrib outside, mostly red to brick
red, sometimes green. Stamens 8; filaments 0.1—0.4
mm; anthers c. 1 mm long. Styles incurved, 0.1—0.3
mm. Fruit erecto-patent to patent, globular to
broadly ellipsoid, 0.7-1 by 0.8-1 mm, not thick-
ened below the sepals, the ribs often + tuberculate,
sometimes with 1(—3) low intercostal tubercles,
glabrous or with incurved and appressed hairs,
surface granular and punctate.
Distr. Indo-China (Cochinchina, Tonkin, sec.
GUILLAUMIN and TARDIEU-BLoT), S. China (also
Kwangtung, Macao, Hongkong, Hainan), through
Fig. 3. Distribution of Haloragis chinensis (Lour.)
MErR.
244 FLORA MALESIANA
[ser. I, vol. 74
Malesia to Micronesia (Carolines: Jap) and
Queensland (Kelsey Creek near Proserpine;
Atherton Mts, introduced in Hawaii (DEGENER
30591); in Malesia: N. Sumatra (Toba-Batak
area), Singapore, N. Borneo (Jesselton, Labuan),
Celebes, Philippines (Luzon, Calamian Is.,
Busuanga, Mindanao), Moluccas (Ceram, Aru Is.),
Lesser Sunda Islands (Sumba, Flores), New
Guinea (also Ferguson I.). Fig. 3.
Ecol. In open spots, between grass, in lalang
wastes, on poor laterites, river banks, poor sandy
dry places, and deforested slopes, from sea-level
up to 2200 m. Fi. fr. Jan.—Dec.
Note. Allied to H. philippinensis but easily
distinguishable. More closely allied to H. tetragyna
Hook. f. and by BENTHAM (1864) even united with
it as var. micrantha BTh. l.c.
3. Haloragis micrantha (THUNB.) R. Br. [in
Flinders, Voy. (1814) App. 550] ex Sires. & Zucc.
Abh. Bay. Ak. Wiss. M.-Ph. Cl. 4? (1844) 133;
repr. Fl. Jap. Fam. Nat. (1845) 25; Hook. f.
Fl. Tasm. 1 (1860) 121; Bru. Fl. Austr. 2 (1862)
482; Hook. f. Handb. Fl. New Zeal. (1867) 66;
Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 100;
CLARKE, FI. Br. Ind. 2 (1878) 430; STAPF, Trans.
Linn. Soc. Bot. II, 4 (1894) 149; Kina, J. As. Soc.
Beng. 66, ii (1897) 310; Batley, Queens]. Fl. 2
(1900) 555; SCHINDL. Pfl. R. Heft 23 (1905) 29;
Gisss, J. Linn. Soc. Bot. 42 (1914) 74; Arfak
(1917) 159; GuILLAUMIN, FI. Gén. I.-C. 2 (1920)
715; Merr. En. Born. (1921) 455; Ripv. Fl. Mal.
Pen. 1 (1922) 692; Merr. En. Philip. 3 (1923) 221;
WENT f. Nova Guinea 14 (1924) 107; STEEN.
Trop. Natuur 21 (1932) 101-102, f. 1-2; ELMer,
Leafil. Philip. Bot. 9 (1933) 3143; STEEN. Bull.
Jard. Bot. Btzg. III, 13 (1934) 218; Back. Bekn.
Fl. Java (em. ed.) 4A (1942) fam. 76, p. 2; STEEN.
Endeavour 21 (1962) 187, f. 3 (map); LARSEN,
Dansk Bot. Ark. 23 (1963) 68; BAcK. & BAKH.
f. Fl. Java 1 (1963) 266; TARDIEU-BLOT, FI. Laos,
Camb. & Vietn. 4 (1965) 122; Caspers, Pac. PI.
Areas 2 (1966) 60, map 31; LARSEN, Dansk Bot.
Ark. 23 (1966) 395, f. 60; KENG, Mal. Nat. J.
23 (1970) 123; MEIDEN, FI. Thail. 2 (1970) 3.
— Gonocarpus micranthus THUNB. Nov. Gen. Pl.
3 (1783) 55; Fl. Jap. (1784) 69, t. 15; WILLp.
Sp. Pl. 1 (1797) 690; KOn. & Sims, Ann. Bot.
1 (1805) 546, t. 12 f. 5; DC. Prod. 3 (1828) 66.
— H. tenella BRONGN. in Duperrey, Voy. Coquille
Bot. (1828) t. 68 f. B. — Goniocarpus rubricaulis
GriFF. Not. Pl. As. 4 (1854) 688, ex descr. —
Hi. paucidentata Hosokawa, Trans. Nat. Hist.
Soc. Form. 30 (1940) 335. — Fig. 1.
Prostrate, glabrous herb, 5-30 cm long; stems
rooting on the nodes, ascending at the apex.
Leaves decussate, obovate, orbicular, ovate, or
oblong, 3-10 by 114-4 mm, the margin on either
side with 1—7 teeth; petiole 0-2 mm. Racemes erect,
simple or little branched, 1-8 cm long. Bracts
lanceolate, acute, 14-114 by 0.2 mm. Bracteoles
filiform or acuminate, 0.1-0.3 mm, with widened
base. Pedicels 14-34 mm. Flowers scattered,
erect then nodding. Sepals ovate-triangular,
0.3-0.6 mm, subsaccate-thickened and shallowly
micrantha
Fig. 4. Distribution of Haloragis
(THuNB.) S. & Z.
cordate at the base, often acuminate. Petals c.
0.9-1.3 mm, red or rosa. Stamens 8; filaments up
to 0.6 mm; anthers 0.3-0.7 mm. Styles incurved,
0.2-0.3 mm. Fruit obovoid-globose to depressed-
globose, c. 0.9-1.1 by 0.7-0.9 mm, smooth.
Distr. S. & E. Asia (E. India, N. Thailand, S.
& E. China, N. Vietnam, Hainan, Taiwan,
Ryukyu Is., Korea, Japan), Australia (New
South Wales, Victoria), Tasmania, New Zealand;
in Malesia: W. Java (Mt Papandajan), N. Borneo
(Mt Kinabalu), Celebes, Philippines (Luzon,
Mindoro, Negros, Mindanao), New Guinea. Fig.
4.
KING recorded a doubtful collection by Scor-
TECHINI in Perak (Malaya) of which only a field
note was in the Calcutta Herbarium; it has never
been found since in Malaya.
Ecol. In marshy mountain turf, moist places
along mountain brooks, on wet ridges, summits and
plateaus, (1000—)1600-3600 m. Fi. fr. Jan.—Dec.
Note. The closest allied species is H. depressa
(A. CUNN.) WALP. from Tasmania. THUNBERG’S
original description obviously contains some
errors (sepals absent, stamens 4).
4. Haloragis acanthocarpa BRONGN. in Duperrey,
Voy. Coquille Bot. (1828) t. 70; Bru. Fl. Austr. 2
(1864) 483; BartLey, Queensl. Fl. 2 (1900) 555;
SCHINDL. Pfi. R. Heft 23 (1905) 35; Ewart &
Davies, Fl. N. Terr. (1917) 214; MeRR. & PERRY,
J. Arn. Arb. 23 (1942) 407; SPECHT, Rec. Arnhem
Land Exp. 3 (1958) 273. — H. leptotheca F. v. M.
TE
—————— <_< Src‘ el
1971]
Fragm. 3 (1862) 32. — H. palauensis TuyAMaA,
J. Jap. Bot. 16 (1940) 283, f. 3—Sa.
Perennial, sometimes woody at the base; stems
decumbent or erect, 40—-75cm_ long, hairy.
Leaves decussate, ovate, elliptic to lanceolate. on
either side with 5-12 teeth, those of the main
stem 1144-414 by 14-2 cm, (often bulbous-based-)
hairy; petiole 1-2!4(-4) mm. Panicle 10-20 by
5-10 cm, hairy. Bracts ovate-lanceolate, 34-114
by 4% mm. Bracteoles ovate-acuminate, 0.2-0.3
mm. Pedicels c. 1 mm, glabrous. Flowers scat-
tered, erect, then nodding. Sepals c. 0.6-0.8 by
0.5mm, glabrous, coriaceous in fruit. Petals c.
114-114 mm, red to brown-red, the midrib dorsally
pubescent. Stamens 8; filaments 14-14 mm; anthers
c. 1mm. Styles curved inward, c. 0.1-0.2 mm.
Fruit ellipsoid-globose, c. 1-2 by 0.8-1.2 mm,
glabrous or appressedly hairy at apex, the 8
longitudinal ribs more or less alternating with 8
rows of 34 blunt spiny tubercles.
Distr. Northern part of Australia, Micronesia
(Carolines: Palau Is.), and Malesia: New Guinea
and Misool I. Fig. 5.
Fig. 5. Distribution of Haloragis acanthocarpa
BRONGN.
Ecol. In steppe-like country and badly drained
savannah forest, also in Melaleuca swamps and
Eucalyptus woodland, surrounding termite mounds,
0-300 m. FI. fr. April—Jan.
5. Haloragis halconensis Merr. Philip. J. Sc. 2
(1907) Bot. 288; En. Philip. 3 (1923) 221. —
H.secunda Rivv. Trans. Linn. Soc. II, Bot. 9 (1916)
41. — H. suffruticosa Gisps, Arfak (1917) 159;
Went f. Nova Guinea 14 (1924) 108-109, incl.
var. ramosa Went f. et var. galioides Went f.
— H. gjellerupii Went f. l.c. 107. — H. nemorosa
Went /. /.c. 107, t. XI B; Merr. & Perry, J. Arn.
Arb. 23 (1942) 408. — H. fruticosa Went f. Nova
Guinea 14 (1924) 106, t. XI A. — H. sanguinea
Merr. & Perry, J. Arn. Arb. 29 (1948) 162. —
H. microphyllum HooGt. Blumea Suppl. 4 (1958)
228
Erect, branched or unbranched, perennial half-
shrubs or shrubs, with often thick, angular stems,
up to 114 m high. Leaves decussate, or in whorls of
3+4, often crowded, patent, erect or reflexed,
HALORAGACEAE (van der Meijden & Caspers) 245
ovate to oblong or sublanceolate, on either side
with 2-13 teeth, hairy on both sides, hairs of the
upper surface bulbous-based, rarely glabrous,
chartaceous to coriaceous, those of the main
stem(s) 44-344 by %-1% cm, of the branches
0.2-1.5 by0.1—lcm;petioles 0-2 mm.J/nflorescence
a wide to contracted panicle of which the lower
branches are axillary in the upper leaf-axils.
Bracts ovate to elliptic, often oblong, 114-4 by
%-1144 mm. Bracteoles 4-1 mm long, with
widened base. Pedicels 14-114 mm, hairy. Flowers
erect, then nodding, often secund. Sepals tri-
angular-acuminate, 34-114 by 14-1 mm, glabrous
(or slightly hairy on the median dorsal side, some-
times only in a few occasional flowers). Petals
2-344 mm long, red, red-brown, or sometimes
white or yellow, the midrib dorsally hairy. Stamens
8; filaments 4-34 mm; anthers 1-214 mm.
Stigmas sessile, globose. Fruit obovoid, globose,
more rarely ovoid, 1144-134 by *4-1 mm, 8-
ribbed, from which 4 more distinct, appressed-
to patently hairy, at least on the ribs.
Distr. Malesia: Philippines (Luzon; Min-
doro: Mt Halcon; Negros: Mt Canlaon; Min-
danao: Mt Apo), E. Celebes (G. Lumut), New
Guinea, and Solomons (Bougainville; New
Georgia Is.: Kolombangara; Guadalcanal). Fig. 6.
Fig. 6. Distribution of Haloragis halconensis
MERR.
Ecol. On both dry and marshy or wet places,
in peaty marshes, edges of swamps, along streams,
both on sand and clayey subsoils, mostly on open
heaths and cliffs in exposed situations, but also in
forest and cloud-forest, 900-3800m. Fl. fr.
Jan.—Oct.
Notes. This species reminds in habit of H.
erecta (MURR.) SCHINDL. and H. exalata F. vy. M.
of Australia and New Zealand, both of which have
developed styles; the first has, moreover, gla-
brous and 4-winged nuts.
H. halconensis shows an astonishing variability,
reflected in the large number of synonyms cited.
These ‘species’ were originally described from
distant places and based on one or a few specimens
from one locality or mountain. With the increase
of collections it appears that the characters on
which they have been based are grading.
The variability is partly due to the very diverse
246
FLORA MALESIANA
[ser. I, voli ge
habitats; specimens found in shaded localities
have a more rank habit with wider inflorescences,
larger leaves and longer internodes than those of
exposed habitats. The latter show a condensed
habit and are frequently unbranched. Besides,
the altitude causes dwarfing of leaves and habit
with increasing height.
Apart from this, the variability must be partly
ascribed to the fact that in all mountain plants
there is a certain trend to local raciation of a wide-
spread population, but on the other hand in se-
veral localities (Mt Arfak, Mt Gautier) branched
and unbranched specimens occur in one locality in
which the unbranched specimens occur in open
places, the branched ones in less open habitat, on
edges of forests and in the forest.
The variability of the species is vegetative and
quantitative (degree of branching, leaf-size, hair-
yness, efc.). I cannot find qualitative differences in
flowers or fruit.
From the description of H. stokesii F. Brown,
from Rapa I., it would appear that it has sessile
stigmas which would point to affinity with H.
halconensis, but it is described with only 2 stigmas
and 4 stamens.
Excluded
Haloragis distichia JAcK; cf. Fl. Mal. I, 5
(1958) 479 = Anisophyllea disticha (JACK) BAILL.
(Rhizophoraceae).
2. LAUREMBERGIA
BerG. Descr. Pl. Cap. (Aug. 1767) 350, t. 5 f. 10; KAN. in Mart. Fl. Bras. 13, 2
(1882) 377, t. 68; SCHINDL. Pfl. R. Heft 23 (1905) 61; A. RAYNAL, Webbia 19
(1965) 683. — Serpicula LINNE, Mant. 1 (Oct. 1767) 16; Mia. Fl. Ind. Bat. 1, 1
(1855) 631. — EFpilithes BL. Biydr. (1826) 734; HAssk. Cat. Hort. Bog. (1844) 86
(‘Epilithos’). — Fig. 8.
Small perennial herbs, prostrate to ascending, often rooting at the nodes, mainly
branched at the base. Stems terete to quadrangular, sometimes with scattered
enatia as are also found on the leaves. Leaves opposite, or in 4, rarely in 2 rows,
rarely subverticillate, simple, entire to dentate. Flowers polygamous, rarely mono-
ecious, 4-merous, in axillary congested clusters of 1-11 flowers, consisting out
of 1 g and 2-10 2 and/or 1 & and 2-10 9, rarely with either 1-3 3 or 1-7 @ flowers.
— ¢$ Flowers: long-stalked; sepals as in 2; stamens 8 or (not in Mal.) 4; ovary
rudimentary, conical, sometimes with some reduced ovules; style often rudimen-
tary. — § Flowers: shortly to long-stalked; petals and anthers as in 3, for the rest
as in 2°. — 9 Flowers: subsessile to shortly stalked; sepals connate at the very base,
mostly triangular; ovary urceolate with (4—)8 ribs, imperfectly 4-celled when young,
later 1-celled with a central columella. Fruit nut-like, pericap hard, enlarging to
fruit-size long before the seed is set.
Distr. Four spp., 1 in S. Africa, 1 in S. Madagascar and Réunion, 1 in Africa, Madagascar and eastern
South America, and 1 in S. India, Ceylon, Sumatra and Java. Fig. 7.
Ecol. The Malesian species is a characteristic mountain plant, generally of poor soils.
Taxon. The genus has been subdivided into 3 subgenera by RAYNAL. I refrain from any subdivision
as the differential characters used for this purpose occur also in Myriophyllum and also in Haloragis in
which they have not been attributed such high classificatory value. I certainly agree with RAYNAL in a
rigid reduction of species proposed by SCHINDLER.
The Indo-Malesian species differs from the others in possessing 8 stamens, and in the central flower
being very long-pedicelled and normally 3.
1. Laurembergia coccinea (BL.) KAN. in Mart. Fl. (1932) 102, f. 2, 163; Bull. Jard. Bot. Btzg III, 13
Bras. 13, 2 (1882) 377, t. 68; SCHINDL. Pfl. R.
Heft 23 (1905) 68; Back. Schoolfl. Java (1911)
477; Koorp. Exk. Fl. Java 2 (1912) 705; FI.
Tjibodas 2 (1923) 217; F. Biey, Diss. Ziirich
(1925) (embryol.) 1-32, f. 1-20, t. 1-2; STEEN.
Arch. Hydrobiol. Suppl. 10 (1932) 315; Doct. v.
LEEUWEN, Verh. K. Ak. Wet. A’dam sect. 2, 31
(1933) 192, f. 37-39, t. 9; STEEN. Trop. Natuur 21
(1934) 218; BAcK. & BAKH. f. FI. Java 1 (1963)
266. — Epilithes coccinea BL. Bijdr. (1826) 734.
— Serpicula brevipes W. & A. Prod. (1834) 338;
Fyson, Fl. Nilg. Puln. Hilltops 3 (1920) 45, t.
330. — Serpicula hirsuta W. & A. Prod. (1834)
338; WiGuT, Ic. 3 (1845) t. 1001; Mia. FI. Ind.
Bat. Suppl. (1861) 328, 128, pro var. incisa Miq.;
TRIM. Fl. Ceyl. 2 (1894) 148; Fyson, FI. Nilg.
t ° matt
Se re
ad
Fig. 7. Distribution of the genus Laurembergia; subg. Serpiculastrum A. RAYNAL (even line), subg.
Laurembergia (broken line), subg. Indolaurembergia SCHINDL. (stippled line) (after RAYNAL, 1965,
slightly corrected).
Puln. Hilltops 3 (1920) 45; Fl. S. Ind. Hill Stat. 1
(1932) 213. — Haloragis oligantha ARN. Nov.
Act. Caes. Leop.-Car. (1836) 18, non W. & A.
1834. — Serpicula epilithes BL. Mus. Bot. Lugd.-
Bat. 1 (1849) 110, nom. illeg.; Mia. Fl. Ind. Bat. 1,
1 (1855) 632. — Serpicula veronicaefolia (non
Bory) Mia. Fl. Ind. Bat. 1, 1 (1855) 632. —
Serpicula javanica Mia. ibid. 1, 1 (1858) 1090;
Back. Schoolfl. Java (1911) 478. — Serpicula
indica Tuw. En. Pl. Zeyl. (1859) 123, nom. illeg.;
CLARKE, FI. Br. Ind. 2 (1878) 431; Fiscner, Rec.
Bot. Surv. India 9 (1921) 78. — Serpicula zeylanica
ARN. ex CLARKE, FI. Br. Ind. 2 (1878) 431, incl.
var. minor THW. ex CLARKE; TRIM. FI. Ceyl. 2
(1894) 147. — L. brevipes (W. & A.) SCHINDL.
Pfl. R. Heft 23 (1905) 67. — L. glaberrima
SCHINDL. Lc. 67. — L. grandifolia SCHINDL. I.c.
66. — L. hirsuta (W. & A.) SCHINDL. Lc. 64,
incl. var. angustifolia SCHINDL., var. rotundifolia
SCHINDL., var. typica SCHINDL., I.c. 65, = var.
hirsuta p.p.; Steen. Arch. Hydrobiol. Suppl. 10
(1932) 315, f. 5; Bull. Jard. Bot. Btzg III, 13
(1934) 218. — L. indica (Tuw.) ScuinDL. Pfl. R.
Heft 23 (1905) 64, nom. illeg. — L. javanica
(Mig.) ScuinpDL. Lc. 69, f. 20; Back. Schoolfl.
Java (1911) 478; Koorp. Exk. FI. Java 2 (1912)
705, f. 89; F. Biey, Diss. Ziirich (1925); Hocur.
Candollea 6 (1936) 467. — L. wangerinii SCHINDL.
Pfl. R. Heft 23 (1905) 65. — L. zeylanica (ARN. ex
CLARKE) SCHINDL. Lc. 68, incl. var. minor (THw.
ex CLARKE) ScuinpL. — L. agastyamalayana
Henry, J. Bomb. Nat. Hist. Soc. 62 (1965) 603,
fig. — Fig. 8.
var. coccinea. — All synonyms, excl. those of
var. zeylanica.
Internodes 3—6(—10) mm, in sterile branches up
to 15mm, glabrous to pilose in two rows or all
round. Stems c. 440 cm long, 1-2 mm g. Lower
leaves always opposite, upper ones mostly ir-
regularly alternate, subsessile, or rarely up to 2
mm long petioled; lamina elliptic to obovate-
oblong, 5-8 by 2—3!4 mm (in sterile branches up
to 18 by 12mm), acute at both ends, almost
entire or with some coarse, obtuse to mucronate,
often apically thickened teeth especially towards the
apex, rather stiff, glabrous to pilose below on the
midrib and along the margin. Clusters with 1 ¢ or
sometimes % flower, and 2-102 ones. — 6
Flowers: pedicel 6—12(—20) mm long after anthesis,
glabrous to thinly pilose, erecto-patent; sepals
narrowly triangular to ovate, 0.4-0.8 by 0.3-0.6
mm, glabrous or with a tuft of short hairs at the
apex; petals c. 214 by 0.8 mm, outside wine-red;
stamens 8; anthers c. 114 by 14 mm; styles 4(-6),
rudimentary. — 9 Flowers: pedicel 1(-1) mm;
sepals as in g. Fruit glabrous to densely pilose,
with 8 whether or not strongly prominent ribs,
episepalous ribs most distinct upwards, alterni-
sepalous ones most distinct below the middle;
between the costae often with thickenings which
are confluent or consist of 2-4 distinct tubercles;
styles 4(—6). Seed globular, c. 0.8-1 by 0.7-0.8 mm,
brownish.
Distr. S. India (Nilgiris, Pulneys), Ceylon
(Adam’s Peak, Hakgalla, Pendrotallagalla) and
Malesia: N. & Central W. Sumatra, and W. to
248 FLORA MALESIANA [ser. I, vol. 72
Central Java (Gedeh to Diéng). Fig. 7 (subg.
Indolaurembergia).
KANITz recorded L. coccinea from Brazil on the
basis of a SCHOTT specimen; I have seen this (in
W). This is certainly erroneously localized as
SCHINDLER already noted on the label.
Ecol. Moist sunny places, often on stony poor
soil, also along brooks, swamps and lake shores,
often periodically partly inundated, only in the
mountains, 1400-3100 m. Fl. Jan.—Dec.
Notes. All Javanese specimens are glabrous,
all Sumatran and Ceylon specimens are hairy, but
the S. Indian specimens are either hairy or gla-
brous. The hairiness is not combined with other
differential characters, hence no taxonomical value
can be attached to it.
The variation in sculpture of the fruit can be
found occasionally in one specimen.
The name Serpicula epilithes BL. is illegitimate
because it is based on Epilithes coccinea BL.;
Serpicula indica THw. is illegitimate because it is
superfluous by the citation of two older names in
synonymy.
var. zeylanica (ARN. ex CLARKE) MEYDEN, var.
nov. — Serpicula zeylanica ARN. ex CLARKE,
FI. Br. Ind. 2 (1878) 431, excl. var. minor THw.
ex CLARKE. — Haloragis oligantha ARN. 1836,
non W. & A. 1834. — Serpicula indica THw. p.p.,
nom. illeg. — L. zeylanica (ARN. ex CLARKE)
SCHINDL., excl. var. minor (THW. ex CLARKE)
SCHINDL. — L. agastyamalayana HENRY.
Leaves numerous, largely in 4 rows, linear-
lanceolate, with mostly only 1 pair of teeth, or
even none, 8-10(—25) by 1-114(-—6) mm. Clusters
as in the type variety, or the upper ones consisting
only of 1-3 g¢ and the lower ones only 9.
Distr. Ceylon (Adam’s Peak) and S. India
(Agastyamalay Hills).
Note. In habit a clear variety, even recorded
from Adam’s Peak, but obviously replacing.
Fig. 8. Laurembergia coccinea (BL.) KAN. a. Plant, Baxinded
x 14, b-d. flowers: b. 2 flower, c. bisexual flower,
anthers removed, lengthwise section, d. 3 flower, Serpicula verticillata L. f. Suppl. (1781) 416 =
all enlarged (after KANITZ, Fl. Bras. 13, 2; the AH ydrilla verticillata (L. f.) RoyLe (Hydrocharita-
hairy form). ceae).
3. MYRIOPHYLLUM
LINNE, Gen. Pl. ed. 5 (1754) 429; Sp. Pl. ed. 1 (1753) 992; DC. Prod. 3 (1828)
68; Bru. Fl. Austr. 2 (1864) 486; SCHINDL. Pfl. R. Heft 23 (1905) 77; MEIDEN,
Blumea 17 (1969) 306. — Fig. 9, 12, 14-16.
Aquatic or terrestrial, glabrous herbs, perennial or (not in Mal.) sometimes
annual. Stems mostly branched, erect or decumbent-ascending, often freely
floating in aquatic forms, sometimes creeping in terrestrial forms, often with
minute outgrowths (‘myriophyllin-glands’) leaving brown circular scars in the
older parts. Leaves often very constantly flanked by 1(-3) ‘stipular’ outgrowths,
which are filiform to subulate, mostly short, becoming dark, caducous; nearly
always dimorphous in aquatic forms; immersed ones nearly always whorled,
1971] HALORAGACEAE (van der Meijden & Caspers) 249
nearly always pinnately divided into long, filiform or ligulate lobes, very rarely
scale-like reduced or absent; aerial leaves mostly whorled in the lower part of the
stem, upwards often alternate, sometimes opposite, mostly wider than the immersed
leaves and more or less entire, not rarely bract-like. Spikes aerial, mostly terminal,
sometimes forming an apical tuft of sterile leaves after fruit setting which may or
may not perpetuate growth in the next season, or pushed aside by an axillary shoot
by which a sympodial structure is emerging, almost always solitary, rarely 14
additional axillary in the upper leaf axils. Flowers mostly sessile, 1 or more in the
axil of a bract or leaf, each with 2 sometimes very inconspicuous bracteoles;
8, often polygamous with the upper ¢ and the lower 2, or plant monoecious, rarely
dioecious. — % Flowers: 4(—2)-merous. Sepals 4 or 0, mostly very small, erect.
Petals 4 or 2, whether or not caducous, in the 2 flowers strongly reduced and
covered by the styles or 0. Stamens 8 or 4 epipetalous, or 1, in the 2 flowers 0;
anthers often mucronate, rather large. Ovary more or less urceolate, nearly always
(mostly alternisepalous) 4-sulcate, 4-2-celled, in 3 flowers reduced or 0; styles 4 or
2. Fruit mostly + urceolate, breaking up into 4 or 2 one-seeded mericarps;
pericarp often + tuberculate, rather thin, the 4 mericarps very hard by the thick
endocarp which has a soft tissue in an upper axial spot.
Distr. Almost ubiquist, except in most of the Arctic and rare in Africa, c. 40 spp. with a distinct centre
in Australia; in Malesia 8 or 9 native spp. from which two endemic.
The distribution of most SE. Asiatic and Australian species is insufficiently known, as such incon-
spicuous aquatics are still ‘under-collected’.
Ecol. Nearly all species are growing in aquatic or at least in swamp or damp habitats, locally gre-
garious, but dry periods are mostly tolerated. Terrestrial forms occur in most species; these are often
more abundantly flowering and bear more ripe fruits than the aquatic forms.
Dispersal for most aquatic species may be vegetative, either by detached parts of stems or in temperate
species by winterbuds. Endozoic dispersal of fruits by ducks and other birds is recorded from M. verti-
cillatum L. and M. spicatum L. of which the fruits float for a short time. The hard and rather thick en-
docarp may stand passage of the guts.
Taxon. In the vegetative parts and habit the species are very variable: plants with immersed leaves
easily produce aerial leaves when the water recedes, vice versa. Sizes of leaves and bracts respectively show
great variation, especially towards the inflorescence. In general it is impossible to identify sterile specimens.
Varieties or even species distinguished merely by vegetative or quantitative characters deserve no sys-
tematic recognition.
SCHINDLER’s subdivision of the genus into 3 subgenera is unsatisfactory; this is in part caused by the
fact that he did not examine material of a number of critical species, and in part by the fact that he used
unreliable characters, paying amongst others too much attention to the number of stamens. Within
Myriophyllum (‘Eumyriophyllum’) he distinguished for example 2 sections, viz Pentapteris DC. em. O.K.
and Tessaronia SCHINDL., differing in the number of stamens, viz 8 and 4 respectively. However, in M.
spicatum of § Pentapteris their number varies from 8 to 2. M. indicumwith 8 and M. tetrandrum with 4
stamens, are certainly very closely allied, but are placed in two different sections, the first in § Pentapteris
and the second in § Tessaronia. In M. dicocum of subg. Dicarium the number of stamens varies from 4 to
8.
An inadmissible procedure has been to subordinate Pelonastes Hook. f. as a subsection to subg.
Myriophyllum in an emended sense, although none of Hooker's original species is attributed to it;
these are arranged by SCHINDLER in the new subg. Brachytheca SCHINDL.
In my opinion it serves no good use to distinguish subgenera or even sections in the genus.
Notes. The sculpture of the fruits becomes only clearly visible in dried material.
Largely because of the supposed incompleteness of the knowledge on the distribution of the species,
I have included 5 extra-Malesian species in the key, wich in this way covers all species from Malesia,
SE. Asia, Madagascar, and Africa.
* For literature on the extra-Malesian species, their distribution and ecology, I refer to my precursory
paper in Blumea 17 (1969) 304-311.
Pre-identification of some species with simple opposite leaves and ¢ flowers (notably M. pygmaeum
and M. pedunculatum) is rather difficult, but they can easily be assigned to the genus by the presence of
dark, subulate pseudostipules and dark-tipped leaves.
250
FLORA MALESIANA
{ser. I, vol. 7}
KEY TO THE SPECIES
1. All leaves alternate to opposite, entire or dentate, or the lower ones pinnate.
2. $ Flower with 2 petals and 1 stamen. Bracteoles minute or 0.
3. 3 Flower stalked. Fruit with 4 mericaps. North Vietnam. Fig. 15d-f. .
3. 3 Flower sessile. Fruit with 2 mericarps.
4. Fruit with rows of distinct spines at the base.
M. bonii TARDIEU-BLOT
5. Spines present over the entire dorsal length of the mericarps. Madagascar. Fig. 15g.
5. Spines absent in the central part of the mericarps. .
4. Fruit with rows of tubercles at the base..
M. mezianum SCHINDL.
11. M. coronatum
10. M. siamense
2.3 Flower with 4 petals and 4 or 8 stamens. Bracteoles small but conspicuous, at least those of the
3 flowers.
ey Leaves 2-15 mm long, always opposite. Stamens 8.
7. Flowers (1—)2(—3) on each branchlet, strictly monoecious, on each branchlet of one sex. Stems
upwardly very much sympodially branched, with very short internodes 0.3-1 mm long. Leaves
caducous. Sepals in ¢ flowers 0.
1. M. pygmaeum
7. More flowers on each branchlet, the upper 8, the ‘lower on or - plant dioecious. Stems with much
longer internodes, monopodially branched. Leaves not caducous. Sepals in 3 flower very con-
spicuous.
6. Leaves 10-30 mm long, mostly alternate. Stamens a
2. M. pendunculatum
8. Flowers (1—)3—5 together, distinctly stalked. Fruit c. 114 by 1 mm, with dorsally rounded mericarps.
S. India. Fig. 14q-+t.
M. oliganthum (W. & A.) F.v.M.
8. Flowers solitary, sessile or nearly | so. . Fruit. C 2" by 2 mm, in transverse section + quadrangular
with sharp ribs and slightly concave sides. .
8. M. tuberculatum
1. At least the lower leaves in whorls, nearly always pinnate.
9. Leaf-base + dilated, those of a whorl, at least in sterile branches, enclosing the stem at the nodes in
the upper parts. Bracteoles subulate, sometimes with | or 2 lateral laciniae. .
4. M. brasiliense
9. Stem always visible on the nodes between the leaf-bases. Bracteoles either not subulate, or with more
laciniae.
10. Bracteoles digitate or pinnate.
11. Fruit cruciform in section, finely tubercled, rarely smooth.
12. Stamens 8. Anther linear, 114-134 mm. Petals 114-2 mm. Ceylon, S. India. Fig. 14d—g.
12. Stamens 4. Anthers oblong, 0.6—-0.8 mm. Petals c. 1(-114) mm long..
11. Fruit in transverse section quadrangular with rounded edges, smooth.
M. indicum WILLD.
7. M. tetrandrum
6. M. verticillatum
10. Bracteoles simple, not dissected: ovate, rhomboid, or lanceolate.
13. Fruit with 2(-4) mericarps, smooth or indistinctly lengthwise lineolate, stalked.
9. M. dicoccum
13. Fruit with 4 mericarps of which sometimes 1-2 are visible but reduced and unfertile, not lineolate.
14. Flowers (1—)2—6 together in the leaf-axil; the middle flower %, shortly but distinctly stalked.
15. Flowers (1—)3—5(—6) together in the leaf-axil. Fruit rounded, shallowly 4-sulcate, tuberculate.
Anthers c. 1 mm long. S. India. Fig. 14q-t.
M. oliganthum (W. &. A) F.v.M.
15. Flowers 1-3 together. Fruit -- cruciform, dorsally narrowly winged, smooth or punctate.
Anthers c. 2 mm long. Madagascar. Fig. 16.
M. axilliflorum BAKER
14. Flowers always solitary, either all sessile, or the flowers stalked.
16. 3 Flower distinctly stalked. Stamens inserted on a 0.1—0.2 mm high androphore. Mericarps
apically spreading by the cushion-like thickened style-bases. .
3. M. propinguum
16. All flowers sessile. Androphore 0. Mericarps not spreading at apex.
17. Mericarps dorsally rounded. Sepals in g flowers distinct, c. 4% by /% mm. Stamens 8. Petals
caducous.
5. M. spicatum
17. Mericarps dorsally acute. "Sepals indistinct. Stamens 4. Petals strongly recurved after anthesis,
persistent until fruit is set.
1. Myriophyllum pygmaeum Matrtr. Bot. Jahrb.
69 (1938) 275; MEIJDEN, Blumea 17 (1969) 311. —
Fig. 9a-b.
Terrestrial or semi-aquatic, growing in dense
cushions. Stems very short, very much branched,
creeping or decumbent with erect branches or erect,
after anthesis sympodially branched, with very
short internodes, 0.3-1 mm. Leaves opposite,
linear to subulate, entire, with a dark thickening on
apex above, erecto-patent, + fleshy, 2-6 by
0.3—14 mm. Flowers solitary in the leaf-axils of very
8. M. tuberculatum
short (5-10 mm long), strictly unisexual branches
which are later sympodially overtopped, each
bearing only (1—-)2(-3) developed flowers and
often besides 2—4 deficient higher flowerbuds. —
6 Flowers: stalked, stalk 4-14 by c. %mm,
erect. Bracteoles oblong, obtuse, finely mucro-
nate, 1 by 0.3mm. Sepals 0. Petals with wide
base and rounded, nearly flat, finely serrate apex,
1-114 mm long, rose to red. Stamens 8, filaments
up to 1mm; anthers elliptic, mucronate, 1-114
by 0.4-14 mm. Rudiment of ovary 0.—2 Flowers:
5 i a ee
Fig. 9. Myriophyllum pygmaeum Matrtr. a. Branch
with ¢ flowers, b. branch with @? flowers. —
“M. pedunculatum Hoox. f. c. g Flower, d. brac-
teole, e. fruit. — M. propinquum CunNn. f. 3 Flower
with bracteoles, g. fruit, A. fruit, transverse
section (a-b HooGLaNp 9972, c-e ROBBINS
381, f-h N. Marcuant s.n.). All 5.
HALORAGACEAE (van der Meijden & Caspers)
251
up to 0.2mm stalked. Bracteoles very small,
narrowly triangular, 0.2-0.4 by 0.2 mm. Sepals 0.
Petals 0. Styles 0.2 by 0.3mm. Fruit truncate
above, shallowly 4-lobed, puncticulate, c. 1 by 1
mm.
Distr. Malesia: E. New Guinea (Morobe
Distr.: Mts Sarawaket and Albert Edward).
Fig. 10.
Ecol. In and along very shallow, boggy,
partly dried up pools in alpine grassland, growing
in dense cushions, c. 3500-3700 m.
2. Myriophyllum pedunculatum Hook. f. in Hook.
Lond. J. Bot. 6 (1847) 474; Fl. Tasm. 1 (1860) 122,
t. 23; Bru. Fl. Austr. 2 (1864) 489; Hook. f.
Handb. New Zeal. Fl. 1 (1864) 67; SCHINDL.
Pfi. R. Heft 23 (1905) 85; CHEESEMAN, Man. New
Zeal. Fl. ed. 2 (1925) 625; Curtis, Stud. FI.
Tasm. 1 (1956) 190; ALLAN, Fl. New Zeal. 1
(1961) 252; MeWDEN, Blumea 14 (1966) 245;
ibid. 17 (1969) 310. — M. longibracteolatum
SCHINDL. Pfl. R. Heft 23 (1905) 84. — Fig. 9c-e.
Terrestrial cr semi-aquatic, whether or not
growing in dense cushions. Stem unbranched to
much branched, decumbent or erect, after anthesis
monopodially accrescent. Leaves opposite, erecto-
patent to reflexed, entire, rarely pinnate, the entire
ones linear, (2—)4~15(—25) by 0.4~-1 mm, obtuse to
mucronate, above at apex with an ovate, darker
coloured cushion-like thickening c. 0.2 mm long.
Flowers solitary in the middle and upper leaf-
axils, the lower often bearing the fruits of the
last season; the upper 2-4(—8) flowers 3, the next
lower 4-8(-12) flowers 2; not rarely dioecious.
Bracteoles lanceolate-oblong to linear-lanceolate,
rarely - elliptic, acute or acuminate, entire or
serrate, 0.6-1.5 by 0.2-0.3(-0.6)mm. — ¢
Flowers: up to 4% mm stalked. Sepals lanceolate,
acute, serrate, c. | by 0.4mm, +- erect, with tiny
whitish tubercles. Petals very concave in the apical
part, the borders nearly flat, and finely serrate,
the apex mucronate and often recurved, in anthesis
erect and 1-114 mm long, after anthesis elon-
gating, up to 2mm long, recurvate, persistent.
Stamens 8; anthers oblong, c. 1-1!4 mm. Ovary
rudimentary, c. 0.3 by 0.3mm. — @ Flowers:
up to 0.2 mm stalked. Sepals lanceolate-oblong,
acute, 0.3-0.8 by 0.2 mm. Petals strongly reduced,
covered by the styles, shorter than 0.1 mm.
Styles with thick, fleshy bases, stigmas after anthesis
strongly ligulately recurved, very long papillose.
Fruit c. 1.2 by 1.2 mm, with apically spreading
mericarps, these terete, with distinct tubercles.
Distr. New Zealand (also Chatham and
Stewart Is.), Tasmania, Australia (Victorian
Alps, New South Wales, also recorded from W.
Australia); in Malesia: New Guinea (E. Papua:
Western Highlands, Morobe Distr.). Fig. 13.
Ecol. In New Guinea in the upper montane
and alpine zone, from 2300-3500 m, in the Austra-
lian region often descending to the lowland. In
swamps, shallow pools, alpine bogs, and inun-
dated river banks.
Notes. In all but one collection I have seen,
the leaves are entire, but in one collection from
FLORA MALESIANA
[ser. I, vologe
Fig. 10. Distribution of Myriophyllum propinquum CunNn. (full line & question mark), M. tetrandrum
Roxs. (black area and dots), M. pygmaeum Mattr. (triangles), and M. coronatum MEISDEN (++).
Victoria (A. C. BEAUGLEHOLE 5887) the medium
and lower leaves are pinnate with few filiform
lobes. This observation clearly shows the great
plasticity of the leaves in this genus.
M. longibracteolatum SCHINDL. differs only in
quantitative characters.
M. votschii SCHINDL. from New Zealand is
closely allied and might also belong to M. pe-
dunculatum.
3. Myriophyllum propinguum CuNN. Ann. Nat.
Hist. I, 3 (1839) 30; ScHINDL. Pfl. R. Heft 23
(1905) 89, incl. var. tenuifolium SCHINDL. I.c.
90; CHEESEMAN, Man. New Zeal. FI. ed. 2 (1925)
624; Ewart, Fl. Vict. (1930) 886, t. 309 excl. c;
BLACK, FI. S. Austr. ed. 2, 3 (1952) 645; CurTIs,
Stud. Fl. Tasm. 1 (1956) 190; ALLAN, Fl. New
Zeal. 1 (1961) 253; MEIsDEN, Blumea 17 (1969)
310. — M. variaefolium Hoox. f. in Hook. Ic.
Pl. 3 (1840) t. 289; Bru. FI. Austr. 2 (1864) 487.
— M. verticillatum L. B ussuriense REGEL, Tent. Fl.
Ussur. (1861) 60. — M. ussuriense (REGEL)
Maxim. Diagn. Pl. 15 (1873) 183; ScCHINDL.
Pfl. R. Heft 23 (1905) 86; Onw1, FI. Jap. (1965)
661. — M. intermedium (non DC.) CLARKE, FI. Br.
Ind. 2 (1878) 433, pro syn. M. variaefolium;
Kirk, Stud. Fl. New Zeal. (1899) 150; CHEESE-
MAN, Man. New Zeal. FI. ed. 1 (1906) 151. —
Fig. 9f-h.
Aquatic or semiterrestrial. Stem branched or
not, erect, ascending or freely floating. Immersed
leaves mostly present, rarely absent, in whorls of
(4-)5-6(-8), 1-214 by 34-114 cm in outline,
with c. 5-10 pairs of 3-25 mm long and c. 0.2 mm
wide filiform lobes. Lowermost aerial leaves in
whorls of (2—)4~-5(-6), pinnate or (less often)
entire, the pinnate ones as the immersed leaves,
the entire ones linear to linear-lanceolate, flat,
1971]
somewhat fleshy, 5-10 by ‘4-1 mm, mostly
patent. Upper aerial leaves entire, dentate or
pinnate, patent to upwards erect, below mostly
punctate, the entire ones 10-20 by 0.7-1.5 mm,
the pinnate ones 15-20 by 2-7 mm in outline,
with 2-7 pairs of erecto-patently incurved, up to
3mm long lobes. Flowers solitary in the upper
leaf-axils, the upper ¢, the lower Q, often & in
between, or dioecious. — ¢ Flowers: stalk up-
and downwards broadened, 0.6—1 by 0.3-0.5 mm.
Bracteoles ovate, acuminate, serrate, c. 1 by 1 mm.
Sepals lanceolate, acute, finely serrate, c. 1 by
Y% mm, soon caducous. Petals soon caducous
after anthesis. Stamens 8, inserted on a 0.1-0.2
mm high, a little narrowed androphore; filaments
at first mobile, later rather stiff, + erect, not
caducous; anthers oblong, obtuse, (1l—)1.2—2.5
by 0.3-0.4 mm. Rudiment of ovary 0. — 2 Flowers:
sessile. Bracteoles + ovate to oblong, concave,
mostly appressed, entire or finely serrate, obtuse,
+ mucronate, 0.3-0.5 by 0.15—0.3 mm, finally
caducous. Sepals 0. Petals 0. Styles spreading at
the base, c. 0.2 by 0.1-0.3 mm. Stigmas after
anthesis strongly recurved, very long papillose,
0.1-0.4mm. Fruit 0.6-1.2 by ‘4-1 mm, the
mericarps terete, apically spreading by the cushion-
like thickened style-bases, with long tubercles at
least at the base, punctate.
Distr. New Zealand, Tasmania, Australia
(S. Australia, Victoria, New South Wales,
Queensland, W. Australia) and E. Asia: NE.
China (Heilong Jang, Whusuli Jiang; Manchuria),
Korea, Taiwan, Japan, S. China (S. Yunnan).
Fig. 10.
Notes. Extremely variable in the Australian
region, especially in vegetative parts. M. pro-
pinquum and M. ussuriense differ only in minor
vegetative characters.
The area is strongly disjunct. One sterile collec-
tion from West New Guinea (Wissel Lakes,
Eyma 4733, at c. 1600m) might belong to this
species or to M. verticillatum.
4. Myriophyllum brasiliense CAmpess. in A. St. Hil-
Fl. Bras. Mer. 13, 2 (1829) 182; ScHINDL. Pfl.
R. Heft 23 (1905) 88; Heyne, Nutt. Pl. ed. 1, 3
(1917) 391; ibid. ed. 3 (1927) 1207; STEN. Arch.
Hydrobiol. Suppl. 10 (1932) 315; Curtis, Stud.
Fl. Tasm. 1 (1956) 190; Mason, FI. Marshes
Calif. (1957) 615, f. 280; Back. & Baku. f. FI.
Java 1 (1963) 266; ALston, Muelleria 1 (1967)
171; Meupen, Blumea 17 (1969) 171. — M.
proserpinacoides Gitt. ex Hook. & ARN. Bot.
Misc. (1833) 313. — Fig. 14a-c.
Aquatic or semiterrestrial. Stem branched
mostly at the base only, perpetuating growth after
anthesis. Lower immersed J/eaves pinnate, seg-
ments thread-like. Upper immersed leaves and
aerial leaves alternately in whorls of 5 (rarely 4),
flanked by 2-3 c. | mm long, mostly recurved
enatia, hydrophobic and often + glaucous.
Leaf-base +. dilated, c. 0.6-1 mm wide, of a
whorl at least in the upper sterile shoots enclosing
the stem at the nodes, blade oblong in outline,
c. 2-4 by 14-1 cm, rather stiffly patent to erecto-
HALORAGACEAE (van der Meijden & Caspers)
253
patent, with 8-18 pairs of subulate, mucronate
lobes, 2-8 by c. 0.2 mm. Mostly monoecious or
+ dioecious, sometimes in the middle with §
flowers. Flowers solitary in the middle and upper
leaf-axils. — ¢ Folwers (not seen in Mal.):
pedicel up to 4 by 0.4mm, up- and downwards
broadened. Bracteoles subulate, 2-3 mm long,
+ flat, often with 1 or 2 lateral laciniae. Sepals
narrowly triangular, acute, serrate towards the
apex, c. 1144-2 by 14 mm, at first erect, finally
recurved. Petals 4, caducous before anthesis,
c. 4mm long. Stamens 8; anthers linear-lanceolate,
c. 4 by 1 mm. Ovary reduced, up to 0.3 mm long,
— ° Flowers: pedicel up to 1 by 0.4 mm, cy-
lindric, erect. Bracteoles subulate with dilated
base, sometimes with 1 lacinia, 1-114 mm long.
Sepals narrowly triangular, acute, entire or scar-
cely serrate, 1-114 by 0.2-0.3 mm, at first erect,
finally recurved. Petals very strongly reduced,
subulate, adnate to the torus, up to 14 mm long.
Styles 0.2-0.3 mm. Fruit (sec. SCHINDLER)
ovoid, 4-sulcate, c. 1.8 by 1.2 mm, shortly pe-
dicelled, papillose.
Distr. Native in South America (E. Brazil,
Uruguay, Argentina, Chile), often cultivated
elsewhere in ponds or aquaria; naturalized in
Japan (“Yamamoko in Settsu’’), in Australia
(New South Wales, Victoria, Queensland; New
Zealand: North I.), in SW. and SE. North
America (casually to New York), and in Malesia:
locally abundant in W. Java.
Ecol. In Java in ditches, ponds and rice-fields,
400-1500 m, freely floating to creeping-ascending.
Uses. In Java cultivated in fish-ponds; tips
are eaten as a vegetable. Easily propagated ve-
getatively.
Vern. Paris, S; diamond-milfoil, parrot-feather, E.
Note. All naturalized specimens are sterile or
have only @ flowers and do not produce fruit.
Also in South America fruits rarely occur. The
only description of the ripe fruit was given by
SCHINDLER. An illustration of an apparently
ripe fruit is found in GLeason, New Britt. &
Brown Ill. Fl. 2 (1952) 601.
5. Myriophyllum spicatum Linné, Sp. Pl. (1753)
992; Maxim. Diagn. Pl. 15 (1873) 183, incl. var.
muricatum; CLARKE, Fl. Br. Ind. 2 (1878) 433;
SCHINDL. Pfl. R. Heft 23 (1905) 90; Koorpb.
Exk. Fl. Java 2 (1912) 708, excl. syn. M. pusillum
BL.; GUILLAUMIN, FI. Gén. L—C. 2 (1920) 717;
Merr. En. Philip. 3 (1923) 221; Ript. Disp.
(1930) 546; Steen. Arch. Hydrobiol. Suppl. 10
(1932) 316; ALLAN, Handb. FI. New Zeal. (1940)
285; Patren, Rhodora 56 (1954) 213; Léve,
Rhodora 63 (1961) 139; Witp, Harmf. Pl. (1961)
42, t. 14d; SuBRAMANYAM, Aq. Ang. (1962) 17;
Tarpieu-BLot, Fl. Laos, Camb. & Vietn. 4
(1965) 125; Ouwt, FI. Japan (1965) 660; RAYNAL,
Fl. Cameroun 5 (1966) 132, fig.; MEUDEN,
Blumea 17 (1969) 309. — M. exalbescens FERN.
Rhodora 21 (1919) 120. — Fig. 12a-c.
Aquatic. Stem much branched. Immersed
leaves in whorls of 4-5, 144-214 by c. 2.cm in
outline, with 7-11 pairs of filiform, obtuse seg-
FLORA MALESIANA
[ser. I, vol. 71
Fig. 11. Distribution of Myriophyllum spicatum L. (after PATTEN, 1954, emended).
ments 6-12 mm long. Aerial leaves mostly sud-
denly much smaller, the lowermost ones always
in whorls of 4, mostly pinnate, as the immersed
ones. Middle and upper leaves entire, rarely
pinnate, obtuse, mostly ovate to obovate, some-
what convex, sometimes spathulate or linear, and
flat, 2-10 by 1-2 mm, patent, recurved in fruit.
Flowers solitary in the middle and upper leaf-
axils, the upper ones g, the lower 9, in between
often %, sessile. Bracteoles broadly ovate, obtuse,
erect, with a brown, + serrate membranous
margin, c. 1.2 by 1mm. — ¢ Flowers: sepals
ovate to triangular, with a brown membranous
mucronate apex, c. 14 by 0.3 mm, erect. Petals at
the base -- auriculate, caducous in anthesis, c.
214 mm long, wine-red. Stamens 8(—2); anthers
linear-lanceolate, c. 2 by 0.4mm. Ovary reduced,
up to 0.3 mm. — ¢% Flowers: below the middle of
the spike with sepals and petals as in 9, with 8
stamens, above the middle of the spike as the 3,
but often poorly setting fruit. — 2 Flowers:
sepals strongly reduced, semiorbicular, with a
brown mucronate apex, c. 0.2 by 0.3 mm, erect.
Petals reduced, ovate, acute, 144 by 0.3mm,
patent, covered by the stigmas, soon caducous.
Styles c. 14 by 0.3 mm. Fruit 144-3 by 114-3 mm,
terete, very deeply and alternisepally 4-sulcate or
deeply 4-cleft; mericarps 4, often only 2 fully
developed, dorsally rounded, with a distinct, often
verrucose marginal ridge, at the back smooth or
verrucose, rarely wholly smooth.
Distr. Almost ubiquist on the northern hemi-
sphere, rare in Africa and in the tropics; in
Malesia: N. Sumatra (Toba Lake) and Philippines
(Mindanao: Lake Lanao, CLEMENS 450, 7.¥.).
Fig. 11.
A record from New Zealand rests according to
Miss R. MASON on an erroneous identification
(Lucy B. Moore, in litt.).
Ecol. Shallow banks of lakes, 670 and 900 m.
Terrestrial forms are rare and sterile (cf. HEGI,
Fl. Mitt.-Eur. ed. 2, 5, 2, 1964, 902, f. 2273)
and might have been found at Lake Toba (LOr-
ZING 7718).
Galls are recorded from the Toba Lake by
RUTTNER c.s. (see ZELLER, Ber. Deut. Bot. Ges.
55, 1937, 473-475, f. 1-2). The stems are malform-
ed and have few reduced and short leaves.
Fruit-setting is often poor although g and 2
flowers are present in the Toba collections.
Notes. Very variable species. Reductions are
not rare in the inflorescence, with the upper part
Fig. 12. Myriophyllum spicatum L. a. 3 Flower,
b. bracteole, c. fruit. — M. dicoccum F. v. M. d.
Bisexual flower with bracteole, e. fruit, f. fruit,
cross-section (a HooKer f. & THOMSON S.n.,
b-c Fl. Turcom. As. 1926, df Brass 28669).
ATES:
1971]
strongly elongating, the lower part compact.
M. exalbescens FERN. is, apart from the stems
which become whitish in drying, in no way
otherwise qualitatively different from the Eur-
opean and Asiatic specimens of M. spicatum L.
Specimens with ‘exalbescens’ characters (PATTEN,
1954; Love, 1961) are found scattered all over the
range of the species, not only in America.
7 |
© a ya: 2 i
Se ee = > |
Fig. 13. Distribution of Myriophyllum pedun-
culatum Hoox. f. (full line), M. tuberculatum
Roxs. (triangles), M. dicoccum F. v. M. (dots),
and M. siamense (CRAIB) TARDIEU-BLOT (+).
6. Myriophyllum verticillatum Lrvné, Sp. PI.
(1753) 992; CrarKeE, FI. Br. Ind. 2 (1878) 87;
Mason, FI. Marshes Calif. (1957) 617, f. 283;
Oxuwt!, Fl. Japan (1965) 660; SincLarir, Gard.
Bull. Sing. 22 (1967) 230; MEUDEN, Blumea 17
(1969) 309. — M. spicatum (non L.) GAERTN.
Fruct. (1788) 331, t. 68. — Fig. 14j-m.
Aquatic, sometimes semiterrestrial. Stems few-
branched or not, often perpetuating growth after
fruit-setting. Immersed Jeaves in whorls of
4-5(-6), 1-4 by 1-4 cm in outline, with c. 8-16
pairs of mostly opposite, filiform segments, 10-25
mm long. Aerial leaves gradually or sometimes
rather suddenly smaller, all pinnate, the lower as
the immersed leaves, the upper lanceolate to
linear-lanceolate with 8-10 pairs of rather stiff,
+ appressed lobes, 5-15 by 2\4-4 mm, mostly re-
curved in fruit. Flowers solitary in the middle and
upper leaf-axils, mostly 9, the upper sometimes 3,
the lower ones often 9°, sessile or subsessile.
Bracteoles + circular in outline, digitate, rarely +-
pinniform, c. 1 mm long. Sepals triangular, acute,
serrate, c. 1 by 0.8mm, erect, in 2 up to 0.3
by 0.4mm. Petals c. 3-4 mm long, rose, erecto-
patent in anthesis, caducous after anthesis,
strongly reduced in 2 flowers and covered by the
styles. Stamens 8; anthers linear-lanceolate, c.
244 by 4 mm. Styles c. 4 by 0.3mm. Fruit
ovoid, in section +. quadrangular with dorsally
rounded, smooth mericarps, c. 3 by 3 mm.
Distr. Europe (north to Lapland, not in Ice-
land and Greenland; including the African Me-
diterranean), Asia (temperate and boreal, east
to Kamchatka and Japan, south to Afghanistan
and Kashmir), North America (Canada, from
HALORAGACEAE (van der Meijden & Caspers)
=
British Columbia to New Foundland, south to
Maryland and California); in Malesia: possibly
found in W. New Guinea (Wissel Lakes, 1600 m.)
Notes. The New Guinean collection (EyMa
4733) is sterile. Identification of vegetative plants
is extremely difficult in this genus. I share the
opinion of SINcLarrR, /.c., that this collection may
belong to M. verticillatum, although it is not im-
possible that it should turn out to belong to M.
propinquum.
Flower characters in the description are taken
from extra-Malesian material.
7. Myriophyllum tetrandrum Roxs. FI. Ind. 1
(1820) 470; DC. Prod. 3 (1828) 69; Roxs. FI.
Ind. ed. Carey 1 (1832) 451; W. & A. Prod.
(1834) 399; ScHINDL. Pfl. R. Heft 23 (1905) 96;
GUILLAUMIN, FI. Gén. I.-C. 2 (1920) 717; Crars,
Fl. Siam. En. 1 (1931) 591; TARDrEU-BLoT, FI.
Laos, Camb. & Vietn. 4 (1965) 125; MEIJDEN,
Blumea 17 (1969) 308; Fl. Thail. 2 (1970) 2. —
M. indicum (non WILLD.) PRAIN, Beng. PI. (1903)
474; Rec. Bot. Surv. India 3 (1908) 210; Sus-
RAMANYAM, Aq. Ang. (1962) 17; S. R. BENNET,
Tax. Stud. Fl. Howrah (Thesis) (1968) 651. —
M. indicum ssp. tetrandrum (ROxB.) MEIDEN,
Ident. Lists 30 (1960) 426, nomen et in sched.;
KENG, Mal. Nat. J. 23 (1970) 123. — Fig. 14h-i.
Aquatic or semiterrestrial. Stem branched, erect.
Immersed /eaves in whorls of (4-)5-6, 2-5 by
144-4cm in outline, with c. 12-24 pairs of c.
10-20 mm long, filiform, acute, brown-tipped
segments. Lowermost aerial leaves mostly pinnate
and rather stiffly spreading with short spreading
lobes. Middle and upper aerial leaves mostly
narrowly triangular in outline, 24 by 14-1 mm,
with 6-12 pairs of erecto-patent, subulate, brown-
tipped, very acute lobes, more rarely lanceolate
to linear-lanceolate, 6-12 by 1-144 mm long,
with few, -- appressed subulate teeth, or almost
entire, at first erecto-patent, in fruit recurved.
Flowers solitary, + sessile in the middle and upper
leaf-axils, 3, or the upper ¢. Bracteoles digitate
or sometimes + pinniform, 0.6-1 mm long, both
with c. 5 subulate lobes. Sepals triangular, 0.1—0.3
by 0.1—0.3 mm, entire or finely serrate, acute, erect.
Petals almost entire, c. 1(-144)mm long, ca-
ducous after anthesis. Stamens 4; anthers +
oblong, 0.6-0.8 by 0.2-0.25mm, obtuse. Fruit
cruciform, c. 2 by 2 mm, mericarps + ovate, with
convex back and + flattened sides, irregularly and
finely tuberculate to smooth.
Distr. N. & E. India (Kashmir, Uttar Pradesh,
Bihar, W. Bengal, Assam, Agartala, Manipur,
Eastcoast from Ganjam southwards), S. Thailand
(Bangtapan Noi: Ta Samet), North Vietnam;
in Molesia: Malay Peninsula (Kangar; Perlis).
Fig. 10.
Ecol. In rather shallow open water of ditches
and canals.
Notes. The species was in the past seldom
recognized by Indian botanists and mostly reduced
to M. indicum, following CLarke (in Fl. Br. Ind.
2, 1878, 433), sometimes to M. tuberculatum.
Although they differ in the number of stamens,
256 FLORA MALESIANA [ser. I, vol. 7?
I believe it to be much closer allied to M. indicum
WILLD. than to M. tuberculatum Roxs., which has
the same number of stamens. M. tetrandrum and
M. indicum can only be distinguished in flowering
material, the fruits and bracteoles showing no
differences. As M. indicum and M. tetrandrum
are replacing, M. indicum being confined to Ceylon
and the South Deccan and M. tetrandrum to
NE. India and Indo-China, I have originally
assigned them subspecific rank, but as according
to PRAGLOWSKI (Grana 10, 1970, 234) their pollen
shows distinct differences, I have concluded that
they are closely allied but distinct species.
A record from Java (ZOLL. Syst. Verz. 2, 1854,
86, ZOLLINGER 883) belongs to Limnophila
(Scrophulariaceae).
8. Myriophyllum tuberculatum Roxs. FI. Ind. 1
(1820) 471; DC. Prod. 3 (1828) 69; Roxs. Fl. Ind.
ed. Carey 1 (1832) 451; Mia. Fl. Ind. Bat. 1, 1
(1855) 635; Kurz, J. As. Soc. Beng. 40, ii (1871)
52; CLARKE, Fl. Br. Ind. 2 (1878) 432; SCHINDL.
Pfi. R. Heft 23 (1905) 96; SINcLAIR, Bull. Bot. Soc.
Beng. 9, 2 (1955) 94; SUBRAMANYAM, Aq. Ang.
(1962) 17; SrncLair, Gard. Bull. Sing. 22 (1967)
230; VASUDEVAN & KEsAN Nair, J. Bomb.
Nat. Hist. Soc. 64 (1967) 391, f. 1-22; MEIJDEN,
Blumea 17 (1969) 308. — M. tetrandrum (non
Roxs.) GRAH. Cat. Pl. Bomb. (1839) 76; STEEN.
Webbia 8 (1952) 435. — M. indicum GRIFF.
Not. 4 (1845) 687, non WILLD. 1805, nom.
illeg. — M. spathulatum Bratt. & HALLBL. J.
Ind. Bot. Soc. 2 (1921) 44, fig. (ex descr.);
SUBRAMANYAM, Aq. Ang. (1962) 17, f. 11. — Fig.
14 n-p.
Aquatic or semi-aquatic. Stems mostly much
branched. Immersed Jeaves in whorls of 4-5,
214-4 by 1-114 cm in outline, with 8-25 pairs of
rather weak, ligulate, acute or shortly mucronate
segments 10-20 mm long. Aerial leaves in the
lower part as the immersed ones, upper ones
finally alternate, with less and shorter lobes, up-
permost almost entire, 5-20 by 1-3 mm, patent or
recurved. Flowers solitary, sessile, in the middle
and upper leaf-axils, mostly all 2, rarely the upper
flowers 3, sometimes the lower 2. Bracteoles +
Fig. 14. Myriophyllum brasiliense CAMBESS. a. 3
Flower with bracteole, b. bracteole of 2 flower,
c. 2 flower. — M. indicum WiLp. d. Flower,
e. bracteole, f. fruit, g. fruit, transverse section. —
M. tetrandrum Roxs. h. & Flower, i. bracteole.
— M. verticillatum L. j. Young % flower, k.
bracteole, /. fruit, m. fruit, transverse section. —
M. tuberculatum Roxs. n. Young § flower with
bracteole, o. fruit, p. fruit, transverse section. —
M. oliganthum (W. & A.) F. v. M. q. Part of in-
florescence with % flowers and young fruits,
r. bracteole, s. fruit, ¢. fruit, transverse section
(a-c O. BUCHTIEN s.n., d-g THwalTes CP 1549,
h-i J. THOMSON s.n., j-m HLB 903.364-343,
n—-p Herb. GRIFFITH 2441, g-t HOHENACKER 1563).
All x5, except g which is X24.
"
1971]
HALORAGACEAE (van der Meijden & Caspers)
257
rhomboid, serrate, acute, c. 1.2 by 0.8mm.
Sepals very small, + orbicular, entire or finely
serrate, erect, 0.1—-0.25 by 0.1-0.25 mm. Petals in
and after anthesis patent and finally recurved-
circinnate, not soon caducous, c. 14-114 mm long.
Stamens 4; anthers elliptic-oblong, c. 14-1 by
0.2-0.3 mm. Styles 0.1-0.2 by 0.2mm, scarcely
recurved, with capitate stigmas. Fruit quadran-
gular, between the sharp and alternisepalous ribs
slightly concave, indistinctly 4-sulcate, aculeate,
finely foveolate, c. 2-314 by 2-314 mm.
Distr. India (Kerala: Alwaye; Mysore;
Bombay: Bombay, Khandala; Madhya Pradesh:
Sagor; Orissa; W. Bengal; Assam; Khasya),
E. Pakistan (Cox’s Bazar), Australia (Northern
Territory: Fog Dam Sanctuary); in Malesia:
N. Malay Peninsula (Kedah, Perlis), SE. Borneo
(Bandjermasin), SW. (Pangkadjene) & SE. Ce-
lebes (Kendari), Moluccas (Halmahera; Sulu Is.:
Mangoli), and Lesser Sunda Islands (W. Flores:
Ruteng). Fig. 13.
Ecol. In shallow, stagnant water at low al-
titude.
Note. See for the differences with M. indicum
and M. tetrandrum the key, the figures, and note
under the preceding species.
9. Myriophyllum dicoccum F. vy. M. Trans. Phil.
Inst. Vict. 3 (1859) 41; Bru. FI. Austr. 2 (1864)
489; F. v. M. Fragm. 8 (1874) 161; ScHINDL. Pfl.
R. Heft 23 (1905) 104; Back. & BAKH. f. FI.
Java 1 (1963) 266; MEIJDEN, Blumea 17 (1969) 307.
— M. humile (non MORONG) SCHINDL. Pfl. R.
Heft 23 (1905) 101, pro coll. ind.; GUILLAUMIN,
Fl. Gén. I.-C. 2 (1920) 718. — M. intermedium
(non DC.) Tarpreu-BLot, Fl. Laos, Camb. &
Vietn. 4 (1965) 126, for the cited BALANSA spe-
cimen only. — Fig. 12d-f.
Aquatic. Stem branched, freely floating. Im-
mersed /eaves in whorls of 4-5, 114-214 by c. 1 cm
in outline, patent to recurved, with 4~-7(-10)
pairs of filiform, rather weak, brown-tipped,
finely mucronate, 5-10mm long _ segments.
Lower aerial leaves verticillate or alternate, pin-
nate, the upper ones obovate-oblong to linear-
lanceolate, above the middle with 2-4(-7) pairs
of dark mucronate teeth, sometimes entire,
patent, upward erecto-patent, 10-25 by 1-3 mm.
Flowers solitary in the middle and upper leaf-
axils, or together with 1 or more reduced, cleisto-
gamous flower(s), %, on a shortly cylindrical, up
to 1% mm long pedicel. Bracteoles + elliptic,
with appressed, brown, serrate teeth, acute, with
1-3 nerves, 114-2 by 0.4-0.8 mm. Sepals + ovate
obtuse or acute, serrate, with a brown margin, erect,
0.2-0.5 by 0.2-0.5 mm. Petals patent to recurved
in anthesis, finally caducous, c. 2 mm long.
Stamens 4, rarely 8; anthers stiffly erect, linear
lanceolate, c. 1144 by 0.3 mm, with brown mu-
cronate apex. Ovary 2-celled, with 1(-2) ovules in
each cell, sometimes 3—4-celled with 1 ovule in
each cell. Styles 2(—4), 0.20.4 by 0.2 mm. Stigma
not or slightly recurved. Fruit c. 144 by 1 mm,
with 2, sometimes 3 or 4 mericarps, smooth or
indistinctly lengthwise lineolate, with a c. 2 by
0.4 mm long, erect or patent stalk. Cleistogamous
flowers: without perianth segments, ovary 2-
celled, + circular, laterally flattened, with 2
ovules, fruit c. 4% by 44 mm, with 2 elliptic seeds;
bracteoles ovate, c. 4 by 0.4 mm; pedicel up to
4 mm in fruit.
Distr. Australia (Northern Territory: Ro-
binson R.; Queensland: Mareeba R.), E. India
(Calcutta Yeels, T. THOMSON), N. Vietnam
(Haiphong, BALANSA 1428); in Malesia: NE. Java
(Madura I.: Burnih Lake). Fig. 13.
Ecol. In lake, at 15 m.
Note. Two specimens from SE. Asia have
erroneously been referred to the North American
M. humile MORONG, which seems more closely
allied to M. dicoccum than suggested by SCHIND-
LER, who places them in different subgenera. The
description and figure of ‘M. intermedium’ by
TARDIEU-BLoT fits M. oliganthum (W. & A.)
F. v. M., but the cited collection of BALANSA
belongs to M. dicoccum.
10. Myriophyllum siamense (CrRA1B) TARDIEU-BLOT,
Adansonia 5 (1965) 37, f. 1-4; Fl. Laos, Camb.
& Vietn. 4 (1965) 128, f. 1-4; MEDDEN, Blumea 17
(1969) 307, f. le; Fl. Thail. 2 (1970) 2. — M.
mezianum var. siamense CRAIB, Fl. Siam. En. 1
(1931) 591. — Fig. 15h.
Semi-terrestrial. Stem much branched, not
more than 5 cm high, rooting at the nodes.
Leaves alternate, rarely subopposite. Immersed
leaves not known. Aerial leaves entire or rarely
+ pinnate with 1-2 filiform segments, 2-4 by
0.2-0.4 mm, with a dark cushion-like thickening
above at apex. Flowers solitary in the upper leaf-
axils, the upper dg, the lower 9. Bracteoles in-
conspicuous or absent. — ¢ Flower nearly sessile.
Sepals 0. Petals 2, oblong to lanceolate, flat, en-
tire, acute, erect before anthesis and up to 1 mm
long, strongly recurved after anthesis. Stamen 1;
filament up to 14 mm; anther elliptic, 0.5-0.9
by 0.3-0.5 mm, mucronate. Rudiment of ovary
0. — 2 Flower sessile. Sepals and petals 0. Styles
shortly conical, distally set with long hairs. Fruit
0.5-0.9mm long. Mericarps 2, oblong, with
mostly 2 distinct dorsal crests of 5—7 tubercles over
the entire length and some rows of lower tubercles
laterally, with a rather indistinct crown of c. 7
short, +- spiny tubercles.
Distr. Vietnam (Prov. Than Hoa; Prov. Lang
Bian: Danhim; Prov. Phu Quoc); in Malesia:
S. Thailand (Nakhon Si Thammarat: Songkhla).
Fig. 13.
Ecol. In small mats on damp, sandy ground at
borders of ponds and marshes near the coast.
11. Myriophyllum coronatum Meupen, Blumea 17
(1969) 305, f. la-c. — Fig. 15a-c.
Terrestrial or semi-aquatic. Stem much bran-
ched. Leaves opposite, the upper ones semi-
opposite to alternate; immersed leaves opposite,
pinnate, 0.5-1.2 by 0.6-1.5 mm in outline, with
2-5 c. 0.1 mm wide, filiform lobes; aerial leaves li-
near, entire or less often + pinnate, 3-10 mm long,
the entire ones 0.3-1.2 mm wide, with a dark
258
FLORA MALESIANA
[ser. I, vol) 7
cushion-like thickening above at apex, flat, patent.
Flowers solitary in the upper leaf-axils, the upper 3,
the lower 2. Bracteoles inconspicuous or 0. —
3 Flowers nearly sessile. Sepals 0. Petals 2, linear,
flat, entire, acute, erect before anthesis and up
to 1 mm long, strongly recurvate after anthesis.
Stamen 1; filament up to 14 mm; anther elliptic,
OFF
Fig. 15. Myriophyllum coronatum ME\SDEN. a. Habit, <7, b. ¢ flower, x25, c. fruit, x25. — M. bonii
TARDIEU-BLOT. d. Habit, <7, e. fruit and ¢ flower, x25, f. cross-section of fruit, x 25. — M. mezianum
SCHINDL. g. Fruit. — M. siamense (CRAIB) TARDIEU-BLOT. A. Fruit, «25 (a—-c RIDSDALE NGF 33585,
d-f Bon 5897, g PERRIER DE LA BATHIE 19310, i BON 5290).
0.2-0.6 by 0.1-0.4 mm, mucronate. Rudiment of
ovary 0. — 2 Flowers sessile. Sepals and petals 0.
Styles shortly conical, distally set with long hairs.
Fruit c. 1 mm long. Mericarps 2, + oblong, at the
base with c. 3 dorsal crests, these with 3—5 thin
long spines curved upwards, and laterally with
some irregular rows of tubercles, upwards slightly
1971] HALORAGACEAE (van der Meijden & Caspers) 259
narrowing, smooth or with some _ tubercles, \\y \\\\ || Wil JAZ ws
widening at the top in an asymmetrical crown of \ J] \ | i} Ll YL
c. 8-12 + patent spines with recurvate tips, which \ rN fal J
are dorsally distinctly longer than laterally, the NX} Ni NGS 4 ap Vez
longest up to 0.8 mm long. Ay \) We Ye fy
: ‘ \\ eR VP Ry | Ay
Distr. Malesia: E. New Guinea (Papua, YA \ A AY iy
Western Distr.: near Weam). Fig. 10.
Ecol. Wet depression in savannah, c. 30m.
Doubtful record
Myriophyllum indicum WILLD.: JUNGH. Java, 2nd
Dutch ed., 1 (1853) 470, ‘Java (Bandung, abun-
dant)’.
No material could be traced; almost certainly a
mis-identification.
Dubious
Ammania pinnatifida LINNE f. Suppl. (1781) 127,
‘Java’.
The name is cited by DC. Prod. 3 (1828) 69, as
possibly synonymous with Myriophyllum_ in-
termedium DC. No material could be found in
Swedish herbaria. See MEIJDEN, Blumea 17
(1969) 311.
Myriophyllum pusillum BL. Mus. Bot. Lugd.-Bat.
1 (1849) 111; Mra. FI. Ind. Bat. 1, 1 (1855) 634;
Suppl. (1860) 328; Sumatra (1862) 128, 328;
Kurz, Nat. Tijd. N. I. 27 (1864) 167; Wars.
Bot. Jahrb. 13 (1891) 395; K. Scu. & Laut. FI.
Schutzgeb. (1901) 483.
Although the description is very brief, the said
provenance said to be ‘Archip. Ind.’, it is not im-
probable that M. pusillum is indeed a Myrio-
phyllum, possibly M. tetrandrum Roxs. Un-
fortunately the type material could even as early
OF OF I
as 1900 not be located. All later records refer to
sterile specimens, those by Miquet (1860) and
Kurz are according to Mr PHILcox (in litt.)
Fig. 16. Myriophyllum axilliflorum BAKER. a.
Habit, <3, b. bisexual flower, 7, c. fruit, x13,
d. cross-section of fruit, x13 (a-d HILDEBRANDT
probably Limnophila (Scrophulariaceae). 4030).
4. GUNNERA
Linng, Syst. Nat. ed. 12 (1767) 597; Mant. 1 (1767) 16; SCHINDL. Pfl. R. Heft. 23
(1905) 104; MATTFELD, Ostenia (1933) 102; BADER, Bot. Jahrb. 80 (1961) 281,
maps. — Pseudogunnera Orstep, Naturh. Fér. Vidensk. Medd. Kjébenhavn
(1857) 193. — Fig. 17-18.
Perennial, + acaulous (sometimes colossal herbs with a stem up to 6m by
20cm @ in extra-Mal. spp.), (in Mal.) stoloniferous, with more or less creeping,
sub- or slightly supraterranean, thick rhizome. Leaves radical (or in non-Mal.
spp. tufted at the end of the stem), reniform-cordate to ovate-truncate, simple to
lobed, crenate to compound-dentate, rarely entire, palminerved, often rugose,
with more or less conspicuous, simple to much divided, sometimes ochrea-like
‘cataphylls’ in the very leaf-axil. Panicle, raceme or spike mostly 1, axillary or
pseudoterminal, simple or compound. Flowers ebracteolate, bisexual or unisexual,
monoecious (or dioecious), if monoecious with ¢ flowers upwards and 9 ones
260 FLORA MALESIANA
(ser. I, volxywt
Fig. 17. Gunnera macrophylla BL. on summit of Mt Kaba, Bencoolen, April 1932 (DE Voocp).
downwards and sometimes with % in between. Sepals 2 (rarely in some flowers 3),
or 0, equal or unequal, often cuspidate, whether or not caducous. Petals 2(-0),
in 2 0, oblong to spathulate, more or less convex outside, glabrous to pilose,
often soon caducous, longer than the sepals. Stamens 2; anthers elliptic to oblong.
Ovary 1-celled, urceolate to compressed-ovoid; styles subulate to filiform, entirely
papillose, exserted in anthesis. Ovule 1. Fruit a drupe or nut-like, coriaceous to
more or less fleshy or juicy, almost globular to 3-gonous; stone crustaceous;
perisperm copious, oily; embryo apical, very small, pear-shaped to obcordate.
Distr. About 30-50 spp., in South and tropical Africa and Madagascar, Malesia, Tasmania, New
Zealand, Antarctica, the Hawaiian Islands, and South and Central America; not in continental Asia
and Australia; in Malesia: not found in the Malay Peninsula and the Lesser Sunda Islands. Fig. 19.
Ecol. Cold or cool, everwet localities, often on marshy soil, in the tropics only in the mountains, in
Malesia above (750—)1000 m.
1971] HALORAGACEAE (van der Meijden & Caspers) 261
Symbiosis with Cyanophyceae. — Gunnera represents a remarkable case of true symbiosis which seems
inherent to all species. Though I am not aware of experiments proving it to be obligatory, this seems
likely. The stem and petiole-bases are provided with mucilage producing glands which offer obviously
the entrance of the Nostoc or Chroococcus species, or hydathodes may serve this purpose and the algal
cells form fairly large and regular intracellular colonies, as REINKE described (Unters. Morph. Veg.-
Organe Gunnera, 1873). See also H. SCHNEGG (Flora 90, 1902, 161—208). Infection seems to take place
very early. For the Malesian G. macrophylla excellent details are described and depicted by BAAS BECKING
(Dodonaea 14, 1947, 93-96). Each of the equitant petiole-bases is provided with three warts, one central
highest and two lateral basal, immediately below which a root emerges; the Nostoc colonies are confined
to the warts.
BAAS BECKING suggested that the phycome contained a substance of ‘auxin’ character, but no later
communication followed. Recently, in a preliminary paper, SILVESTER & SMITH (Nature 224, 1969, 1231)
found that Nostoc in cultures is able to fix nitrogen and that the Gunnera-Nostoc symbiosis is capable of
nitrogen fixation. It is made certain that the nitrogen absorbed by the Gunnera (arenaria) plants in pure
cultures on nitrogen-free media came from Nostoc. Gunnera plants raised from seed and grown on media
free of both Nostoc and nitrogen compounds showed at the 6 leaf stage yellow chlorotic leaves and poor
growth. No experiments were yet made to grow Nostoc-free Gunnera on media containing combined
nitrogen to check whether the symbiosis is obligarory for Gunnera; SILVESTER (in litt.) suspects
perhaps it is not. As Nostoc is ubiquist, all Gunnera plants in nature contain Nostoc and this may be fa-
vourable indeed for its growth on nitrogen deficient soils.
Morph. The evaluation of the so-called ‘ligule’ or ‘stipules’ is still under dispute. Their development
is various but specific; in G. magellanica they are very large, in the New Zealand species they are absent
(SCHNEGG, /.c.). SKOTTSBERG (Svensk Bot. Tidskr. 22, 1928, 392-415) regarded them as cataphylls. The
latter opinion seems unlikely as they are not contained in the leaf spiral as remarked by MATTFELD (Os-
tenia 1933, 109), but axillary.
Taxon. SCHINDLER subdivided the genus into 5 subgenera, to which MATTFELD added a sixth mono-
typic one from E. South America, Ostenigunnera MATTFELD (1933, l.c.) which is very distinct and deserves
clearly subgeneric rank. The others are Misandra in South America, Panke in South America, Juan
Fernandez, and Hawaii (20-30 spp.), Milligania in New Zealand and Tasmania (9 spp.), Pseudogunnera
in Malesia (1 sp.), and Gunnera (subg. Perpensum) in Madagascar and East Africa (1 sp.).
The characters SCHINDLER used for this classification are the occurrence of stolons, the distribution of
the sexes, the structure and size of the ‘cataphylls’, and the size of the plant. It seems to me that, Os-
tenigunnera excepted, the mutual similarity is so large that distinguishable taxa at most deserve sectional
rank, the more so as distinctions are less sharp than presented by SCHINDLER. For example: within
Milligania occur both species with monoecious and with monoecious or dioecious flowers. In Perpensum
with bisexual flowers sometimes the upper are 3, whereas in Pseudogunnera between the upper 3 and
lower 2 sometimes & flowers are found. As to size of leaves Pseudogunnera (G. macrophylla) displays an
enormous variability in New Guinea, from 2!4—70 cm, depending either on altitude or habitat, so that
small specimens of G. macrophylla from New Guinea so much resemble Milligania that RIDLEY, who
described such small specimens as a separate species, G. reniformis, promptly declared them to be allied
to the Tasmanian G. cordifolia HooK. f. of subg. Milligania.
I believe that the subdivision of Gunnera could be improved if only 2 subgenera are recognized:
subg. Gunnera and subg. Ostenigunnera. The first might then be distinguished into two or three sections,
viz sect. Gunnera, including subg. Perpensum, Pseudogunnera, and Milligania, and occurring in Africa,
Madagscar, Malesia, Tasmania, and New Zealand, and sect. Panke, including subg. Panke, occurring in
Central and South America, Juan Fernandez, and the Hawaiian Is. The affinity of the South American
subg. Misandra is uncertain to me, it may be kept separate, or may be included within sect.
Gunnera.
Nomencl. The name Panke which is still used as the name of a South American subgenus of Gunnera
already occurs in a work by L. FeurLiée (Journ. Obs. 1, 1714, t. 30), as part of a phrase name. This was
post-Linnean literatim translated into German by G. L. Hutu (Beschreib. Arnzey Pfl. 1, Niirnberg,
1756, 42, t. 30). In this rare work all plants are beautifully depicted and accompanied by phrase names
preceding their description. There is of course no question that FeurLLfe adhered to the binomial system,
i.e. that he distinguished genera and species. The German literatim translation does neither, and from the
introduction by HuTH appears that he merely published it as a translation, without himself having the
intention to make any changes towards framing it according to the binomal nomenclature. Therefore,
this translation has no nomenclature standing.
It must be admitted, though, that not long ago a ‘generic’ name from this work was rejected nomen-
claturally, viz Urceolaria, obviously because D. L. DeNHAM (Taxon 10, 1961, 247) stated in his proposal
that Hutu’s names are effectively and validly published and ‘must’ be accepted. The members of the
.Committee on nomina generica conservanda have obviously not verified DeNHAm’s statement on this
rare work and were thus misled. In Hutn’s translation neither genera nor species were indicated as such;
each single species had a different phrase name, or better, was indicated by a phrase indication. Nomen-
claturally this translation cannot be used for post-Linnean validation and should be discarded. Con-
sequently we have refrained from proposing Panke to be rejected in favour of Gunnera.
eg J ide ‘
te». =
" Ma ae od
Bs ee ie ee
hE nt Bt Ce
“a ek Fe OF aKa
+
FLORA MALESIANA
(ser. I, vol, 7?
Fig. 18. Gunnera macrophylla BL. group above waterfall Tjibeureum, Mt Gedeh, W. Java, 1750m
(VAN STEENIS).
1. Gunnera macrophylla BL. Bijdr. (1826) 513;
BENN. Pl. Jav. Rar. (1828) 70, t. 15; Mia. PI.
Jungh. (1851) 70; Fl. Ind. Bat. 1, 1 (1856) 769,
incl. var. sumatrana Mia. I.c. 770; BL. Mus. Bot.
Lugd.-Bat. 2 (1856) 101; DC. Prod. 16, 2 (1868)
598; WARB. Bot. Jahrb. 16 (1893) 15, 24, var.
papuana WARB.; Koorp. Minah. (1898) 451;
K. Sco. & Laur. FI. Schutzgeb. (1901) 483;
SCHINDL. Pfi. R. Heft 23 (1905) 114, f. 33; BAcK.
Schoolfl. Java (1911) 477; Ripv. J. Fed. Mal. St.
Mus. 8, 4 (1917) 33; Herne, Nutt. Pl. ed. 3 (1927)
1207; C. T. Wuite, Proc. R. Soc. Queens]. 34
(1922) 483; Koorp. FI. Tjibodas 2 (1923) 217;
BAKER, J. Bot. (1924) Suppl. 35; Went, Nova
Guinea 14 (1924) 105; JocHEMs, Trop. Natuur
15 (1926) 68, f. 4; DE Vooap, ibid. 22 (1933) 226;
Exum. Leafi. Philip. Bot. 9 (1933) 3143; STEEN.
Bull. Jard. Bot. Btzg III, 13 (1934) 217; Trop.
Natuur 30 (1941) 172, f. 4; Bull. Jard. Bot. Btzg
III, 17 (1948) 463; Back. & BAKu. f. Fl. Java 1
(1963) 266. — G. erosa BL. Verh. Bat. Gen. 10
(1825) 81, nom. nud.; DC. Prod. 16, 2 (1868) 599.
— Pseudogunnera macrophylla (BL.) ORSTED,
Naturh. Fér. Vidensk. Medd. Kjébenhavn (1857)
599. — Sarcospermum petasites REINW. MS.,
in De Vriese, Reinw. Reise (1858) 576, nomen. —
G. reniformis RipL. Trans. Linn. Soc. Lond. II,
9 (1916) 40, ex descr.; WENT, Nova Guinea 14
(1924) 105; STEEN. Bull. Jard. Bot. Btzg UI,
13 (1934) 217. — Fig. 17-18.
Perennial, stoloniferous, pilose herb. Stem very
short, c. 12cm g, pilose; rhizome sublignose,
elongated, dark. Stolons often numerous, +
terete, the internodes very long, up to c. 40 cm,
c. 3mm g, with 2 opposite, caducous, small bud-
scales, soon producing a new flowering plant at
each node. Leaves reniform to cordate, angular,
rarely irregularly (2—)3-lobed, acutely irregularly
sphacelate-dentate, bullate, venation strongly pro-
minent, reticulate beneath, 214-70 by 214-70 cm;
1971]
petiole up to 70 cm, 3-10 mm g, + terete, widely
channelled above, costate, -+ sheath-like at
base, shortly decurrent, each of the lcwer equitant
petiole-bases provided with 3 warts, 1 central
highest and 2 lateral lower, below which an
adventitious root emerges. Panicles 1(-3) sub-
terminal, shortly peduncled or sessile, often hidden
under the leaves, in fruit accrescent to 25-60 cm;
lateral spikes very numerous, each in the axil ofa
bract 44-144 by %4-1 cm, the lower ones 214-9
cm, with 2 flowers, the upper ones with ¢ or in-
termixed with § flowers. — ¢ Flowers: sepals
HALORAGACEAE (van der Meijden & Caspers)
263
triangular to broadly triangular, with acuminate or
cuspidate, sphacelate apex, erect, 0.6—-1.2 by c. 1
mm, glabrous; petals spathulate, mucronate,
glabrous or pilose outside, glabrous inside, +
densely pilose on the margin, caducous before
anthesis; filaments up to c. 2 mm, c. 0.3 mm @,
erecto-patent, stiffish; anthers +- elliptic, obtuse or
shortly mucronate, 114-2 by 1144-144 mm. —
° Flowers: sepals as in 3; petals 0; stigmas sessile,
subulate, up to 2 mm long. Drupe + globose,
glabrous, juicy, c. 2 mm; stone flattened globular,
c. 1 mmo.
Fig. 19. Distribution of Gunnera macrophylla BL.
Distr. Malesia: Sumatra, W. & Central Java,
N. Borneo (Mt Kinabalu), Philippines (Luzon,
Negros), NE. Celebes (Mt Klabat), Sangihe L.,
New Guinea (incl. also Goodenough I. & New
Britain: Mt Talawe), and Solomon Is. (Kolom-
bangara). Fig. 19.
Ecol. In the mountains from (750—)1250-3000
m. Wet or damp places, along brooks, river banks,
near waterfalls, wells and seepage, mostly on open
or lightly shaded places in disturbed habitats,
pioneering on bare lands, along paths and in light
places in secondary forests, often gregarious.
Vern. Tératé gunung, S; Philippines: balai,
baloi, Ig., Bon.
Morph. The so-called sphacelate teeth on the
leaves and the inflorescence-scales are probably
hydathodes.
Numerous semiglobular gland-like outgrowths
are found especially on the young leaves, on and
near the nerves.
Coalescence of parts of the inflorescence occurs
frequently.
Notes. The small specimen described from Mt
Carstensz in New Guinea as G. reniformis RIDL.
is presumably an early flowering juvenile in-
dividual developed on a stolon; such specimens
have also been collected elsewhere in New Guinea.
Sterile specimens have sometimes erroneously
been referred to Petasites (Compositae) by re-
semblance of the leaf (VAN STEENIS, 1948, /.c.).
Excluded
Hydrospondylus (submersus) Hassk. Flora 25
(1842) Beibl. 2, 33 = Hydrilla verticillata (L. f.)
RoyLe (Hydrocharitaceae).
FLORA MALESIANA
under the Auspices of
Lembaga Biologi Nasional (Botanic Gardens of
Indonesia), Bogor, Indonesia,
and the Rijksherbarium, Leyden, Holland,
executed by Foundation Flora Malesiana
Scientific Communications
concerning the Flora Malesiana should be addressed to
the General Editor, Dr. C. G. G. J. VAN STEENIS
SUBSCRIPTION REQUEST AND RELATED CORRESPONDENCE
SHOULD BE DIRECTED EXCLUSIVELY TO
WOLTERS-NOORDHOFF PUBLISHING
P.O. Box 58, Groningen,{The Netherlands
For sale only - Not for exchange
Series I - Spermatophyta (Flowering Plants)
Volume 1. Cyclopaedia of collectors & collections. 1950. pp. clii + 639
Volume 2. Malesian vegetation. _ (In preparation)
Volume 3. Malesian plant geography (In preparation)
Volume 4. General chapters and revisions. 1948-1954. pp. ccix + 631
Volume 5. Bibliography, specific delimitation, & revisions. 1955-1958. pp.
cccxlii + 596
7
Volume 6. Systematic revisions. 5 parts, 1960-1967.
pp. 914 (final part in press).
ANI
WOLTERS-NOORDHOFF PUBLISHING, GRONINGEN, THE NETHERLANDS
PRINTED IN THE NETHERLANDS
ISBN 90 01 31813 4
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