Flora Malesiana

FLORA
MALESIANA

SERIES I- SPERMATOPHYTA
Flowering Plants

Vol. 8, part 2

Revisions

INDEX TO REVISED FAMILIES

Aceraceae. . . 4: 3, 592
Actinidiaceae s. str. . .4: 37
Aizoacese: 5 574. 5 sais 8; 207
Alismataceae Be Sh 62915
Amaranthaceae
4: 69, 593; 6: 915
Anacardiaceae . . In press
Ancistrocladaceae . . .4: 8
Aponogetonaceae 4: 11; 7: 213
Balanophoraceae. . . . 7: 783
Basellaceae .... . .5: 300
Batidacede ... .-. .5: 414
Betulaceae Se 207 2.62917
Bignoniaceae Peart rope
Bixaceae’sstrs | 222... ../45239
Burmanniaceae A 132592
Burseraceae
5: 209, 567; 6: 917; 7: 820
Butomaceae :43.5.°-' .. - 52118
Byplaaceaeioen es ds
@allitrichaceae’. . . . . 4: 251
Campanulaceae 6: 107, 928
Cannabinaceae .. . .4: 223
Capparidaceae. .... 6: ‘61

Caprifoliaceae
4: 175, 598; 6: 928

Cardiopteridaceae . . .7: 93
Celastraceae 6: 227, 389, 930
Centrolepidaceae. . . .5: 421

Ceratophyllaceae. . . .4: 41
Chenopodiaceae

4: 99, 594: 6: 932

Glethracedes =. 4. +4. TRIBE)
Cochlospermaceae . . .4: 6]
Combretaceae

4: 533; 5: 564; 6: 932

Connaraceae 5: 495; 6: 933

Convolvulaceae 4: 388, 599;

5: 558; 6: 936; 7: 823

@omaceae:) of, 24.’ 8: 85
Corynocarpaceae

4: 262; 5: 557

Crassulaceae ..... 4: 197

Crypteroniaceae . . . .8:187

WRPETAGCAG Wh)... e 7: 435

Datiscaceaes ogc. 4: 382

Dichapetalaceae 5: 305; 6: 941
Dilleniaceae . . 4: 141; 7: 824
Dioscoreaceae. ... . 4: 293

Dipsacaceae.. t+ ane . 4: 290
Droseraceae. . 4:.3/72)52/557
Elatinaceae: 2 — See 4: 203
Epacridaceae .«.)... . 62422
Ericaceae . . . . 6: 469, 943
Erythroxylaceae . . . .5: 543
Fagaceae hs as taet = perce
Ficoidaceae =. 2 -3/)..7)-: 4: 267

Flacourtiaceae
5: 1, 565; 6: 943; 7: 827

Flagellariaceae. . .. . 4: 245
Geraniaceae.. ... . 6: 445
Gnetaceae. . . 4: 336; 6: 944
Gonystylaceae. .... 4: 349

Goodeniaceae
5: 335, 567; 6: 949; 7: 827

Haemodoraceae . . . .5: 111
Haloragaceae . 2... 7: 239
Hamamelidaceae. . . .5: 363
Hippocrateaceae . . . . 6: 389
Hydrocaryaceae ... .4: 43
Hydrocharitaceae

5: 381; 6: 952
Hydrophyllaceae . . 4: 207
Hypericaceae aeons oe. $2 1
Icacinaceae 7 e414 eee
Inidaceae wane wee es hae,
Juglandaccae eas sae 6: 143
JUUCACEde 2. et eee 4: 210
Juncapinacedelie. + es). 4: 57
Labiatae Sho scott TD DYESS
Recdceten ais eink DOES,
emnacedey fy. "aise. 2; 2A9
Lentibulariaceae . SPA
Loganiaceae. . 6: 293, 953
Lophopyxidaceae FCI OD
Malpighiaceae. . ... Sen 25
Martyniaceae ..... 4: 216
Molluginaceae. ... . 4: 267
Moringacede 20 acute 4: 45
Myoporaceae ..... 4: 265
Mryricaceaer 2) 8225. 4: 277
INajadaceae eS 6: 157
Nyctaginaceae. .... 6: 450
INyssacede>- | a ack ee. 4: 29
Ochnacedey sare Me
Onaeracedey ko st dee O8
Oxalidareal> hss 53 A. IAS
Papaveraceae. 8 te 5: 114

LIBRARY

Passifioraceae . .\..%, - 7: 405
heaanacede.. «36 a: 4: 216
Pentaphragmataceae . . 4: 517
Pentaphylacaceae 531s
Philydraceae. . ... . ; 4: 5
Phytolaccaceae 4: 229
Pittosporacese’. 2 3. 5. | 5: 345
Plumbaginaceae . . . . 4: 107
Podostemaceae

4: 65; 6: 963

Polemoniaceae. . 4: 195
Pontederiaceae. . 4: 255
Portulacaceae ..... 72 YZ
Primiulacedé=- . .. 2.) 6073
Protedceden! <i) = Fie 5: 147
Punicaceaey 3 i: 4: 226
Restionaceae :..... 5: 416

Rhizophoraceae 5: 429; 6: 965

SANIGACEAE eet et here 5: 107
Salvadoraceae ..... 4: 225
Sarcospermaceae. . . .4: 32
Saururaceae,. . 2% : . 4: 47

Scyphostegiaceae

5: 297; 6: 967
Simaroubaceae 6: 193, 968
Sonneratiaceae

4: 280, 513; 6: 973

Sparganiaceae..... 4: 233
Sphenocleaceae 4: 227
Stackhousiaceae . . 4:35
Staphyleaceae ..... 2°. 6: 49
Siviidiacede 5 sni. se 4: 529
Siyracacede./5 (4 fa au) 4: 49
Symplocaceae ..... 8: 205
iBaccdceae i>. Gis cei 7: 806
Thymelaeaceae

4: 349; 6: 1, 976; 7: 830
Mrapacese <3. a a 4: 43
Mnvonmiaceae | .7.-. ave 4: 59
AIRTHEACeEAG 26. ar een 4: 235
MiypHacede 4. . S25 tame 4: 243
Wimacede 253. ie owe S$: 34
Umbelliferae 4: 113, 595;

5: 555: 6: 983: 7: 830

Valenianaceae 0. 5c 4: 253
Violaceae . 7: 179, 831
Xyridaceae 4: 366, 598
Zygophyllaceae ie

a

NICAL GARDEN
BRONX, NEW YORK 10458

—As

-
- «

ULMACEAE (E. Soepadmo, Kuala Lumpur) .
Monoecious or dioecious (?), evergreen, deciduous or semideciduous shrubs or
trees, (in Mal.) unarmed and often buttressed. Growth habit (in Mal.) flush-wise,
except for Trema and Parasponia. Indumentum of simple, bulbous-based unicellular
hairs and/or multicellular glandular hairs. Stipules caducous or rarely rather long
persistent and completely enclosing the bud, extrapetiolar or intrapetiolar, basally
attached or rarely peltately attached to the nodes, free or connate. Leaves simple,
(in Mal.) alternately arranged, petioled, pinnately nerved or triplinerved at base,
often asymmetrical at base, entire or variously serrate. Inflorescences |-many-
flowered, 3, 2, S2, or $Y, axillary, subterminal, or borne on leafless, older branch-
lets or on short, lateral, leafless new shoots, paniculate, racemose, thyrsoid, cymoid,
or rarely capitate, bracteate; bracts minute, caducous. Flowers functionally 3, 2, or
8. — 3 Flowers solitary or in condensed cymoid clusters along the rachis, sessile
or short-pedicelled; perianth (4~)5(—7)-lobed, lobes free or variously connate,
imbricate or induplicate-valvate in bud; stamens as many as tepals, attached to the
base of and opposite the perianth lobes, straight or incurved in bud; anthers ovoid,
ellipsoid or subreniform, apiculate or non-apiculate, initially tetrasporangiate,
later becoming 2-celled, dehiscing lengthwise, introrse or extrorse; pistillode
present or absent, if present either rather well developed or rudimentary, densely
whitish to silvery, soft or hirsute pubescent. — 2 & & Flowers sessile or stalked,
solitary in the axils of the upper new leaves or arranged in various types of in-
florescences ; perianth herbaceous or thin-coriaceous, (4-)5(—7)-lobed, lobes always
imbricate in bud and connate at base, (in Mal.) long persistent; staminodes or
stamens as many as perianth lobes or absent; ovary superior, 2-carpellate, (in
Mal.) 1-celled, sessile or stipitate; style 1, tubular, short or absent, stigmatic arms
2, slender, often bifid to deeply lobed at the tip, adaxially papillose-stigmatic for
their entire length; ovule 1, anatropous to hemi-anatropous, subapical, pendulous,
bitegmic. Fruit a drupe or a samara, faintly angular or flat and winged. Seed
mostly exalbuminous; embryo large, straight or curved; cotyledons flat-convex,
fleshy, straight or variously folded, often foliaceous. Germination mostly epigeal.

Distribution. There are 15 genera, c. 200 spp., widely distributed in the tropics, subtropics,
and temperate regions of Europe (as far north as 70°, Scandinavia), Africa (South of Sahara),
continental Asia, Malesia, Australia (Queensland and New South Wales), Pacific Islands (as far
as Tahiti; incl. also Hawaii and the Galapagos Is.), North, Central, and South America (as far
south as 40°, Argentina). Fig. 1.

Fossils. Various macrofossils (woods, drupes, and leaf-impressions) and microfossils (pollen
grains) attributed to Ulmaceae have been reported by different authors from various localities in
Alaska, North America, Europe, continental Asia, and Japan. Amongst the older records are
pollen grains of the Ulmus-Zelkova-type from Golden Valley Formation in North Dakota and
Rocky Mountains, U.S.A. (Paleocene) and wood and leaf-impressions attributed to U/mus from
Rocky Mountains and to Planera from Alaska (Late Paleocene). By the middle of the Eocene and
throughout the Miocene and Pliocene fossils of Ul/maceae become more abundant (common) in
the northern hemisphere, especially in Europe and North America. However, very little is known
about the geological history of the family in Asia.

Of the Malesian genera belonging to the tribe Ce/toideae, fossil records of Gironniera (identifi-
cation very doubtful) go back to Middle Eocene (Alaska), that of Ce/tis to Miocene (Japan), and
Trema to Upper Oligocene (Japan). — Literature: Berry, Tree Ancestor (1923) 146; WATARI,
Jap. J. Bot. 11 (1941) 385; J. Fac. Sc. Un. Tokyo III, 6 (1952) 97; La Morte, Mem. Geol. Soc.
Am. 51 (1952) 112, 260, 346, 360; PrakAsH & BARGHOORN, J. Arn. Arb. 42 (1961) 185, 347;
Greouss, Tert. Angios. Hung., Ak. Kiado Budapest (1969) 83; Fercuson, Verh. Kon. Ned.

(31)

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32 FLORA MALESIANA [ser. I, vol. 8?

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Fig. 1. Approximate range of Ulmaceae.

Ak. Wet. sect. II, 60 (1971) 103; LEopotp & MacGiniTiE in Graham, Flor. & Paleofl. Asia &
N.E. America (1972) 147; WoLFE, /.c. 200; TANAI, /.c. 235; WOLFE, Brittonia 25 (1973) 334.

Ecology. In Malesia species of the Ulmaceae may be found in both primary and secondary
forests, from sea-level up to 2000 m; they are more common in the lowlands and hills.

Apart from Ulmus lanceaefolia, Celtis rigescens, Gironniera nervosa and G. subaequalis which
may attain up to 45 m in height and 100 cm in diameter, all species are understorey shrubs or
trees. Of the 6 genera occurring in Malesia, species of Aphananthe, Celtis, Gironniera and Ulmus
are basically primary forests inhabitants, though they may survive and thrive in secondary
forests as well. Of these, Ce/tis and Gironniera species are the most widely spread and may be
found growing on various types of soils, including those derived from limestone.

In Malesia Ulmaceae are found under both everwet and more seasonal climatic conditions,
but some show preference for one or the other. In Celtis two groups of species may be dis-
tinguished, viz the group of C. tetrandra, timorensis and rubrovenia, and that of C. philippensis.
The former is either semideciduous or deciduous and found mainly under a drier and more
seasonal climate, while the latter is evergreen and grows better in more humid environmental
conditions. Aphananthe and Ulmus species are mainly found in regions subject to a rather distinct
seasonal climate, and they are either semideciduous or deciduous. Members of the genera
Parasponia and Trema are pioneer plants preferring and thriving well in newly opened up
habitats, e.g. forest clearings, thickets, roadsides, flood-plains, on volcanic ashes, efc. Fig. 10, 11,
17. They are usually short-lived (at least in Malaya, 5-7 years) and soon will die out, particularly
when over-grown by the other more aggressive and long-lived pioneer plants, e.g. species of
Macaranga, Mallotus, and Grewia, etc. For this reason species of Trema and perhaps also those
of Parasponia are usually not or rarely found in old secondary forest. It also may be noted here
that most Trema species grow, thrive, and are more widely spread in the western parts of Malesia,
while Parasponias are more common in the eastern parts, especially in New Guinea.

The structure and position of the inflorescence and flowers, particularly the amount of pollen
grains produced and the structure of the stigmas, and also the absence of nectary, seem to suggest
that pollination is most likely affected by wind, though insects may not be ruled out altogether as
possible agents for pollination.

Except for Ulmus, which produces a dry, flat, winged fruit, the other Malesian genera have
various types of fleshy drupes which turn to bright yellow, orange, or deep-red in colour when
ripe. These drupes are most probably dispersed by various species of frugivorous birds or arbo-
reous mammals. Alternatively, at least in some species, e.g. Celtis philippensis var. wightii which
is very common in coastal vegetation, fruit dispersal may be carried out by water currents. In
Ulmus the winged fruits are easily dispersed by wind.

1977] ULMACEAE (Soepadmo) 33

It should be noted here that there is a very high percentage of seed abortion in Malesian genera
for reasons unknown. This is made good by the production of a great number of flowers and
fruits, produced regularly throughout the year or at least twice a year. Except for U/mus, the
embryo is protected by a strong, hard and durable endocarp. In all genera endosperm is usually
scanty or absent.

Anatomy. For general surveys also covering the older literature see SOLEREDER, Syst. Anat.
Dicot. Stuttgart (1899) 860-865 and ibid. (1908) 295; METCALFE & CHALK, Anat. Dicot. Oxford
(1950) 1271-1278; Sweirzer, J. Arn. Arb. 52 (1971) 523-585. Additional selected references:
Denay, l’Appareil conducteur foliaire des Urticacées, des Moracées et des Ulmacées (Ordre des
Urticales), Arras (1934); JANssonius, Mikr. 6, Leiden (1934) 1-308 (wood anatomy, under Urti-
caceae); LEROY, Bull. Mus. Nat. Hist. Nat. Paris sér. 2, 18 (1946) 118-123 & 180-184 (taxonomy,
and anatomy of Aphananthe); DEN BERGER, Determinatietabel Malesié, Veenman, Wageningen
(1949) (wood identification); RAo, GovINDU & THIRUMALACHAR, J. Indian Bot. Soc. 29 (1950)
224-226 (aerial roots, Trema) ; JANSSONIUS, Blumea 6 (1950) 407-464 (wood anatomical affinities) ;
Descu, Mal. For. Rec. 15 (1954) 618-620 (wood); JutrE, Nova Guinea n.s. 10 (1959) 241-278
(wood); MosELEY, Brittonia 25 (1973) 356-379 (anatomy and relationships).

SWEITZER’s study (/.c.) is the most up-to-date survey.of leaf and wood anatomy of the Ulma-
ceae. Although his extensive research materials included very few Malesian species his general
conclusions are probably largely applicable to the Malesian species as well.

The wood anatomy is indicative of the mutual affinities of all Ulmaceous genera. Shared
characters are: predominantly simple vessel perforations, short vessel members, alternate
intervessel pits. Fibres with simple to slightly bordered slit-like pits. Parenchyma at least partly
vasicentric. Genera of the tribe U/meae (in Malesia only represented by U/mus lanceaefolia) have
exclusively homocellular rays. In Ce/teae (in Malesia all other genera) at least part of the ray
tissue is heterocellular.

The wood of U/mus lanceaefolia differs from all species described in literature in lacking the
ring porosity and the typical ulmiform arrangement of vessel clusters (original observation).
Instead, its vessel distribution resembles the diffuse porous group of tropical Celtis species. In
Celtis the very striking differences in vessel distribution between tropical and extratropical
species are well documented (cf. Sweitzer, /.c.). Although from SwEITZErR’s and other publica-
tions some quantitative and qualitative differences between Malesian genera of Celteae can be
deduced, our knowledge is still based on too limited materials to allow conclusions on diagnostic
and systematic implications.

The leaf anatomy of U/maceae at the same time supports its coherence as a family and provides
an interesting diversity, of great potential diagnostic and systematic value. All U/maceae share
the dorsiventral leaf architecture. The stomata are confined to the abaxial epidermis and are of
the anomocytic type. The indumentum includes bulbous-based unicellular trichomes the walls of
which are usually silicified. Mineral inclusions of calcium carbonate or silica in cystoliths (with or
without pegs) are of common occurrence. The trichome-complement, presence or absence of
mucilage cells, crystal complement, loose or compact structure of the spongy tissue, petiole and
midrib vasculature show a considerable diversity. SWEITZER’s data and other reports from the
literature do not yet allow a leaf anatomical characterization of the individual Malesian taxa,
but preliminary studies are indicative that this will be possible if more material is studied.

The entire evidence from vegetative anatomy supports the traditional placement of U/maceae
in Urticales. — P. BAAs.

Palynology. Based on size, sculpturing of exine and number of pores, pollen grains of
Ulmaceae may be divided into two major morphological types, namely the U/mus-type and the
Celtis-type. In the Ulmus-type the pollen are oblate to subspherical, amb convex or straight;
(4-)5(-7)-porate, 20-30 by (26—)28-38(-—51) um, pore circular to elliptic, c. 2-3 by 3-4 um,
slightly thickened around its margin; exine rugulate-reticulate. Genera with this type of pollen
grains are: Ampelocera, Hemiptelea, Holoptelea, Phyllostylon, Planera, Ulmus, and Zelkova. In
the Celtis-type the pollen is suboblate to spherical, amb convex; (2—)3—4(—5)-porate, pores
circular or elliptic (elongated towards the poles), often annular and protruding, c. 2-3 by
3-4 um; 17-25 by 19-30 um; exine more or less smooth but for very fine (1-1.5 um) scabrae.
This type is found in Aphananthe, Celtis, Chaetacme, Gironniera, Lozanella, Parasponia, Ptero-

34 FLORA MALESIANA [ser. I, vol. 82

celtis and Trema. It may be noted here that pollen grains of Gironniera, Parasponia and Trema
are usually slightly smaller and have finer exine sculpturing than those of other genera with
Celtis-type of pollen, while the pollen grains of the Malesian species of Trema and those of
Parasponia parviflora MiQ. are predominantly diporate. As for pollen grains, Ulmaceae are
very closely allied to Moraceae and Urticaceae, particularly to the former. According to NAIR
(1967) the pollen type found in Ulmaceae, Moraceae and Urticaceae is derived from a tricolpate
type of Ranalean stock. — Literature: ERDTMAN, Pollen Morph. & Taxon. 1 (1956) 442; IkusE,
Pollen Grains of Japan (1956) 62; PRAGLOWsKI, Grana Palyn. 3 (1962) 45-65; KUPRIANOVA,
Kom. Bot. Inst. Ac. Sc. USSR 1 (1965) 54-58; Nair & SHARMA, Bot. Notis. 118 (1965) 177-
186; STRAKA, Pollen et Spores 8 (1966) 241-264; Nair, Rev. Palaeobot. & Palyn. 3 (1967)
81-91; MALLIK & CHAUDHURI, Bull. Bot. Soc. Beng. 22 (1968) 105-108; TsuKADA, Bot. Mag.
Tokyo 81 (1968) 385-395; Rao & Leer, Pacif. Sc. 24 (1970) 255-268; HUANG, Pollen Fl. Pl.
Taiwan (1972) 235; SowuNnmi, Grana Palyn. 13 (1973) 145-186; ADAmMs & Morton, Atl.
Pollen Trees & Shrubs Canad. & U.S. 9 (1974) pl. 17; STOCKMARR, Grana Palyn. 14 (1974)
103-107; Kepves & PARDUTZ, Acta Biol. Szeged. 20 (1974); HAMILTON, Pollen et Spores 18
(1976) 54-57.

Embryology. Apart from several species of U/mus and Holoptelea very little is known about
the sporogenesis and embryogenesis of the U/maceae. From a very limited information so far
published it appears that the anthers are initially tetrasporangiate but become bisporangiate just
before anthesis through the breakdown of the adjoining wall between the locules. The anther-
wall development conforms with the so-called basic-type in which the parietal cells divide both
anticlinally and periclinally to form the endothesium layer, two (Trema and Ulmus) or three to
four (Holoptelea integrifolia) middle-layers and glandular tapetum. Simultaneous cytokinesis in
the microspore mother-cells follows meiosis and as a result the pollen grains are initially arranged
in either tetrahedral or decussate tetrads. At anthesis the pollen grains are either 2-celled (Holop-
telea and Trema) or 3-celled (U/mus). In Celtis, Holoptelea and Trema up to 80% of the pollen
grains produced are sterile or imperfectly developed. The ovule is anatropous to hemianatropous,
bitegmic, crassinucellar or tenuinucellar (in a few species of U/mus) with the micropyle formed by
both integuments (Celtis and Trema) or by the inner integument only (Holoptelea and Ulmus). In
Holoptelea and Trema the megaspore mother-cell divides into 4 daughter cells arranged in a
linear tetrad, and of these only the chalazal megaspore develops into Polygonum-type of embryo-
sac. In U/mus, however, the embryo-sac is tetrasporic and either belongs to Adoxa- or Drusa-type
or variation of these two types with 4-12 antipodal cells. The pollen tube enters the ovule either
through the micropyle, the integuments or the chalaza. Endosperm formation is nuclear and
the tissue is either diploid or triploid and later becomes cellular. Embryo development conforms
with the Onagrad-type in Holoptelea and Solanad-type in U/mus. Polyembryony is a common
phenomenon, especially in U/mus. The mature embryo is straight with broad, flat or plano-
convex, equal or slightly unequal cotyledons in Holoptelea, Planera, Phyllostylon, Ulmus, and
Zelkova, or curved with ascending hypocotyle and narrow, incurved or induplicate-plicate or
variously folded cotyledons which are mostly unequal in length in Ampelocera, Aphananthe,
Celtis, Gironniera, Parasponia, Pteroceltis and Trema. — Literature: SHATTUCK, Bot. Gaz. 40
(1905) 205-223; LELIVELD, Rec. Trav. Bot. Néerl. 32 (1935) 543-573; Capoor, Beih. Bot.
Centralbl. 57 (1937) 233-249; WALKER, Am. J. Bot. 37 (1950) 47-52; HJELMQvisT & GAZzZI, Bot.
Notis. 118 (1965) 329-360; Davis, Syst. Embryol. Angiosp. (1967) 266-267.

Chromosomes. From various published data it seems that the chromosome number in the
Ulmeae (Holoptelea, Ulmus, and Zelkova) isn = 14 and 2n = 28, 42, and 56, though reports of
n = 15 and 30 have been made on U/mus americana. In the Celtideae the number seems to be less
constant varying from n = 10, 2n = 20, 28, 40 in Celtis (9 spp.); n = 30, 2n = 84 in Chaetacme
(2 spp.); ton = 10, 10 + B, 18, 20, and 80 in Trema (3 spp.). It may be noted here that as for
chromosome number, U/maceae seems to be closely related to Moraceae where n = 12-16,
2n = 24, 26, 28, 42, 56, and 84, and to Urticaceae of which n = 14, 28 and 2n = 22, 24, 28, 52,
and 84. — Literature: KRAUSE, Ber. Deut. Bot. Ges. 48 (1930) 9-13; Planta 13 (1931) 29-84;
WALKER, Science 75 (1932) 107; SAx, J. Arn. Arb. 14 (1933) 82-84; BowbDeEN, Am. J. Bot. 32
(1945) 195; DARLINGTON & WyLiE, Chromos. Atlas Fl. Pl. (1955) 182-183; MANGENOT &
MANGENOT, Bull. Jard. Bot. Brux. 28 (1958) 315-329; ArorA, Bull. Bot. Surv. India 2 (1960)

1977] ULMACEAE (Soepadmo) 35

305; GAJAPATHY, Bull. Bot. Surv. India 3 (1961) 49-51; GRUDZINSKAJA & ZAKHARYEVA, Bot.
Zhurn. 52 (1967) 641-651; Hsu, Taiwania 13 (1967) 117-129; MEHRA & GILL, Taxon 17 (1968)
574-576; J. Arn. Arb. 55 (1974) 663-677; Feporov (ed.), Chromos. Numb. FI. Pl. (1969) 710-
711; GADELLA c.s. Acta Bot. Neerl. 18 (1969) 74-83; MEHRA & Hans, Taxon 18 (1969) 310-315;
TATAYUK & TURCHANINOVA, Tsitologia & Genetika 4 (1970) 397-401 ; HANs, Cytologia 36 (1971)
341-345; Niewaus, Taxon 20 (1971) 355; MeHRa, Nucleus 15 (1972) 64-83; SARKAR, Taxon 22
(1973) 652.

Chemotaxonomy. SOLEREDER mentioned the more or less general occurrence of cystoliths
and cystolith-like structures (SiO, + CaCO;) in Ulmaceae. The tendency to accumulate carbon-
ate of lime seems to be very strong in this family ; CaCO, is deposited in wall structures (e.g. hairs,
cystoliths) and in cell lumina (e.g. in heartwood of Ul/mus and Celtis; in seed coat cells of Celtis).
Often oxalate of lime is also present in large amounts; solitary and clustered crystals occur in the
family. Anatomically easily detectable internal excretion comprises also mucilage production.
The mucilage is deposited in epidermal cells (many taxa) or in mucilage idioblasts in the meso-
phyll of some genera and in barks and flowers of most species of U/mus. The bark of Ulmus
rubra Muu. (‘Slippery Elm’) was used formerly as a mucilaginosum in official medicine. In
mucilage-rich elm barks large mucilage idioblasts may develop to lysigenous mucilage cavities.
Chemically elm bark mucilages are characterized by a high content of galacturonic acid, galac-
tose, 3-0-methylgalactose and rhamnose. U/maceae are moderately strong accumulators of
polyphenolic compounds. Derivatives of caffeic acid, catechins, pro-anthocyanidins (formerly
leucoanthocyanidins), flavonols (especially glycosides of kaempferol and quercetin) and con-
densed (= flavanoid) tannins seem to occur more or less ubiquiteous in leaves, fruits, barks and
woods. According to LEBRETON flavonoid constituents with a trihydroxylated B-ring (in casu
myricetin and prodelphinidin), an assumedly primitive feature, are restricted to Celtideae.
(+)-Catechin was definitely identified in leaves, twigs and barks of European elms and its
7-xyloside was isolated from the stem-bark of U/mus americana L. C-Glycoflavons (tremasperin)
occur in leaves of Trema aspera BL., and the wood of Zelkova serrata (THUNB.) MAKINO contains
large amounts of the fungistatic 6-C-glucoflavonoids keyakinin and keyakinol. Tannin contents
of woods, barks, leaves and fruits are moderate (mostly less than 10%). There is only one report
in literature indicating a possible co-occurrence of galli- and ellagitannins with condensed tannins
in Ulmaceae; bark and wood of Celtis australis L. contain gallic acid and derivatives of ellagic
acid according to CHARI c.s. (1968).

Much chemical work was performed with elm barks and especially elm woods in connection
with ‘Dutch Elm Disease’. Cadinane-type oxigenated sesquiterpenes seem to be present in the
young wood of every species. On aging (heartwood formation) or after fungal infection, synthesis
and accumulation of fully aromatic (cadalenal, hydroxycadalenal) and (or) o-quinonoid (the
mansonones) cadinane derivatives take place in American elm species belonging to the sections
Trichoptelea, Microptelea and Chaetoptelea; they seem to be absent from the sections Blepharo-
carpus and Madocarpus in which all European elms are included. It deserves mentioning that the
antifungal cadalenals and mansonones represent phytoalexin-like stress compounds in U/mus,
and occur at the same time as normal heartwood constituents in U/mus and Zelkova (but not in
Celtis); they are chemically identical with, or biochemically closely related to the gossypol-
mansonone-group of constituents of many Malvaceae, Bombacaceae and Sterculiaceae (man-
sonones were first detected in the wood of Mansonia altissima A. CHEV.). It was recently shown
that hemigossypol, the precursor of the long-known gossypol, is a phytoalexin in many malva-
ceous plants and that p-quinonoid derivatives of hemigossypol are engaged in the plants resistance
against attack by several phytophagous insects (J. R. GRAY c.s. J. C. S. Chem. Commun. 1976,
109; J. A. VeEcu c.s. l.c. 144). As far as ecological chemistry (defensive substances) is concerned,
Ulmaceae much resemble members of the order Malvales. Leaf, bark and wood waxes were
investigated by several authors in recent time. They seem to consist mainly of alkanes, long-chain
fatty acids, wax alcohols and phytosterins. Additionally pentacyclic triterpenes are often present;
6-amyrin (i), lupeol (ii), betulin (iii), friedelin (iv), friedelanol (v), moretenol (vi), simiarenol (vii)
and simiarenon (viii) were reported from leaves and (or) barks of Celtis australis L. (iii), C.
laevigata WiLLD. (vi), Holoptelea integrifolia PLANCH. (iv, Vv), Trema guineensis FICALHO (reported
as T. orientalis BL.; vii, viii), Ulmus americana L. (ii, esterified with cerotinic acid) and Zelkova

36 FLORA MALESIANA [ser. I, vol. 82

serrata MAKINO (iv). The heartwood of Holoptelea integrifolia PLANCH. yielded 2«-hydroxy-3-
epioleanolic acid (G. MisrA c.s. Planta Medica 27, 1975, 290); this is the only triterpenic acid
isolated hitherto from U/maceae. Seeds of Ulmaceae seem to store predominantly proteins and
fatty oils. The oils have linolic (Celtis, Chaetacme, Trema), oleic (Holoptelea) or capric (Ulmus,
Zelkova) acids as main fatty acid. Species of Ce/tis and Pteroceltis accumulate small amounts of
quebrachitol in leaves; this cyclitol could not be detected in leaves of species of Ulmus and
Zelkova (Hemiptelea included). Alkaloid-like compounds are recorded in literature from
members of Ampelocera, Aphananthe, Celtis, Gironniera, Trema and Ulmus, but only in the case
of Ampelocera ruizii KLOTZSCH an alkaloid-like compound isolated from leaves was chemically
identified; it proved to be an «-pyridone derivative related to trigonelline (R. H. BURNELL c.s.
Lloydia 38, 1975, 444). The foetid smell of some Celtis woods of India, Indonesia (‘kaju tai’) and
Africa is caused by skatol. Several species of U/maceae are reported to be toxic in literature.
GRESHOFF isolated a toxic bitter principle from the leaves of Aphananthe aspera (THUNB.)
PLANCH. (= Homoioceltis aspera BL.) which he compared with his streblide (from Streblus asper
Lour.; strebloside is now known to be a cardenolide). Leaves of Trema cannabina Lour.
(= Sponia virgata PLANCH.) and of T. aspera BL. (= T. cannabina) were reported to be cyano-
phoric; both species, however, are polymorphic with regard to cyanogenesis if the botanical
identification of all plant samples investigated hitherto was correct. Leaves of T. aspera (= T.
cannabina) contain another toxic principle called trematoxin; its chemical structure is not yet
known.

From the taxonomic point of view three facts deserve special mentioning: (1) U/maceae are
generally included in Urticales; their chemistry agrees rather well with such a classification as is
indicated by patterns of mineralisation and phenolic compounds. (2) The chemistry of Ulmaceae
resembles members of Malvales in several respects: chemistry of stress compounds; mucilages
with high contents of galactose, rhamnose and galacturonic acid; some features of the poly-
phenolic and triterpenic patterns. (3) The classification of Ulmaceae in Ulmoideae and Celti-
doideae (ENGLER’s Syllabus 2, 1964) or Ulmeae and Celtideae (HUTCHINSON, General of Flowering
Plants 2, 1967) is not very satisfactory from the chemical point of view (see cadinane-type
sesquiterpenes including mansonones and capric acid as main fatty acid in seed oils in U/mus and
Zelkova, but not in Celtis).

For more phytochemical details and references see my ‘Chemotaxonomie der Pflanzen’ 6
(1973) 545-554, 762-763, 791, 796. — R. HEGNAUER.

Taxonomy. The family name Ul/maceae was first introduced and defined by MirBEL in 1815,
at which time it included only Ce/tis and U/mus. Link (1831) proposed splitting Ulmaceae into
two separate families, i.e. U/maceae to include U/mus and related genera, and Celtidaceae com-
prising Celtis and its allies, an opinion which was supported by GRUDZINSKAYA (1967). However,
all contemporary taxonomists generally agree to regard Ulmaceae as a natural taxon closely
related to Moraceae and Urticaceae and to include these families in the order Urticales. Any
difference of opinion is usually restricted to the inclusion or exclusion of a few genera in the
family. In the most recent treatise, HUTCHINSON (1967) divided the family into two tribes, namely
the Ulmeae (fiowers bisexual, fruit not drupaceous, embryo straight, cotyledons flat or longitudi-
nally folded) to include: Holoptelea, Planera, Phyllostylon, and Ulmus, and the Celtideae (flowers
unisexual or sometimes bisexual, fruit drupaceous, embryo curved, cotyledons mostly variously
folded) comprising Ampelocera, Aphananthe, Celtis, Chaetacme, Gironniera, Hemiptelea,
Lozanella, Mirandaceltis, Parasponia, Pteroceltis, Trema and Zelkova. This subdivision was
supported by SwWEITZER (1971) who studied the anatomy of leaf and wood. However, as has been
mentioned under Embryology and Palynology, the embryo of Ze/kova is straight, and the pollen
(also of Ampelocera and Hemiptelea) belongs to the Ulmus-type (see also ERDTMAN, 1956).
Furthermore in many species of Celtis the flowers are bisexual, and in U/mus lanceaefolia and
U. parvifolia the flowers are either functionally male or female. This seems to indicate that the
tribal subdivision as proposed by HUTCHINSON is not a clear cut case, but that Ulmaceae is a
natural taxon. It should be noted further that the Mexican genus Mirandaceltis is in the present
study regarded as congeneric with Aphananthe.

As for phylogenetic relationship, there seems to be two different opinions. Bessey (1915) and
THORNE (1968, 1973) placed Ulmaceae along with Moraceae and Urticaceae in the superorder

1977] ULMACEAE (Soepadmo) 37

PI
ae

< ES
\ |

Fig. 2. Ulmus lanceaefolia Roxs. ex WALL. a. Habit, nat. size, b. fruit, x 2, c. persistent cup-shaped
perianth, x 2, d-e. embryo, nat. size, g. flowering twig, x 2/,, h. cluster of ¢ flowers, x 8, i. ¢ flower,
x 14, j. cluster of 3 flowers, x 2, k-l. 3 flower, x " ad HANSEN c.s. 11265, g-i ScuMuTz 3024, j—/ LisTeR

38 FLORA MALESIANA [ser. I, vol. 8?

Malviiflorae, and considered them as families having a very close affinity to or derived from the
Malvales. On the other hand, authors such as HUTCHINSON (1967), CRONQUIST (1968), TAKHTAJAN
(1969), SweITzER (1971), efc., are of the opinion that Ulmaceae, Moraceae, and Urticaceae are
closely allied to or have been derived from the Hamamelidales. — Literature: MIRBEL, Elém. Phys.
Veg. Bot. (1815) 905; Link, Handb. 2 (1831) 445; Bessey, Ann. Mo. Bot. Gard. 2 (1915) 109-164;
ERDTMAN, Pollen Morph. & Pl. Tax. (1956) 442-443; GruDZINsKAyYA, Bot. Zhurn. 52 (1967)
144-150; HUTCHINSON, Genera of Flowering Plants 2 (1967); CRoNquist, Evol. & Class. Fl. Pl.
(1968) 166-167; THORNE, Aliso 6 (1968) 57-66; Brittonia 25 (1973) 395-405; TAKHTAJAN, FI. PI.
Orig. & Disp. (1969) 210-212; Sweitzer, J. Arn. Arb. 52 (1971).

Uses. 1. Timber. Throughout the north temperate regions the tough, strong and durable
wood with attractive appearance and excellent bending quality of many species of Celtis and
Ulmus is extensively used for various purposes including shipbuilding, panelling, furniture, boxes,
crates, veneers, etc. and that of Zelkova and Phyllostylon for making weaver’s shuttles, scales,
piano-keys, efc. In Central America timber of Chaetoptelea (= ?Ulmus) is used for railway
sleepers, frames and wheels of vehicles. In Africa and India wood of Holoptelea is utilized for
various building purposes. In Malesia and neighbouring countries except Aphananthe cuspidata,
Celtis rigescens, C. hildebrandii, C. tetrandra, Gironniera nervosa, Ulmus lanceaefolia and a few
others, the trees seldom reach timber size, and as a consequence very little is known about their
usage. Of these species the timber is locally used for making planks in house-building and other
light constructions. The soft wood of Trema and other species of Gironniera is used locally for
making tea-chests and match-sticks, for firewood and charcoal.

2. Bark. Due to the high content of mucilagenous substances, decoction of barks of Holop-
telea, Parasponia, Trema and Ulmus mixed with some other ingredients is used in local folk
medicines to cure ailments such as inflammation of mucous membrane, rheumatism, etc. The
tough fiber is known to be used locally for making ropes.

3. Root. Decoction of roots of Gironniera and Trema species mixed with other substances is
used to cure sore mouth, diarrhoea, and also applied as protective medicine after child-birth.

4. Leaves. Especially of Trema species leaves are used as fodder, though due to the presence of
glucocides they could be poisonous if consumed in a large quantity.

5. Fruits. In India fruits of Celtis and Holoptelea are known to be eaten.

6. Shade trees. Trema has been used for shade in coffee and cocoa plantations in various parts
of Asia.

7. Soil conservation. In South Africa Trema has been planted to protect soils against erosion
(SCHEEPERS c.s.). As both Trema and Parasponia species come up in dense seral stands on eruptiva,
on fresh volcanic ash, are sometimes pioneers on lavastreams, and are almost invariably an
important constituent of thickets, seral regrowths, and secondary forest, I would emphasize that
they may represent an untapped cheap source for soil conservation for poor, eroded soils and old
mining lands. They have all the favourable qualities of pioneer plants, indifference to soil, pro-
ducing abundant seed, and that already at a very early age, and furthermore they are available
almost throughout the year. Curiously I do not know of experiments by the Indonesian Forestry
Service in this respect. — Literature: BURKILL, Dict. Econ. Prod. Mal. Pen. (1935) 513-514,
1088-1089, 2213-2214; MeTCALFE & CHALK, Anat. Dicot. (1950) 1277; SCHEEPERS c.s. Tijd.
Natuurwet. S. Afrika Akad. Wet. & Kunst. 8 (1968) 105-120; Sweitzer, J. Arn. Arb. 52(1971)
52a.

KEY TO THE GENERA

1. Flowers always borne on bare older branches, and organized in a condensed cluster on short leafless
lateral shoots; perianth cup-shaped, 5-7-lobed; ovary (fruit) stipitate. Fruit a dry, flat, winged samara.
Embryo straight es wee ie dl ee eee APS eee 1. Ulmus

1. Flowers axillary, or rarely borne in a condensed capitate thyrse on older branches (Gironniera celtidi-
folia); perianth 4~5-lobed, with the lobes free from one another except for their base; ovary sessile.
Fruit a fleshy drupe. Embryo variously curved.

2. Leaves triplinerved at base, or if pinnately nerved the stipules do not leave a circular scar around the
node; lateral nerves less than 5 pairs.
3: Stipules intrapetiolar, conate-s° +): "0G Oe ae ee 6 Lee ee 2. Parasponia
3. Stipules extrapetiolar, free.

1977] ULMACEAE (Soepadmo) 39

4. Female flowers borne in condensed, multiflowered raceme. Perianth of male flower induplicate-
valvate. Fruit compressed, elliptic lens-shaped in cross-section .......... 3. Trema

4. Female flowers solitary in the axils of leaves or borne in a cymoid cluster of 2-3. Perianth lobes of
male flowers imbricate. Fruit faintly 3—5-angular in cross-section.

5. Male flowers borne in a 2-3-flowered cymoid inflorescence or in a much-branched paniculate,
subterminal inflorescence. Female flowers borne in a racemose cluster of 2-10 or in a $8, much-
branched racemose inflorescence; staminodes mostly present. Cotyledons broad, variously folded
SES Ir et eS , k ounies oe aoe ue eS. Bey bdo ee 2 4. Celtis

5. Male flowers organized in a condensed, multiflowered raceme. Female flowers always solitary or
rarely borne in a 2-3-flowered racemose, $2 inflorescence; staminodes always absent. Cotyledons

2 LOT OTe Tay 27 8 RS Ge ee, ee 5. Aphananthe

2. Leaves pinnately nerved; lateral nerves more than 5 pairs; stipules free but overlapping each other,

m falling leaving circular scar around the node. . ......4.+..4+ +56. 6. Gironniera
1.ULMUS

LINNE, Gen. Pl. ed. 5 (1754) 106; ENDL. Gen. Pl. (1837) 276, Suppl. 2 (1842) 29;
PLANCH. Ann. Sc. Nat. III, 10 (1848) 259; in DC. Prod. 17 (1873) 154; BAILL. Hist.
Pl. 6 (1877) 137; B. & H. Gen. Pl. 3 (1880) 351; Hook. f Fl. Br. Ind. 5 (1888) 480;
ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 62; BERNARD, Bull. Herb. Boiss. II, 5
(1905) 1097; ibid. 6 (1906) 23; SCHNEIDER, Oest. Bot. Z. 66 (1916) 21, 65; in Sargent,
Pl. Wils. 3 (1917) 238; GaGnep. Fl. Gén. I.-C. 5 (1927) 674; TuTin, FI. Europ. 1
(1964) 65; Hutcu. Gen. FI. Pl. 2 (1967) 147; Touw & STEEN. Blumea 16 (1968) 84.
— Fig. 2, 4-6.

Deciduous or semideciduous trees or shrubs. /nnovations densely set with greyish
to brownish simple hairs, glabrescent. Buds ovoid-conical or obovoid-globose,
scales imbricate, hard and tough, glabrous. Stipules extrapetiolar, caducous. Leaves
pinnately nerved, variously serrate to crenate, thin- to thick-coriaceous and rigid,
glabrous or variously sparsely hairy at least beneath. Flowers % but of two kinds,
one functionally g and the other functionally 2, variously stalked and spirally
arranged in fascicles of 3-15 on short lateral shoots. Perianth mostly campanulate,
variously 4-8-lobed. Anthers glabrous, reniform, extrorse. Ovary compressed,
sometimes stipitate; style short. Ovule 1, anatropous to amphitropous. Fruit a dry
and compressed nutlet surrounded by a membranous reticulate-venose wing.
Seed: endosperm absent, embryo straight with planoconvex cotyledons. Germina-
tion epigeal.

Distr. About 20-25 spp., distributed in Europe (as far north as 68°), W. & SW. Russia, N. & NE.
India, Burma, China, Korea, Japan, Formosa, Indo-China, N. Thailand, and in North America from
N. Mexico to the U.S.A. east of the Rocky Mts as far north as 60°. In Malesia: 1 sp. so far known from a
few localities in N. Sumatra, the Lesser Sunda Is. (Flores), and Central & S. Celebes.

As has been indicated by SCHNEIDER, /.c., there seem to be three centres of distribution, i.e. the Euro-
em crmtre (5-6 spp.), the Indian-E. Asian centre (10-15 spp.), and the North American centre (4-5 spp.).

ig. 3.

Fossils. Numerous fossils (pollen grains, leaf-impressions, and wood fragments) have been reported
from various late Cretaceous and Tertiary deposits in Europe, Russia, China, Japan, North America, and
Greenland. Fig. 3.

Ecol. In Malesia the genus is so far known only from areas more or less subject to a seasonal climate
at 200-1450 m.

Taxon. Currently there is not a single worldwide monograph of the genus available for reference. The
latest and perhaps the most comprehensive revision since PLANCHON’s work (1873) is that by SCHNEIDER
(1916). He distinguished 26 spp. and recognized 5 distinct sections in the genus based on morphological
characters derived from inflorescence, flowers, and fruits.

When more specimens from China become available for further studies, | believe the number of species
occurring in the Indian-E. Asian centre will have to be reduced considerably.

40 FLORA MALESIANA

[ser. I, vol. 8?

Fig. 3. Approximate range of U/mus L. with number of spp. in each of the three centres, Malesian localities

belonging to a species of the Asian centre. Fossil localities outside the present range indicated by dots;

adopted from BERNARD, /.c.; GREGUss (Tert. Angios. Hung., Ak. Kiado Budapest, 1969, 83), and La
Motte (Mem. Geol. Soc. Am. 51, 1952, 346).

Fig. 4. Peeling bark of Ulmus lanceaefolia RoxB. ex
WALL., x 1/, (Photogr. ScHMuTz, 5 Nov. 1972,
Flores, Nunang).

1. Ulmus lanceaefolia Roxs. ex WALL. Pl. As. Rar.
2 (1831) 86, t. 200; Roxs. Fl. Ind. ed. Carey 2
(1832) 66 (‘lancifolia’); PLANCH. Ann. Sc. Nat. III,
10 (1848) 281; in DC. Prod. 17 (1873) 162; Kurz,
For. Fl. Burma 2 (1877) 473; GAMBLE, Man. Ind.
Timb. ed. 1 (1881) 342; Hook. f. Fl. Br. Ind. 5
(1888) 480; Hems-. J. Linn. Soc. Bot. 26 (1894) 447;
PRAIN, Beng. Pl. (1903) 718; BRANDIS, Ind. Trees
(1906) 594; SCHNEIDER, Oest. Bot. Z. 66 (1916) 32;
in Sargent, Pl. Wils. 3 (1917) 263; Merr. Contr.
Arn. Arb. 8 (1934) 44; Touw & STEEN. Blumea 16
(1968) 84; MELVILLE & HEYBROEK, Kew Bull. 26
(1971) 24 (‘lanceifolia’). — U. hookeriana PLANCH.
in DC. Prod. 17 (1873) 162; ENGL. in E. & P. Nat.
Pfi. Fam. 3, 1 (1888) 62. — U. tonkinensis GAGNEP.
Fl. Gén. I.-C. 5 (1927) 674. — Fig. 2, 4-6.

Small to large tree up to 48 m, 70cm @, often
with fluted trunk. Bark rough, pustulate, with large
warty lenticels. Branchlets initially densely set with
greyish to brownish curly simple hairs, later glab-
rous and sparsely warty lenticellate. Buds obovoid-
globose, c. 2-3 mm @; bracts dark brown. Stipules
linear-lanceolate acute, c. 4-5 by 1-11/, mm, soon
caducous. Leaves thin- to thick-coriaceous, lanceo-
late to ovate-lanceolate, (2-)4-6(-9) by (1-)2-3
(—31/,) cm (index 2-21/,), broadest at or slightly
below the middle, more or less glabrous, glossy;
base rounded to attenuate-acute, unequal;
margin serrulate to serrulate-crenulate; apex acute
with blunt tip; midrib raised beneath and flattish to
impressed above, as the petiole initially densely
greyish, curly hairy on both surfaces, glabrescent;
nerves (6~)10-12(-14) pairs, subparallel, often
rather irregularly spaced, slightly raised beneath,
flattish to impressed above, often forked near and
towards the leaf-margin; reticulations fine, areo-
late; petiole (2~-)3—-4(—6) by '/,-1 mm. Flowers in
fascicles of 3-10. — Functionally 3 flowers globose
before anthesis, 1!/,-2 mm @, subglabrous; lobes
5-6, obovate-lanceolate, c. 2 by 1mm; ts

1977]

Fig. 5. Ulmus lanceaefolia Rox. ex WALL. with old
leaves at Nunang (Photogr. ScHmMutTz, 15 Oct.
1972, Flores).

glabrous, slender, c. 1 mm; anthers c. 1 by '/, mm,
glabrous; pistillode compressed obovate-elliptic,
glabrous. — Functionally 2 flowers (as seen under a
very young fruit): perianth campanulate, lobes
5-6, rounded-elliptic, hairy along the margin;
filaments slender, glabrous, 3-5 mm, anthers as in
3 flower; ovary stipitate, glabrous, + obovate-
elliptic. Fruit obovate-elliptic, glabrous, including
the wing 2-3'/, by 1'/,-2cm, stalk 5-10 mm,
articulate, lower part hairy.

Distr. China (?), India (E. Himalaya, Sikkim,
Bhutan, Khasia Hills, Manipur, Assam), Bangla-
desh, Burma (Hukong Valley, Chittagong Hills),
Thailand (northern parts), Laos, Vietnam (Mt
Bavi); in Malesia: N. Sumatra (Gajo- & Karo-
Batak Lands), Lesser Sunda Is. (Flores), and Cele-
bes (Poso; Bonthain). Fig. 7.

Ecol. Scattered tree in lowland to submontane
forest, 200-1450 m. In Thailand it is confined to
forests along streams and in Flores it has been
found on limestone. Fl. fr. in the northern hemis-
phere Febr.—April; in Flores Nov.

Taxon. U. lanceaefolia is very closely allied to
U. parvifolia Jacq. from China and Japan. It differs
from the latter by its narrow leaves with a shorter
petiole, serrulate to serrulate-crenulate margin,
and fewer lateral nerves, obovoid-globose buds,
the campanulate perianth of the functionally ?
os and the reticulate venation of the fruit; see

2

ULMACEAE (Soepadmo) 4]

Uses. Very little is known about the usage of
this species, but judging from the enormous size it
can attain it must have been a useful timber in
house-building, construction, efc., at least to the
local inhabitants.

Vern. Sumatra: péngki(h), poki, Karo-Batak,
pongki, Toba-Batak; Lesser Sunda Is.: ngguling,
nggulung, Flores; Celebes: mota, Bonthain.

€

ny

Fig. 6. Ulmus lanceaefolia Roxp. ex WALL.,
leafless, in flower, + x 2/3, at Nunang (Photogr.
ScHMUTZ, 5 Nov. 1972, Flores).

Fig. 7. Range of Ulmus lanceaefolia Roxs. ex WALL.

42 FLORA MALESIANA [ser. I, vol. 8?

|

i
Y

y
|

Fig. 8. Parasponia rigida MeRR. & PERRY. a. Habit, x 2/,, b. twig-tip with stipules, c. connate stipules,
from inside, d. detail of leaf undersurface, all x 3, e. ¢ flower, f. ditto in LS. g. young 9 flower, h. mature
2 flower, i. ditto in LS, all x 12, j. fruit, k. ditto in LS, both x 12 (af ANU 6463, g—k SCHODDE 4828).

1977] ULMACEAE (Soepadmo) 43
2. PARASPONIA

Mia. Pl. Jungh. (1851) 68; Fl. Ind. Bat. 1, 2 (1859) 218; BL. Mus. Bot. 2 (1856) 65;
PLANCH. in DC. Prod. 17 (1873) 194; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888)
65; J. J. SMITH in K. & V. Bijdr. 12 (1910) 662; BAcK. & BAKH. f. FI. Java 2 (1965)
12; Hutcu. Gen. FI. Pl. 2 (1967) 149. — Fig. 8, 10-11.

Shrubs to medium-sized trees. Bark grey-brown, smooth to finely fissured ; inner
bark fibrous and tough. /nnovations with dense appressed, silvery to greyish hairs.
Stipules intrapetiolar, connate into a bifurcate unit and together enclosing the
terminal bud, caducous. Leaves (in Mal.) triplinerved at base, concolorous, above
non-scabrous to variously scabrous, mostly glabrous except for the midrib and
lateral nerves, lower surface variously pubescent. Inflorescences axillary, 3, 9, or
32, much-branched, many-flowered, paniculate or thyrsoid, including the bracts
densely short greyish appressed-pubescent. Flowers 5-merous. — ¢ Flower +
globose, perianth lobes imbricate in bud; stamens glabrous, introrse; filaments
subulate, glabrous; anthers reniform to subglobose, sub-basifixed, glabrous; pistil-
lode obovoid-conical, compressed, surrounded by hirsute hairs at its base. — 9
Flower ovoid-conical; staminodes absent; ovary ovoid, slightly compressed; stig-
matic arms short, simple; ovule anatropous. Drupe ovoid, slightly compressed
pericarp fleshy and fibrous, endocarp hard and stony. Seed: endosperm scanty or
copious; embryo curved, cotyledons equal, hypocotyle ascending.

Distr. 5 spp., in Polynesia (Tahiti) and Melanesia (Fiji, New Hebrides, Solomons); in Malesia: New

Guinea (incl. New Britain), Moluccas (Ternate, Banda), Philippines, Celebes, Lesser Sunda Is. (Lombok,
Bali), Java, and S. Sumatra (Palembang). Fig. 9, 12.

Fig. 9. Approximate range of the genus Parasponia Mia. (line); localities of P. andersonii (PLANCH.)
PLANCH. (dots) and P. melastomatifolia J. J. S. (triangles).

Ecol. In New Guinea and the Pacific islands the genus is found as a pioneer plant invading and occupy-
ing newly available habitats from the lowland up to 2000 m. In Java the same situation was described b
E. W. CLAson from natural regeneration on volcanic ash of Mt Kelud, together with Trema (Bull. Jard.
Bot. Btzg III, 13, 1935, 509). Fig. 10, 11. Recorded as a ery on lavastreams of Mt Batur in Bali by
De Voocp (Trop. Natuur 29, 1940, 48, f. 12). Grows well in all types of soils, including volcanic ash and
limestone, very often gregariously together with Trema spp. and forming a dense thicket on ridges, hills
and along river-banks.

44 FLORA MALESIANA

[ser. I, vol. 8?

TAXON. Parasponia is morphologically very similar to Trema but can easily be distinguished from the
latter by its imbricate perianth lobes of the male flowers and intrapetiolar, connate stipules enclosing the

terminal bud.

KEY TO THE SPECIES

1. Basal nerves running up throughout the length of the leaf or nearly so.
2. Leaf thick-coriaceous, lower surface densely set with soft erect hairs; margin distinctly serrate;

reticulations prominent beneath. Inflorescence 3 or 2

petiole... oh

, at anthesis condensed and shorter than the
1. P. rigida

2. Leaf chartaceous to thin- -coriaceous, lower surface glabrous or sparsely appressed- hairy; margin
finely serrulate to subentire; reticulations obscure. Inflorescence 3 or $9, at anthesis lax and longer

than the petiole.
Basal nerves running up to ee

. 2. P. melastomatifolia

2/, the length of the leaf.

"5. Leaf thick-coriaceous, upper surface strongly rugose and scabrous, lower surface densely pubescent;

midrib, nerves, and reticulations prominent beneath

. P. rugosa

3. Leaf chartaceous to thin-coriaceous, upper surface not or hardly rugose nor ‘scabrous, lower surface
sparsely appressed pubescent or glabrous; midrib, nerves, and reticulations only slightly raised

beneath.

4. Leaf elliptic-lanceolate, more or less glabrous; nerves more than 4 on each side, straight and ascend-
ing at a narrow angle (less than 40°) from the midrib. Inflorescences mostly 32, very rarely g or 2

4. P. parviflora

4. Leaves ovate to ovate-elliptic, underneath sparsely appressed pubescent; nerves less than 4, usually 3
on each side, arcuating at a wider angle (c. 45—60°) from the midrib. Inflorescences 3 or 9, "rarely 32
=

1. Parasponia rigida MeRR. & PERRY, J. Arn. Arb.
22 (1941) 254. — Fig. 8.

Small tree, up to 10m, 10cm @. Branchlets
initially densely silver-appressed-hairy, glabrescent
and sparsely warty lenticellate. Stipules ovate-
lanceolate, 8-10 by 2-3mm. Leaves elliptic to
ovate-lanceolate, thick-coriaceous, (5—)8—10(—12)
by (11/,—)2-31/,(—4!/.) cm (index 2!/,-3'/,), broadest
at or below the middle; base rounded to subcordate,
mostly symmetrical; margin serrate; apex acute to
acuminate; above more or less glabrous, rugulose
and scabrous, beneath densely soft-hairy; midrib
and nerves strongly raised beneath, flattish to im-
pressed above; reticulations subscalariform, dis-
tinct beneath; petiole 10-12 by 2-3 mm, terete.
Inflorescences 3 or 2, very rarely $2, 10—S0-flowered,
at anthesis condensed, c. +/,-1 cm long, usually
shorter than the petiole. — ¢ Flowers 1-2 mm @,
perianth lobes narrow-ovate, concave, c.2 by 1 mm,
appressed-pubescent outside; filaments c. 1 mm;
anthers ellipsoid, c. 1 by */, mm, pistillode obovoid-
conical, c. 1 by 1/, mm. — 9 Flower ovoid, c. 2 by
1 mm; perianth lobes ovate-acute, c. 3/, by 1/, mm,
sparsely pubescent outside; ovary c. 2 by 1 mm,
stigmatic arms spreading, long-papillose, c. +/,-
1 mm. Drupe ovoid-globose, c. 3-4 mm @, turning
orange to red when ripe. Endosperm copious.

Distr. Malesia: New Guinea. Fig. 12.

Ecol. Dominant pioneer tree in secondary
vegetation on ridges, also in mossy forest and on
limestone hills, 1000-2000 m. FI. fr. Jan.—Dec.

Vern. Bésukan, Hattam lang., golan, Finsch-
hafen dial., Morobe Distr.

2. Parasponia melastomatifolia J. J. SmitH, Nova
Guinea 8, 2 (1914) 891, t. 158. — P. simulans MERR.
& Perry, J. Arn. Arb. 22 (1941) 255.

Shrub to small tree, up to 6 m, with spreading
and brittle branches. Branchlets initially densely
appressed-silvery-hairy, glabrescent and sparsely
lenticellate. Stipules ovate-elliptic, 4-5 by 2-3 mm
Leaves elliptic-lanceolate to ovate-elliptic, (5-)6-8

P. andersonii

(-10) by (1'/,-)2"/,-3"/,(-41/.)cm (index 2-3),
broadest at or slightly below the middle; charta-
ceous to thin-coriaceous, above more or less
glabrous, hardly scabrous and often with mineral
deposits on the epidermis, beneath sparsely ap-
pressed-hairy especially on midrib and nerves, or
completely glabrous; base more or less rounded,
symmetrical; margin finely serrulate to + entire;
apex acute; midrib and nerves slightly raised
beneath, flattish to impressed above; nerves 1-2
pairs, the basal ones running throughout the length
of the leaf or nearly so, at an acute angle of less than
45°; reticulations subscalariform, inconspicuous
on both surfaces; petiole (S—)8—12(-15) by 1 mm,
sulcate, sparsely appressed-pubescent. Jnflores-
cences 3 or 3g, paniculate, 10—20-flowered, at
anthesis lax, 2-3 cm long and across, axes c. 1 mm
@, densely appressed-pubescent; bracts ovate,
c. 11/, by 1mm, sparsely appressed-pubescent
outside. — 3 Flowers c. 1-2 mm @, perianth lobes
ovate-acute, c. 1/,-1 by 1/, mm, sparsely appressed-
hairy outside; filaments c. 1mm, anther sub-
globose, c. 1 by !/, mm, pistillode obovoid, c. 4/,-1
by ?/,mm. — @ Flower ovoid, c. 1-2 by 1 mm;
perianth lobes ovate-acute, c. +/,-1 by +/,—/, mm,
sparsely appressed-hairy outside; ovary ovoid, c. 2
by 1 mm; stigmatic arms c. 1 mm, incurved. Drupe
ovoid-globose, c. 3 by 2mm. Endosperm scanty.

Distr. Malesia: New Guinea. Fig. 9.

Ecol. Common and dominant in seral vegetation
in gullies and river-banks, 200-1000 m. Fi. fr
Jan.—Dec.

Vern. Kwatoro, Onjob lang., Northern Distr.,
E. New Guinea.

3. Parasponia rugosa BL. Mus. Bot. 2 (1856) 66. —
P. aspera BL. l.c. 66; Miq. Fl. Ind. Bat. 1, CA ge
218. — Trema vulcanica MERR. Philip. J. Sc.
(1912) Bot. 260. — Trema philippinensis Fine,
Leafi. Philip. Bot. 9 (1934) 3218. — P. parviflora
(non Mia.) STEEN. Philip. J. Sc. 91 (1962) 507. —
Fig. 10-11.

1977]

ULMACEAE (Soepadmo) 45

Shrub or medium-sized tree, up to 20 m, 30 cm
@, with spreading branches. Bark smooth, grey-
brown; inner bark tough, brownish. Branchlets
densely, silvery, appressed, long-hairy, subglab-
rescent. Stipules ovate-lanceolate, (6—)8(—10) by
2-3 mm. Leaves ovate-lanceolate, (7—)8—10(—12)
by (2-)2!/,-31/,(—5) cm (index 2'/,-3), broadest at
or below the middle, thick-coriaceous; above
rugose and scabrous, sparsely hairy especially on
midrib and nerves, beneath densely silvery hairy by
soft, erect hairs; base rounded to cordate, equal to
slightly unequal; margin serrate, apex acute to
acuminate, the acumen up to 2cm; midrib and
nerves strongly raised and prominent beneath,
flattish to impressed above; nerves 2-4 pairs,
arcuate and ascending, the basal ones running up to
1/,7/, the length of the leaf; reticulations dense,
areolate, prominent beneath; petiole 7-10 by
2-3 mm, subterete, densely appressed-pubescent.
Inflorescences 3 or $2, 10-50-flowered, densely sil-
very appressed-hairy, at anthesis condensed, axes
c. */,-1"/, cm long, 1-2 mm @; bracts ovate-acute,
c. 1*/,-1 mm. — ¢ Flower c. 1-2 mm @; perianth
lobes ovate-acute, c. 1-2 by 1 mm, sparsely hairy
outside; filaments 1-1!/, mm, anthers ovoid-reni-
form, c. 1mm @; pistillode ovoid, compressed,
c. 1-2 by 1 mm. — 2 Flower ovoid-conical, c. 1—14/,
by 1 mm; perianth lobes narrow ovate-acute, c. 1-
1*/, by 1 mm, sparsely appressed-hairy outside;
ovary ovoid, c. 1-2 by 1 mm, stigmatic arms c.
1mm, spreading. Drupe ovoid, 2-3 by 2mm,
turning red when ripe. Endosperm copious.

Distr. Malesia: East Java (Mts Kelud &

Lamongan), Lesser Sunda Is. (Bali, Lombok),
Philippines (Luzon, Leyte, Mindanao), Celebes
(near Makassar; Tondano, Menado), Moluccas
(Ternate, Banda), New Guinea (W. & E. High-
lands and Morobe Distr., incl. New Britain).
Fig. 12.

Fig. 10. Pioneer vegetation on the volcanic ash of
Mt Kelud, East Java, of Saccharum spontaneum
and Parasponia rugosa BL. (Photogr. CLASON).

Ecol. Rather common and often dominant or
co-dominant pioneer plant in seral vegetation on
various types of soils including volcanic ash, 50—
1900 m. Fi. fr. Jan.—Dec. Fig. 10, 11.

Uses. Strips of the inner bark are used as ropes
in house and fence building by local inhabitants.

Vern. Java: anggring, anggris, J; Philippines:
analdung, If.; Moluccas: kayu kuli, Banda; New
Guinea: wanep, Enga lang., W. Highlands Distr.,
la karabi, W. Nakanai, New Britain.

Fig. 11. Older pioneer forest on Mt Kelud of Parasponia rugosa BL., Trema, Cyathea contaminans, and
Amomum (Photogr. CLASON).

46 FLORA MALESIANA

4. Parasponia parviflora Mig. Pl. Jungh. (1851)
69; Fl. Ind. Bat. 1, 2 (1859) 218, t. 16; BL. Mus.
Bot. 2 (1856) 65, f. 35; PLANCH. in DC. Prod. 17
(1873) 194; J. J. SmirH in K. & V. Bijdr. 12 (1910)
663, p.p excl. syn. P. aspera BL.; BACK. & BAKH.
f. Fl. Java 2 (1965) 12. — P. similis BL. Mus. Bot. 2
(1856) 66.

Small to medium-sized tree, up to 15 m. Branch-
lets initially densely silvery or grey appressed-hairy,
glabrescent, smooth. Stipules ovate, 5-10 by
2-4 mm, sparsely hairy outside. Leaves lanceolate
to narrow ovate-lanceolate, (3—)5-8(-10) by
(1-)2-3(—3'/,) cm (index 3-4), broadest at or below
the middle; chartaceous to thin-coriaceous, above
+ glabrous, not scabrous, beneath initially
appressed-hairy, later glabrous except for the
midrib and nerves; base rounded, more or less
equal; margin finely serrate, apex acute; midrib
and nerves slightly raised beneath, impressed and
inconspicuous above; nerves 4-6 pairs, straight,
ascending and parallel, at a narrow angle (30—40°),
basal ones running up to + half the length of the
leaf; reticulations fine, subscalariform, indistinct
on both surfaces; petiole terete, densely appressed-
hairy, 5-10 by 1 mm. Inflorescences 3, 2, or 38,
5-30-flowered, at anthesis condensed, shorter than
or as long as the petiole, as the bracts densely
short-hairy ; bracts ovate-acute, 1—11/, by '/,-1 mm.
— 3 Flowers glabrous, 1-2 mm @; perianth lobes
c. 1-11/, by ?/, mm; filaments 1/,-1 mm, anthers
subglobular, c. 1 by +/,mm; pistillode ovoid-
conical, compressed, 1-1'/, by ‘'/,mm. — 9
Flowers ovoid-conical, c. 2 by 11/, mm, + glabrous
except for the inner base of the perianth lobes;
perianth lobes ovate-acute, c. 1 by '/, mm; ovary
ovoid, c. 11/, by 1 mm; stigmatic arms spreading,
c. 1/,-1 mm. Drupe ovoid-conical, slightly com-
pressed, 11/,-2 by 11/, mm. Endosperm scanty.

Fig. 12. Localities of Parasponia rugosa BL. (dots),
P. parviflora Mia. (triangles), and P. rigida MERR.
& PERRY (squares).

Distr. Malesia: S. Sumatra (Palembang, very
rare), Java (common). Fig. 12.

[ser. I, vol. 8?

Ecol. In secondary or seral vegetation on ex-
posed habitats, also in teak forest, often rather
common and dominant locally especially on soils
derived from volcanic ash, 500-2000 m. Fi. fr.
Jan.—Dec.

Vern. Kurai, k. lélaki, k. tiangkréng, S, anggring,
anggris, anggrung, J

5. Parasponia andersonii (PLANCH.) PLANCH. in
DC. Prod. 17 (1873) 193. — Sponia andersonii
PLANCH. Ann. Sc. Nat. III, 10 (1848) 336; See-
MANN, FI. Vit. (1867) 235; PARHAM, Pl. Fiji Isl.
(1972) 133. — P. paucinervia MERR. & PERRY, J.
Arn. Arb. 20 (1939) 324.

Shrub to medium-sized tree, up to 15m and
30cm @. Branches spreading and drooping,
initially densely set with erect but soft, silvery hairs,
subglabrescent and sparsely warty lenticellate.
Bark smooth to nodular, grey-brown; inner bark
fibrous, tough, orange to brownish. Stipules ovate-
acute, sparsely hairy outside, 6-10 by 3-4 mm.
Leaves ovate to elliptic, thin-coriaceous, (5—)8—12
(-14) by (2-)3-4(-6)cm (index 2-3), broadest
below or at the middle; above subglabrous, scab-
rous, often covered with mineral deposits, beneath
sparsely set (rarely rather densely) with short and
soft hairs especially on midrib and nerves; base
rounded to subcordate, equal, rarely unequal;
margin serrate, apex acute to acuminate; midrib
and nerves slightly raised beneath and impressed
above; nerves 3-4 pairs, arcuating and ascending
at an angle of 45-60°, basal extending up to c. 2/,
the length of the leaf; reticulations fine, subscalari-
form, rather distinct below; petiole (7—)10—15(—20)
by 1-2 mm, densely set with silvery, soft, erect
hairs, flat or sulcate. Inflorescences 3, 2, or rarely
32, 10-30-flowered, at anthesis condensed or lax,
shorter than or as long as the petiole, including the
bracts densely silvery, soft-hairy; bracts ovate-
acute, c. 1 by !/, mm. — ¢ Flowers c. 11/,-2 mm @;
perianth lobes ovate-elliptic, c. 1 by +/,mm;
stamens glabrous; filaments c. 1mm, anthers
subreniform to subglobular, c. 1 by +/,mm;
pistillode subovoid-conical, c. 11/, by !/, mm. —
Flowers ovoid-ellipsoid, c. 11/, by 1 mm; perianth
lobes ovate-acute, c. 1 by !/,mm; ovary ovoid,
slightly compressed, c. 1 by '/, mm; stigmatic
arms c. ?/,mm, spreading and short-papillose.
Drupe ovoid, slightly compressed, 2-4 by 2-3 mm.
Endosperm copious.

Distr. Polynesia (Tahiti), Melanesia (Fiji, New
Hebrides, Solomons, very common), ? New
Caledonia (no specimen seen but cf. GUILLAUMIN,
Fl. Nouv.-Caléd. 1948, 94); in Malesia: New
Guinea (several islands off Madang and Milne
Bay) and New Britain. Fig. 9.

Ecol. Primary as well as secondary forests, on
various types of soils including limestone,
0-1500 m. Fi. fr. Jan.—Dec.

Uses. In the Solomons the bark is reputed to
have medicinal properties.

Vern. New Britain: ip, ivu; Solomons: bulasisi,
bulsisi, Kwara lang.; Fiji: ndroi, ndrou.

1977] ULMACEAE (Soepadmo) 47

3. TREMA

Lour. Fl. Coch. 2 (1790) 562; BL. Mus. Bot. 2 (1856) 58; Bru. Fl. Austr. 6 (1873)
157; B. & H. Gen. Pl. 3 (1880) 355; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 65;
BERNARD, Bull. Herb. Boiss. II, 6 (1906) 31, maps 19-21; J. J. SMirH in K. & V.
Bijdr. 12 (1910) 649; RENDLE, FI. Trop. Afr. 6, 2 (1917) 10; De Wrrt, Bull. Bot.
Gard. Btzg III, 18 (1949) 184; Hutcu. Gen. FI. Pl. 2 (1967) 148; Extras, J. Arn. Arb.
51 (1970) 37, f. 2; SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973) 420. — Sponia
COMMERS. ex LAMK, Dict. 4 (1795) 138; ENDL. Gen. Pl. 4 (1837) 276; PLANCH.
Ann. Sc. Nat. II, 10 (1848) 264; in DC. Prod. 17 (1873) 195. — Fig. 13, 16-17.

Trees or shrubs, often buttressed and with spreading and drooping branches,
monoecious. Jnnovations variously and densely set with simple bulbose-based hairs
or/and with short multicellular capitate-glandular hairs. Terminal buds ovoid-
conical, enclosed by overlapping but free extrapetiolar, caducous stipules. Leaves
penninerved ; above + glabrous, variously scabrate, beneath glabrous, subglabrous,
or variously densely set with bulbous-based hairs and/or with short multicellular
glandular hairs; base triplinerved, cordate to acute, often unequal-sided; margin
variously serrate or denticulate; apex acute to acuminate or caudate; petiole sul-
cate. Inflorescence axillary, paniculate or thyrsoid, many-flowered, condensed or
lax at anthesis, 3, 2, g2, densely and variously pubescent; bracts minute, ovate-
acute, caducous. — ¢ Flower globular; perianth 4~S-lobed, lobes induplicate-
valvate in bud, boat-shaped; stamens glabrous, introrse; filament subulate, glab-
rous, incurved in bud; anthers subglobular to reniform, glabrous, dorsifixed near
the base; pistillode present, hirsute at base. — 9 Flower ovoid; perianth 4~5-lobed;
staminodes absent or very rarely present; ovary ovoid, (in Mal.) glabrous, slightly
compressed, sessile; style short; ovule ana- to amphitropous. Drupe ovoid or sub-
globose, (in Mal.) slightly compressed, glabrous; exocarp fleshy and fibrous,
endocarp stony and very hard. Seed with a rather scanty or copious endosperm;
embryo curved or nearly involute; hypocotyle ascending; cotyledons equal.
Germination epigeal.

Distr. About 10-15 spp., widely distributed throughout the tropics and subtropics. In Asia (with 6-7
spp.) from the warmer parts of the Himalayas, extending north-eastwards to China (incl. Hainan, Hong-
kong, Formosa) and S. Japan and south and south-eastwards through India, Burma, Thailand, Indo-
China, and Malesia to the tropical and subtropical parts of Australia and the Pacific islands as far east as
Tahiti (31° N-37° S). In Africa (with 3-4 spp.) it occurs south of the Sahara to S. Africa and Madagascar
(22° N-28° S). In America (with 4-5 spp.) the genus is known from Central & S. Florida and Mexico,
extending south-eastwards through Central America, Bermuda, and the Bahamas, the Greater Antilles
and southwards to South America as far south as the northern parts of Argentina (26° N-25° S). In
Malesia: 4 spp., widely spread. Fig. 14, 15.

Ecol. Throughout its range of distribution the genus seems to grow well and often gregariously in
newly opened up habitats on various types of soils ranging from heavy laterite to limestone soils and soils
derived from volcanic ash (fig. 17), from sea-level up to 2000 m.

Pollination is probably affected by wind and small insects.

The fruits which turn orange, red or black when ripe are dispersed by various species of bulbuls.

In East Java CLASON (Bull. Jard. Bot. Btzg III, 13, 1935, 509, f. IIT) reported that following the frequent
eruptions of Mt Kelud, Trema spp. together with Parasponia spp. formed a dominant association in the
regrowths on volcanic ash.

Taxon. The genus is homogeneous and closely related to Parasponia and Celtis. This is corroborated
by the anatomy of the wood and leaves. Reports on the cytology are, however, suggesting that the number
of chromosomes is not constant. svit

Embryology. Very little is known about the sporogenesis and embryogenesis of the genus. A preliminary
study carried out recently on Trema cannabina and T. tomentosa in the Malay Peninsula indicates that the

development of the anther and microspores follow the so-called dicotyledon-type, and that of the embryo-
sac conforms with the Polygonum-type.

48 FLORA MALESIANA [ser. I, vol. 8?

en
exer

‘

Fig. 13. Trema orientalis (L.) BL. a. Habit, with functionally 3 flowers, x 2/3, b-e. 3 flowers at various

stages of development, all x 8, f. 2 inflorescence, x 2/3, g. 2 flower, h. older 2 flower, 2 tepals removed,

i. ditto in LS, j-k. mature fruit, |. ditto, exocarp halfway removed, showing stone, m. ditto in LS,n.embryo,
0. detail of lower leaf surface, all x 8 (a-e BW 13889, fi BW 7019, j-n Brass 6496).

1977] ULMACEAE (Soepadmo) 49

meets

q84q
Wi!
Nt

Ss
Bw
=

AN :

9

Fig. 15. Species density of Trema Lour. in Indo-Australia; above the hyphen the number of endemic spp.,
below it the number of non-endemic spp.

Chromosomes. A few counts on the chromosome number which have been reported by various cytolo-
gists suggest that cytogenetically the genus is rather variable. In Trema politoria from India n = 10+B
(Mewra & GILL, Taxon 17, 1968, 574; J. Arn. Arb. 55, 1974, 663); in T. orientalis n = 18 (ArorA, Bull.
Bot. Surv. India 2, 1960, 305), or n = 20 (GaAJAPaATHY, ibid. 3, 1961, 49; Hsu, Taiwania 13, 1967, 117), or
n = 10 (MeHRA & Hans, Taxon 18, 1969, 310; HANs, Cytologia 36, 1971, 341); and in 7. tomentosa
(cited as T. amboinensis) n = 10 or 80 (Hans, /.c.; MEHRA, Nucleus 15, 1972, 64).

Specific delimitation has proved to be difficult and has led to more than 50 names in the genus. This was
partly due to the various interpretations of the early described species. There is still no unanimity of
opinion about the number of good species in the continents. In Africa, for example, ENGLER (Pfl. Welt Afr.
3 (1), 1915, 11) estimated the number for Africa at 5-7, following BLUME (1856), but ReNDLE (1917) and
piemare th (Kew Bull. 19, 1964, 143) accept only one, either under the specific name 7. guineensis or T.
orientalis.

50 FLORA MALESIANA [ser. I, vol. 8?

In absence of a critical, reliable world monograph there is a similar uncertainty about the number of
species in the neotropics and in Indo-Malesia. For Malesia out of 20-25 published names of species and
varieties, only 4 spp. are recognized here.

The proliferation of name giving in Malesia is mainly due to the fact that Trema spp. have a growth
habit of continuously producing lateral and terminal new shoots on which flowers and fruits are borne.
Many specimens collected were from these young shoots in which the indumentum and leaf-shape is often
different from that of mature leaves. For accurate identification leaves, inflorescences, and fruits of mature
specimens are essential. Besides, the indumentum was in earlier descriptions mostly derived from low
magnification observations, but to differentiate sterile material of T. orientalis and T. tomentosa the
difference in the indumentum becomes only clear under at least 40 x magnification. It is impossible to
name young sterile specimens.

KEY TO THE SPECIES
(Based on mature leaves, inflorescences, and fruits)

1. Petiole (7—)10-15(-25) mm. Leaves broad ovate-elliptic, rarely narrow lanceolate (but then either
glabrous or silvery brown tomentose beneath), (3—)5—10(—13) by (11/.-)2-4(-5'/,) cm (and then glab-
rous or sparsely pubescent beneath) or (5—)10—15(-19) by (2—)3-7(—10) cm. Inflorescence 3, 2, or $9,
at anthesis lax, 10—100-flowered, 11/,-5 cm long. Mature fruit 3-5 by 2-4 mm.

2. Leaves (3—)5—10(-13) by (11/,-)2-4(-5'/,) cm, glabrous or rarely sparsely pubescent beneath, char-
taceous to thin-coriaceous; base rounded, rounded-attenuate, truncate, or very rarely subcordate,
mostly symmetrical; nerves 2-4(-5) pairs. Inflorescence always with a slender axis. Mature fruit
c. 3 by 2 mm, orange or red in colour 1. T. cannabina

2. Leaves (S—)10-15(-19) by (2-)3-7(-10) cm, densely and variously hairy beneath, thin- to thick-
coriaceous; base cordate, subcordate or rounded, mostly asymmetrical; nerves 4-8 pairs. Inflores-
cence with a stout or slender axis. Mature fruit black, 3-5 by 3-4 mm.

3. Leaves beneath (fig. 16) completely covered with matted glaucous to silvery straight appressed or
curly, erect hairs and short, multicellular capitate-glandular hairs (epidermis invisible even under
high magnification), very often distinctly discolorous with the upper surface darker in colour and
weakly scabrate. 3 Inflorescence up to 2!/, cm. Fruit + globular. ....... 2. T. orientalis

3. Leaves beneath (fig. 16) densely or sparsely (but not completely) set with velvety greyish-brown erect
hairs only (epidermis clearly visible between the hairs even under low magnification), mostly con-
colorous, dark-brown to blackish in dried specimens; upper surface strongly scabrate. ¢ Inflores-

ea oe te eS. eee Eee? de) ie neem, Sele Bem ld

cence up to 5 cm. Fruits ovoid, compressed .

CAG pay ese Me ely sy pf CS ae) e: .

T. tomentosa

1, Petiole (2-)3-6 (-8) mm. Leaves narrow ovate-lanceolate, (4-)5—6(-11) by (11/,-)2-3(-4) cm, beneath
densely tomentose by short, matted, rufous, 1-celled and multicellular glandular hairs (hoary).
Inflorescence $ or $9, at anthesis condensed and shorter or as long as petiole, 5—15-flowered. Mature

fruit subglobose, 2-3 by 2 mm.

1. Trema cannabina Lour. Fl. Coch. 2 (1790) 563;
Me_rr. Trans. Am. Phil. Soc. 24, 2 (1935) 131; DE
Wit, Bull. Bot. Gard. Btzg III, 18 (1949) 184;
Back. & BAKH. f. Fl. Java 2 (1965) 12; SoepADMO
in Whitmore, Tree Fl. Mal. 2 (1973) 421. — Celtis
amboinensis WILLD. Sp. Pl. 4, 2 (1805) 997. — Celtis
commersonii BRONGN. in Duperrey, Voy. Bot. Coq.
Phan. (1829) 215. — Sponia commersonii (BRONGN.)
Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834)
498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 317; in
DC. Prod. 17 (1873) 198. — Sponia timorensis
Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834)
498; PLANcH. Ann. Sc. Nat. III, 10 (1848) 318; in
DC. Prod. 17 (1873) 196; Mia. Fl. Ind. Bat. 1, 2
(1859) 216. — Sponia amboinensis (WILLD.)
Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834)
498, quoad nomen. — Sponia virgata PLANCH. Ann.
Sc. Nat. III, 10 (1848) 316; in DC. Prod. 17 (1873)
195, incl. var. major PLANCH. lI.c. 196; Miq. FI.
Ind. Bat. 1, 2 (1859) 216; Kurz, For. Fl. Burma 2
(1877) 469. — Sponia glabrescens PLANCH. Ann. Sc.
Nat. III, 10 (1848) 317; Mia. Fl. Ind. Bat. 1, 2
(1859) 217. — Sponia viridis PLANCH. Ann. Sc.
Nat. IIT, 10 (1848) 319. — T. commersonii (BRONGN.)
BL. Mus. Bot. 2 (1856) 60. — T. timorensis (DECNE)
BL. /.c. 60; Hoox. f. Fl. Br. Ind. 5 (1888) 483;
Hems-. J. Linn. Soc. Bot. 26 (1894) 452; Laur.
Bot. Jahrb. 50 (1913) 317, incl. var. carinata (BL.)

4. T. angustifolia

Laut. et var. pallida (BL.) LAuT.; HAND.-MAZz.
Symb. Sin. 7 (1929) 107. — T. virgata (PLANCH.)
BL. Mus. Bot. 2 (1856) 59; J. J. SMirH in K. & V.
Bijdr. 12 (1910) 652; Laur. Bot. Jahrb. 50 (1913)
313, incl. var. pubigera (BL.) Laut. lc. 315;
SCHNEIDER in Sargent, Pl. Wils. 3 (1917) 289;
RIDL. Fl. Mal. Pen. 3 (1924) 319; GAGNepP. Fl. Gén.
I.-C. 5 (1927) 686; HAND.-Mazz. Symb. Sin. 7
(1929) 106; CorNER, Ways. Trees (1940) 694; Li,
Woody FI. Taiwan (1963) 109. — T. amboinensis
(WILLD.) BL. Mus. Bot. 2 (1856) 61, guoad nomen,
excl. syn. et sched.; MERR. Int. Rumph. (1917) 187.
— T. viridis (PLANCH.) BL. Mus. Bot. 2 (1856) 58.
— T. glabrescens (PLANCH.) BL. /.c. 58. — T. cari-
nata BL. l.c. 59. — T. pallida BL. l.c. 60. — T.
pubigera BL. l.c. 60. — T. morifolia BL. l.c. 59;
Laut. Bot. Jahrb. 50 (1913) 318. — Sponia pallida
(BL.) Mia. Fl. Ind. Bat. 1, 2 (1859) 215; PLANCH.
in DC. Prod. 17 (1873) 196. — Sponia carinata
(BL.) Mia. FI. Ind. Bat. 1, 2 (1859) 215; PLANCH. in
DC. Prod. 17 (1873) 202. — Sponia pubigera (BL.)
Mia. FI. Ind. Bat. 1, 2 (1859) 216; PLANcH. in DC.
Prod. 17 (1873) 197. — Sponia morifolia (BL.)
PLANCH. in DC. Prod. 17 (1873) 196. — Sponia
vieillardii PLANCH. I.c. 201. — Sponia aspera var.
viridis (PLANCH.) BTH. Fl. Austr. 6 (1873) 158. —
T. orientalis var. amboinensis (WILLD.) KuRZ, For.
Fl. Burma 2 (1877) 469, guoad nomen. — T. vieil-

1977]

ULMACEAE (Soepadmo) 51

lardii (PLANCH.) SCHLTR, Bot. Jahrb. 36 (1905)
31.

Shrub or small much-branched tree up to 6m,
15cm @. Bark smooth, grey-brown. Branchlets
slender, spreading, often drooping, initially densely
silvery-hairy, glabrescent and sparsely lenticellate.
Stipules linear-lanceolate, 5-7 by 1-2 mm. Leaves
chartaceous to thin-coriaceous, narrow ovate-
caudate to broad ovate-acute, or elliptic-lanceolate,
(3-)5-10(-13) by (11/,-)2-4(-5"/,) cm (index 2-3
(4)), broadest below or at the middle; base
rounded to attenuate and acute, rarely subcordate,
slightly contracted and more or less symmetrical;
margin serrulate to denticulate for its entire length;
apex with a sharp tip; above glabrous and variously
scabrate, beneath glabrous or sparsely appressed-
hairy; midrib and nerves raised beneath, impressed
above; nerves (2—)3—4(—5) pairs, arcuate and sub-
parallel, basal ones running up to + 2/, the length
of the leaf; reticulations fine, subscalariform,
obscure to visible beneath; petiole (S—)8—12(-15)
by 1-2 mm, glabrescent. Inflorescence 3 or 39,
with slender axes, 10—15-flowered, at anthesis lax,
c. 1-2'/, cm long, densely greyish appressed-hairy ;
bracts ovate-acute, c. 2-3 by 1 mm. — ¢ Flowers
c. 1-2 mm @, outside sparsely hairy, glabrescent;
perianth lobes 4-5, membranous, oblong-lanceo-
late, c. 1-1'/, by */,-1 mm; filaments c. 1 mm, an-
thers c. 1 by !/, mm; pistillode obovoid, compressed,
c. 1 by '/, mm. — 9 Flowers c. 11/,-2 by 1-1'/, mm;
perianth lobes mostly 5, membranous, glabrous,
ovate-acute, c. 1—1"/, by !/, mm; staminode absent;
ovary c. 1 by '/, mm; stigmatic arms spreading or
incurved. Drupe 2-3 by 2 mm, turning deep-orange
or red when ripe. Endosperm copious.

Distr. Burma, China, Formosa, Hainan, Indo-
China, Thailand, common throughout Malesia to
Australia, Melanesia (Solomons, New Caledonia,
New Hebrides), W. Polynesia (Fiji, Samoa), and
Micronesia.

Ecol. Common as a pioneer in newly opened up
habitats along roadsides, edges of forests, re-
growths, thickets, and in young secondary vegeta-
tion, from sea-level up to 1200 m. FI. fr. Jan.—Dec.
At least in Malaya pollination is affected by wind
and by small insects (diptera). Ripe fruits are dis-
persed by various species of bulbuls.

Taxon. In Malesia there seem to be three rather
but not completely distinct entities. These can be

as follows: (i) specimens which have a
completely glabrous, chartaceous and narrow-
ovate leaf of (3—)5—8(—10) by (1!/,-)2-3(-4) cm with
an index of 2'/,-3, more or less non-scabrate upper
surface, and 2-3 lateral nerves (7. cannabina and
T. virgata); (ii) specimens with a thin-coriaceous,
narrow ovate-lanceolate to elliptic-lanceolate leaf
of (6-)7-10(-12) by (1'/,-)2-3(-3'/,) cm, with an
index of 4-5, slightly scabrate upper surface and
sparsely hairy lower surface, and 4-5 lateral nerves
which are straight and ascending and forming a
narrow angle (less than 40°) with the midrib (7.
timorensis, T. virgata var. scabra, and T. cannabina
var. scabra) ; (iii) those with a broad ovate and coria-
ceous leaf of (8—)9-11(-13) by (3-)4-4'/,(—5'/,) cm,
with an index of 2'/,-3, rugose and slightly scab-
rate upper surface and sparsely pubescent beneath,
and 3-4 lateral nerves forming a broad angle (more
than 45°) with the midrib (7. glabrescens, T. viridis,
and T. vieillardii). Various intermediates are how-

ever present, making it difficult to recognize them
as distinct infra-specific taxa.

Vern. Malaya: ménérong, méngkirai, M; S.
Sumatra: délung, M, Palembang; Java: anggrung,
J; N. Borneo: bintanong, Murud; Lesser Sunda Is. :
rédong kué, Flores, pépaka, Alor: Moluccas:
loli-sawu, Halmaheira; Solomons: bulasisi, Kwara.

2. Trema orientalis (L.) BL. Mus. Bot. 2 (1856) 62;
Bru. FI. Austr. 6 (1873) 158; Hook. f. Fl. Br. Ind. 5
(1888) 484; J. J. SmMirH in K. & V. Bijdr. 12 (1910)
655, p.p., excl. syn. T. commersonii et T. griffithii;
Laut. Bot. Jahrb. 50 (1913) 320, p.p., incl. var.
rigida (BL.) Laut. l.c. 322, excl. var. viridis et var.
amboinensis; CORNER, Ways. Trees (1940) 694,
pl. 211, p.p. excl. syn.; DE Wit, Bull. Bot. Gard.
Btzg III, 18 (1949) 189, p.p., incl. var. bicornis DE
Wir, /.c. 190, excl. var. bicolor et syn. T. angusti-
folia, T. imbricata et T. velutina; BACK. & BAKH. f.
Fl. Java 2 (1965) 12, p.p., excl. syn. T. amboinensis
auct. non (WILLD.) 'BL.; SOEPADMO in Whitmore,
Tree Fl. Mal. 2 (1973) 421. — Celtis orientalis
LINNE, Sp. Pl. 2 (1753) 1044; Roxs. Fl. Ind. ed.
Carey 2 (1832) 65. — Celtis rigida BL. Bijdr. (1825)
486. — Celtis discolor BRONGN. in Duperrey, Bot.
Voy. Coq. Phan. (1829) 215, pl. 47B. — Sponia
discolor (BRONGN.) DECNE, Nouv. Ann. Mus. Hist.
Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III,
10 (1848) 324; in DC. Prod. 17 (1873) 201. —
Sponia orientalis (L.) DECNE, Nouv. Ann. Mus.
Hist. Nat. III, 3 (1834) 498; PLANCH. Ann. Sc. Nat.
III, 10 (1848) 323; in DC. Prod. 17 (1873) 200. —
Sponia rigida (BL.) DECNE, Nouv. Ann. Mus. Hist.
Nat. III, 3 (1834) 498; PLANcH. Ann. Sc. Nat. III,
10 (1848) 336; Mra. FI. Ind. Bat. 1, 2 (1859) 217. —
Sponia argentea PLANCH. Ann. Sc. Nat. III, 10
(1848) 323; in DC. Prod. 17 (1873) 201. — Sponia
wightii PLANCH. Ann. Sc. Nat. III, 10 (1848) 322;
WicuT, Ic. 6 (1853) t. 1971. — T. argentea
(PLANCH.) BL. Mus. Bot. 2 (1865) 58. — T. bur-
mannii BL. I.c. 62. — T. rigida (BL.) BL. /.c. 61. —
T. scaberrima BL. l.c. 63. — T. wightii (PLANCH.)
BL. /.c. 58. — T. discolor (BRONGN.) BL. /.c. 58;
Laut. Bot. Jahrb. 50 (1913) 319. — Sponia scaber-
rima (BL.) Mig. Fl. Ind. Bat. 1, 2 (1859) 217;
PLANCH. in DC. Prod. 17 (1873) 202. — Sponia
burmannii (BL.) PLANCH. in DC, 17 (1873) 200. —
Fig. 13, 16.

Shrub to large tree, 3-36 m, 10-90 cm @. But-
tresses, if present, up to 1'/,m. Bark smooth to
finely fissured, lenticellate, grey-brown or whitish-
grey. Branchlets, stipules, petioles, and inflores-
cences densely set with appressed and matted or
erect silvery to glaucous 1-celled hairs and short
multicellular glandular hairs. Stipules linear-
lanceolate to ovate-acute, 3-4 by 1-2 mm. Leaves
thin- to thick-coriaceous, often rigid and brittle,
ovate, ovate-lanceolate to narrow elliptic, lan-
ceolate, (6-)10—-15(-18) by (1'/,-)2'/,-6(-10) cm,
index (2-)3—4(—5'/,), broadest at or mostly below
the middle, mostly discolorous, above dull grey-
brown or grey-green in dried specimens, scabrate
and sparsely set with bulbous-based hairs, beneath
densely oamsds by a combination of silvery,
glaucous or grey-brown, appressed 1-celled hairs
and shorter multicellular glandular hairs (fig. 16);
base cordate, rounded, or sometimes truncate,
often contracted, asymmetrical or symmetrical ;

margin serrate to denticulate for its entire length;

52

FLORA MALESIANA

[ser. I, vol. 8?

apex acute to acuminate-caudate; midrib and
nerves raised beneath and impressed above; nerves
4-6(-8) pairs, the lowest pair arcuating and run-
ning up to !/,—*/, the length of the leaf; reticulations
subscalariform to subareolate, sometimes strongly
raised and distinct beneath; petiole (7—)10—15(-18)
by 1-2 mm, densely short pubescent. Inflorescences
either ¢ or 2 borne on separate vegetative branches,
a much-branched panicle or thyrse, at anthesis
lax or condensed, axes 1-2 mm thick; bracts ovate-
acute, 2-3 by 1mm. Flowers 5-merous. — ¢
Inflorescences up to 3—5 cm long, 20-100-flowered ;
3 flower c. 11/,-2 mm @; perianth lobes ciliate,
11/,-2 by 1mm; filaments 1-1'/,mm, anthers
c. 1 by 1/, mm, pistillode obovoid-conical, com-
pressed, 1-1!/, by '/,-1 mm. — 2 Inflorescences
5-15-flowered, 11/,-2!/,cm long, axes 1-2 mm
thick; 2 flower c. 2-3 by 1-2 mm; perianth lobes
ovate acute, c. 1-1!/, by 1/, mm, ciliate and densely
short pubescent, glabrescent; staminode absent;
ovary ovoid-conical, c. 2 by 1 mm; stigmatic arms
slender, c. 1-11/,mm., spreading. Drupe 3-5 by.
2-4mm, turning black when ripe. Endosperm
scanty to copious.

Distr. ?Tropical Africa, SE. Asia (Ceylon,
India: from W. Himalayas to Bombay and Mala-
bar; Burma, Thailand, Indo-China, China, also
Hainan, Formosa, to S. Japan), through Malesia
to Queensland, Melanesia (Solomons), Micronesia
(Marianas), and Polynesia (Fiji, Tonga, Tahiti).
In Malesia: Malay Peninsula and Sumatra (rather
rare), Java (rather common in the hills and sub-
montane regions), Lesser Sunda Islands (rare),
Borneo (common), Philippines (rather common),
Celebes (rather rare), Moluccas (rare), and New
Guinea (incl. New Britain, rare).

Ecol. In W. Malesia and continental Asia the
species is more common in the hills and montane
regions between 600-2000m, whereas in E.
Malesia, Australia, and Pacific Islands it is more
common in the lowlands. The ripe fruits which
turn to deep purple or black are dispersed by,

~\.s Le . i — —"

various species of birds, particularly bulbuls.
Fl. fr. Jan.—Dec.

Taxon. Three rather but not completely distinct
entities may be recognized. These are: (i) specimens
from continental Asia and W. Malesia which have
been variously identified as T. orientalis, rigida,
argentea, and wightii by previous authors. They
are characterized by: thick-coriaceous, broadly
ovate to ovate-elliptic leaves with grey-brown to
glaucous indumentum, slightly asymmetrical to
symmetrical cordate, subcordate or rounded base,
rugose upper surface, and acute to acuminate
apex; and by a relatively larger fruit of c. 4-5 by
3-4 mm and stouter inflorescence axes.

(ii) Specimens from S. Japan, Formosa, Hainan,
the Philippines, New Guinea, Micronesia, Mela-
nesia, and Polynesia, and Australia, which have
been included in the so-called T. discolor, charac-
terized by: thin-coriaceous, narrow-ovate leaves
with strongly asymmetrical cordate base, hardly
scabrate upper surface, short and matted silvery to
grey-brown indumentum, lax inflorescence with
slender axes, and fruits c. 3-4 by 2-3 mm.

(iii) A few specimens from scattered localities in
S. China, Thailand, Sumatra, and Borneo, which
have been described by DE Wir (1949) as T. orien-
talis ssp. bicornis, characterized by: very narrow,
ovate-lanceolate thin-coriaceous leaves with silvery
appressed and matted dense indumentum on the
lower surface and non-scabrate upper surface,
6-8 pairs of nerves, and the shorter and few-
flowered inflorescence.

Several intermediates are present however,
making formal infraspecific distinction not ad-
visable.

Vern. ndérung, Karo,

Sumatra: endrung,

indarung, Pajakumbu, bandorung, Tapanuli, lando-
jung, Simelungun, éndélung, Palembang, Bencoo-
len, magélong, nélung, Bencoolen, nériung, Lam-
pong. Java: gorai, kuraj, S, anggrung, njampu, J.
Lesser Sunda Is.: /énggung, Bali, rédong, Flores,
tabélah,

W. Sumbawa. Borneo: randagong,

x */ a
- ; ‘i oS

*
Dey La

Fig. 16. Indument of lower leaf-surface, strongly enlarged. Left: only unicellular hairs of Trema tomentosa

(Roxs.) HARA, between which the epidermis is visible. Right: 7. orientalis (L.) BL., with long unicellular
hairs and crowded multicellular crisped hairs covering the epidermis.

1977]

tandago, Dusun; béngkirai, E. Kutai. Philippines:
anadgong, Bis. Celebes: ngawoi, Malili, mawa,
Bonthain, kantu, Toradja, tajapu. Moluccas: rufu,
Ternate, /aei, Tidore, soka soka, E. Ceram. West
New Guinea: bésuwai, Hattam lang., karara,
Ambai, kKaniem, mier, Kebar lang.

3. Tremia tomentosa (RoxB.) HARA, Fl. E. Himal.
2 (1971) 19; So—PADMo in Whitmore, Tree Fl. Mal.
2 (1973) 423. — Celtis orientalis (non L.) BL. Bijdr.
(1825) 485. — Celtis amboinensis (non WILLD.)
BRONGN. in Duperrey, Bot. Voy. Coq. Phan.
(1829) 212, pl. 47A, p.p., excl. specim. ex Ventenat,
Amboina. — Celtis tomentosa Roxs. Fl. Ind. ed.
Carey 2 (1832) 66. — Sponia amboinensis (WILLD.)
Decne, Nouv. Ann. Mus. Hist. Nat. III, 3 (1834)
498, quoad specim.; PLANCH. Ann. Sc. Nat. III, 10
(1848) 321; Mia. Fl. Ind. Bat. 1, 2 (1859) 216;
PLANCH. in DC. Prod. 17 (1873) 198. — Celtis lima
(non Sw.) BLANCO, FI. Filip. 2 (1837) 139.— Sponia
griffithii PLANCH. Ann. Sc. Nat. III, 10 (1848) 324.
— Sponia tomentosa (ROxB.) PLANCH. /.c. 336. —
Sponia velutina PLANCH. I.c. 327, p.p., excl. specim.
Cuming 1232 ex Luzon. — Sponia blancoi PLANCH.
l.c. 327; Mia. Fl. Ind. Bat. 1, 2 (1859) 218. — 7.
griffithii (PLANCH.) BL. Mus. Bot. 2 (1856) 58. —
T. blancoi (PLANCH.) BL. /.c. 58. — T. imbricata
BL. /.c. 63. — T. velutina (PLANCH.) BL. l.c. 58;
GaGnepP. Fl. Gén. I.-C. 5 (1927) 689; L1, Woody FI.
Taiwan (1963) 109. — T. amboinensis (WILLD.)
BL. Mus. Bot. 2 (1856) 61, quoad specim.; BTH. FI.
Austr. 6 (1873) 159; Hook. f. Fl. Br. Ind. 5 (1888)
484; K. Sco. & Laut. Fl. Schutzgeb. (1900) 264;
J. J. SmirH in K. & V. Bijdr. 12 (1910) 659, p.p.,
excl. syn. Celtis amboinensis WILLD. et Trema
burmannii Bi.; Merr. En. Born. (1921) 217;
Ript. Fl. Mal. Pen. 3 (1924) 319. — Sponia
imbricata (BL). PLANCH. in DC. Prod. 17 (1873)
199. — T. orientalis var. amboinensis (WILLD.)
Kurz, For. Fl. Burma 2 (1877) 469, quoad specim.;
Laut. Bot. Jahrb. 50 (1913) 321. — T. orientalis
(non (L.) BL.) MERR. Sp. Blanc. (1918) 121. — T.
dielsiana HAND.-MAzz. Symb. Sin. 7 (1929) 106;
P’e1, Bot. Bull. Ac. Sin. 1 (1947) 289. — Fig. 16.
Shrub to medium-sized tree of 5—15(—24) m,
5—30(-50) cm @. Bark grey-brown, smooth to
finely fissured, lenticellate. Branchlets, inflores-
cences, petioles, stipules, and underside of leaves
densely and thickly set with greyish, erect, velvety
hairs. Stipules linear-lanceolate, c. 5 by 1 mm.
Leaves thin- to thick-coriaceous, broadly ovate to
ovate-elliptic, (5—)8-15(-19) by (2-)4~7(-9) cm,
index 2'/,-3, broadest mostly below the middle;
more or less concolorous, drying dark-chocolate
brown to blackish brown; above strongly scabrate;
base cordate, rarely subcordate or rounded, mostly
strongly asymmetrical, rarely symmetrical; margin
serrate throughout, apex acute to acuminate-
caudate, acumen sharp, 1-3 cm; midrib and nerves
raised beneath (often very strongly), impressed and
hairy above; nerves pairs, ascending and
solicavatiel, at an angle of + 45°, the lowest pair
running to + */,/, the length of the leaf ; reticu-
lations subscalariform to subareolate, often rather
distinct beneath; petiole 1-1'/,cm by 1-2 mm,
densely pubescent. Inflorescences 3,9, or 39, either
on the same or on different vegetative branches;
bracts ovate-acute, c. 1 by '/, mm. — At anthesis
dé and 3° axes of the inflorescences lax, 2'/,-4'/, cm

ULMACEAE (Soepadmo) 53

long, 20—100-flowered; 3 flower c. 1'/,2mm @;
perianth lobes mostly 5, elliptic, c. 11/, by 1 mm;
filaments c. 1 mm, flat, glabrous, anthers c. 1 by
‘/,mm; pistillode obovoid-ellipsoid, compressed,
1'/, by '/, mm. — 2 Inflorescence 1-2 cm long, axes
1-2 mm thick, 5—15-flowered; 2 flower c. 2 by
1 mm; perianth lobes 4-5, ovate-acute, c. 1 by
‘/,mm; staminode mostly absent, if present
strongly reduced in size and non-functional; ovary
c. 1"/, by '!/,-1 mm, stigmatic arms slender, c. 1 mm,
spreading. Drupe c. 3 by 2mm, maturing black.
Endosperm copious.

Distr. East tropical Africa, Madagascar, SE.
Asia: Pakistan, India, Bangladesh, Burma, Thai-
land, Indo-China, China (incl. Hainan), Hong-
kong, Formosa, Ryu Kyu Is. (Okinawa), through-
out Malesia to Queensland, Melanesia (New
Caledonia), Micronesia, and Polynesia (Fiji,
Tonga, and Hawaii).

Ecol. Common in the lowlands and hills, at sea-
level up to 1000 m, as a pioneer plant invading and
occupying newly opened up habitats on all kind of
soils, including limestones. F/. fr. Jan.Dec. At
least in Malaya pollination is affected by wind and
small insects (diptera). The ripe black fruit is dis-
persed by various species of birds.

Taxon. Evidently, 7. tomentosa is closely allied
to T. orientalis, and it is possible that, when more
field data become available in the future, the former
may prove to be only representing a juvenile
ontogenetical form of the latter. Except for a few
specimens from the Philippines (e.g. WHITFORD
681, BS 37313, 48355, ELMER 8417) and from New
Guinea (e.g. ANU 2075, 2752, 6240, HARTLEY
10937, MANNER & STREET 270, NGF 29353,
SCHODDE 1419, and BW 16510) in which the leaves
are thick-coriaceous and with a more or less sym-
metrical base and pale grey-brown in colour,
specimens of 7. tomentosa can be easily dis-
tinguished from those of T. orientalis by the charac-
ters mentioned in the key. Fig. 16. It is also interest-
ing to note that according to HANs (Cytologia 36,
1971, 341) and MEHRA (Nucleus 15, 1972, 64) the
chromosome number in T. tomentosa is n = 10 or
80, whereas that of T. orientalis isn = 10, 18, or 20.

Vern. Malay Peninsula: ménarong, méndarong,
méngkirai, M. Sumatra: béngkirai, Gajo, éndélung,
Palembang, eémaha, Enggano, hana(w)e, Batak,
kamésén silai, Simalur, mangkirai, Pajakumbu,
manghirei, ménkirei, Lingga, mudén sabu, Djambi,
(n)dér(r)ung, Karo-Batak, randurung, Toba, sang-
kiraja, Batak, tindjau, Riouw. Java: anggrung, J,
kuraj, k. awéwena, S. Lesser Sunda Is.: rédong,
damot, Flores, ruka parak, Sumba. Borneo: Sara-
wak: murieng, Bidajuh, kérénénog, Iban, tuku baroh,
Land Dayak; N. Borneo: anjalakat, Kedayan;
Brunei: balék balék angin jantan, balik angin,
rundagong, Brunei, bintanong, Murut, damai,
Suluk, entimon, Iban, lindagong, Kedayan, landa-
gong, ‘Dusun Tambato & Kayan, /undagong, sali-
muak, Dusun, randagong, Tenggara, réndagong,
Dusun Labuk; E. Borneo: bangérai, bangkirai,
tjalundung, E. Kutai; W. Borneo: éngkirai, butu.
Philippines: anaginong, Mang., anugdon, Tag-Bis.,
anabiong, hanagdong, Tag., ‘arangyan, karayang-
yang, Tagb. Moluccas: mandalirung’ a, Talaud,
pohon rupong, Banda, rufut, Buru. West New
Guinea: fidukwa, Manokwari, hormas, Sorong;
East New Guinea: komukai, Maring name, natua,

54 FLORA MALESIANA

|
Fig. 17. Young blukar (regrowth or secondary forest) on an abandoned tea estate near Tapos, West Java,

c. 1000 m, consisting of three layers: 2 m high stand of Eupatorium inulifolium, 5-6 m high tree ferns of
Cyathea contaminans, above which is an open canopy of Trema orientalis (L.) BL. (Photogr. VAN STEENIS).

Kainantu, seraun, Daga-Bonenau, wanip, Enga
lang., wantip, Medlpa, Wahgi, wan’um, Mendi.

4. Trema angustifolia (PLANCH.) BL. Mus. Bot. 2
(1856) 58; Hook. f. Fl. Br. Ind. 5 (1888) 484;
GAGNEP. FI. Gén. I.-C. 5 (1927) 686; HAND.-MAzz.
Symb. Sin. 7 (1929) 108. — Sponia angustifolia
PLANCH. Ann. Sc. Nat. III, 10 (1848) 326; Mia. FI.
Ind. Bat. 1, 2 (1859) 215; PLANcH. in DC. Prod. 17
(1873) 202. — Sponia acuminatissima MiQ. Sumatra
(1861) 410; PLANcH. in DC. Prod. 17 (1873) 202. —
Sponia sampsonii HANCE, Ann. Sc. Nat. V, 5 (1866)
242. — T. acuminatissima (MIQ.) BOERL. Handl. 3
(1900) 358. — T. lanceolata MerR. Lingn. Sc. J. 7
(1931) 302. — T. sampsonii (HANCE) MERR. &
CHUN, Sunyatsenia 5 (1940) 40. — T. orientalis var.
ut DE WIT, Bull. Bot. Gard. Btzg III, 18 (1949)

Shrub or small tree with spreading and drooping
branches, 3-7 m, 5-15 cm @. Branchlets densely set
with rufous multicellular glandular hairs and
glaucous short and matted 1-celled hairs, sub-
glabrescent. Stipules linear-lanceolate, 3-4 by
1mm. Leaves chartaceous to thin-coriaceous,
narrow ovate-lanceolate to lanceolate, (3—)5—8(-10)
by (1-)2-3(-4) cm, index 3-3/,, broadest below or
at the middle; discolorous, upper surface strongly
scabrate, dark chocolate-brown to blackish-brown,

[ser. I, vol. 82

lower surface densely set with short and matted
rufous to glaucous 1-celled and multicellular
glandular hairs; base rounded to attenuate, sym-
metrical; margin finely serrate throughout; apex
acute to acuminate; midrib and nerves slightly
raised beneath, impressed above; nerves 4-5 pairs,
straight, ascending at 30-40°; reticulations fine,
subscalariform to subareolate, obscure above and
faintly visible beneath; petiole (2—)3-6(-8) by
1 mm, densely short hairy. Inflorescences 3 or 39,
densely set with a short rufous indumentum, much-
branched, (5—)10-15(—30)-flowered, at anthesis
condensed, shorter than or as long as the petiole;
bracts narrow ovate-acute, c. 4/,-1 by 1/,—'/, mm.
— 6 Flowers c. 1-11/,mm @; perianth lobes 5,
elliptic, c. 1-11/, by 1 mm; filaments c. 1 mm long,
c.1/,-1 mm @; pistillode obovoid-ellipsoid, strong-
ly compressed, c. 4/,-1 by 1/,mm. — 9 Flowers
ovoid-conical, c. 2 by 1 mm; perianth lobes 5,
narrow-lanceolate, acute, 1/,-1 by 1/,—'/,mm;
staminode absent; ovary c. 1-2 by 1 mm; stig-
matic arms c. 1/,-1 mm, spreading or incurved.
Drupe c. 1/,-2 mm @, turning orange to red when
ripe. Endosperm copious.

Distr. China (Yunnan, Hainan), Thailand, and
Indo-China; in Malesia: Malay Peninsula (com-
ae Sumatra (rare), Borneo (rare), Celebes (very
rare).

1977] ULMACEAE (Soepadmo) 55

Ecol. Scattered in newly available habitats in Excluded
the lowlands to submontane regions, from sea-level : er
rate eae . Trema (Parasponia) lancifolia Ripv. J. Mal. Br.
to 1200 m. Fi. fr. Jan.—Dec. Ripe fruits are dis- R. As. Soc. 1 (1923) 91 = Debregeasia longifolia

persed by various species of bulbuls. ‘
Vern. Malaya: ménarong, méngkirai, M; (Burm. f.) WEDD. (Urticaceae).

Sumatra: kayu anggurung, M, Eastcoast, mang-
kirai kétjil, M, Palembang.

4.CELTIS

LINNE, Gen. PI. ed. 5 (1754) 467; Sp. Pl. 2 (1753) 1043; PLANCH. Ann. Sc. Nat. III,
10 (1848) 262; BL. Mus. Bot. 2 (1856) 70; Mia. Fl. Ind. Bat. 1, 2 (1859) 220;
PLANCH. in DC. Prod. 17 (1873) 168; B. & H. Gen. Pl. 3 (1880) 354; ENGL. in E. &
P. Nat. Pfl. Fam. 3, | (1888) 63; BERNARD, Bull. Herb. Boiss. II, 5 (1905) 1112,
maps 9-15; J. J. SMitH in K. & V. Bijdr. 12 (1910) 639; Leroy, Fl. Madag. et Com.
Fam. 54 (1952) 3; POLHILL, Kew Bull. 19 (1964) 139; Hutcu. Gen. FI. Pl. 2 (1967)
147; Extas, J. Arn. Arb. 51 (1970) 32; SOEPADMO in Whitmore, Tree Fl. Mal. 2
(1973) 414. — Solenostigma ENDL. Prod. Fl. Norf. (1833) 41; BL. Mus. Bot. 2
(1856) 66; Mia. Fl. Ind. Bat. 1, 2 (1859) 219. — Fig. 18, 20, 22-23.

Small to large monoecious or polygamo-monoecious trees, often buttressed.
Bark smooth or finely fissured, often conspicuously warty lenticellate. Branches
(in Mal.) unarmed, initially densely yellow-brown or rufous-hairy, glabrescent;
hairs l-celled. Buds enclosed by the overlapping stipules or naked. Stipules thick
and tough, peltately attached or free and scarious, caducous. Leaves entire dr not,
3-nerved at base, semideciduous or persistent. /nflorescenes 3, %, or 3%, branched
racemes or panicles, few- to many-flowered, axillary or subterminal on the new
shoot; staminate inflorescences borne on the lower and leafless part or in the axil of
leaves of the new shoot; in the ¢% inflorescence the &% flowers are borne on the
distal ends of the axes; bracts minute, caducous. — Staminate (3) flowers globular,
pedicelled or sessile; perianth lobes 4-5, imbricate in bud, membraneous, boat-
shaped, outside sparsely pubescent, at anthesis recurved, caducous; stamens
glabrous, inserted on the densely pilose receptacle; filaments subulate, incurved in
bud and spreading elastically, exserted at anthesis; anthers ovoid to subreniform,
dorsifixed just above the emarginate base, extrorse; pistillode present or absent. —
% Flowers ovoid, pedicelled; perianth lobes 4-5, imbricate in bud, connate at base,
membranous, outside sparsely pubescent, boat-shaped, at anthesis recurved, cadu-
cous; stamens well-developed and functional or rudimentary, other characters as in
3 flowers; ovary ovoid-ellipsoid, sessile, style short or + absent; stigmatic arms
elongate, divergent, the tips entire to deeply bifid; ovule anatropous. Drupe fleshy,
ovoid, ellipsoid or globose; exocarp thick and firm, mesocarp thin and fleshy, con-
taining slimy substances; endocarp hard and persistent, smooth or variously ridged
or pitted. Seed: coat membranous, chalazal area broad, dark-coloured and close to
the minute hilum; endosperm scanty or wanting, oily or gelatinous, nearly enclosed
between the folds of the cotyledons. Embryo curved, cotyledons broad, foliaceous,
equal or unequal in thickness, flat or conduplicate, variously folded, incumbent on
or embracing the short superior and ascending radicle. Germination epigeal.

Distr. About 50-60 PP: widely distributed in tropical and temperate regions of the world, the majority
of species (30-40) in the Old and New World tropics, throughout Malesia (9 spp.). Fig. 19, 21.

56 FLORA MALESIANA [ser. I, vol. 8?
a A eee

Fig. 18. Celtis philippensis BLANCO. a. New shoot bearing flowers,
x 10, f. fruits, x 2/5, g. fruit, exocarp in LS, x 11/3,
and chalaza, nat. size, j. ditto in LS showing folded
in LS, x 1*/;. — C. tetrandra Roxs. 1. New shoot bearing flowers, x 2/3,

flowers, x 10 (a—c MERRILL 52, d-e BS 1920, f-k BARTLETT 15071, -o Scumutz 1666).

x 2/3, b-c. 3 flowers, x 8, d-e. 3 flowers,
h. stone, endocarp in LS, x 11/3, i. seed showing testa
cotyledons, shaded oily endosperm, x 1'/3, k. embryo
m-n. § flowers, x 8, 0. ¢.

1977] ULMACEAE (Soepadmo) 57

Fig. 19. Approximate range of Celtis L. Fossil records are indicated by dots.

Fossils. Numerous fossilized wood-fragments, leaf-impressions, drupes and pollen grains have been
discovered in various localities in Europe, the U.S.A., and in Asia. According to ELtas /.c. the first
(oldest) records are apparently from the early Eocene in Wyoming and the late Eocene in Georgia, U.S.A.
Continuing through the Oligocene, Celtis spp. are best known from the Miocene, fossilized material of
younger age is relatively sparse at least in the U.S.A. — References: A. GRAHAM (ed.), Floristics and
Paleofloristics of Asia & Eastern North America (1972) 147; GreGuss, Tert. Angios. Hung., Ak. Kiado
Budapest (1969) 83; La Motte, Mem. Geol. Soc. Am. 51 (1952) 112. Fig. 19.

Ecol. Malesian species of Celtis may be classified into two rather distinct ecological groups, i.e. the
C. philippensis var. philippensis and C. tetrandra groups. The first group, which includes also C. hilde-
brandii, C. latifolia, C. luzonica, C. paniculata, and C. rigescens, is found mainly in the lowland forests,
both primary and secondary, and is an important constituent of the understorey tree community in moist
areas. The C. tetrandra group, which includes C. rubrovenia, C. timorensis, and C. philippensis var. wightii,
is confined to areas which are subject to a rather pronounced seasonal climate, or if they occur in wetter
regions, they grow on strongly drained substrates, e.g. rocky shores, limestone, efc. In tune with this
environmental preference, the second group shows a more prominent flush-wise growth habit and its
species are completely or partly deciduous. In Malaya, species of Celtis are producing flowers around
July-September, while fruit ripens January—March.

How pollination is affected is not certain, but judging from the structure and position of the inflores-
cence, some species (e.g. C. philippensis var. philippensis, C. hildebrandii, and C. latifolia) may be pollinated
Hd = while others (C. tetrandra, C. timorensis, C. rubrovenia, and C. rigescens) may be pollinated

wind.

The ripe fleshy drupes which turn to orange, red or bluish-black may be dispersed by birds, or alter-
lose they may be dispersed by water as the embryo is protected by the hard, persistent and durable
endocarp.

Morph. Except for C. paniculata the stamens of the pistillate flowers are well-developed and functional.
In the male or staminate flowers the pistillode is rudimentary or completely absent in C. tetrandra,
C. timorensis, and C. rubrovenia. In the other species the pistillode is present and relatively rather well-
developed though non-functional.

Chromosomes. The chromosome numbers reported are: n = 10 (2n = 20) (C. australis var. eriocarpa,
C. inguana, C. laevigata, C. occidentalis, C. sinensis, and C. timorensis (under C. cinnamomea)); 2n = 22
(C. spinosa); 2n = 28(C. occidentalis);2n = 40(C. australis and C. tupalangi). — References: Sax, J. Arn.
Arb. 14 (1933) 82; Bowpen, Am. J. Bot. 32 (1945) 195; DARLINGTON & WyLie, Chromos. Atlas (1955)
182; MeHra & GILL, Taxon 17 (1968) 574; GADELLA et al. Acta Bot. Neer]. 18 (1969) 74; MEHRA & HANs,
Taxon 18 (1969) 310; Feporov (ed.), Chromos. Numbers Flow. Pl. (1969) 710; MeHra & GILL, J. Arn.
Arb. 55 (1974) 663. :

According to Sax I.c. there seems to be at least in C. occidentalis a high degree of pollen sterility and a
high incidence of meiotic irregularity. This may be one of the causes why in Celtis there is a very high
percentage of barren seeds production, even among tropical species. f

Embryology. No detailed study on the microsporogenesis, megasporogenesis and embryogenesis of
Aye aig has ever been carried out. In Malesia the solitary ovule is bitegmic, anatropous and inserted
just below the apex of the locule. After fertilization both integuments develop into thin membranous
seed coats with a broad, dark-coloured, more or less circular chalaza. The endocarp becomes woody and
very hard and impregnated by mineral deposits. It is persistent and becomes variously sculptured (ridged,

58 FLORA MALESIANA [ser. I, vol. 8?

pitted, or nearly smooth). The embryo is strongly curved with the hypocotyle superior and ascending,
situated in between or nearly enclosed by the broad, thick, foliaceous cotyledons. The cotyledonar lobes
are somewhat unequal in thickness, and they are either induplicate or variously folded. Endosperm is very
scanty to absent and either gelatinous or oily. Especially in C. paniculata and C. tetrandra, at least 70-80%
of the fruits produced are barren. Though the fruits are developed normally, the embryo fails to grow and
becomes shrivelled. As a result the fruits are empty.

KEY TO THE SPECIES
(Measurements of leaf and fruit based on fully mature material)

1. Leaves entire or nearly so. ¢ Inflorescence a much-branched many-flowered panicle with up to 150

flowers. Pistillode rather well-developed, c. 1—-11/, by 1/,-1 mm. Stigmatic arms bilobed or bifid at the

tip.

2. Leaves rugose, brittle, sparsely pubescent beneath; midrib and lateral nerves strongly raised beneath.
Stipules not peltately attached, free from one another. Fruit densely appressed-hairy. 1. C. rigescens
2. Leaves not rugose, not brittle, glabrous; midrib and nerves only slightly raised beneath. Stipules

peltately attached, overlapping. Fruit glabrous.

3. Leaves with (2—)3—5 pairs of nerves.

4. Leaves elliptic-orbicular or elliptic-oblong, index (11/,—)1*/,(—2); midrib and nerves slightly raised
beneath; lowest pair of nerves running to 7/,—*/, the length of the leaf, upper pairs of nerves ascend-
ing and arcuating. Stigmatic arms shallowly bilobed at the tip. Fruit globose, 16-20 by 14-18 mm.

2. C. luzonica

4. Leaves elliptic or ovate-eliptic, index (11/,—)2—2'/,(—3); midrib and nerves flattish beneath; lowest
pairs of nerves running up to !/;—'/,(—?/,) the length of the leaf; upper pairs of nerves weak, sub-
horizontal. Stigmatic arms deeply bifid at the tip. Fruit ovoid or ellipsoid, 7-12 by 5-8 mm.

5. Nerves 3-5 pairs; lowest pair running to '/;-'/, the length of the leaf. Inflorescence mostly d or ¥.
Stamens of § flower rudimentary, non-functional. Fruit ovoid, feebly 4-5-angular in CS; endo-

carp with reticulate ridges... .... .

PES ee ee 3. C,

paniculata

5. Nerves 1—2(—3) pairs; lowest pair running up to 2/, the length of the leaf. Inflorescence g or ¢ 4.
Stamens of % flower well-developed and functional. Fruit ellipsoid, + terete; endocarp smooth.

3. Leaves with 1 pair of nerves.

4b;).C. philippensis var. wightii

6. Leaf symmetrical. Inflorescence ¢ or 38. 3 Inflorescence 15—-40-flowered. Cotyledons curved but

not folded.

7. Leaves (4-)8-14(-18) by (2—)3—6(-8) cm; reticulations fine, dense. Ovary glabrous. Fruit 8-15 by

7-12 mm; endocarp smooth

4a. C. philippensis var. philippensis

7. Leaves (8-)15—18(—25) by (6—)8—-12(-18) cm; reticulations coarse, wide-spaced. Ovary densely

appressed-pubescent. Fruit 15-25 by 10-18 mm; endocarp pitted. .....

5. C. latifolia

6. Leaf oblique. Inflorescence f and %. 3 Inflorescence 60-150-flowered. Cotyledons cure and many
6.

times folded.

hildebrandii

1. Leaves serrulate to denticulate at least in the upper half. g¢ Inflorescence racemose, 5—20-flowered.
Pistillode strongly reduced in size or absent. Stigmatic arms entire.
8. Leaf not strongly oblique in outline; nerves 1—2(—3) pairs. Inflorescence 3 and 3%; 3 flower 5-merous.

9. Leaves (6-)10—12(-17) by (21/.-)4-S(-8) cm, index 11/,-2'/. g

Inflorescence 10—20-flowered. Ovary

glabrous. Infructescence 4-5 cm long. Fruit ovoid, strongly beaked, 5-10 by 3-6 mm.

7. C. timorensis

9. Leaves (3—-)4-5(-6!/,) by (11/,-)1!/,-2'/,(-3'/,) cm, index 2-3.3. 3 Inflorescence 5—7-flowered. Ovary
densely hairy. Infructescence c. 1'/, cm. long. Fruit globose, not beaked, c. 3 by 3 mm

8. Leaf strongly oblique; nerves 3-4 pairs. Inflorescence gj and %; ¢ flower 4-merous..

1. Celtis rigescens (M1Q.) PLANCH. in DC. Prod. 17
(1873) 182; So—EPADMO in Whitmore, Tree Fl. Mal.
2 (1973) 416. — Solenostigma rigescens Mia.
Sumatra (1861) 411. — Solenostigma sumatrana
Mia. /.c. 411. — C. sumatrana (MiQ.) PLANCH. in
DC. Prod. 17 (1873) 181. — C. nymanii K.Scu. in
K.Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905)
240; Laur. Bot. Jahrb. 50 (1913) 311. — C. asperi-
folia MeRR. Philip. J. Sc. 17 (1920) 246; En. Philip.
2 (1923) 32. — Fig. 20e, 23a-b.

Large tree up to 45 m, 1 m ©. Buttresses up to
6m tall, 3m out, 5cm thick. Bark grey-brown,
smooth, finely fissured to pustulate and lenticellate.
Innovations densely yellowish brown to rufous
simple hairy. Older twigs glabrous, finely striate

8. C. rubrovenia
9. C. tetrandra

and sparsely minute lenticellate. Terminal buds
ovoid-conical, c. 3 by 2 mm, scales densely yellow-
ish-brown tomentose. Stipules linear-lanceolate,
4-5 by 1-1'/, mm. Mature /eaves thick-coriacecus,
strongly rugose, stiff and brittle when dry, ovate-
elliptic to elliptic-oblong, (5S—)8-12(-15) by (2%/,-)
3—5(-6!/,) cm (index 11/,-2'/,), broadest at or
slightly below the middle; above glabrous, shining,
beneath sparsely yellowish-brown pubescent es-
pecially on midrib and nerves; base rounded to
subcordate, symmetrical, rarely attenuate-rounded
and slightly asymmetrical; margin undulate, en-
tire or distantly serrate in the upper half, very often
incurved; apex rounded-acute to acuminate;
midrib and nerves strongly raised beneath, flattish

1977]

ULMACEAE (Soepadmo) 59

Fig. 20. Celtis paniculata (ENDL.) SPACH. a. 2 Inflorescence, x 8, b. flower, <x 14, c. fruit, x 11/3, d. shoot

apex showing terminal bud above 2 pairs of overlapping stipules, x 5. — C. rigescens (MIQ.) PLANCH.

e. Embryo, folded cotyledons, x 11/,. — C. timorensis SPAN. f. Embryd in LS, x 2. — C. hildebrandii

SOEPADMO. g. Pitted endocarp, x 3, h-i. embryos, x 4. — C. Jatifolia (BL.) PLANCH. j. Fruit, nat. size

(a—b Kornass! 463, c. KOSTERMANS & SOEGENG 352, d PLEYTE 73, e BLOEMBERGEN 4231, f Forbes 1073,
g—i CLEMENS 8375, j BSIP 11768).

to deeply impressed above; main nerves (2-)3(-4)
pairs, ascending and arcuating, anastomosing
along the margin, the lowest pair running up to
1/,/, the length of the leaf; reticulations sub-
iform to subareolate, distinct beneath;
petiole 6-10 by 1-2 mm, shallow-sulcate, densely
appressed yellowish-brown pubescent, glabrescent.
Inflorescence 3 or % or rarely mixed, 1'/,—5 cm long;
ovate, c. '/,-1 by '/, mm. — ¢ Inflorescence

a much-branched, multiflorous (up to c. 70)
icle up to 5cm long, axes slender, flexuous,
rne on the lower and leafless part of the new
shoot or in the axils of lower new leaves; ¢ flower
1'/,-2mm @, subsessile ; perianth lobes (4-)5, elliptic,
c. 1-1"/, by 1 mm; filaments 1—1'/,mm long, anthers
reniform, c. 1 by '/, mm; pistillode minute, sur-
rounded by dense Fp yellowish-brown hirsute
hairs. In a mixed inflorescence, the few ¢ flowers
are situated near the tip of the axis. — % Inflores-
cences borne in the axils of leaves of the new shoot,
1-1'/,cm long with up to 5 flowers; % flower
ovoid, c. 2 by 1 mm; perianth lobes (4-)5, elliptic-
rounded, c. 1'/, by 1 mm, stamens (4-)5; filaments
c. 1 mm, anthers c. '/, 1/,mm,; ovary slightly
compressed, densely pale yellowish-brown ap-
pressed-hairy; stigmatic arms spreading, bifurcate
at the tip. Infructescence up to 2-3 cm long, axes
up to 2mm thick, with (1-)2(-3) fruits. Fruit

ellipsoid, faintly S-angular, c. 15 by 10mm,
appressed-pubescent, glabrescent; exocarp 2-3 mm
@, rather woody, occasionally lenticellate, turning
deep-red when ripe, containing slimy substances
when boiled. Embryo curved, cotyledons folia-
ceous, folded, equal. Endosperm scanty.

Distr. Solomons (common); in Malesia: New
Guinea (incl. New Britain, common), Moluccas
(Ceram, Buru, Sula, Morotai), NE. Celebes (Mina-
hasa), E. Borneo (W. Kutei), W. Java (Bantam),
Central & S. Sumatra, Anambas Is., and Malaya
(Perak, Selangor, Pahang, Johore).

ies density of Celtis L. in Malesia;
above the hyphen endemic spp., below it the non-
endemic ones.

Fig. 21. S

FLORA MALESIANA [ser. I, vol. 82

es

oA aes

z wd .
2 ‘<a

WIND, Jan. 1925).

1977]

Ecol. In primary and _ secondary forest,
0-1500 m, on various types of soils including lime-
stone. Often rather common locally (W. New
Guinea and Solomons). Fi. (March-April) Sept.,
Dec., fr. Dec.—Aug.

Vern. Malay Peninsula: mémpélas bulan, Abor.,
rémpélas, M. Sumatra: klutum, marsékan, rém-
pélas, M. Palembang, asin asin, Anambas. West
New Guinea: séhiega, sériega, Manikiong lang.,
wennémaram, Berik lang., Hollandia, bégék,
Oransbari, Hattam lang., pieh(h), Kebar, bipa,
Wandammen lang., Adi I., ainam, Key lang.,
hajanggwij, manji, sékika, Manokwari, pimar,
Sidei lang., warén, siempai, wiempai, Biak lang.,
walik, walis, Mooi lang.; East New Guinea:
gairama, watot, Waria dial., Moresby, Madang
Distr., koru, Bambi, suri, Madang, bagibib,
Kaigorin, aban, bison, sam, Bilia, baigu(p), Amele,

ip, kara, sungung, susuik, Dumpu, goga,
mutum, sawan, Faita, tapang, Washuk, Sepik
Distr.; ailee, Urin, S. New Britain; gwalafalisi,
lausi, Kwara lang., Solomons.

Note. Sterile specimens, especially those with
young leaves only, may easily be mistaken as
belonging to Grewia or Microcos (Tiliaceae) or to
Ziziphus angustifolius (BL.) HATUus.

2. Celtis luzonica WaARB. in Perkins, Fragm. FI.
Philip. — 164; Merr. En. Philip. 2 (1923) 32.
Tree up to 30 m, 90 cm 2. Bark smooth yellow-
ish-grey. Innovations sparsely set with minute
simple hairs, glabrescent. Terminal bud ovoid, c. 6-
10 by 3-5 mm. Stipules ovate, thick, glabrous,
peltately attached and overlapping, c. 5-10 by 3-5
mm. Leaves broad elliptic to elliptic-oblong, thick-
coriaceous, glabrous, (8—)12—16(-19) by (5—)8-10
(-12)cm, index (1*/,-)1*/,(-2); base rounded,
symmetrical; margin entire, undulate, apex
rounded with blunt and sometimes emarginate tip;
midrib and nerves raised beneath, flattish to shal-
lowly impressed above; nerves 2-3 pairs, ascending
and arcuating, at least at an angle of 55° with the
midrib, anastomosing and looped along the margin,
the lowest pair running up to 2/,—*/, the length of
the leaf; reticulations irregular to subscalariform,
sparse, distinct beneath or obscure on both sur-
faces; petiole 10-20 by 1'/,-2(-3)mm, flat or
shallowly sulcate. Flowers 5-merous. Inflorescences
3 or 3 ¥, much-branched panicles or racemose, up
to 150-flowered, up to 10 cm long, axillary or sub-
terminal, including the bracts sparsely puberulous,
glabrescent; bracts ovate, minute, c. 1 by '/, mm.
— 3 Flower c. 1*/,-2 mm @; perianth lobes c. 1'/,-
2 by 1 mm; filaments 1'/,-2 mm, anthers subreni-
form, c. 1-1'/, by '/,-1 mm; pistillode ovoid-
conical, —— c. 1 by '/, mm. — § Flowers
ovoid-coni borne at the distal parts of the
inflorescence, 2'/,-3 by 2-2'/, mm; perianth lobes
c. 1*/,-2 by 1-1'/, mm; filaments c. 1-1'/, mm,
anthers c. */, by */,mm; ovary + compressed,
a pubescent apically, c. 2-2'/, by 1-1'/, mm;
gmatic arms eeeenes or incurved, broadened
ee shallowly bifid at the tip. Infructescence with a
stout axis 2-3 mm thick, bearing up to 10 fruits.
Fruit globose, glabrous, 16-20 by 14-18 mm, +
terete, reddish-brown when ripe; endocarp 4-lobed
and splitting at maturity; embryo curved, hypoco-
tyle ascending, cotyledons foliaceous, equal,

ULMACEAE (Soepadmo) 61

folded; endosperm very scanty to nearly absent.

Distr. Malesia: Philippines (Luzon, Surigao,
Mindanao, Mindoro, Samar, Masbate).

Ecol. In thickets and forests at low altitude.
Fl. fr. March-July.

Vern. Philippines: malaikmo, malaitmo, Tag.,
daloo, Todaya.

3. Celtis paniculata (ENDL.) PLANCH. Ann. Sc.
Nat. III, 10 0 (1848) 305; in DC. Prod. 17 (1873) 182;
Bru. Fl. Austr. 6 (1873) 156; NADEAUD, En. Pl.
Tahiti (1873) 42; Laur. Bot. Jahrb. 50 (1913) 310;
KANEH. Fl. Micron. (1933) 82, f. 14; FRANCIs,
Austr. Rain-forest Trees ed. 2 (1951) 67. — Sole-
nostigma paniculatum ENpD.L. Prod. Fl. Norf.
(1833) 42. — C. pacifica PLANCH. Ann. Sc. Nat. III,
10 (1848) 308; in DC. Prod. 17 (1873) 184. —
C. ingens F.v.M. Fragm. 4 (1864) 88. — C. vitiensis
A. C. SmitH, Bull. Torr. Bot. Cl. 70 (1943) 536;
J. Arn. Arb. 31 (1950) 150; PARHAM, PI. Fiji Isl.
(1964) 88. — Fig. 20a-d, 23i.

Small to large tree, up to 36m, 70cm @.
Buttresses occasionally present, up to 11/, m. Bark
smooth to finely fissured, light- to dark-brown.
Innovations sparsely appressed-puberulous, glab-
rescent. Older twigs glabrous, sparsely lenticellate.
Stipules overlapping, embracing the twig and
enclosing the bud, broad-ovate-acute, thick,

c. 4-5 by 3-4 mm. Leaves elliptic to ovate-elliptic,
68. 113) by (2'/,-)3-S(-6) cm, index (11/,-)2
(-2/,); coriaceous, glabrous, dull grey-green when
dried ; base attenuate-rounded, mostly symmetrical,
margin entire, often undulate, apex bluntly acute
to rounded-acute; midrib and nerves only slightly
raised beneath, flattish to shallowly impressed
above; main nerves 3-5 pairs, ascending and ar-
cuating at an angle of over 50° with the midrib,
anastomosing and looped along the margin, the
lowest pair running up to "/3;-"/, the length of the
leaf; reticulations fine, irregular, obscure on both
surfaces; petiole 6-15 by 1—-11/, mm, flat to + sul-
cate. Flowers 5-merous. Inflorescences 3 or 2 or
rarely mixed, axillary or subterminal, much-
branched, 5-60-flowered, including the bracts
sparsely appressed-puberulous; bracts ovate-acute,
c. 2 by 1 mm. — ¢ Inflorescences up to 4cm long,
borne on the lower and leafless part or axillary on
the lower leaf of the new shoot, paniculate, 30—60-
flowered; ¢ flowers subsessile, c. 1-1'/,mm @;
perianth lobes transparent, c. 1—-1'/, by '/, mm;
filaments c. 1 mm, anthers subreniform, c. 3/,—1 by
1/, mm; pistillode minute, compressed. — Mixed
or 2 inflorescences axillary on the upper parts of the
new shoot, racemose, 5-10-flowered; 2 flower
ovoid-conical, c. 2 by 1 mm; perianth lobes ovate-
acute, c. '/,-1 by '/, mm, subglabrous; staminodes
rudimentary, very much shorter than the perianth;
ovary + compressed, subglabrous, c. 1'/,-2 by
1 mm; stigmatic arms spreading, bifid at the tip.
Fruit ovoid, faintly 4-S-angular, beaked, glabrous,
7-12 by 5-8 mm, bluish to glaucous when ripe,
mostly sterile.

Distr. Australia (Queensland, Norfolk L.),
Melanesia (Solomons, New Hebrides, New Cale-
donia), Polynesia (Fiji, Tonga, Tahiti, Cook Is.,
Pitcairn, Mangarawa I., Tuamotus, Marquesas),
Micronesia (Marianas); in Malesia: Lesser Sunda
Islands (Wetar), Borneo (Mt Kinabalu, very rare),
Moluccas (Morotai, Ceram, Ambon, Tanimbar),

62 FLORA MALESIANA

[ser. I, vol. 8?

eee (common in West, apparently rare in
East).

Ecol. Primary and secondary forest, 0-900 m,
on well drained soils including coral limestone,
very often common locally (W. New Guinea and
Solomons). Fi. fr. July—May.

Vern. New Guinea: wiempai,
Manikiong; Solomons: J/ausiasi,
marasa, Sabalu.

4. Celtis philippensis BLANCO, FI. Filip. (1837) 197;
PLANcH. Ann. Sc. Nat. III, 10 (1848) 306; in DC.
Prod. 17 (1873) 184 (‘philippinensis’); Bru. FI.
Austr. 6 (1873) 156; VIDAL, Rev. PL. Vasc. Filip.
(1886) 248; Merr. Philip. J. Sc. 1 (1906) Suppl. 42;
Sp. Blanc. (1918) 122; En. Philip. 2 (1923) 32;
Leroy, Bull. I.F.A.N. 10 (1948) 212, incl. var.
consimile (BL.) LEROY; Fl. Madag. Fam. 54 (1952)
3; SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973)
416. — C. wightii PLANCH. Ann. Sc. Nat. III, 10
(1848) 307; Wicur, Ic. Pl. (1853) t. 1969; PLANCH.
in DC. Prod. 17 (1873) 184; Hook. f. Fl. Br. Ind. 5
(1888) 483; BRANDIS, Ind. Trees (1906) 594; J. J.
SMITH in K. & V. Bijdr. 12 (1910) 647; GAGnep.
Fl. Gén. I.-C. 5 (1927) 683; PoLHILL, Kew Bull. 19
(1964) 141; BAck. & BAKH. f. Fl. Java 2 (1965) 11.
— C. strychnoides PLANCH. Ann. Sc. Nat. III, 10
(1848) 306; in DC. Prod. 17 (1873) 185; Wars.
Bot. Jahrb. 13 (1891) 287; Laur. in K.Sch. & Laut.
Fl. Schutzgeb. (1900) 264. — C. mauritiana
PLANCH. Ann. Sc. Nat. III, 10 (1848) 307; in DC.
Prod. 17 (1873) 184. — Sponia_ strychnifolia
TeysM. & BINN. Nat. Tijd. N. I. 4 (1853) 394; Ned.
Kruidk. Arch. 3 (1855) 392. — Solenostigma
brevinerve BL. Mus. Bot. 2 (1856) 67. — Solenos-
tigma laurifolium BL. I.c. 68; Miq. Fl. Ind. Bat. 1, 2
(1859) 220, incl. var. constricta MiQ. — Solenos-
tigma hasseltii BL. Mus. Bot. 2 (1856) 68. — Sole-
nostigma consimile Bu. l.c. 68. — Solenostigma
djungiel BL. I.c. 69. — Solenostigma philippinensis
(BLANCO) Mia. FI. Ind. Bat. 1, 2 (1859) 220. —
Solenostigma wightii (PLANCH.) Mig. I.c. 220.— C.
brevinervis (BL). PLANCH. in DC. Prod. 17 (1873)
183. — C. laurifolia (BL.) PLANCH. l.c. 185. —
C. hasseltii (BL.) PLANCH. I.c. 185. — C. djungiel
(BL.) PLANCH. /.c. 185. — C. mindanaensis ELMER,
Leafi. Philip. Bot. 8 (1915) 2842. — C. collinsae
CraIB, Kew Bull. (1918) 370; Rm. Fl. Mal. Pen. 3
(1924) 322. — C. multifolia ELmer, Leafl. Philip.
Bot. 10 (1939) 3796, angl., inval. — Fig. 18a—k, 22.
Small to large tree, up to 30m, 80cm @.
Buttresses if present up to 2!/, m tall, 2 m wide and
10 cm thick. Bark smooth to finely fissured, pale
grey to grey-brown. Innovations initially sparsely
to densely set with yellowish-brown appressed
or/and woolly hairs, glabrescent. Stipules ovate-
acute, 6-10 by 2-4 mm, thick, peltately attached,
overlapping and enclosing the bud. Leaves thick-
coriaceous, glabrous, full grey when dried, elliptic-
oblong to suborbicular, (4~)8-14(-18) by (2-)
3-6(-8) cm, index (13/,-)2 —3; base rounded or
attenuate-rounded, mostly symmetrical ; margin
entire often undulate (immature leaves very rarely
distantly serrulate at the upper half); apex rounded
to acute; midrib and nerves raised beneath, im-
pressed to flattish above; main nerves 1 pair,
ascending, arcuating and running throughout the
length of the leaf (var. philippensis) or 1-3 pairs,
the lowest pair ascending, arcuating, and running

Biak, séhiega,
Kwara; Fiji:

up to about 2/; the length of the leaf and then
anastomosing with the 1-2 weaker and more or
less horizontal upper nerves (var. wightii); reticula-
tions fine, dense, subscalariform or subareolate,
usually rather distinct beneath; petiole 6-15 by
1-2 mm, sulcate. Inflorescences 3 or 3%, much-
branched panicles, many-flowered, including the
bracts densely yellow-brown to rufous soft-hairy ;
bracts ovate-acute, c. 3 by 1 mm. In the ¢¢ in-
florescence the ¥% flowers are borne on the distal
parts of the inflorescence. — 3 Inflorescences borne
on the lower part of the new shoots, 2-4 cm long,
with up to 40 flowers; ¢ flowers c. 2mm @;
perianth lobes ovate-elliptic, c. 11/,-2 by 1 mm;
filaments 1-11/,mm long, anthers subreniform,

c. 1/,-1 mm by '/, mm; pistillode ovoid-cylindrical,
caiinceasedl Ce; Ele by '/,.mm. — Mixed in-
florescence up to 5cm long, up to 50-flowered,
borne on the upper part of the new shoots; %
flowers ovoid, c. 2—2'/, by 2mm; perianth lobes
ovate-elliptic, c. 2-2'/, by 1 mm; filaments 1-2 mm,
anthers +/.-lmm @; ovary ovoid-cylindrical,
c. 2-21/, by 11/,-2 mm, glabrous except at the base;
stigmatic arms spreading, c. 1-1'/,mm_ long,
bilobed to bifid at the tip. Infructescence up to
4-5cm long, carrying 1-3 fruits, axes 1-2 mm
thick. Fruit ovoid, glabrous, 8-15 by 7-12 mm,
beaked when young; exocarp less than 1 mm @,
sometimes lenticellate, turning orange to red when
ripe; endocarp -+- smooth; embryo curved, hypo-
cotyle ascending, cotyledons broad, foliaceous,
unequal in thickness, not folded; endosperm oily,
scanty to absent.

Distr. Tropical Africa to Madagascar, Indian
Ocean (Réunion, Mauritius, etc.), India, Burma,
? SE. China, Hongkong, Taiwan, Indo-China,
Thailand, throughout Malesia to NE. Australia
and the Solomons.

Taxon. A rather variable, widely spread species
with two rather but not completely distinct
varieties. These are:

a. var. philippensis, characterized by larger
leaves of (7—)9-12(-18) by 4-8 cm with one pair of
nerves usually running more or less throughout the
length of the leaf, subscalariform reticulation, and
larger fruit of 10-15 by 8-12 mm;

b. var. wightii (PLANCH.) SOEPADMO, comb. nov.
(basionym: C. wightii PLANCH. I.c. supra). Fig. 22.
Characterized by smaller leaves, (4-)5—7(-9) by
(2-)3-4(—5!/,) cm with 1-3 pairs of nerves and the
lowest pair mostly running up to 2/; the length of
the leaf, and slightly smaller fruit, 8-12 by 6-10 mm.

It should be noted, that the distinguishing charac-
ters mentioned above should be applied in com-
bination; if taken individually they may not be
clearly well defined. For example, there are several
specimens (e.g. GARDNER 5.n., THWAITES CP 50
from Ceylon; KING s.n. and BROWNE s.n. from
India; PARKINSON 214 from the Andamans
UNEsco 214 from Malaya; JAcoss 4709, 4711, at
KosTERMANS 23061 from Java; KosTERMANS &
WIRAWAN 61 from the Lesser Sunda Is.; NGF .
19100 & 30787 from New Guinea; MERRILL Sp.
Blanc. 52 from the Philippines, etc.) which have
both types of venation. As for the fruit, the smaller
size in var. wightii may be due in part to the fact
that they are not fully ripe, as the majority of them
are empty (without embryo). Furthermore, it was
also noticed that most specimens of var. wightii

1977]

ULMACEAE (Soepadmo) 63

have been collected from localities under a strong
seasonal climate or from trees growing on well-
drained and poor soils (rocky or sandy beach,
limestone hills, etc.).

Ecol. Understorey tree in primary and secon-
dary forests, on various types of soils, at low alti-
tudes (0-650m); often gregarious and very
common locally. Fi. fr. mostly July—April,. The
fruits which turn to orange or red when ripe may be
dispersed by birds, but in the case of var. wightii,
which mostly grows in the very coastal forest, they
may be dispersed by sea-water as well; (the endo-
carp is woody, hard, and persistent).

Uses. Though not durable, the wood is locally
used for house-building.

Vern. Java: ki-éndog, ki-howé, S, W. Java,
kéraja, pusutan, séntok, sépat, sépréh, tjéngkék,
wuluh, J, Central & E. Java; N. Borneo: nyelepi;
Philippines: malaitmo, narabagsay, Tag.; Celebes:
kao lulu, Malili; Moluccas: horo, Morotai;
Lesser Sunda Is.: menulang, Sumba, nemu, Flores;
New Guinea: piéh, Kebar, marmar, Tor, Berik,
sehiega, Manikiong, bépiejét, Hattam, warén,
Biak, ikai, ikoi, Kemtuk, mélawar, Mooi, etc.

5. Celtis latifolia (BL.) PLANCH. in DC. Prod. 17
(1873) 186; Wars. Bot. Jahrb. 13 (1891) 287; Laur.
in K. Sch. & Laut. Fl. Schutzgeb. (1900) 264; Bot.
Jahrb. 50 (1913) 311. — Solenostigma latifolium
BL. Mus. Bot. 2 (1856) 67; Mia. FI. Ind. Bat. 1,
2 (1859) 219. — Solenostigma zippelii BL. Mus. Bot.
2 (1856) 67. — C. zippelii (BL.) PLANCH. in DC.
Prod. 17 (1873) 186. — C. kajewskii MERR. &
Perry, J. Arn. Arb. 22 (1941) 254. — Fig. 20j, 23k.

Tree up to 35 m, 80 cm @. Buttresses plank-like,
up to 2 m tall, 2'/, m out and 6 cm @. Bark smooth
to finely fissured, pustulate-lenticellate, light-
brown to grey-brown. Innovations densely yellow-
ish-brown hairy, glabrescent. Young twigs blackish
when dry, older ones greyish, glabrous and sparsely
lenticellate. Terminal buds ovoid-conical, acute,
10-15 by 4-5 mm. Stipules peltately attached, over-
lapping, thick, ovate-acute, c. 10 by 5 mm. Leaves
thick-coriaceous, glabrous or sparsely pubescent
beneath, especially on midrib and nerves, broadly
ovate to elliptic-orbicular, (8-)15-18(-25) by
(6—)8—-12(-18) cm, index 1'/,-2'/,; base rounded to
subcordate, symmetrical to + asymmetrical;
margin entire, undulate, often recurved; apex
bluntly rounded or acute to acuminate; midrib and
nerves strongly raised beneath, impressed above;
nerves 1-2 pairs, ascending and arcuating, the
lowest pair running through */, of the length of the
leaf; reticulations coarse, wide-spaced, subscalari-
form, distinct beneath; petiole 10-20 by 2-3 mm,
glabrous, shallowly sulcate. Flowers 5-merous.
Inflorescence 3 or 3%, axillary or borne on the
lower part of the new shoot, 10-30-flowered, in-
cluding the bracts densely yellowish-brown
appressed-hairy; bracts ovate, c. 2 by 1 mm. — g
Inflorescence (not fully developed) up to 2 cm long,
15-30-flowered, paniculate; ¢ flowers c. 1'/,-2 mm
@; perianth lobes c. 1'/,-2 by 1 mm; filaments
c. 1mm, anthers c. '/,-1 by '/;mm,; pistillode
compressed ovoid, c. 1 by '/, mm. — 3% Inflores-
cence racemose, 5-10-flowered, slender, up to
7 cm long, few-branched; % flower ovoid-ellipsoid,
borne on the distal part of the inflorescence,
c. 2-3 by 2mm; perianth lobes ovate-lanceolate,

c. 2-21/, by 1 mm, at anthesis recurved; filaments
up to 11/, mm, anthers c. !/,—3/, by #/. mm; ovary
ovoid-cylindrical, c. 2-3 by 1-1'/, mm, initially
densely appressed-hairy, glabrescent except for
the basal part; stigmatic arms spreading, shallowly
bilobed at the tip. Infructescence up to 5 cm long,
carrying 1-5 fruits, axes sturdy c. 2-3 mm thick.
Fruit ovoid, faintly 4—S-angular, glabrous, 1?/,-
21/, by 1—-13/4 cm, exocarp occasionally lenticellate,
up to 2 mm @, turning to orange or deep-red when
mature. Embryo curved, cotyledons fleshy, un-
equal in thickness, hypocotyle ascending; endo-
sperm absent.

Distr. Solomons (very common); in Malesia:
Philippines (Palawan), Moluccas (Morotai,
Tidore), and New Guinea (in West very common in
the vicinity of Manokwari and Hollandia; in East
in Sepik and Morobe Districts; New Britain).

Ecol. Primary and secondary forests on sandy
clay soils, 0-400 m. FI. fr. mostly Jan.—Aug.

Vern. Moluccas: tohu, Morotai; New Guinea:
sehiega, Manikiong, marmar, Berik, bepiet,
Hattam; Solomons: /ae-lae, Kwara.

6. Celtis hildebrandii SOEPADMO, sp. nov. — Fig.
20g-i, 231.

Species valde affinis C. philippensi var. philippensi
et C. latifoliae, sed ab eis differt folio asymmetrico,
inflorescentia 3 valde ramosa multiflora, et cotyle-
donibus multiplicatis. T: BW 7936.

Arbor magna usque ad 45 m alta et 1 m diam.
Folia tenuiter coriacea, oblique ovato-elliptica,
(5-)8-11(-14) x (3-)4-6(-8) cm, ind. 1,5-2, glabra,
nervis lateralibus uniparibus usque ad */, partem
laminae longitudinis ascendentibus, reticulatione
laxa subscalariformi; petiolus applanatus vel +-
sulcatus, 8-15 x 1-1,5 mm. Inflorescentiae 3 et §;
3 multiramosae, 60-150-florae; % 5-10-florae.
Fructus ovoideo-globosus, + 4- vel 5-—angulatus,
10-12 x 8-10 mm; endospermium nullum; cotyle-
dones multiplicati; embryo curvatus.

Large tree up to 45 m, 100 cm @. Buttresses up
to 23/, m tall, 2m out and 10cm @. Bark smooth
to finely fissured, often pustullate, light-brown to
grey-brown. Innovations densely rufous to yellow-
ish-brown appressed-hairy, glabrescent. Terminal
buds ovoid-conical, acute, c. by 3mm.
Stipules ovate-acute, peltately attached and
overlapping, thick, c. 5 by 3mm. Leaves thin-
coriaceous, obliquely ovate-elliptic, (S—)8—11(—-14)
by (3-)4-6(—8) cm, index 1'/,-2; glabrous; often
discolorous, upper surface dull grey-green, lower
surface dull chocolate- or grey-brown; base
attenuate-rounded, mostly asymmetrical; margin
entire, often undulate; apex acute to acuminate;
midrib and main lateral nerves raised beneath,
impressed or flattish above; nerves one pair,
ascending and arcuating, running up to */, or the
entire length of the leaf; reticulations lax, sub-
scalariform, rather distinct beneath; petiole
glabrous, 8-15 by 1-1'/,mm, flat, + sulcate.
Inflorescence 3 or %, axillary or subterminal on
new shoots, much-branched, many-flowered,
including the bracts rather densely yellowish-brown
to rufous app -hairy, glabrescent; bracts
ovate-acute, c. 2-3 by 1I-1'/,mm. Flowers 5-
merous. — 6 Inflorescences much-branched
panicle, 60-150-flowered; ¢ flower c. 2mm @;
perianth lobes c. 1'/,-2 by 1 mm; filaments 1-1"/,

[ser. I, vol. 8?

FLORA MALESIANA

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ATL ISS ARS
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1977]

mm, anthers subreniform, c. 3/,-1 by +/,mm;
pistillode ovoid-cylindrical, compressed, c. 3/, by
1/,mm. — 8&8 Inflorescence a much-branched
raceme, up to 4-5cm long, 5-10-flowered; %
flower ellipsoid, c. 3 by 2 mm; perianth lobes ovate-
acute, c. 2-3 by 1 mm; filaments 1-11/, mm, an-
thers c. 1/, by '/, mm; ovary ovoid-ellipsoid, c. 2 by
1 mm, slightly compressed, initially densely hairy,
glabrescent; stigmatic arms initially curved, later
spreading, c. 1-1'/,mm long, broadened and
shallowly bilobed at the tip. Infructescence 3-5 cm
long, axes glabrous, c. 1-2 mm @, bearing 2-5
fruits. Fruit ovoid-globose, glabrous, + 4-5-
angular, slightly beaked, 10-12 by 8-10mm;
exocarp thin, endocarp pitted; embryo curved,
hypocotyle ascending, cotyledons broad, folia-
ceous, folded; endosperm absent.

Distr. Solomons (common); in Malesia:
Moluccas (Buru, rare), New Guinea (W. & E.
parts, common; New Britain).

Ecol. In both primary and secondary forests at
0-1000 m; often very common and gregarious
locally; on various types of soil. Male flowers
appear with new shoots around Nov.—Dec., and %
ones around Jan.—Feb. ; fruits mature by May-June.
The ripe fruits, which turn deep purple or bluish
black, may be dispersed by birds.

Vern. New Guinea: béhég, Hattam, walik,
Mooi, sehiega, Manikiong, piéh, Kebar, biempai,
Biak; Solomons: /aussi, lai-lai, Kwara.

7. Celtis timorensis SPAN. Linnaea 15 (1841) 343;
PLANCH. Ann. Sc. Nat. III, 10 (1848) 315; BL.
Mus. Bot. 2 (1856) 71; Mig. Fl. Ind. Bat. 1, 2
(1859) 22; PLANCH. in DC. Prod. 17 (1873) 180. —
C. cinnamomea LINDL. ex PLANCH. Ann. Sc. Nat.
III, 10 (1848) 303; BL. Mus. Bot. 2 (1856) 72;
PLANCH. in DC. Prod. 17 (1873) 181; Kurz, For.
Fl. Burma 2 (1877) 472; Hook. f. Fl. Br. Ind. 5
(1888) 482; PRAIN, Beng. Pl. 2 (1903) 719; BRANDIS,
Ind. Trees (1906) 596; J. J. SmitH in K. & V. Bijdr.
12 (1910) 644; Merr. En. Philip. 2 (1923) 32;
Gacnep. Fl. Gén. I.-C. 5 (1927) 682; Back. &
Baku. f. Fl. Java 2 (1965) 11. — C. reticulosa Mia.
Pl. Jungh. (1851) 69; Fl. Ind. Bat. 1, 2 (1859) 222.
— C. hamata BL. Mus. Bot. 2 (1856) 72; PLANCH.
in DC. Prod. 17 (1873) 180. — C. waitzii BL. Mus.
Bot. 2 (1856) 71; Mia. Fl. Ind. Bat. 1, 2 (1859) 221;
PLANCH. in DC. Prod. 17 (1873) 180. — C. dyso-
doxylon Taw. En. Pl. Zeyl. (1861) 267. — C.
crenato-serrata MerRR. Philip. J. Sc. 5 (1910) Bot.
174. — Fig. 20f, 23f-h.

Medium-sized tree up to 20 m, 30cm @. Bark
smooth, grey. Innovations densely rufous-hairy.
Branchlets glabrous, rather densely lenticellate.
Terminal buds ovoid-conical, c. 3-4 by 2mm.
Stipules linear-lanceolate, 5-10 by 1-2 mm. Leaves
thin-coriaceous, ovate-elliptic to elliptic-oblong,
(6-)10-12(-17) by (2'/,-)4—5(-8) cm, index 1'/,-
2'/,; except for the midrib and nerves glabrous,
discolorous, upper surface dull grey to blackish,
lower surface chocolate-brown; base rounded to
subcordate, mostly asymmetrical, margin undulate,
distantly serrulate to crenate at least for the upper
half; apex acute to acuminate; midrib and nerves
flattish to impressed above, raised beneath; nerves
1-2(-3) pairs, arcuating and ascending, the lowest
pair running to (*/,-)*/,(—*/,) the length of the leaf,
anastomosing along the margin; reticulations

ULMACEAE (Soepadmo) 65

subscalariform, sparse, rather distinct beneath;
petiole S—15 by 1-2 mm, sulcate. Flowers 5-merous.
Inflorescence 3 or 3%, racemose, lax, axes slender,
always longer than the petiole, including the bracts
sparsely rufous-hairy; bracts narrow ovate-acute,
c. 3-5 by 1-2 mm. — ¢ Inflorescences borne on the
lower and leafless parts of the new shoots, much-
branched, 10—20-flowered, up to 3cm long; ¢
flower c. 2 mm @; perianth lobes c. 1!/,-2 by 1 mm;
filaments 1-1'/,mm, anthers reniform, c. 1 by
1/, mm; pistillode present but strongly reduced in
size. — § Inflorescences borne in the axils of new
leaves, 4-7-flowered, up to 2-3 cm long; % flower
c. 2-3 mm @; perianth lobes c. 11/,-2 by 1 mm;
filaments c. +/,-1mm, anthers’ subreniform,
c. +/,/, by '/, mm; ovary ovoid-ellipsoid, c. 2 by
1 mm; stigmatic arms c. 1-1'/,mm, spreading.
Infructescence up to 4-5 cm long. Fruit ovoid,
terete or 4-angular, beaked, 5-10 by 3-6 mm,
glabrous. Embryo curved, hypocotyle ascending;
cotyledons broad, folded, equal; endosperm scanty.

Distr. Ceylon, India, Bangladesh, Burma,
Thailand, Indo-China; in Malesia: Central Suma-
tra (rare), Java (common; incl. Christmas I.),
Lesser Sunda Islands (Flores, Timor), N. Borneo
(Mt Kinabalu), Philippines (Luzon).

Ecol. Outside Malesia the species grows in
evergreen forests or along streams in deciduous
forests. In Malesia it is more common in areas
with a rather prominent seasonal climate,
0-1500 m. FI. fr. Nov.—April.

Vern. Central Sumatra: ki tondok, si tjakik;
Java: jeungil, ki tamiang, S, tjéngkék, tjitik, J;
Philippines: takulao, Ibn., malabutulan, Tag.

8. Celtis rubrovenia ELMER, Leafl. Philip. Bot. 2
(1908) 464; Merr. En. Philip. 2 (1923) 33. — C.
similis MERR. & PERRY, J. Arn. Arb. 22 (1941) 253.
— Fig. 23}.

Deciduous tree up to 30 m, 60 cm @. Bark grey-
brown, smooth. Innovations densely rufous
appressed-hairy, glabrescent. Older branchlets
glabrous, sparsely lenticellate. Buds ovoid-conical,
c. 3 by 2 mm, bracts ovate, imbricate, c. 1 by '/,mm.
Stipules subulate, c. 3-4 by 1 mm. Leaves (fully
developed ones) thick-coriaceous, rigid, brittle,
ovate to ovate-elliptic, (3—)4~-S5(-6'/,) by (1'/4-)
11/,—21/,(—3'/,) cm, index 2—3.3; sparsely appressed-
hairy especially on the midrib and nerves, glabres-
cent; base attenuate-rounded to rounded, sym-
metrical to asymmetrical; margin of young leaves
serrate to crenate at least at the upper half, that of
old leaves entire or distantly serrate in the upper
half only; apex acute, acuminate to caudate, tip
usually very sharp; midrib and nerves strongly
raised beneath (reddish in fresh specimens),
impressed above; nerves (1—)2(—3) pairs, arcuating,
ascending, at a narrow angle with the midrib less
than 45°, lowest pair running to c. ?/, the length of
the leaf; reticulations fine, subareolate, obscure on
both surfaces; petiole (3-)4-6(-8) by 1mm,
deeply sulcate. Flowers 4-merous. Inflorescences
3 or 3%, racemose, 1-2 cm tong, 4-7-flowered. —
3 Inflorescence borne on the lower and leafless
parts of the new shoot, 5—7-flowered; ¢ flower
c. 2mm @; perianth lobes c. 1'/, by 1 mm, ciliate;
filaments 1-1'/,mm _ long, anthers reniform,
c. 1 by */4mm; pistillode minute. — Mixed (3%)
inflorescences 4-5-flowered, borne in the axil of

66

FLORA MALESIANA

[ser. I, vol. 8?

new leaves; §% flower borne on the upper part of
the inflorescence, c. 2-2'/, mm @; perianth lobes
c. 11/,-2 by 1 mm, ciliate; stamens slightly smaller
in size than those of the ¢ flower; ovary ovoid,
11/,-2 by 1mm, densely hirsute; stigmatic arms
spreading, 11/,-2 mm long. Infructescences up to
11/,cm long, carrying (1-)2(-3) fruits. Fruit
globose, c. 3 by 3mm, glabrous, + 4-angular.
Endosperm scanty, oily, transparent. Embryo
curved, hypocotyle ascending; cotyledons equal,
foliaceous, folded.

Distr. Malesia: Philippines (Luzon), New
Guinea (Kebar Valley, Baliem R.; Morobe Distr.).

Ecol. In primary forests on hills, ridges between
500-1800 m. In New Guinea sometimes rather
common locally. Fl. fr. Sept.-March.

Vern. Philippines: palek; W. New Guinea:
dotjoni, nitjoni, Kebar.

9. Celtis tetrandra Roxs. Fl. Ind. ed. Carey 2
(1832) 63; PLANCH. Ann. Sc. Nat. III, 10 (1848)
300; in DC. Prod. 17 (1873) 179; Kurz, For. Fl.
Burma 2 (1877) 472; GamBLE, Man. Ind. Timb.
ed. 1 (1881) 344; Hook. f. Fl. Br. Ind. 5 (1888) 482,
incl. var. hamiltonii Hook. f. et var. mollis (PLANCH.)
Hook. f.; PRAIN, Beng. Pl. 2 (1903) 719; BRANDIS,
Ind. Trees (1906) 596; J. J. SmirH in K. & V.
Bijdr. 12 (1910) 641; Gacnep. FI. Gén. I.-C. 5
(1927) 681; BACK. & BAKH. f. Fl. Java 2 (1965) 11.
— C. trinervia Roxs. FI. Ind. ed. Carey 2 (1832) 65,
non LAMK, 1797. — C. acata HAMILT. Trans. Linn.
Soc. 17 (1834) 211; PLANcH. Ann. Sc. Nat. III,
10 (1848) 299. — Sponia tetrandra (ROxB.) VOIGT,
Hort. Suburb. Calc. (1845) 294. — C. napalensis
PLANCH. Ann. Sc. Nat. III, 10 (1848) 298. — C.
glabra PLANCH. I.c. 298. — C. roxburghii PLANCH.
Lc. 302. — C. hamiltonii PLANCH. I.c. 301; in DC.
Prod. 17 (1873) 179. — C. mollis PLANCH. Ann. Sc.
Nat. III, 10 (1848) 297, p.p., quoad specim. ex
Wallich 7203; in DC. Prod. 17 (1873) 179. — C.
serotina PLANCH. Ann. Sc. Nat. III, 10 (1848)
301; WiGHT, Ic. 4, 4 (1850) t. 1570. — Fig. 18l-0,
23d-e.

Semi-deciduous tree up to 40 m and 100cm @.
Bark smooth to rough, grey-brown. Innovations
densely rufous-hairy, glabrescent. Terminal bud
ovoid-globose, 2-3 mm @. Stipules linear, c. 5 by
1mm. Leaves coriaceous, oblique ovate-elliptic,
(4-)6-10(-13) by (12/,-)2#/2-34/2(-5) cm, index 2-3,
broadest at or below the middle; slightly dis-
colorous, above glabrous, dull grey in drying,
beneath glabrous or sparsely rufous-pubescent
especially on midrib and nerves, yellow grey-green
in drying; base rounded-attenuate, strongly asym-

metrical; margin denticulate or distinctly serrate
at the upper half, or subentire; apex acute, acumi-
nate, to caudate; midrib and nerves flat to im-
pressed above, raised beneath; nerves 3-4 pairs,
ascending and arcuating, anastomosing near the
margin, the lowest pair running to c. +/;-*/, the
length of the leaf; reticulations fine, subscalariform,
rather distinct beneath; petiole 5-12 by 1-2 mm;
slightly sulcate. Inflorescence 3 or %, cymoid, in-
cluding the bracts densely rufous-hairy. — 3 In-
florescence borne on the lower and leafless part of
the new shoot, 2—5-flowered, occasionally several
of them are clustered together on leafless short
lateral new shoots giving rise to a raceme of cymes;
3 flower c. 2-3mm @; perianth lobes 4(—5),
c. 11/,-2 by 1 mn, ciliate; filaments +/,-1 mm,
anthers reniform, c. 1 by ‘/, mm, sparsely seri-
ceous; pistillode very much reduced. —
Inflorescences 2—5-flowered, axes slender, in the
axils of new leaves; % flower c. 2mm @; perianth
lobes ciliate, c. 2 by 1 mm; stamens as in the g
flower; ovary ovoid, c. 2 by 1 mm, sparsely minute
pubescent; stigmatic arms c. 2 mm long, ?/, mm
broad, spreading. Fruit + globose, 5-8mm @,
glabrous, turning deep-red or black when ripe.
Endosperm scanty.

Distr. India, Bangladesh, Burma, Thailand,
Indo-China; in Malesia: N. Sumatra (Gajo and
Karo Lands) Java (West: G. Galunggung; Central:
G. Muria; East: Bodjonegoro, Mts Kawi & Idjen,
Besuki, etc.), Lesser Sunda Islands (Bali, Sumbawa,
Flores).

Ecol. In primary and secondary forests, 0-2000
m. In Burma and Thailand very often in evergreen
or semi-deciduous forest along river-banks. In
Malesia the preference seems to be largely to areas
subject to a seasonal climate. Fl. fr. Aug.—April.

Vern. Sumatra: ndokum, sigar, Karo, bitatar,
Toba, témung, M; W. Java: ki djeungkil, ki
tamiang, S, téritih, tritih, J; Lesser Sunda Is.: pusu,
Sumbawa, namut, Flores.

Excluded

Celtis grewioides WarRB. Bot. Jahrb. 13 (1891) 287
= Ziziphus angustifolius (MiQ.) Hatus. Nova
Guinea Bot. n. 3 (1960) 13.

Solenostigma angustifolium Mig. Sumatra (1861)
412. — C. angustifolia (MiQ.) PLANCH. in DC.
Prod. 17 (1873) 186, non LinDL. ex WALL. Cat.
(1831) n. 3691, nomen = Ziziphus angustifolius
(Mig.) Hatus. Nova Guinea Bot. n. 3 (1960) 13.

5. APHANANTHE, nom. gen. cons.

PLANCH. Ann. Sc. Nat. III, 10 (1848) 265, non Link, 1821; Mia. FI. Ind. Bat. 1, 2
(1859) 218; PLANCH. in DC. Prod. 17 (1873) 207; Hook. f in B. & H. Gen. Pl. 3
(1880) 355; ENGL. in E. & P. Nat. Pfl. Fam. 3, | (1888) 66; BERNARD, Bull. Herb.
Boiss. II, 6 (1906) 34; GaGnep. Fl. Gén. I.-C. 5 (1927) 690; Leroy, Bull. Mus.
Hist. Nat. Paris II, 18 (1946) 118, 180; Fl. Madag. Fam. 54 (1952) 12; J. Agr.
Trop. Bot. Appl. 8 (1961) 72; Li, Woody FI. Taiwan (1963) 105; HUTCH. Gen. FI.

1977] ULMACEAE (Soepadmo) 67

Pl. 2 (1967) 149. — Homoioceltis BL. Mus. Bot. 2 (1856) 64. — Galumpita BL. l.c.
73; Mia. Fl. Ind. Bat. 1, 2 (1859) 223. — Gironniera subg. Galumpita Hook. f. in
B. & H. Gen. Pl. 3 (1880) 356; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 66. —
Mirandaceltis A. J. SHARP, Bol. Soc. Bot. Mex. 23 (1958) 38, f. 1-4. — Fig. 24.

Monoecious, deciduous or semideciduous shrubs or trees, often buttressed.
Innovations densely or sparsely, whitish-grey to rufous, appressed-pubescent.
Indumentum consisting of bulbous-based, unicellular, finely tuberculate hairs and
multicellular, glandular hairs. Older branches glabrous, lenticellate, bearing lateral
and terminal buds. Stipules lateral, extrapetiolar, subulate, caducous. Leaves
alternate, petioled, glabrous, coriaceous, triplinerved at base or pinnately nerved.
Inflorescences 3, 2, or very rarely g9, axillary; bracts minute, caducous. — ¢
Inflorescences a condensed, multi-flowered raceme, borne on the lower parts of the
new shoots; ¢ flowers short-stalked, globular, 4-5-merous; perianth lobes mem-
branous, imbricate in bud, sparsely appressed pubescent outside; stamens glabrous,
filaments subulate, inflexed in bud, anthers ovoid-subreniform, non-apiculate,
introrse; pistillode absent, replaced by a cluster of whitish to silvery, erect, soft,
simple hairs. — 2 Flowers solitary in the axil of the upper leaves of the new shoot,
or borne in a 2-3-flowered mixed (g9) racemose inflorescence; long-stalked;
perianth lobes 4-5, long-persistent; staminode absent; ovary sessile, ovoid-ellip-
soid, terete to angular; stigmatic arms tubular; ovule anatropous. Drupe fleshy,
ovoid-globose, faintly 3-5-angular or + terete, glabrous; endocarp hard and per-
sistent. Seed exalbuminous, coat membranous, few cells thick; embryo curved,
hypocotyle ascending, cotyledons more or less equal, involute. Mode of germina-
tion unknown.

Distr. About 4-5 spp. Mexico, Madagascar, Ceylon, India, Burma, China (also Hainan), Korea,
Japan, Taiwan, Hongkong, Indo-China, Thailand, Andamans, through Malesia to Australia (Queensland
and New South Wales) and Solomons. Throughout Malesia (except Malaya and Moluccas): 2 spp. Fig. 25.

Ecol. In Malesia mainly found in areas subject to a rather strong seasonal climate, on various types of
— = the coastal lowlands, hills, and gallery forests, 0-750 m, locally often abundant and forming dense

ckets.

In the north temperate and subtropical regions the species flower in April-May and drupes ripen in
July-August. Tropical species produce flowers twice a year, viz around March-April and Sept.—Oct. and
fruit ripens in June-July or Nov.—Dec.

deciduous or semideciduous habit, flush-wise mode of growth, structure, size, colour of the in-
florescence and flowers suggest that pollination i is affected by wind. The drupes which turn to a deep red
colour when ripe are possibly dispersed by frugivorous birds.

KEY TO THE SPECIES

1. Leaves ovate, elliptic or obovate, (2—)4—6(—10) by (1—)2—3(-4) cm, triplinerved at base, margin variously
serrate or dentate, ug fh rey subentire; nerves (3—)4—5(—6) pairs, at a 30-45° angle with the midrib.
Mature fruit 6-10 by 4-7 mm, beak ONO. ANd ens caf dust ah. omhe hee 1. A. philippinensis

1. Leaves ovate-elliptic or elliptic-oblong, (S—)10-14(—20) by (2-)3—6 (-8) cm, pinnately nerved, margin
entire or very rarely distantly, minute serrulate at the upper half; nerves (5— \7-8 (—10) pairs, at an angle
of more than 60° with the midrib. Mature fruit 15-20 by 8-12 mm, beak up to5 mm. . 2. A. cuspidata

1. o_o philippinensis PLANCH. Ann. Sc. Nat. Shrub to medium-sized tree up to 28 m, 40 cm @.
Ill, 10 (1848) 337; Mig. Fl. Ind. Bat. 1, 2 (1859) Trunk often fluted, low-buttressed, occasionally
219: PLANCH. in DC. Prod. 17 (1873) 208; BrH. producing suckers. Bark smooth to finely fissured,
Fl. Austr. 6 «1873) 160; Hook. f. Icon. III, 2 ( 1876) peeling off into rectangular flakes, lenticellate.
65, t. 1741; Merr. En. Philip. 2 (1923) 34; Francis, Young parts densely or sparsely greyish-brown or
Austr. Rain-forest Trees ed. 2 (1951) f. 24 & 25. rufous, short, simple pubescent. Stipules subulate,
— Taxotrophis rectinervia F.v.M. Fragm. 6 (1863) 2-3 by '/,mm. Leaves glabrous, thin- to thick-
192. — A. rectinervia (F.v.M.) PLANCH. in DC. coriaceous, ovate, elliptic, or obovate, (2—)3—6(—10)
Prod. 17 (1873) 208. — Fig. 24j-n. by (1-)2-3(-4) cm, index 2-2.6; base attenuate

68 FLORA MALESIANA [ser. I, vol. 8?

Fig. 24. Aphananthe cuspidata (BL.) PLANCH. a. Habit, nat. size, b. LS of mature fruit, x 11/3, c. basal

view of fruit, nat. size, d. flowering young shoot, nat. size, e. 3 flower before anthesis, x 6, f. ditto in

section, x 9, g. g flower at anthesis, x 13, h. 2 flower, x 8, i. ditto in section, x 16. — A. philippinensis

PLANCH. j. mature fruit, x 2, k. ditto, basal view, x 12/3, I-n. variation of leaf-form, x 2/3 (a-c KOORDERS
21330, d-i Koorpers 30071, j, k, m, n BORDEN FB 1286, / Ramos BS 27383).

. - ae —_—— oa =

ULMACEAE (Soepadmo) 69

Fig. 25. Range of Aphananthe PLANCH. — A. cuspidata (BL.) PLANCH. (dots), A. philippinensis PLANCH.
(stars), A. sakalava Leroy (square), A. aspera ENGL. (circles), A. monoica (HEMSL.) LEROY (triangles).

or rounded, more or less equal-sided; margin
serrate, dentate, or rarely subentire, ends of serra-
tion occasionally developing into sharply mucro-
nate structures; apex acute, or rounded, tip blunt;
midrib and nerves strongly raised beneath,
flattish above; nerves (3—)4-S(—7) pairs, ascending,
straight or arcuating, subparallel, at 30-45° with
the midrib, not anastomosing near the margin;
reticulations fine, lax, subscalariform, faintly
visible beneath; petiole (2-)3-S(-7) by 1mm,
adaxially flat or shallowly sulcate. — 3g Inflores-
cence 1-3cm @, 10—50-flowered; bracts linear-
acute or narrow ovate-acute, '/,-1 by '/,mm; gd
flowers 1—1'/, mm @; perianth lobes ovate-lanceo-
late, 1-1'/, by ‘'/,-1 mm; filaments '/,-*/, mm,
anthers c. 3*/, by '/,mm. — @ Flowers always
solitary, ovoid-ellipsoid, 1-1!/, by 1 mm, c. 5mm
pedicelled; perianth lobes narrow ovate-acute,
1/,-1 by '/,mm; ovary 1-1'/, by 1 mm, densely
appressed-hairy; stigmatic arms 2-3 mm. Fruit
ovoid-globose, 6-8 by 46mm, 3-4-angular,
sparsely appressed-pubescent, glabrescent, pedicel
c. 5-10 mm.

Distr. Australia (Queensland and New South
Wales) and Solomons; in Malesia: New Guinea
and Philippines (Luzon). Fig. 25.

Ecol. In primary and secondary forest subject to
a rather strong seasonal climate, 0-750 m. In New
Guinea it is often found in semi-deciduous
ey. or mixed Eucalyptus-forest, occasionally
orming dense thickets especially on hillsides. Fi.
Apr.—May and Sept.—Oct., fr. mature in July-Aug.
and Nov.-Dec.

2. gard cuspidata (BL.) PLANCH. in DC.
Prod. 17 (1873) 209. — Cyclostemon cuspidatum
BL. Bijdr. (1825) 599. — Galumpita cuspidata (BL.)
Bui + 2a Bot. 2 (1856) 73; Mig. Fl. Ind. Bat. 1, 2

(1859) 224. — Gironniera nitida Btu. Fl. Hongk.
(1861) 324. — Gironniera reticulata THw. En. Pl.
Zeyl. 1 (1861) 268; Hook. f. Fl. Br. Ind. 5 (1888)
486. — Gironniera lucida Kurz, For. Fl. Burma 2
(1877) 470; Hook. f. Fl. Br. Ind. 5 (1888) 486. —
Gironniera cuspidata (BL.) KuRZ, For. Fl. Burma 2
(1877) 470; J. J. SMiTH in K. & V. Bijdr. 12 (1910)
688; Merr. En. Philip. 2 (1923) 35; Back. &
BAKH. f. Fl. Java 2 (1965) 12. — Gironniera
curranii MerR. Philip. J. Sc. 4 (1909) Bot. 251. —
A. lissophylla GAGNEP. Bull. Soc. Bot. Fr. 72 (1925)
804; Fl. Gén. L-C. 5 (1927) 690. — Gironniera
thompsoni KING ex A. M. & J. M. Cowan, Trees
N. Bengal (1929) 122. — ?Gironniera yunnanensis
Hu, Bull. Fan Mem. Inst. Biol. Bot. Ser. 10 (1940)
150. — ?A. yunnanensis (HU) GrupDz. Nov. Syst.
Pl. Vasc. USSR (1964) 66. — Fig. 24a-i.

Small to medium-sized tree up to 30 m, 60 cm @.
Buttresses up to 1 m tall, and 2 m out. Bark surface
rough, grey-brown, often flaky. Young branchlets
sparsely, minutely, appressed-pubescent, glab-
rescent; older branchlets sparsely lenticellate.
Stipules narrow ovate-acute, 2-3 by | mm. Leaves
coriaceous, glabrous, ovate-elliptic to elliptic-
oblong, (S5—)10—14(-20) by (2-)3-6(—8) cm, index
2'/,-3; base rounded, subcordate, or attenuate,
equal-sided or occasionally slightly unequal;
margin entire, rarely distantly, minute serrulate in
the upper half, often undulate; apex acute, acumi-
nate, or cuspidate, acumen up to 2cm, sharp-
tipped; midrib strongly raised and prominent
beneath, impressed to flattish above; nerves (5—)7—
8(-10) pairs, slightly raised beneath, flattish above,
subparallel, arcuating, at an angle of more than
60° with the midrib, weakly anastomosing near the
margin; reticulations fine, indistinct on both
surfaces, subscalariform to sub-areolate; petiole
glabrous, (5—)8—12(—15) by 1-2 mm, sulcate. — ¢

70 FLORA MALESIANA

[ser. I, vol. 8?

Inflorescence up to 4cm, 10-30-flowered; bracts
ovate-acute, */,—'/, by '/, mm; ¢ flower 1'/,-2 mm
@, short-stalked; perianth lobes obovate-lanceo-
late, c. 2 by 1 mm;; filaments 1-1'/, mm, anthers
ovoid-subreniform, c. 1mm @. — @ Flower
solitary or borne in a 2-3-flowered mixed inflores-
cence, ovoid-ellipsoid, c. 2 by 1 mm; pedicels up to
10mm; perianth lobes coriaceous, Ovate-acute,
c. 2 by 1 mm; ovary ovoid, glabrous, c. 1'/, by
1mm; stigmatic arms 2-3mm. Fruit ovoid,
glabrous, including the beak 15-20 by 8-12 mm,
beak up to 5 mm; pedicel up to 3cm, 1 mm @.

Distr. Ceylon, India, Burma, Andaman Is., ?
China, Hainan, Hongkong, Indo-China, Thailand;
in Malesia: NE. Sumatra (Sibolangit, doubtfully
native), Java (mainly N. Central & E. parts),
Lesser Sunda Islands (Bali, Sumbawa, Flores,
Timor, rather common), Borneo (very rare,
Kinabalu area), Philippines (Mindanao, rare),
Celebes (rare). Fig. 25.

Ecol. In primary lowland to submontane forest,
0-1300 m. In Malesia it is mainly confined to low-

land forests subject to a rather pronounced seasonal
climate. In Thailand it occurs mainly in the ever-
green or semi-deciduous forests along streams.
Outside Malesia flowering in March-April and
fruits mature in July-August. In Malesia it flowers
twice yearly, viz in Febr.-March and Sept.—Oct.,
fruits maturing in June-July and Nov.—Dec.

Vern. Java: suluh, wuluh; Lesser Sunda Is.:
sulu, Bali, k. belikat, Sumbawa, k. loko, Flores.

Note. Though I have not been able to examine
any of the specimens cited by Hu (1940), judging
from the description given it is likely that Giron-
niera yunnanensis Hu must be referred to this
species.

Excluded

Aphananthe negrosensis ELMER, Leafl. Philip.
Bot. 2 (1909) 575. — Calaunia negrosensis (ELMER)
Grubz. Nov. Syst. Pl. Vasc. USSR (1964) 54 =
Streblus glaber (MERR.) CORNER, Gard. Bull.
Sing. 19 (1962) 221 (Moraceae).

6. GIRONNIERA

GAUDICH. Voy. Bonite (1844) t. 85; PLANcH. Ann. Sc. Nat. III, 10 (1848) 338;
BL. Mus. Bot. 2 (1856) 72; Mia. Fl. Ind. Bat. 1, 2 (1859) 222; PLANCH. in DC.
Prod. 17 (1873) 205; Hook. f. in B. & H. Gen. PI. 3 (1880) 356; Fl. Br. Ind. 5 (1888)
485; ENGL. in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 66; BERNARD, Bull. Herb. Boiss. II,
6 (1906) 34, map 24; J. J. SmiTH in K. & V. Bijdr. 12 (1910) 665; Hutcu. Gen. FI.
Pl. 2 (1967) 149; PHUPHATHANAPHONG, Thai For. Bull. 6 (1972) 49; SOEPADMO in
Whitmore, Tree Fl. Mal. 2 (1973) 417. — Nemostigma PLANCH. Ann. Sc. Nat. III,
10 (1848) 265, cf. p. 338. — Helminthospermum Tuw. in Hook. J. Bot. Kew Misc. 6
(1854) 301, t. 9c. — Fig. 26.

Shrubs or large-sized trees, very rarely buttressed. Bark smooth to finely fissured,
grey-brown, often lenticellate. Jnnovations densely or sparsely set with golden
yellow or yellowish-brown indumentum consisting of simple bulbous-based, finely
tuberculate, appressed or erect hairs and multicellular capitate-glandular hairs.
Stipules extrapetiolar, free but overlapping each other and completely enclosing
the bud, on falling leaving a circular scar around the node. Leaves pinnately
nerved, nerves parallel, regularly well-spaced. Inflorescence 3 or 9, very rarely 39,
axillary or borne on older, leafless branchlets, 1-many-flowered, paniculate, race-
mose, thyrsoid, or capitate; bracts ovate-acute, minute, caducous, but rather long
persistent in the 9 inflorescence. — $ Flower globular, sessile or short-stalked along
the axes; perianth lobes 5, imbricate in bud; stamens glabrous, anthers introrse;
filament subulate, inflexed; anthers ovoid-reniform, apiculate, sub-basifixed;
pistillode present, rather well-developed or very rudimentary, densely set with
whitish or silky erect hairs. — 9 Flower ovoid-ellipsoid, compressed ; perianth lobes
4-5, usually unequal in size, long persistent; staminodes absent; ovary ovoid-
ellipsoid, strongly compressed, sessile, densely or sparsely appressed-hairy,
glabrescent; stigmatic arms up to 1!/, cm long, curled in bud, later spreading, rather
long persistent; ovule anatropous. Drupe ovoid-globose, convex elliptic lens-
shaped; exocarp thin, strongly adnate to the hard and persistent endocarp. Seed

1977] ULMACEAE (Soepadmo) 71

Fig. 26. Gironniera celtidifolia Gaupicu. a-b. 2 Flowers, one in CS, c-g. ¢ flowers, various details. —

G. subaequalis PLANCH. h. 3 Flower, m. young infructescence, p-r. fruits, one in LS. — G. nervosa

PLANCH. i-j. ¢ Flowers. — G. parvifolia PLANCH. k. 3 Inflorescence, n. young infructescence. — G.

rhamnifolia BL. \. 3 Inflorescence. — G. hirta RIDL. o. Young infructescence. a, c, m-r x 3/,,b x 3, d-e,

g-h x 7,f x 9,i x 3/5,j «x 2,k-l x 1/, (a-b BS 48861, cg MALIWANAG 119, A RAHMAT SI BoggA 9311,

i-j KEP/FRI 14582, k KEP/FRI 18017, i ZIPPELIUS 5.n., m MOH. SHAH 519, n SF 34908, 0 bb 30873, p-r
KEP/FRI 99766).

72 FLORA MALESIANA [ser. I, vol. 8?

coat membranous, a few cells thick; endosperm absent; embryo curved, hypo-
cotyle ascending; cotyledons fleshy, narrow, equal. Mode of germination unknown.

Distr. 6 spp., SE. Asia: Ceylon, Andaman Is., Burma, Thailand, Indo-China, China (Yunnan,
Kwangtung, Canton, Hainan, Hongkong), Micronesia (Palau and Ponape Is.), Melanesia (Solomons),
Polynesia (Samoa, Fiji); in Malesia: throughout the region except for the eastern part of Java, the Lesser
Sunda Islands, and the SE. Moluccas. Fig. 27.

Fig. 27. Approximate range of the genus Gironniera GAUDICH. with the number of species in each island or
partial area. There are no endemic species.

Fossils. Wo.Fe (in Graham, Flor. & Paleoflor. Asia & N.E. America, 1972, 200, pl. IV, f. 2) reported
leaf-impressions attributed to Gironniera from the Early Ravenian Flora of Alaska (Middle Eocene).
However, in examining the published photograph (p. 211), Iam very much in doubt whether the specimen
belongs to the genus at all.

Ecol. Understorey shrubs or trees in both primary and secondary forests, at 0-1300 m, often abundant
and gregarious; on various types of soil, including those derived from limestone. In areas where the
seasonal climate is prominent mainly found in the evergreen forest along streams.

Judging from the structure of the flower and inflorescence, it is inferred that the pollination may be
affected by wind. The drupes which turn to bright yellow or orange in colour when ripe are mostly barren,
and they may be dispersed by various species of frugivorous birds. Direct observations in the field are,
however, wanting.

Notes. Gironniera spp. have a continuous, flush-wise growth habit and have the ability to produce
flowers and fruits at a very young (sapling) stage (2-3 m tall). Since the plants are often very common
locally both in the primary and secondary forests, produce flowers and fruits regularly, and are very easy
to collect, most of the examined specimens were gathered from these young plants. The presence of so
many specimens collected from juvenile plants hampers proper identification even when they are fertile.

In the present revision, the characters used in the key were taken from specimens collected from mature
or fully grown trees, while those mentioned in the description of each species include also data from speci-
mens collected from the young plants, thus to include the total morphological variability.

On the material and field notes so far available it is impossible at this stage to determine whether the
genus is strictly monoecious or dioecious. In most cases, the specimens display only fruits or ¢ inflores-
cences, thus giving the impression that the genus is dioecious. However, there are a few collections (in all
species but G. hirta) which have both ¢ influorescences and infructescences attached to the same branch-
let, or they are borne on separate branchlets belonging to a single collecting number.

Mrs. PHUPHATHANAPHONG /.c. accommodated the Malesian specimens into two species, G. nervosa and
G. celtidifolia, without argumentation. I cannot agree with this view.

1977] ULMACEAE (Soepadmo) 73

KEY TO THE SPECIES
(Based on specimens from mature trees)

1. ¢ & 9 Inflorescence and infructescence a condensed, capitate thyrse or panicle, axillary or borne on
older leafless branchlets. ¢ Flower with a rather well-developed pistillode. Leaves thick-leathery,
distantly serrate or subentire. Terminal bud (stipules) up to 4"/, by'/,cm. . . . . 1. G. celtidifolia

1. ¢ & 2 Inflorescence and infructescence a lax, branched or unbranched panicle, raceme, or thyrse,
always axillary. ¢ Flower with a strongly rudimentary pistillode. Leaves chartaceous to thick leathery,
in Mal. specimens mostly entire. Terminal bud less than 3 cm by less than 0.3 cm.

2. Mature leaf densely pubescent beneath. ¢ Inflorescence a much-branched thyrse; 3 flowers arranged
in condensed cymoid clusters of 5-10 along the axes.

3. Young branchlets densely short appressed-hairy. Leaves elliptic-lanceolate to elliptic-oblong, index
21/,-3, broadest at the middle; nerves (12-)15(—17) pairs, parallel, straight, 5-8 mm spaced, not or
only weakly anastomosing along the margin. Well-developed 9 inflorescence (thus also the infructes-
cence) mostly an unbranched panicle, c. 3 cm long, bearing less than 10 flowers (fruits).

2. G. nervosa

3. Young branchlets densely set with long, erect hairs. Leaves mostly ovate-elliptic, rarely elliptic-
oblong, index c. 2, broadest below or rarely at the middle; nerves (8—)10—12(-14) pairs, subparallel,
10-15 mm spaced, arcuating and anastomosing towards the margin. Well-developed 9 inflorescence
(cq. infructescence) a mostly much-branched panicle 5-10 cm long, bearing up to 20 fruits

3. G. hirta

2. Mature leaf glabrous. ¢ Inflorescence unbranched or a branched raceme or panicle; ¢ flowers

solitary along the axes.

4. Mature leaf thick-coriaceous, broad-ovate, elliptic, or elliptic-oblong, broadest below or at the
middle, index 2-2?/,, base unequal-sided. Terminal bud 2-3 by 0.2-0.4cm. Well-developed ¢
inflorescence a much-branched panicle, carrying 40-100 flowers. Well-developed 2 inflorescence
(cq. infructescence) carrying (2—)5—10(—15) flowers (fruits). Mature fruits 10-12 by 9 by 5 mm.

4. G. subaequalis

4. Mature leaf chartaceous to thin-coriaceous, elliptic-lanceolate or elliptic-obovate, broadest at or
slightly above the middle, index (2'/,-)3(-3'/2), base more or less equal-sided. Terminal bud 2 by
1/, cm. Well-developed ¢ inflorescence an unbranched panicle carrying 15-30 flowers. Well-developed
2 inflorescence (cq. infructescence) carrying 1-3(—5) flowers (fruits). Mature fruits 7-8 by 5-6 by

mm,
5. Nerves (5-)6-8(-9) pairs. ¢ Flowers short-stalked. Axes of 9 inflorescence (infructescence) slender

and thin (less than 0.3 mm), 4-10 cm long, carrying 1—3(—5) flowers (fruits). . .

5. G. parvifolia

5. Nerves 10-12 pairs. ¢ Flowers sessile along the axes. Axes of 2 inflorescence (infructescence)

1/,-1 mm thick, 3-4 cm long, carrying (1-)3-5 flowers (fruits).

1. Gironniera celtidifolia GAUDICH. Voy. Bonite
(1844) t. 85; PLANcH. Ann. Sc. Nat. III, 10 (1848)
340; Mia. Fl. Ind. Bat. 1, 2 (1859) 223; Seem.
Fl. Vit. (1865) 236; PLANcCH. in DC. Prod. 17
(1873) 207; Laut. Bot. Jahrb. 50 (1913) 326;
Merr. En. Philip. 2 (1923) 35; PARHAM, PI. Fiji
Is. (1964) 88. — G. sibuyanensis Emer, Leafl.
Philip. Bot. 5 (1913) 1845; Merr. En. Philip. 2
(1923) 35. — G. grandifolia Merr. & PERRY,
J. Arn. Arb. 20 (1939) 325. — G. retinervia MERR.
& Perry, /.c. 326. — Fig. 26a-g.

Shrub or medium-sized tree up to 18 m tall and
25cm @. Bark greyish-brown to dark-brown,
smooth or finely fissured. Terminal buds c. 4—4'/,
by +/,cm; stipules narrow lanceolate-acute,
c. 4-4'/, by */,/,cm. Leaves thick-coriaceous,
elliptic-, lanceolate-, or obovate-oblong, (8—)15-
25(-42) by (4-)6-12(-19)cm, index 2!/,-3'/,,
broadest at or above the middle; base unequal-
sided, attenuate, rounded, or cordate-auriculate;
margin distantly serrate at least for the upper half
or subentire; apex rounded, acute, or acuminate-
caudate, acumen up to 4 cm long; glabrous above,
sparsely appressed-hairy beneath especially on
midrib and nerves, glabrescent; midrib and nerves
strongly raised beneath, flattish to shallowly im-
Sood. above; nerves (8~)12—16(-19) pairs,
anastomosing along the margin; reticulations
subscalariform, evenly spaced, distinct beneath;
petiole (3-)5-12(-15) by (1-)I"/,-2"/,(-4) mm,

6. G. rhamnifolia

terete or adaxially flat near the base, strigose,
glabrescent. Inflorescences a condensed, capitate,
much-branched panicle or thyrse, axillary or borne
on older, leafless branchlets, many-flowered, ¢ or
2, very rarely $9, including the bracts densely
appressed-hairy; bracts 2-4 by 2 mm. — ¢ Flower
c. 11/,-2 mm @, sessile or short-stalked along the
axes, perianth lobes ovate-elliptic, concave,
c. 1/,-2 by 1-1'/,mm; filaments 1'/.-2 mm,
anthers 1-1'/, by 1mm; pistillode cylindrical,
11/,-2 by */,mm, at base surrounded by dense,
whitish, erect hairs. — 9 Flower 2-3 by 2mm;
perianth lobes ovate, 1-1'/, by '/.-1 mm; ovary
2-3 by 1'/,-2mm, densely appressed-hairy;
stigmatic arms 1-1'/,cm, slender. Fruit ovoid-.
ellipsoid, sparsely appressed-hairy, glabrescent,
5-8 by 4-6 by 3-4 mm, beak up to 5 mm.

Distr. Micronesia (Palau, Ponape), Melanesia
(Solomons), Polynesia (Samoa, Fiji); in Malesia:
Borneo (doubtful record), Philippines (common
and widespread), Moluccas (Morotai, Halma-
heira), New Guinea (common in W. & E.; also in
Admiralty, Misima, Sudest, and Rossel Is.).

Ecol. In both primary and secondary forests,
0-1200 m; often locally abundant as understorey
shrub or tree. Fl. fr. Jan.—Dec.

Vern. Philippines: ablang, Mang., dita, Tag.,
mangabau, Bis., tanguntungan, Mbo,; Moluccas:
koko, Morotai; New Guinea: aimarwirieh, Biak,
taun, Wapil lang., Sepik Distr.; Solomons:

74 FLORA MALESIANA

aisulia, Kwara; Fiji: sisisi; Samoa:
pua-pua, puluvao.

Notes. In most specimens examined, the g
inflorescence shows abnormal development and
produces a malformed structure consisting of
numerous superimposed bracts in the axils of
which are found sterile structures resembling in
size and shape that of the pistillode in the normally
developed and functionally ¢g flowers. Well-
developed and functionally ¢ flowers are to be
found only at the distal parts of these abnormally
developed inflorescences. In a 39 inflorescence,
which is very rarely present, the functionally 9
flowers are borne on the central main axis and they
are flanked by two or more lateral, functionally ¢
flowers. Most of the fruits so far examined (more
than 95%) are barren.

Size and shape of the leaf are rather variable. In
general, specimens collected from a higher altitude
or from a more exposed habitat have a smaller
leaf and less pairs of nerves than those gathered
from lowland and shady localities. Intermediates
are, however, not uncommon, and for this reason
G. sibuyanensis, G. grandifolia, and G. retinervia
are here reduced.

masivau,

2. Gironniera neryosa PLANCH. Ann. Sc. Nat. III,
10 (1848) 338; BL. Mus. Bot. 2 (1856) 74; Mia.
Fl. Ind. Bat. 1, 2 (1859) 222; PLANcH. in DC.
Prod. 17 (1873) 206; Kurz, For. Fl. Burma 2
(1877) 469; Hook. f. Fl. Br. Ind. 5 (1888) 485;
Merr. En. Born. (1921) 216; Rip. FI. Mal. Pen. 3
(1924) 320; CorNER, Ways. Trees (1940) 688;
SOEPADMO in Whitmore, Tree Fl. Mal. 2 (1973)
419, f. 2. — G. penangiana GANDOG. Bull. Soc.
Bot. Fr. 66 (1919) 289. — G. sponioides GANDOG.
l.c. 289. — Fig. 26i-j.

Small to large-sized tree up to 40 m, 60cm @.
Buttresses sometimes present, low. Bark smooth or
finely fissured, grey-green to dark grey-brown,
often hoop-marked and lenticellate. Terminal bud
1-2 by +/,—'/,cm; stipules 11/,-2 by 12/,—'/, cm.
Leaves thick-coriaceous, rigid, elliptic-lanceolate
to elliptic-oblong, (61/,-)10—15(-18) by (2'/,-)4-6
(-8)cm, broadest at the middle, index 21/,-3;
except for the midrib and lateral nerves glabrous
above, densely set with yellowish-brown, soft,
slender hairs beneath; base rounded or attenuate,
unequal-sided ; margin entire, often recurved; apex
rounded to acute; midrib and nerves strongly
raised beneath, flattish or impressed above; nerves
(12-)14-16(-17) pairs, 5-8 mm spaced, arcuating
but not anastomosing near the margin, forming an
angle of up to 60° with the midrib; reticulations
dense, regularly spaced, scalariform or subscalari-
form, strongly raised and prominent beneath,
obscure to faintly visible above; petiole 5-10 by
1-2 mm, subterete, densely yellowish-brown ap-
pressed, pubescent. Inflorescences 3 or 9, axillary,
borne on separate shoots, including the bracts
densely yellowish-brown, appressed-pubescent;
bracts narrow ovate-acute, c. 1-2 by 1mm. — g
Inflorescence a slender, lax, pendent, branched
panicle of condensed cymes, up to 7cm long,
bearing 20-100 flowers; 3 flowers in clusters of
5-10 along the axes, c. 2 by 2 mm; perianth lobes
broad ovate-acute, c. 1-1'/, by 1 mm, densely, short
appressed-hairy outside; filaments 1-1'/, mm,
anthers ovoid, c. 1mm @; pistillode strongly

[ser. I, vol. 8?

rudimentary. — 2 Inflorescence a simple or
branched panicle, up to 2!/,cm long, 5-10-
flowered; 2 flowers sessile along the axes, com-
pressed ovoid-conical, 2-3 by 1'/,-2 mm; perianth
lobes ovate-acute, densely appressed-hairy outside,
1'/,-2 by 1mm; ovary densely appressed-hairy,
1'/,-3 by 1-1'/,mm; stigmatic arms 5-10 mm.
Infructescence up to 3 cm long, bearing (2—)4—6(-8)
fruits. Fruit subglobose or ovoid, densely
appressed-pubescent, 5-8 by 4-6 by 34mm,
short-beaked.

Distr. Thailand; in Malesia: Malay Peninsula
(incl. Singapore; common), Sumatra (rather rare),
Borneo (common).

Ecol. In primary and secondary forests, 0-1300
m, mostly below 500 m; often common locally as
understorey tree in lowland forests. In Thailand it
occurs mainly in the evergreen forest along streams.
Fl. fr. Jan._Dec., but mainly during July—Dec.

Vern. Malaya: médang berbulu, m. kasap, tapis,
M; N. Borneo: /uazon, Kadasan, hugot-hugot,
Dusun; Kalimantan: kayu ruas, Bandjar, gagas,
Bassap.

3. Gironniera hirta Rip. J. Str. Br. R. As. Soc. n.
82 (1920) 194; Fl. Mal. Pen. 3 (1924) 321; Sor-
PADMO in Whitmore, Tree Fl. Mal. 2 (1973) 417,
f. 2. — Fig. 260.

Shrub to medium-sized tree up to 30 m, 30cm @.
Bark smooth, light to grey-brown. Young branch-
lets, petiole, stipules, terminal bud densely set with
golden yellow, long, soft, erect hairs. Terminal
buds ovoid-conical, 11/,-3 by +/,-1/, cm; stipules
11/,-3 by 1/,cm. Leaves thick-coriaceous, rigid,
ovate-elliptic or rarely elliptic-oblong, (6—-)14-18
(—23) by (3—)5—8(—12) cm, index c. 2; base rounded,
subcordate or attenuate, unequal-sided; margin
entire, rarely distantly serrulate in the upper half;
apex rounded, acute to acuminate; except for the
midrib and nerves which are densely or sparsely
set with long, appressed or erect, soft hairs, glab-
rous above, densely soft-pubescent beneath; mid-
rib and nerves raised and distinct beneath, flattish
or impressed above; nerves (8—)10—12(-14) pairs,
10-15 mm spaced, arcuating towards and anasto-
mosing near the margin; reticulation subscalari-
form to subareolate, well-spaced, distinct beneath,
obscure above; petiole 2-10 by 1-3 mm, terete or
flat above near the base. Inflorescences 3 or 2,
axillary, borne on separate branches, including the
bracts densely golden yellow pubescent; bracts
ovate-acute, 1—-1!/, by !/,-1 mm. — ¢ Inflorescence
slender, lax, branched, thyrsoid, up to 8cm long,
bearing up to 100 flowers; 3 flower 11/,-2 mm 2,
sessile along the axes, in cymoid clusters of 3-10;
perianth lobes 11/,-2 by 1 mm, densely appressed-
hairy outside; filaments 1—-11/, mm, anthers ovoid-
reniform, c. 1 mm @; pistillode strongly rudimen-
tary. — 2 Inflorescence (as seen in a very young
infructescence) a branched panicle, up to 5cm,
carrying 2-20, sessile or short-stalked flowers;
perianth lobes narrow ovate-acute, c. 2 by 1 mm,
densely appressed short-hairy outside; ovary
(young fruit) ovoid, densely short-, appressed-
hairy, c. 3 by 2 mm; stigmatic arms up to 1 cm.
Infructescence up to 10 cm long, axes c. 1 mm thick,
bearing (2-)5—15(—20) fruits. Drupe ovoid-com-
pressed, densely short-appressed-pubescent, 8-10
by 6-8 by 3-5 mm, short-beaked.

1977]

ULMACEAE (Soepadmo) 15

Distr. Malesia: Malaya (rather rare), Sumatra
(rare), Borneo (common), Moluccas (rare), New
Guinea (rare, mainly in W.).

Ecol. Scattered as an understorey shrub or tree
in lowland forests, 0-700 m. Fi. fr. Jan.—Dec.

Vern. Malaya & Sumatra: hampas tébu,
hampélas burung, médang berbulu, m. kasap, M;
Kalimantan: kayu ruas, lémpung bulu, Bandjar;
Sarawak: puloh, Iban; New Guinea: warpis, Biak.

Note. Closely related to G. nervosa, but readily
distinguished from the latter by the characters
mentioned in the key. Specimens from Malaya
(mainly from the Kluang area in Johore and
Pahang), including the type, have much thinner
leaves with a distantly serrulate margin than those
from Sumatra, Borneo, and New Guinea. Since in
most cases the field notes of the Malayan speci-
mens indicate that the height of the tree was never
more than 3 m, it is assumed that these specimens
have been collected from saplings.

4. Gironniera subaequalis PLANCH. Ann. Sc. Nat.
III, 10 (1848) 339, p.p., excl. var. ceylanica; BL.
Mus. Bot. 2 (1856) 73, incl. var. brevistylis BL.,
var. scabrida BL. et var. serrulata BL. l.c. 74; Mia.
Fl. Ind. Bat. 1, 2 (1859) 222; PLANcH. in DC.
Prod. 17 (1873) 206; Hook. f. Fl. Br. Ind. 5 (1888)
485; Hemsv. J. Linn. Soc. Bot. 26 (1894) 452;
BRANDIS, Ind. Trees (1906) 596; J. J. SMitHin K. &
V. Bijdr. 12 (1910) 666; Nova Guinea 8 (1912) 892,
incl. var. papuana J. J. S.; Laut. Bot. Jahrb. 50
(1913) 326; Merr. En. Born. (1921) 217; En.
Philip. 2 (1923) 35; Ripv. Fl. Mal. Pen. 3 (1924)
320; GAGNEP. Fl. Gén. I.-C. 5 (1927) 678; CorNER,
Ways. Trees (1940) 690; Back. & Baku. f. FI.
Java 2 (1965) 12; SoEPADMO in Whitmore, Tree FI.
Mal. 2 (1973) 419, f. 2. — Sponia annulata TeyYsM.
& Brinn. Ned. Kruidk. Arch. 3 (1855) 408. — G.
costata Mia. in Zoll. Syst. Verz. (1855) 88; Fl. Ind.
Bat. 1, 2 (1859) 223. — G. chinensis Bru. FI.
Hongk. (1861) 324. — G. nervosa var. subaequalis
(PLANCH.) Kurz, For. Fl. Burma 2 (1877) 470. —
G. amboinensis Laut. Bot. Jahrb. 50 (1913) 326. —
G. longifolia Crats, Kew Bull. (1918) 371. — G.
sumatrana GANDOG. Bull. Soc. Bot. Fr. 66 (1919)
288. — G. blumei GANDOG. I.c. 288. — G. borneen-
sis GANDOG. l.c. 288. — G. ferruginea GANDOG.
Lc. 289. — Fig. 26h, m, p-r.

Small to large-sized tree up to 40 m, 60cm @.
Bark smooth to finely fissured, pustular or lenti-
cellate. Terminal bud 2-3 by '/,—'/, cm; stipules
linear-lanceolate, 1'/,-2'/, by */,—'/,cm. Leaves
thick-coriaceous, broad ovate-elliptic or elliptic-
oblong, (6-)12-16(-21) by (3'/,—)5-8(-13) cm,
index 2-2'/,, except for midrib and nerves glabrous;
base attenuate or rounded, unequal-sided; margin
entire or occasionally, especially when young,
distantly serrulate at least for the upper half; apex
rounded or attenuate-acute; midrib and nerves
raised beneath, flattish above, sparsely or densely
yellowish-brown appressed short-hairy beneath;
nerves (6-)8(-10) pairs, 10-15 mm spaced, at more
than 60° with the midrib, arcuating and anasto-
mosing along the margin; reticulations subscalari-
form, fine, dense or rather well-spaced, slightly
and clearly visible beneath, flattish and faintly
visible above or obscure; petiole 5-15 by 1-2 mm,
terete or adaxially flat near the base, sparsely or
densely appr , simple, short, yellowish-brown

pubescent, glabrescent. Inflorescence $ or Q,
axillary, borne on separate branchlets or rarely on
the same branchlet, including the bracts sparsely
to densely short, yellowish-brown, appressed-hairy,
glabrescent; bracts ovate-acute, 1-2 by */,-1 mm.
— 6 Inflorescence paniculate, pendent, much-
branched, 40-100-flowered, axes up to 3-7cm
long, '/.-1 mm thick; ¢ flowers 11/,-2!/,mm @,
sessile and solitary along the axes or in clusters of
3-5 on short, condensed secondary branches of the
panicle; perianth lobes sparsely short appressed-
pubescent, glabrescent, broadly ovate-rounded,
2-2'/, by 11/,-2 mm; filaments 11/,-2 mm, anthers
ovoid-subreniform, c. 1 by 1mm; pistillode
strongly rudimentary. — 2 Inflorescence racemose,
unbranched or more commonly branched, axes
3-6 cm long, 1-1'/, mm thick, bearing (2-)5-10
(—15) flowers; 2 flowers solitary and short-stalked
along the axes, 2-4 by 2mm; perianth lobes
broadly ovate-acute, 1'/,-2 by 1'/, mm, sparsely
appressed-pubescent outside; ovary 2-3 by 2 mm,
densely appressed-pubescent, glabrescent; stig-
matic arms up to 2 cm. Jnfructescence with a sturdy
axis up to 2mm thick, 5—10cm long, bearing (2—)5-8
(—10) fruits. Fruit 1-11/, by 8-9 by 5-6 mm, sparsely
appressed-pubescent, glabrescent, beak 2—5 mm.

Distr. A rather variable species widely distri-
buted in the Andaman Is., Burma, China (Yunnan,
Kwangtung, Canton, Hainan), Hongkong, Indo-
China, Thailand, throughout Malesia (except the
Lesser Sunda Is.).

Ecol. Understorey shrub or tree in primary and
secondary forest, 0-1300m, more commonly in
the lowland between 200-500 m. Fi. fr. Jan.—Dec.

Vern. Malaya: hampas tébu, médang kasap, M;
Sumatra: silu, siluk, M; W. Java: ki bulu, S; N.
Borneo: kuayun, ruwayon, Dusun, untoh bulu,
Iban; Kalimantan: katul, Bulungan; Anambas
Is.: pupoh, M; W. New Guinea: gawa, géwa,
mégawa, Mooi, nadjun, nitjun, Kebar, bobohufeka,
Manikiong, warpis, Biak.

5. Gironniera parvifolia PLANCH. Ann. Sc. Nat.
III, 10 (1848) 338 (‘parvifolium’); Mig. Fl. Ind.
Bat. 1, 2 (1859) 223; PLANcH. in DC. Prod. 17
(1873) 206; Hook. f. Fl. Br. Ind. 5 (1888) 486;
RID. Fl. Mal. Pen. 3 (1924) 321; Corner, Ways.
Trees (1940) 689; SogPADMO in Whitmore, Tree
Fl. Mal. 2 (1973) 419, f. 2. — G. subaequalis var.
ceylanica PLANCH. Ann. Sc. Nat. III, 10 (1848)
339; THw. En. Pl. Zeyl. (1861) 268; Hook. f/.
Fl. Br. Ind. 5 (1888) 485. — Helminthospermum
scabridum Tuw. in Hook. J. Bot. Kew Misc. 6
(1854) 303, t. 9c. — G. paucinervia Merr. J. Str.
Br. R. As. Soc. nm. 77 (1917) 189; En. Born.
(1921) 217. — G. zeylanica GANDOG. Bull. Soc.
Bot. Fr. 66 (1919) 288. — G. scabrida (THw.)
ALSTON in Trimen, Fl. Ceyl. 6 (1931) 267. — Fig.
26k, n.

Shrub or small-sized tree up to 15 m, 20cm @.
Bark smooth to finely fissured, lenticellate, grey-
green or grey-brown. Terminal bud (5—)8—15(—18)
by 1-3 mm; stipules linear-acute, 8-15 by 2-3 mm.
Leaves chartaceous to thin-coriaceous, rarely
coriaceous, elliptic-lanceolate or elliptic-obovate,
very rarely ovate-elliptic, (4—)8-12(-16) by (1'/,-)
3--4(--5) cm, index 3—3'/,, glabrous, ; base attenuate
or rounded, more or less equal-sided; margin
entire or sometimes minutely and distantly serrulate

76 FLORA MALESIANA

in the upper half (extra-Mal. and young speci-
mens); apex attenuate-acute or rounded-
acuminate; midrib and nerves slightly raised
beneath, flattish above; nerves (5—)6—8(-9) pairs,
7-10 mm spaced, at up to 60° with the midrib,
arcuating and anastomosing along the margin;
reticulations subareolate, fine, dense, visible
beneath, obscure above; petiole 5—7 by 1—1'/, mm,
sulcate. Inflorescences 3 or 9, axillary, borne on
separate branchlets, including the bracts sparsely,
short, appressed-pubescent, glabrescent; bracts
Ovate-acute, membranous, c. 1 by !/,mm. —
3 Inflorescence a slender, pendulous, simple or
branched raceme, bearing 15-30 flowers, axes up
to 5 cm long, 0.2-0.3 mm thick; 3 flowers solitary
along the axes, short-stalked, 1'/,2mm @;
perianth lobes broad ovate, c. 1!/.-2 by 1—-1"/, mm;
filaments 1-1'/, mm long, anthers ovoid-reniform,
3/,-1 mm @; pistillode strongly rudimentary. — 9
Inflorescence a slender, 1—3(—5)-flowered raceme,
4-10 cm long, axes 0.2—0.3 mm thick; 2 flower 2-3
by 2 mm, short-stalked; perianth lobes mostly 4,
unequal in size, 11/,-2!/, by 1-2 mm; ovary sparsely
short appressed-pubescent, 2-3 by 2mm; stig-
matic arms up to 14/, mm. Jnfructescences slender,
up to 10 cm long, bearing 1—3(—5) fruits, axes thin-
ner than 1 mm. Fruit c. 8 by 6 by 4mm, sparsely,
short appressed-pubescent, glabrescent; beak up
to 5 mm.

Distr. Ceylon; in Malesia: Malay Peninsula
(incl. Singapore; very common), Sumatra (rare),
Borneo (common).

Ecol. In primary as well as in secondary forest,
0-1300 m, but mostly in the lowland, on various
types of soil including those derived from lime-
stone. F/. Jan.—April, fr. Oct.—Dec.

Vern. Malaya: hampas tébu, médang kasap, M;
Sarawak: tépadé, Kelabit.

Note. As in other species of the genus, most of
the specimens so far available suggest that the
plant is monoecious. However, in MAINGAy K.D.
1470 from Malaya both the ¢ inflorescence and
infructescence are found on separate branchlets of
the same collection number. Assuming that these
branchlets were collected from the same tree, it
would suggest that the species is monoecious but
producing ¢ and 2 flowers at different stages of its
growth. More field work is required to determine
its breeding system.

6. Gironniera rhamnifolia BL. Mus. Bot. 2 (1856)
(4t. 255 Vio Hl Inds Bats W262 (1859)5223-
PLANCH. in DC. Prod. 17 (1873) 206; LAuT. Bot.
Jahrb. 50 (1913) 326. — Fig. 261.

[ser. I, vol. 8?

Shrub to medium-sized tree, up to 25 m, 40 cm
@. Bark smooth, pale grey-brown. Branchlets
initially densely set with simple, yellowish-brown,
soft, erect hairs, glabrescent. Terminal buds 1-1'/,
by 0.2-0.3 cm; stipules narrow elliptic-lanceolate,
c. 1#/, by 4/,cem. Leaves chartaceous to thin-
coriaceous, glabrous, elliptic-lanceolate or rarely
ovate-elliptic, (5—)8-15(-17) by (2-)4-6(-8) cm,
index 21/,-31/,; base rounded or attenuate, +
equal-sided; margin entire; apex acute to acumi-
nate; midrib and nerves slightly raised beneath,
flattish to slightly impressed above, often densely,
short, appressed-hairy; nerves 10-13 pairs, at an
angle of over 60° with the midrib, arcuating and
anastomosing towards the margin; reticulations
fine, lax, subscalariform to subareolate, faintly
visible beneath; petiole 5-10 by 1-1'/,mm,
glabrous, shallowly furrowed or flat above near the
base. Inflorescences 3 or 2, axillary, including the
bracts sparsely short, yellowish-brown appressed-
pubescent; bracts narrow ovate-acute, c. 1 by
1/,mm. — 6 Inflorescence an unbranched or
branched panicle, 3-5cm long, bearing 15-30
flowers; 3 flowers sessile and solitary along the
axes, c. 1-l'/,mm 2%; perianth lobes ovate,
11/,-2 by 1mm, sparsely short, appressed-
pubescent outside; filaments 11/,-2 mm, anthers
ovoid-reniform, +/,-1mm @; pistillode strongly
rudimentary. — 2 Inflorescence an unbranched,
(1-)3-5-flowered panicle, 3-4cm long with the
axes c. !/,-1 mm thick; 2 flowers 2-3 by 2 mm;
perianth lobes 5, ovate, 11/,-2 by 1 mm; ovary 2-3
by 2mm, densely short, appressed-pubescent,
glabrescent; stigmatic arms up to 11/, cm. Infruc-
tescence up to 5 cm long, axes c. 1 mm or thicker,
bearing (1—)3—5 fruits. Fruit + glabrous, c. 7 by 6
by 3 mm, short-beaked.

Distr. Malesia: N. Moluccas (Morotai), New
Guinea (incl. Biak, Japen, and Polima Is.).

Ecol. Primary and secondary forests, locally
often very common in rocky or stony habitats
including limestone, 0-1000 m. FI. fr. Jan.—Dec.

Vern. W. New Guinea: warpis, Biak, tamanpara,
Japen, ramé, Iko.

Excluded

Gironniera glabra MERR. Philip. J. Sc. 1 (1906)
Suppl. 42 = Paratrophis glabra STEEN. J. Bot. 72
(1934) 8 = Chevalierodendron glabrum Leroy,
C. R. Ac. Sc. Paris 227 (1948) 146 = Streblus
glaber (MERR.) CORNER, Gard. Bull. Sing. 19
(1962) 221 (Moraceae).

IRIDACEAE (D. J. L. Geerinck, Bruxelles)!

Perennial herbs, often with bulbs, tubers or rhizomes, sometimes undershrubs.
Leaves simple, equitant (except in Crocus), with parallel nerves. Inflorescences
terminal or axillary, in cymes, spikes or panicles, sometimes very contracted or
flowers solitary, bracteate and with | or 2 spathes. Flowers bisexual, actinomorphic
to zygomorphic, often marcescent. Tepals free or united into a tube, in 2 whorls,
the inner ones rarely inconspicuous (Patersonia). Stamens 3 or exceptionally 2 with
1 staminode (in the Australian Diplarrhena), free or united into a tube, basifixed
or dorsifixed, opposite to the outer tepals. Ovary inferior (or superior in the
Tasmanian IJsophysis), 3-celled with axillary placentas; style entire or trifid,
sometimes tepaloid; stigmas 3 or 6, terminal or sometimes axillary, alternating
with or opposite to the outer tepals; ovules generally numerous. Fruit capsular,
dehiscing loculicidally, apically or irregularly. Seeds angular, flat or globose, some-
times winged.

Distribution. Cosmopolitan, with c. 60 genera and c. 800 spp., predominantly in the tropics and the
southern hemisphere. In Malesia: only two Australasian genera each with 1 sp., and four exotic ones
introduced and naturalized.

Many are cultivated in botanic gardens and occasionally in private gardens; see for an elaborate
treatment BACKER, Handb. FI. Java 3 (1924) 116-130 and BACKER & BAKH. f. Fl. Java 3 (1968) 144-154.

Ecology. Both native species are characteristic mountain plants.

Morphology. This family is usually herbaceous, but in a few genera (Klattia, Nivenia, Patersonia,
Witsenia) stems may be woody at the base. The leaves are equitant and are laterally compressed, the two
halves are free at the sheathing base and gradually fused to the top. The flowers are actinomorphic to
distinctly zygomorphic with intermediate forms, sometimes in the same genus.

Uses. Belamcanda chinensis and Eleutherine palmifolia are used for medicinal purposes, probably mainly
eet 9s of their glucosides; cf. HEYNE, Nutt. Pl. (1927); BURKILL, Dict. (1935); QUISUMBING, Med. PI.

ilip. (1951).

KEY TO THE GENERA

1. Flowers all sessile. Tepals united into a tube. Capsules included in bracts or spathes.
2. Flowers actinomorphic. Inner tepals inconspicuous. Stamens united into an undivided or trifid tube.
Caespitose or rhizomatous herbs or undershrubs . . . . . 2... ee ee ee 1. Patersonia
2. Flowers + zygomorphic. Tepals unequal. Stamens free. Cormogenous herbs. . . . 5. Gladiolus
1. Flowers all pedicelled. Tepals free or nearly so. Capsules exserted from bracts or spathes.
3. Tepals clawed. Stamens free.
4. Cormogenous herbs. Stamens appressed against the back of the style-arms. Ovary not beaked.

eae with bilobed arma... |. sc. a) Roe ee An ie og ee es ce 6. Trimezia
4. Rhizomatous herbs. Stamens not appressed against the back of the style-arms. Ovary beaked.
Peres Sith UNGIVIOSD BTM: ey end. ey arte lin ale Pl spa Ni ae 3. Belamcanda

3. Tepals not clawed.
5. Tepals shortly connate, subequal. Stamens united into a tube, rarely nearly free. Caespitose to
RICE TPIS. foe Dy oe rh ee eae Ys a ea ey eee a Re ns 2. Sisyrinchium
5. Tepals free, the inner ones smaller. Stamens free. Bulbous herbs. . ..... . 4. Eleutherine

1.PATERSONIA

R.Br. ex KER-GAWL. Bot. Mag. (1807) t. 1041, nom. cons.; Prod. Nov. Holl.
(1810) 304; Bru. FI. Austr. 6 (1875) 400; GreRINCK, Bull. Jard. Bot. Nat. Belg. 44
(1974) 41. — Genosiris LaBiLL. Nov. Holl. Pl. Sp. 1 (1804) 13, t. 9. — Fig. 1-3.
Caespitose to rhizomatous herbs or undershrubs. Inflorescences terminal, in
few-flowered contracted cymes, each with 2 spathes. Flowers actinomorphic,
sessile, bluish to purple, exceptionally yellow or whitish, Tepals dimorphic, united
into a long and filiform tube at the base, the inner lobes inconspicuous. Stamens 3,

(1) With co-operation by the General Editor.
(77)

FLORA MALESIANA [ser. I, vol. 8?

Fig. 1. Patersonia lowii StapF on Mt Losir, Gajolands, N. Sumatra, at c. 2400 m altitude (Photogr. DE
WiLpe-Duyries, April 1975, n. 16390).

1977]

IRIDACEAE (Geerinck) 79

united into an undivided or trifid tube. Ovary cylindrical, lanate; style entire;
stigmas 3, subfoliaceous, alternating with the outer tepals. Capsules loculicidal,

included. Seeds angular or ellipsoidal.

Distr. Australia and Tasmania (12 spp.), and Malesia (1 sp.).

Ecol. Open, low shrubberies, heaths and sedge-lands, 2000-3500 m.

Note. In sterile state the habit of Patersonia is strikingly resembling that of the sedge genus Machaerina.
A specimen mentioned by WENT f. (Nova Guinea 14, 1924, 114) as Patersonia from Mt Goliath (DE Kock
50) belongs to Machaerina, as corroborated anatomically by Dr P. Baas.

1. Patersonia lowii STAPF, Trans. Linn. Soc. Bot.
II, 4 (1894) 241, t. 20, f. 7-9; MerR. Philip. J. Sc. 2
(1907) Bot. 268; En. Born. (1921) 119; En. Philip.
1 (1923) 220; Not. Nat. Ac. Nat. Sc. Philad. n. 47
(1940) 2. — P. borneensis STAPF, Trans. Linn. Soc.
Bot. II, 4 (1894) 241; Grsss, J. Linn. Soc. Bot. 42
(1914) 165; Merr. En. Born. (1921) 119. — P. novo-
guineensis GipBs, Arfak (1917) 101; WENT f. Nova
Guinea 14 (1924) 114, incl. var. auriculata WENT,
l.c.; HaTus. Bot. Mag. Tokyo 56 (1942) 426. —
Fig. 1-3.

Tufted herb, 15-60cm high. Leaves basal to
subbasal, flat to + biconvex, 5-60 cm by 3-6 mm,

er

Fig. 2. Patersonia lowii Starr. a Habit, b. capsule,
both nat. size, c. seed, x 5 (a VAN ROYEN &
SLeumer 7102, b—-c Brass 22259).

reddish or rarely whitish tomentellous to glab-
rescent along the margins towards the top, +
glaucous. Inflorescences equalling the leaves or
nearly so; peduncle 8-50 cm long, glabrous, the
lower part surrounded by a persistent central leaf;
spathes suboval to narrowly suboval, 2'/,-5 cm by
7-12 mm, dark brown-orange, greyish when
growing old, distinctly striate, with a red-hairy line
on the keel to glabrous. Flowers bluish to pale
mauve or purple, sometimes whitish; perigone-
tube 2-2'/, cm long, the outer lobes 8-16 by 6-10
mm. Staminal tube entire; anthers yellow. Ovary
c. 5 mm long. Capsules 2-3 cm long; valves 3-4 mm
wide. Seeds c. 2 mm, black.

Distr. Malesia: Sumatra (Gajolands: Mt Losir),
Borneo (Mts Kinabalu and Murud, Kalabit
Highlands), Philippines (Mindoro), New Guinea
(Tamrau Range, Arfak Mts, Mamberamo River,
Central to Milne Bay Districts). Fig. 4.

Ecol. Open shrubby vegetation or open forests,
sedge meadows and heaths, on stony or impervious

Fig. 3. Patersonia lowii Starr. Same locality as in
fig. 1

80 FLORA MALESIANA [ser. I, vol. 8?

clay soils, often gregarious, 2000-3500 m. Fi. fr. A variable species concerning the indument; the
Dec.—Aug. Flowers open early in the morning but disjunct populations are not uniform. It seems to
become soon marcescent. be allied to the widely distributed Australian

Vern. New Guinea: atetdzjii, Mt Arfak, P. fragilis (LABILL.) ASCHERSON & GRAEBNER,
Manikiang lang. which differs by glabrous leaves and spathes,

Notes. The sizes of the tepals and of the seeds _ inflorescences much shorter than the leaves and the
have been taken from the original descriptions of lower part of the peduncles never surrounded by a
the synonymous taxa. central leaf.

p\—I—*= Bene < 28 A

5 . Pin. 2%

oT RRaStieerS
SS

ae

Fig. 4. Range of Patersonia lowii STAPF.

2. SISYRINCHIUM

LINNE, Sp. Pl. (1753) 954; JoHNsTON, J. Arn. Arb. 19 (1938) 376; FosTER, Contr.
Gray Herb. 166 (1948) 28. — Renealmia R.Br. Prod. (1810) 592, pro parte, non L.
f. 1781. — Fig. 5-6.

Caespitose to rhizomatous herbs. /nflorescences axillary or terminal in panicles
of fan-shaped and few-flowered cymes or of many-flowered clusters, each with 1-2
spathes. Flowers actinomorphic, pedicelled, bluish or yellowish. Tepals subequal,
shortly connate at the base. Stamens 3, united into a tube at the base, rarely nearly
free. Ovary: style trifid, the lobes filiform; stigmas 3, small, alternating with the
outer tepals. Capsules exserted. Seeds small, globose.

Distr. About 100 spp. in Central and South America, 1 native sp. in New Zealand, Australia and East
Malesia (New Guinea); also one species introduced.

The Papuan-Australasian species was mostly arranged in the closely allied genus Libertia Spr. How-
ever, the tepals are not dimorphic but about similar so that it must be arranged in Sisyrinchium.

1977]

IRIDACEAE (Geerinck) 81

KEY TO THE SPECIES

1, Leaves cauline, distichous. Inflorescences in 1—2-flowered, contracted cymes, a few at a cauline leaf.

Flowers c. 5 mm long. Stamens nearly free, glabrous

1. S. pulchellum

1. Leaves mostly basal. Inflorescences in 3-6-flowered cymes, generally 2 at a cauline leaf. Flowers c. 7 mm

long. Stamens united in a hairy tube, trifid at the top

1. Sisyrinchium pulchellum (R.Br.) F.v.M. Fragm.
Phyt. Austr. 7 (1870) 92; Trans. R. Soc. Victoria 1
(1889) 34; GEERINCK, Bull. Jard. Bot. Nat. Belg. 44
(1974) 59. — Renealmia pulchella R.Br. Prod.
(1810) 592, to replace S. pulchellum R.Br. I.c. 305.
— Libertia pulchella Spr. Syst. Veg. 1 (1824) 169;
LANE-POOLE, For. Res. (1925) 77; Laur. Bot.
Jahrb. 62 (1929) 462; STEEN. Bull. Jard. Bot.
Btzg III, 13 (1934) 220; HooGL. Blumea 4 (1958)
235; BaLGooy, Pac. Pl. Areas 2 (1966) 286;
L. Moore, New Zeal. J. Bot. 5 (1967) 267. — Fig.
Sa.
Glabrous herb, 10-35 cm high. Leaves cauline,
distichous, linear, 4-22 cm by 1-10 mm. Inflores-
cences in 1—2-flowered, contracted cymes, a few ata
cauline leaf, each cyme with 1 spathe: this 4-15
(440) by 2-6mm. Pedicels to 4cm. Flowers c.
5mm long. Tepals white to yellowish, c. 4 by
11/,-2 mm. Stamens nearly free, filaments c. 4 mm,
anthers 1 mm long. Ovary ellipsoid, c. 1 mm long;
style with undivided part 1-2 mm, the lobes 1-2
mm long. Capsules globular, 2-5 mm @; valves
c. 2mm wide. Seeds black, 1 mm @.

Distr. New Zealand, Australia (New South
Wales, Victoria, Tasmania), and East Malesia:
New Guinea (Lake Habbema area and Mt Antares
in West, many localities in East). Fig. 6.

Fig. 5. Sisyrinchium pulchellum (R.BR.) F.v.M. a.

isecicence. nat. size. — S. micranthum Cav.

b. Inflorescence, nat. size (a KALKMAN 4497, b VAN
ROvEN 16028).

2. S. micranthum

Ecol. Open forests and shrubby vegetation, in
tree fern heath and alpine grassland, common on
Mt Sarawaket in Libocedrus-Dacrydium forest
(LANE-PooLe), 2400-3700 m. F/. mostly Jan.—Aug.

Note. Moore (/.c. 255-275) studied the variation
in New Zealand and distinguished three species
among which are two polyploids.

Fig. 6. Range of Sisyrinchium pulchellum (R.BR.)
F.v.M.

2. Sisyrinchium micranthum Cav. Diss. Bot. 6
(1788) 345, t. 191; Back. Handb. Fl. Java 3 (1924)
125; JoHNsTON, J. Arn. Arb. 19 (1938) 390;
Foster, Contr. Gray Herb. 166 (1948) 31; WILLIs,
Handb. FI. Victoria 1 (1962) 335; STEEN. Blumea 15
(1967) 154; Back. & Baku. f. Fl. Java 3 (1968)
150. — Fig. 5b.

Slender herb, 5-25cm high, glabrous, with a
flat stem. Leaves few, mostly basal, linear, 3-12 cm
by 1-5 mm. Inflorescences in 3-6-flowered cymes,
generally 2 at a cauline leaf, each cyme with 2
spathes, outer spathe 20-30mm long, inner
spathe 15-25 mm long, both 1-2 mm wide. Flowers
c. 7mm long. Tepals yellow with red or brown
markings, c. 6 by 1 mm. Stamens united in a hairy
tube, trifid at the top; filaments c. 1 mm long;
anthers '/, mm long. Ovary ellipsoid, laxly hairy,
c. 1 mm long; style with the undivided part c. 1 mm
long, the lobes c. */,-*/,mm long. Capsules
globose, 2-3mm long; valves c. 2mm _ wide.
Seeds black, 1 mm @.

Distr. Southern to Central America; naturalized
in Australia, New Caledonia, Fiji, New Zealand,
and also in Malesia: W. Java (Tjibodas), E. New
Guinea (Morobe Distr.: Edie Creek), perhaps
elsewhere.

Ecol. In the vicinity of the Tjibodas Botanic
Garden as a weed in grassland and waste places, at
c. 1450 m, certainly escaped from the garden. In
Papua at c. 1950 m almost certainly introduced
from Australia where it was first recorded about
1870. Poisonous to stock.

82 FLORA MALESIANA [ser. I, vol. 8?

3. BELAMCANDA

ADANS. Fam. 2 (1763) 60.

Rhizomatous herbs. /nflorescences terminal, in panicles of flowered and corym-
biform cymes, each with 2 spathes. Flowers actinomorphic, pedicelled, yellowish to
orange. Tepals subequal, shortly connate, clawed. Stamens free. Ovary beaked;
style trifid, the lobes short; stigmas 3, small, alternating with the outer tepals.
Capsules exserted. Seeds large, globose.

Distr. Monotypic, native of China and Japan, cultivated in Malesia and many other countries and

sometimes naturalized.

1. Belamcanda chinensis (L.) DC. in Redouté,
Liliac. 3 (1805) t. 121; Koorp. Minah. (1898) 313;
C. B. Ros. Philip. J. Sc. 6 (1911) Bot. 196; Koorp.
Exk. Fl. Java 1 (1911) 312; Merr. Fl. Manila
(1912) 152; Philip. J. Sc. 11 (1916) Bot. 260; En.
Philip. 1 (1923) 220; Back. Handb. Fl. Java 3
(1924) 124; Heyne, Nutt, Pl. (1927) 461; GAGNEP.
Fl. Gén. I.-C. 6 (1934) 675; Burk. Dict. (1935) 315;
STEEN. Fl. Sch. Indon. (1949) 147; Quis. Med.
Pl. Philip. (1951) 181; HENDeERs. Mal. Wild FI.
Monoc. (1954) 168, f. 98; OHw1, Fl. Japan (1965)
316; BAcK. & BAKH. f. Fl. Java 3 (1968) 149. —
Balem-canda schularmani RHEEDE, Hort. Mal. 11:
73, t. 37. — Ixia chinensis LINNE, Sp. Pl. (1753)
36. — Pardanthus chinensis KER-GAWL. in KGnig
& Sims, Ann. Bot. 1 (1805) 247; BL. En. Pl. Jav.
(1827) 26; Zoi. Syst. Verz. 1 (1854) 70; FILet,
Pl. Bot. Tuin Weltevreden (1855) 13; Mia. FI. Ind.
Bat. 3 (1859) 579; BLANco, FI. Filip. ed. 3 (1877-
83) t. 376.

Corymbosely branched, glabrous herb, 1-11/, m
high. Leaves distichous, basal and cauline, broadly
linear, 30-60 by 2-4cm, glaucous. Inflorescences
in 6—12-flowered cymes, once or twice branched;
spathes membranous, c. 10 by 4mm. Pedicels
24cm, persistent. Tepals yellowish to orange,
with red spots, unguiculate, obovate, 25-35 by

c. 7mm, outer ones largest. Filaments filiform;
anthers c. 10 mm long. Ovary ellipsoid, c. 5mm
long; style-arms gradually thickened upwards.
Capsules obovate, 15-20 mm long; valves 8-12 mm
wide. Seeds shining black, 5mm @.

Distr. Native in China and Japan; in Malesia
introduced and cultivated, locally naturalized
(Sumatra, Java, S. Celebes, Philippines, Moluccas:
Morotai, Banda, Ceram); cultivated and locally
naturalized in many tropical and subtropical coun-
tries, e.g. Hainan, Taiwan (Formosa), Tonga, etc.

Ecol. In Java it is mainly naturalized in the
eastern part between 750 and 2100 m, occurring in
thickets and forest edges, and said not to grow
well at low altitude.

The flowers open in the forenoon and have
withered by midday.

Uses. HEyneE and BurRKILL (//.cc.) mention usage
for several minor medicinal purposes, the dried
rhizome being used as a purgative and for com-
plaints of the chest and liver, efc.

Vern. Sumatra: piso-piso, Batak. Java: akar
tjamaka, djamaka, gégébangan brodjo lintang,
suliga, S, sémprit, wordi, J. Celebes: kariménga
kulo, katna, kétép, kétéw, kiris, Minahasa, Alf.
lang., tagari, Bonthain. Philippines: abinaco.

4. ELEUTHERINE

HERBERT, Bot. Reg. 29 (1843) t. 57, nom. cons.

Bulbous herbs. Inflorescences axillary in few-flowered and contracted cymes,
each in 2 spathes. Flowers actinomorphic, pedicelled, whitish. Tepals free, the inner
ones smaller. Stamens 3, free. Style deeply trifid; stigmas 3, small, alternating with
the outer tepals. Capsule loculicidal, exsert. Seeds ellipsoidal to angular.

Distr. In America 2 spp. and according to GAGNEPAIN (FI. Gén. I.-C. 6, 1934, 676) 2 spp. in Indo-
China. Introduced in Malesia, and locally naturalized.

1. Eleutherine palmifolia (L.) MERR. Philip. J. Sc. 7
(1912) Bot. 233; Fl. Manila (1912) 153; Sp. Blanc.
(1918) 104; En. Philip. 1 (1923) 220; Quis. Med.
Pl. Philip. (1951) 182; BAck. & BAKH. f. Fl. Java 3
(1968) 150. — Sisyrinchium palmifolium LINnE,
Mant. 1 (1767) 122. — Sisyrinchium bulbosum
MILL. Gard. Dict. ed. 8 (1768) n. 3. — Ixia
americana AUBL. Pl. Guian. 1 (1775) 33. — Moraea
plicata Sw. Fl. Ind. Occ. 1 (1797) 82. — Antholyza
meriana (non L.) BLANCO, FI. Filip. (1837) 24; ed. 3,

1 (1877) t. 100; MerRR. Publ. Gov. Lab. Philip. 27
(1905) 85. — E. plicata HERBERT, Bot. Reg. 29
(1843) t. 57. — E. bulbosa (MILL.) URBAN in
Fedde, Rep. 15 (1918) 305; LEONARD, Bull. Soc.
R. Bot. Belg. 84 (1951) 55. — E. americana MERR.
ex Heyne, Nutt. Pl. Ned. Ind. ed. 2, 1 (1922) 502;
Back. & SLoot. Handb. Thee (1924) 91, t. 91;
Back. Handb. FI. Java 3 (1924) 126; HEyNg, Nutt.
Pl. (1927) 462.

Herb, 30-60 cm high, glabrous; bulb red, ovoid,

1977]

2'/,-5 cm long. Leaves basal 3-4 from each bulb,
and cauline, narrowly elliptic, plicate-nerved,
25-60 by 1-2!/, cm. Inflorescences in 4-10-flowered
cymes; spathes 12-16mm long, green. Flowers
very fugacious, white. Tepals obovate, c. 15 mm
long. Stamens yellow to orange, 8-10 mm long.
Ovary ellipsoidal, c. 2mm _ long; style-arms
filiform, yellow; stigmas white. Capsules globose,
c. 6mm long. Seeds dark brown, c. 2mm @.

Distr. Native in tropical America, cultivated
and naturalized in tropical Africa and in Malesia:
W. Java, W. Borneo, and the Philippines (Luzon,
Leyte, Negros, Mindanao); in Java already noticed
+ 1820.

Ecol. A weed, finally tufted, in estates and waste
places, which multiplied by its tubers; c. 150-1500
m. In Java the scentless flowers open at about 5
o’clock in the afternoon, but have already wilted at
about 7 o’clock.

IRIDACEAE (Geerinck) 83

Uses. According to QUISUMBING /.c. in the
Philippines macerated bulbs are applied on the
stomachs of children to relieve gas pains, and a
decoction is diuretic. According to HEYNE /.c. this
finally strongly stooling, tufted plant is cultivated
and its bulbs have various applications in native
medicine: diuretic, purgative, emetic, against
dysentery, jaundice, etc.

Vern. Vijfuursbloem, D. Java: babawangan, b.
beureum, bawang sabrang, b. sieum, S, bawang
kapal, M, brambang sabrang, luluwan sapi, téki
sabrang, J. Philippines: ahos-dhos, C. Bis., bakong
sa Persia, mala-bauang, rosas sa Siam, Tag., hagu-
sahis, S. L. Bis., palmilla, Spanish.

Note. The tepals are sometimes numerous, up to
15; the number of the stamens is sometimes up to
8; the ovary is sometimes 4—11-locular with the
same number of style-arms.

5. GLADIOLUS

LINNE, Sp. Pl. (1753) 36; GEERINCK, Bull. Jard. Bot. Nat. Belg. 42 (1972) 269;
Lewis & OBERMEYER, J. S. Afr. Bot. Suppl. 10 (1972).

Cormogenous herbs. /nflorescences terminal or sometimes axillary in spikes,
rarely secund. Flowers + zygomorphic, sessile, various in colour, each in a single
spathe. Tepals unequal, united into an often curved tube. Stamens 3, often arched,
free. Style entire; stigmas 3, obovate, alternating with the outer tepals. Capsule
loculicidal, included. Seeds often winged.

Distr. About 180 spp. in Africa, South Europe and West Asia, one locally naturalized in Malesia.

1. Gladiolus natalensis (ECKLON) REINW. ex Hook.
Bot. Mag. (1831) t. 3084; GeerINCK, Bull. Jard.
Bot. Nat. Belg. 42 (1972) 281; Lewis & OBERMEYER,
J.S. Afr. Bot. Suppl. 10 (1972) 44.

var. natalensis.

Stems 50-150 cm. Leaves almost basal, broadly
linear, up to 30 by 4-5 cm. Inflorescences terminal
in 2-25-flowered spikes; spathes 4-8 cm long.
Flowers yellowish to pinkish, often with brown

markings. Perigone-tube curved, 2-5cm _ long,
lobes unequal, the upper 4'/,-Scm, the outer
laterals 3—4'/, cm, the inner laterals 2-3 cm and the
lower 2'/,-3'/, cm long. Filaments curved, 4'/,-6
cm; anthers 1'/,-13/, cm. Ovary ellipsoid, 5-8 mm
long; style curved, 2-7cm; stigmas 5-7 mm.
Capsule ellipsoid, 2-5 cm long; valves 7-10 mm
wide. Seeds flat, winged, golden brown, 7-10mm @.

Distr. Tropical and southern Africa, naturalized
in Malesia: Philippines (Luzon).

6. TRIMEZIA

SALISB. ex HERBERT, Bot. Reg. 30, Misc. (1844) 88; Dievs, Pfl. Fam. ed. 2, 15a
(1930) 497; Foster, Rhodora 64 (1962) 307.

Cormogenous herbs. Inflorescences axillary in few-flowered cymes, each in 2
spathes. Flowers actinomorphic, pedicelled, yellowish to reddish. Tepals free,
clawed, the inner ones shorter and narrower with recurved tops. Stamens free,
against the back of the style-arms. Style trifid, the lobes broadly flat, bilobed;
stigmas small, opposite to the outer tepals. Capsules apically dehiscent, with 3 pores
exserted beyond the spathe. Seeds globose to angular.

Distr. A few species in Central and tropical America, one introduced in Malesia (Malaya, West Java).

84 FLORA MALESIANA

1. Trimezia martinicensis (JACQ.) HERBERT, Bot.
Reg. 30, Misc. (1844) 88; BAckK. Handb. FI. Java 3
(1924) 121 (‘Trimeza’); HENDERS. Mal. Wild FI.
Monoc. (1954) 168, f. 98; Foster, Rhodora 64
(1962) 308; BAcK. & BAKH. f. Fl. Java 3 (1968)
148. — TJris martinicensis JACQ. En. Pl. Carib.
(1760) 12. — T. lurida Saiss. Trans. Hort. Soc. 1
(1812) 308; HENpDerRs. Gard. Bull. S. S. 4 (1928)
341. — Cipura martinicensis KTH in H. B. K. Nov.
Gen. Sp. 1 (1816) 320.

Glabrous herb, 100-150 cm. Leaves basal to
cauline, linear, 20-100 cm long and 8-12 mm wide.
Inflorescences in 3—6-flowered cymes, solitary or
binate; spathes 2-2!/,cm long, 10-25 cm
peduncled. Pedicels 1'/,-31/, cm. Flowers yellow,
brownish at the base. Outer tepals obovate, erect

[ser. I, vol. 8?

to patent, 19-25 mm long and 10-13 mm wide;
inner tepals narrower, S-shape curved. Stamens
3-4 mm long. Ovary ellipsoidal ; style-arms 5-7 mm
long, shortly bilobed. Capsule ellipsoid, 13-20 mm
long. Seeds brown, superficially ribbed.

Distr. Native of Mexico, cultivated and locally
naturalized in Malesia: Malaya, West Java.

Ecol. In sunny or slightly shaded localities,
between grass, originally in Malaya at Kuala
Lumpur, but now not uncommon in Malaya
(HENDERSON), in West Java at Bogor on and around
a native cemetery (BACKER), in both cases escaped
from a Botanic Garden, below 250m. Flowers
expand in the forenoon and have withered by
midday.

Vern. Forenoon yellow flag, E.

CORNACEAE (kK. M. Matthew, Tiruchirapalli, India)!

In the past century Cornaceae were mostly delimited in a wide sense and they
represented a fairly heterogeneous assemblage. HARMs (Ber. Deut. Bot. Ges. 15,
1897, 28 and in E. & P. Nat. Pfi. Fam. 3, 8, 1898, 255) distinguished 7 subfamilies.
Of these Garryoideae were later mostly recognized as a separate family Garryaceae,
Alangioideae as Alangiaceae, Nyssoideae and Davidioideae together as Nyssaceae,
leaving Cornaceae with the remaining three subfamilies Cornoideae, Curtisioideae
(monotypic, South Africa) and Mastixioideae (monotypic, Indo-Malesian tropics).
Cf. WANGERIN, Pfl. Reich Heft 414 (1910) 18.

In recent years, however, the other genera (6) of the Cornoideae, besides Cornus,
have also been recognized as monotypic families, with the exception of Corokia
which was transferred to Saxifragaceae-Escallonioideae. Notably TAKHTAJAN
(Proiskh. Prokruitosem. Rast. : 89, non vidi) is in favour of these monotypic families.
In his ‘Flowering Plants’ (ed. C. JEFFREY; 1969: 227) he accepted 7 segregate
families besides Cornaceae sens. str. (omitting mention of two Madagascan genera,
one of which he had formerly also raised to family rank, according to SHAW,
1973). These 7 families he arranged, together with Araliaceae and Umbelliferae, in
the order Cornales, a phylogenetic construction of affinity not much different from
earlier conceptions. The general impression is thus that the distinction of the
segregate families is largely an inflation in rank.

We have not followed this tendency towards inflation advocated by a few con-
temporary systematists and have accepted Cornaceae in the wide sense. We do not
feel that inflation has the merit of improving scientific insight in the mutual syste-
matical affinities, which remain as they were, either as tribes or as subfamilies,
representing together one phylogenetical whole. In addition the disadvantage of the
inflation is that the multiplication of family names becomes unnecessarily a real
challenge to our capacity to memorize, and deflates firmly established family
concepts.

We briefly mention that further relationships are sometimes suggested with
quite remote groups. RENDLE (Class. Fl. Pl. 2, 1952, 422) suggested alliance with
Caprifoliaceae, e.g. Viburnum; affinity has also been suggested with Saxifragaceae-
Escallonioideae. \t falls beyond the scope of the present account to elaborate further
the extensive literature on the subject.

Cornaceae are in great majority northern extratropical, in which zone also many fossils are known.
There are some stray genera on the southern hemisphere. Mastixia is tropical but was found in abundance
in the Tertiary in the subtropics and warm-temperate regions of the northern hemisphere. See under the
genus.

Note. Besides the native genus Mastixia the family is represented in Java by Aucuba japonica THUNB.
which is sometimes cultivated in the mountains. Cf. BAck. & BAKH. f. Fl. Java 3 (1965) 159. — Ed.

1. MASTIXIA

BLuME, Bijdr. (1826) 654; HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1898) 262;
WANGERIN, Pfl. Reich Heft 414 (1910) 19; HALL. f. Beih. Bot. Centralbl. 34, 2
(1916) 40; DANser, Blumea | (1934) 47; MATTHEW, Blumea 23 (1976) 51, f. 1-6. —
Fig. 1, 3.

(1) Composed from the precursory revision in Blumea 23 (1976) 51-93 by the General Editor.

Thanks are due for the financial assistance by the Netherlands Organization for the Advancement of
Pure Research (Z.W.O.).

(85)

86 FLORA MALESIANA [ser. I, vol. 8?

Unarmed, resinous, evergreen trees up to 40(—60) m; branchlets with pith. Leaves
simple, exstipulate, entire, acute, alternate or (sub)opposite to decussate, sometimes
with domatia. Thyrses terminal on the main shoots, sometimes also on the laterals,
up to 4(—8) times branched, the branches of the first order either (sub)opposite
(‘Oppositae’) or spirally arranged (‘Alternae’); further branchings with a tendency
towards decussate arrangement and terminated by cymes; cymes with the central
flower most often sessile and ebracteolate, lateral flowers pedicelled and bracteo-
late. Bracts and bracteoles ovate to triangular, connate or free, lower bracts
sometimes gradually becoming foliaceous. Flowers bisexual, greenish to yellowish.
Calyx 4-5(-6-7)-toothed or -lobed, persistent. Petals valvate, 4-5(-6), thick, ovate
to oblong-elliptic, inflexed at apex and 2-dentate or fimbriate, sometimes with a

Fig. 1. Mastixia kKaniensis MELCH. ssp. kaniensis. a. Habit, x 2/3, b. terminal cymes, x 3, c. flower, d.
ditto in LS, e. stamen in dorsal and frontal view, f. receptacle containing ovary, disk, and style, all x 6, g.
fruit, x 2/3, h. CS of fruit, x 2/3, i. embryo, x 6 (a-b BSIP 3080, c—f CLEMENS 1890, g-i BSIP 2809).

1977] CORNACEAE (Matthew) 87

median ridge inside, spreading or reflexed. Stamens 4-S(-6), or 8, alternating with
the petals, erect in bud; when 8 in 2 alternate whorls of 4; filaments subulate,
flattened; anthers cordate, dorsifixed, abutting on and alternating with the disk
lobes, latrorse; connective + protruding. Ovary inferior, turbinate, 1-celled, sur-
mounted by a prominent, fleshy, persistent disk c. !/,; the height of the receptacle;
invaginations of the disk abaxially 4-5 (fitting the filaments) and adaxially 8 or 10
(fitting the thecae), becoming shallower with age; style stout, ribbed; stigma
punctiform, sometimes deeply 2-fid or 4~S-lobed, lobes sometimes reflexed. Ovule
1, pendulous laterally from the roof of the cell. Drupe subglobose to oblong, sur-
mounted by calyx and disk; pericarp thin or thick, dark purple to blue when ripe;
endocarp woody, sulcate on one side externally and internally deeply protruding
into the fruit cavity as a wedge-shaped or swollen incomplete septum. Seed fitting
the fruit cavity; testa membranous; endosperm copious; embryo small; cotyledons
foliaceous; radicle elongate.

Distr. About 13 spp. in SE. Asia (Western Ghats & Ceylon, NE. India, Bhutan, Burma, Thailand,
Indo-China, S. Yunnan, Hainan) through Malesia to New Britain and the Solomon Islands. Fig. 2.

Ecol. Primary and secondary forest, often in moist habitats, from sea-level up to 1800(-2400) m.

Fossil endocarps of Mastixioids are found in quantity in the warmer Tertiary in Europe, Great Britain
and North America. Cf. KIRCHHEIMER, Die Laubgewachse der Braunkohlzeit (1957) and D. H. Mat,
Palaontol. Abhandl. Deut. 2 (1) (1964). The Pleistocene Glacial Epoch is held responsible for the contrac-
tion of the range, similarly as happened to Symplocos, Meliosma, and so many other genera of the Tertiary
mixed mesophytic forest on the northern hemisphere.

Taxon. Mastixia was subdivided into two subgenera by WANGERIN (1910) on the 4- and 5-merousness
of the flowers respectively. Though this character is still used for discrimination of species. it seems

Fig. 2. Range of the living species of the genus Mastixia BL. For each district, island or island group the
number of species is given, above the hyphen the endemic ones, below the hyphen the non-endemic ones
(occurring in more than one district).

88 FLORA MALESIANA [ser. I, vol. 8?

artificial for subgeneric rank. Instead, I have proposed another subdivision (1976) into two subgenera,
in one of which (subg. Manglesia) the stamens number 8 and are arranged into 2 whorls, while in subg.
Mastixia the stamens number 4—5(—6) and stand in 1 whorl. Other differential characters support this
subdivision; see also the key.

Anatomy. For general anatomical surveys also giving the older literature see SOLEREDER, Syst. Anat.
Dicot. Stuttgart (1899) 487-495 and ibid. (1908) 171-172; MercaLre & CHALK, Anat. Dicot. Oxford
(1950) 735-741. Additional selected references: Mott & JANssonius, Mikr. 3 (1918) 722-737 (wood
anatomy); ADAMS, J. Elisha Mitchell Sci. Soc. 65 (1949) 218-244 (comparative wood anatomy); JANs-
sONIUS, Blumea 6 (1950) 424 (wood anatomical affinities); VERSTEEGH, Acta Bot. Neerl. 17 (1968) 151-
159 (wood anatomy).

The wood of Mastixia like that of most other Cornaceae is primitive. It has diffuse, exclusively solitary
vessels with scalariform perforations (many-barred), fibre-tracheids, diffuse parenchyma, and hetero-
geneous rays. MOLL & JANssontus /.c. reported vertical intercellular canals in Mastixia rostrata and
M. trichotoma. The latter are absent from M. tetrapetala studied in Leiden. The leaf and twig anatomy of
Mastixia is characterized by the occurrence of secretory canals. This important feature is absent from the
other genera of the Cornaceae. Their presence in Mastixia can be used as an argument to stress the
affinities of Cornaceae with Araliaceae and Umbelliferae of the Cornales for which families they are
typical. — P. BAAS.

Galls. Only two galls have been described by DocTers vAN LEEUWEN (Ned. Kruidk. Arch. 51, 1941,
207) in the species where they most occur, viz M. rostrata and M. trichotoma, both caused by aphids. They
occur, however, rather random in many species and varieties, with preponderance in ser. Oppositae.
None have been found yet in species of subg. Manglesia. There are four kinds: on the stem, the leaf, the
an ORESCE CS and the fruit. Sometimes they can be quite large, as has been cited under the species. See

oS:

Uses. Although trees may reach a considerable size, the scattered occurrence does not contribute to
general use as timber; besides, the timber is not of good quality and is only used for minor purposes. OE
BURKILL, Dict. (1935) 1428.

Notes. In key and descriptions the width of the submature flower is that of the corolla.

About the use of the term ‘merousness’ of the flower it should be remarked that this cannot be used in
the strict sense, as 4- and 5-merous flowers often occur in one inflorescence. If it is said ‘basically 4-merous’,
this means that at least 80°% of the flowers are 4-merous and the same holds for basically 5-merous
flowers, so that the prevalent pattern is obvious.

Moreover it should be remarked that the number of sepals frequently tends to be higher than that of
petals and stamens.

In exceptional cases identification of sterile or immature material must remain uncertain.

Unfortunately no separate key can be provided for fruiting material.

KEY TO THE SPECIES

1. Stamens 8, in 2 whorls of 4. Inflorescence branches 4-angular (at least when young). Calyx subtruncate
with minute, acute teeth. Bracts caducous. Pedicels of lateral flowers of terminal cymes over 5 mm,
slender. Septum of endocarp swollen to at least '/; of the diameter of the fruit. Branchlets subterete.
Domatia occasional, suborbicular. SUBG. MANGLESIA ........... . . 1. M.octandra

1. Stamens 4-5(-6), in one whorl. Inflorescence branches terete. Calyx distinctly lobed. Bracts subper-
sistent. Septum of endocarp wedge-shaped. SuBG. MASTIXIA.

2. Inflorescence branches of the first order (sub)opposite or decussate. Branchlets and leaves generally
(sub)opposite or decussate; nodes flattened. Fruits generally ovoid. Ser. Oppositae.
3. Flowers basically 5-merous.
4. Sepals less than half as long as wide. Inflorescence subglabrous to puberulous. Fruit ovoid to
oblong, with inconspicuous persistent sepals ............... 2. M. kaniensis
4. Sepals almost as long as wide. Inflorescence velutinous to woolly. Fruit elongate-ovoid, with con-
spicuous persistent sepals . ............. . . 3a. M. trichotoma var. korthalsiana

6. Leaves (sub)opposite or alternate; petioles slender. Leaves chartaceous to subcoriaceous; nerva-

tion rather weak, without intermediary nerves. Inflorescence slender with bracts all under 3 mm.

Byatt Wipe PO UREN A, ws ke ar nse ss we hs soa Bie ite Cee A nt oie

2. Inflorescence branches of the first order scattered. Branchlets and leaves scattered; nodes terete.
Fruit generally ellipsoid or oblong. Ser. Alternae. :

7. Branchlets woolly. Leaves 13-30 by 5!/,-15 cm, with midrib and nerves (even veinlets) woolly to

villous; petioles stout, 4 cm or longer, woolly. Fruit over 4 by 2 cm. Flowers 5-merous
6. M. macrocarpa

1977]

CORNACEAE (Matthew)

89

7. Branchlets not woolly. Petioles up to 4 cm. Fruit up to 4cm long.

8. Flowers basically 4-merous.

9. Leaves glaucous and waxy below, thick-coriaceous, with intermediary nerves; apex apiculate.

Sepals as long as wide

M. glauca

9. Leaves not glaucous and waxy below, without intermediary nerves ; apex ‘other than apiculate.

Calyx teeth at most half as long as wide.

10. Leaves crowded at apices of branchlets, thick-coriaceous; acute to acuminate. Inflorescence

branches stout, compact. Fruit ellipsoid, 1!/, cm @.

. 8. M. tetrapetala

10. Leaves evenly spread, chartaceous to subcoriaceous; apex ‘caudate (over 1 cm). Inflorescence

branches rather slender
8. Flowers basically 5-merous.

5. M. rostrata

11. Leaves abruptly cuspidate (over 1 cm); nerves arcuate, clearly impressed above. Inflorescence
raceme-like, seldom branched more than twice. Petals densely silky outside. Branchlets slender.

Fruit oblong, 1*/,-2 by 0.8-l1cm .

9. M. cuspidata

11. Leaves other than abruptly cuspidate; nerves not arcuate but mostly sharply prominent, veins
mostly distinct. Inflorescence usually branched twice or more, not terminating into a dichasium.
Petals glabrous to appressed hairy. Fruit ovoid to oblong, 2!/4,-3'/, by 1-11/, cm

10. M. pentandra

1. Subgenus Manglesia

MATTHEW, Blumea 23 (1976) 64, f. 1 (map) & 2.
Branchlets and leaves decussate. Stamens 8, in 2 whorls. Inflorescence branches
4-angular. Calyx subtruncate. Fruit with swollen septum.

Distr. 2 spp., in NE. India, N. Burma, NW. Thailand, Central Sumatra.

. Mastixia octandra MATTHEW, Blumea 23
(i976) 65, f. 3 (map).

Tree up to 25 m; d.b.h. up to 90 cm. Branchlets
slender, decussate, terete, glabrous. Leaves decus-
sate, ovate to elliptic, 4-8 by 11/,-3 cm, chartaceous,
glabrous; base cuneate; apex acuminate; nerves
6-8 pairs, with intermediary ones; veins distinct on
both surfaces; an occasional subcircular doma-
tium at the axil of nerves; petiole 1-1'/, cm,
slender. Inflorescence up to 15cm, slender, glab-
rous, branched up to 5 times; branches of the first
order decussate; pedicels of lateral flowers of
terminal cymes over 5 mm, slender. Bracts ovate,

under 3mm, glabrous. Submature flower bud
3mm @. Calyx subtruncate, thin; teeth 4, minute,
acute, thin. Petals 4, thick, glabrous outside.
Stamens 8. Ovary glabrous. Fruit turbinate, 1 cm.

Distr. Malesia: West Central Sumatra, once
found.

Ecol. Mountain forest, 1700-1800 m.

Notes. Easily distinguished from the continen-
tal Asian M. euonymoides PRAIN by smaller,
chartaceous leaves, suborbicular domatia, more
slender, lax and elongate inflorescence parts, the
thin calyx with acute teeth, and the generally
pedicelled middle flower of the cymes.

2. Subgenus Mastixia

Cf. MATTHEW, Blumea 23 (1976) 66, f. 1 (map) & 2. — Mastixia subg. Tetra-

mastixia et Pentamastixia WANGERIN, Pfl. Reich Heft 414 (1910) 21, 25.
Branchlets and leaves scattered or (sub)opposite. Stamens 4-5(-6), in one whorl.

Inflorescence branches terete. Calyx lobed. Septum of the fruit wedge-shaped.

Distr. 11 spp., covering the entire range of the genus.

1. Series Oppositae

MATTHEW, Blumea 23 (1976) 66.

Inflorescence branches of the first order (sub)-
opposite or decussate. Branchlets and leaves ditto;
nodes flattened. Fruit usually ovoid.

Distr. Throughout Malesia, in continental Asia
only in Pensinsular Thailand.

2. Mastixia kaniensis Metcn. Bot. Jahrb. 60
(1925) 172; Danser, Blumea 1 (1934) 52. — M.

ledermannii Mevcn. Bot. Jahrb. 60 (1925) 173. —
M. pentandra (non Bu.) DANsER, Blumea 1 (1934)
50, p.p.; MATTHEW, Blumea 23 (1976) 67. — Fig.
yl 3a-b (galls).

Tree up to 31m; d.b.h. up to 75(-90) cm.
Branchlets stout or slender, (sub)opposite, sub-
glabrous to velutinous. Leaves (sub)opposite,
elliptic, obovate, oblong or oblanceolate, (3'/,-)
4'/,-18 by 2-8 cm, chartaceous to thick coriaceous,
subglabrous, rarely densely velutinous; base
attenuate to truncate; apex acuminate to caudate;

90 FLORA MALESIANA [ser. I, vol. 8?

a

Fig. 3. Galls of Mastixia. a. M. kaniensis MELCH. spp. ledermannii (MELCH.) MATTHEW, b. M. kaniensis

MELCcH. ssp. kaniensis, c. M. pentandra BL. ssp. chinensis (MERR.) MATTHEW, d-e. M. pentandra BL. ssp.

Dhilippinensis (WANGERIN) MATTHEW, f. M. rostrata BL. ssp. caudatifolia (MERR.) MATTHEW, g. M. tricho-

toma BL. var. korthalsiana (WANGERIN) DANSER, h. M. trichotoma BL. var. rhynchocarpa DaANsER. All nat.

size, except g < 2 (a CLEMENS 5361, b SCHLECHTER 17703, c LACE 5641, d WENZEL 1150, e FB 2201,
Ff KOsTERMANS 12573, g KOSTERMANS 7316, h KOSTERMANS 7620).

1977]

nerves 3—9(-11) pairs, sometimes arcuate; veins
usually obscure; petiole 1-2(-4)cm, stout or
slender. Inflorescence up to 8 cm, stout or slender,
subglabrous to puberulous, branched up to 3-(4)
times, at times terminating in a dichasium, branches
of the first order (sub)opposite. Bracts triangular
to lanceolate, up to 4mm, puberulous to veluti-
nous. Submature flower bud 1'/,-2'/,mm @.
Sepals (4-)5(-6-7), broader than long. Petals
(4-)5(-6), thick or thin, glabrous or puberulous
outside. Stamens (4—-)5(-6). Ovary sparsely puberu-
lous. Fruit ovoid to oblong, 1'/,-2!/, by 1-11/, cm,
dull or shining when dry; persistent disk incon-
spicuous to prominent; sepals inconspicuous.

Distr. Malesia: Moluccas, New Guinea, New
Britain, and the Solomon Islands. Fig. 4.

Note. Two replacing subspecies are distin-
guished. The maximum degree of fluctuation in the
number of flower parts occurs in the New Guinea
—Solomons area.

KEY TO THE SUBSPECIES

1. Leaves chartaceous to subcoriaceous; branch-
lets and inflorescence axes slender; petals thin,
glabrous outside. ; a. ssp. kaniensis

1. Leaves thin- to thick-coriaceous; branchlets and
inflorescence axes usually stout; petals thick,
puberulous outside. b. ssp. ledermannii

a. ssp. kaniensis. — Fig. 1, 3b (galls).

Branchlets slender, subglabrous to velutinous.
leaves (31/,- )4"/,-141/, by 2-6 cm, chartaceous to
subcoriaceous; base attenuate to cuneate; nerves
3-7 pairs, sometimes arcuate, prominulous below;
petiole 1-2 cm. Inflorescence up to 6 cm, slender,
branched 2(-3) times, lax. Flowers relatively small.
Bracts triangular to lanceolate, under 3 mm. Sub-
mature bud 1'/,mm @. Petals (4-)5, relatively

+ kaniensis Melchior ssp. kaniensis
sep. ledermannii (Melch.) Matthew
+ @ugenioides Matthew
+ rostrata Blume sap. rostrata
sep. caudatifolia (Merr.) Matthew

CORNACEAE (Matthew) 91

thin, glabrous outside. Stamens (4-)5. Fruit ovoid
to oblong, 2-2!/, by 12/,-1"/, cm.

Distr. Malesia: East New Guinea, New Britain,
and Solomon Islands. Fig. 4.

Ecol. Common in primary forests from low
altitude up to 1200 m. FI. fr. Jan._Dec. Leaf- and
fruit-galls occur.

Note. Occasional specimens in New Guinea are
densely velutinous, others are less so. The basal
branches of the inflorescence of the first order are
at times subtended by foliage leaves.

b. spp. ledermannii (MELCH.) MATTHEW, Blumea 23
(1976) 67. — M. ledermannii MELCH. — Fig. 3a
(galls).

Branchlets stout, often rusty puberulous when
young, subglabrous later. Leaves 6-18 by 2!/,-8
cm, thin- to thick-coriaceous; base attenuate to
truncate; nerves 3—9(-11) pairs, sometimes arcuate,
obscure to prominent below; petiole 11/,-2(—4) cm.
Inflorescence up to 8cm, stout, branched 3-(4)
times, compact. Flowers relatively large. Bracts
lanceolate below, up to 4mm. Submature flower
bud 2!/, mm @. Petals (4-)5(-6), thick, puberulous
outside. Stamens (4-)5(-6). Fruit ovoid, 1%/,-
21/2 by 1-11/, cm.
ae Malesia: Moluccas and New Guinea.

ig. 4.

Ecol. Primary and secondary forests, 100-1800
m. FI. fr. Jan.—Dec. Leaf-galls occur.

Vern. New Guinea: bie, bon, Muju, labak-
— Mooi, masjiw, Wandammen, samuwin,
Biak.

Notes. In some specimens a dense indumentum
is found.

Though the two subspecies are clearly replacing,
some specimens of ssp. ledermanii occur in the area
occupied by ssp. kaniensis, but at higher altitude
than ssp. kaniensis in this area.

Fig. 4. Localities of three species and two subspecies of Mastixia.

92 FLORA MALESIANA

(ser. I, vol. 8?

3. Mastixia trichotoma BL. Bijdr. (1826) 655; DC.
Prod. 4 (1830) 275; BL. Mus. Bot. 1 (1850) 257,
f. 58; Miq. Fl. Ind. Bat. 1, 1 (1856) 772, incl. var.
laxa Mia. l.c.; K. & V. Bijdr. 5 (1900) 90; WaAn-
GERIN, Pfl. Reich Heft 414 (1910) 24, f. 1A-E;
Koorp. Atlas 1 (1913) t. 190; Fl. Tjib. 2 (1923);
DANSER, Blumea 1 (1934) 57, incl. var. tenuis,
acuminatissima, clarkeana, korthalsiana, maingayi,
benculuana, rhynchocarpa et simalurana DANSER,
lc. 61-65; BACK. & BAKH. f. FI. Java 2 (1965) 159;
Meer, Bot. News Bull. Sandakan 8 (1967) 65;
ibid. 10 (1968) 179, illus.; MATTHEW, Blumea 23
(1976) 68. — M. laxa Bl. Mus. Bot. 1 (1850) 257,
incl. var. angustifolia BL. l.c.; WANGERIN, Pfl.
Reich Heft 414 (1910) 24. — M. acuminatissima
BL. Mus. Bot. 1 (1850) 258; Mra. Fl. Ind. Bat. 1, 1
(1856) 772, (1858) 1095; WANGERIN, Pfl. Reich
Heft 414 (1910) 22, f. 1F. — M. caesia BL. Mus.
Bot. 1 (1850) 258. — M. kimanilla BL. l.c. 258;
Mia. Fl. Ind. Bat. 1, 1 (1856) 772, (1858) 1095,
incl. var. caesia Mia. l.c. 772; K. & V. Bijdr. 5
(1910) 94; WANGERIN, Pfi. Reich Heft 414 (1910)
25. — M. maingayi CLARKE, FI. Br. Ind. 2 (1879)
746; Kina, J. As. Soc. Beng. 71, ii (1902) 74, incl.
var. subtomentosa KING, l.c. 75; WANGERIN,
Pfil. Reich Heft 414 (1910) 22. — M. junghuhniana
(non Mia.) CLARKE, FI. Br. Ind. 2 (1879) 746. — M.
clarkeana KinG, J. As. Soc. Beng. 71, ii (1902) 75 &
var. macrophylla K1NG, |.c.; WANGERIN, Pfl. Reich
Heft 414 (1910) 24; HALL. f. Beih. Bot. Centralbl.
34, 2 (1916) 40. — M. korthalsiana WANGERIN in
Fedde, Rep. 4 (1907) 335, incl. var. macrophylla
WANGERIN, /.c. 336 et Pfl. Reich Heft 414 (1910)
25, 26; HALL. f. Beih. Bot. Centralbl. 34, 2 (1916)
40. — M. propinqua RIpDL. J. Fed. Mal. St. Mus. 4,
1 (1909) 25; Fl. Mal. Pen. 1 (1922) 890. — Vitex
premnoides ELMER, Leafl. Philip. Bot. 8 (1915) 2874.
— M. premnoides HA .v. f. Beih. Bot. Centralbl.
34, 2 (1916) 41; Merr. Philip. J. Sc. 13 (1918) 43;
En. Philip. 3 (1923) 242. — M. rostrata (non BL.)
RIDL. FI. Mal. Pen. 1 (1922) 890. — Fig. 3g-h (galls).

Tree up to 40m; d.b.h. up to 50(—-150) cm;
branchlets stout or slender, opposite, puberulous
to woolly. Leaves opposite, ovate, elliptic to
oblong, 5—24(-28) by 2-12cm, thin- to thick-
coriaceous, subglabrous to villous below; base
cuneate, obtuse or attenuate; apex acute to acu-
minate; nerves 5-15 pairs, impressed above,
prominulous to prominent below, at times arcuate;
veins prominulous to prominent below; petiole
11/,—21/,(—31/,) cm, stout or slender. Inflorescence
up to 15cm, stout or slender, compact or lax,
puberulous to woolly; branched up to 5 (or 6)
times; branches of the first order opposite; higher
order bracts triangular, more or less connate,
villous to woolly; lower bracts lanceolate, up to
10 mm, villous to woolly. Submature flower bud
2-31/, mm @. Sepals 4 or 5, as long as wide, thick,
puberulous to villous. Petals 4 or 5, thick, puberu-
lous to villous outside. Stamens 4 or 5. Ovary
puberulous to villous. Fruit ovoid to elongate,
acute, 1!/,—31/, by 1/,-2 cm; persistent disk incon-
spicuous to prominent; sepals prominent.

Distr. Peninsular Thailand and throughout
Malesia, except New Guinea. Fig. 5.

Notes. DANSER (/.c. 59-61) adequately dis-
cussed variations within the species; most of the
vernacular names he listed (/.c. 72-73) belong to the
present species.

Fig. 5. Localities of Mastixia trichotoma BL. and
its varieties.

Figure 1 of WANGERIN (/.c. 23) is rather sche-
matic. The position of ovule attachment and the
length of the sepals are inaccurately drawn.

KEY TO THE VARIETIES

1. Inflorescence villous to woolly.

2. Twigs woolly. Leaves 9-20 cm long, thick-
coriaceous, villous to woolly; nerves often
arcuate; base obtuse to truncate. Fruit ovoid,
more than 1!/,cm @ . . . .¢. var. maingayi

2. Twigs not woolly. Leaves 5-15cm long,
chartaceous to subcoriaceous, subglabrous to
puberulous; nerves seldom arcuate; base
attenuate to cuneate. Fruit elongate-ovoid, up
to 1!/,cm @.

3. Inflorescence compact. Sepals, petals, and
Stamens.4:, Ses “eae a. var. trichotoma
3. Inflorescence very lax. Sepals, petals, and
Stamens! 52> b. var. korthalsiana

1. Inflorescence subglabrous to puberulous.

4. Fruit with prominent persistent disk, over 21/,
by 11/, cm. Leaves 10-24 by 5-12 cm. Inflores-
cence robust, up to 15cm, branched 5 (to 6)
times, not terminating in a dichasium.

d. var. rhynchocarpa

4. Fruit without prominent persistent disk, up to
2 by 1 cm. Leaves usually 5-12 by 2-4'/, cm.
Inflorescence slender, up to 10 cm, branched
3(-4) times, often terminating in a dichasium

e. var. cClarkeana

a. var. trichotoma — M. laxa BL. — M. trichotoma
BL. var. laxa Mig. — M. laxa BL. var. angustifolia
BL. — M. acuminatissima BL. — M. trichotoma
BL. var. acuminatissima DANSER — M. caesia BL.
—M. kimanilla Bt. incl. var. caesia Miq.
Branchlets rather slender, yellowish, puberulous
to velutinous. Leaves elliptic to oblong, 5-15 by
3-8 cm, chartaceous to subcoriaceous, sub-
glabrous to puberulous; base cuneate to attenuate;
apex acute to acuminate; nerves 5-8 pairs, promi-
nulous to prominent below, seldom arcuate;
petiole 11/,—2(—2!/,) cm, rather slender. Inflores-
cence up to 8 cm, compact, branched 3(-4) times,
villous to woolly; basal bracts under 5mm;
terminal bracts often deeply boat-shaped. Sepals 4.
Petals 4. Stamens 4. Fruit elongate-ovoid, 11/.-3 by

1977]
1-11/, cm; persistent disk inconspicuous; sepals
prominent.

Distr. Malesia: N. Sumatra, W.-E. Java,

Borneo, Lesser Sunda Is. (Bali). Fig. 5.

Ecol. In primary forest from low altitude up to
1800 m. Fi. May—Jan., fr. July—March. Galls occur
on stem and fruit.

Vern. Java: djérét, huru hiris, h. minjak, kéndu,
kibéntéli, kibunting, kidédak, kilumlum, kilun glum,
kiménjan, kiténjo, (huru) méhmal, mémah, palaglar
minjak, ténggau, ténju.

b. var. korthalsiana (WANGERIN) DANSER, Blumea |
(1934) 63; MATTHEW, Blumea 23 (1976) 70. —
M. korthalsiana WANGERIN. — Fig. 3g (galls).

Branchlets rather slender, yellowish, subglab-
rous to velutinous. Leaves subovate-elliptic, 10-12
by 3-5 cm, subcoriaceous; base long attenuate;
apex acuminate; nerves 5-6 pairs, seldom arcuate;
petiole 1/,—-13/,cm, rather slender. Inflorescence
up to 10cm, very lax, branched 3(-4) times, few-
flowered, velutinous to woolly; bracts under 3 mm.
Sepals 5. Petals 5. Stamens 5. Fruit elongate-ovoid,
2'/,-3 by 1-1'/, cm, persistent disk inconspicuous,
sepals prominent.

Distr. Malesia: Sumatra, Borneo. Fig. 5.

Ecol. In primary forest from low altitude up to
640 m, often scattered. Rather prone to galls.

Vern. Sumatra: médang kladi. E. Borneo:
sérgam pipit, Sangkulirang I.

Note. The lax and few flowered inflorescence,
the 5-merous flowers, and elongate fruit distinguish
this variety from var. trichotoma to which it is
closely allied.

ce. var. maingayi (CLARKE) DANSER, Blumea 1
(1934) 63; MATTHEW, Blumea 23 (1976) 70. — M.
maingayi CLARKE, incl. var. subtomentosa KING —
M. junghuhniana (non MiQ.) CLARKE — M. rostrata
(non BL.) Rip_. — M. propinqua RIDL.

Branchlets very stout, yellowish, woolly. Leaves
ovate to elliptic, 9-20 by 4-11 cm, thick coriaceous,
tough and stiff; base obtuse to truncate; apex acute
to acuminate; nerves 5-6 pairs, deeply impressed
above, very prominent below, often arcuate,
numerous parallel intermediary veins conspicuous;
petiole 1'/,-2'/,cm, stout, villous to woolly.
Inflorescence up to 15cm, compact, branched
4(-5) times, velutinous to golden woolly; basal
bracts up to 1 cm, persistent. Sepals 4. Petals 4.
Stamens 4. Fruit ovoid, 2'/,-3'/, by 11/,-2.cm;
persistent disk inconspicuous, sepals prominent.

Distr.Malesia: Sumatra, Malay Peninsula (also
Penang), Banka, Borneo. Fig. 5.

Ecol. In primary forests with Dipterocarps; also
in secondary or marsh forests, from the lowland to
1400 m. Fl. Febr.-Au . (Nov. ), Sr. April—Dec. Galls
occur on leaves and fruit.

Vern. Malaya: karu nuri, kayu béngkal bukit,
kayu maura, médang. Banka: médang puntung,
m. pusér. Borneo: médang kanigara.

Note. Easily recognized by the woolly indumen-
tum, large and stiff leaves with prominent veins,
and the numerous massive fruits.

d. var. rhynchocarpa DANseR, Blumea | (1934) 64;
MATTHEW, Blumea 23 (1976) 71. — M. trichotoma
. var. benculuana et var. simalurana DANSER. —

Fig. 3h (galls.

CORNACEAE (Matthew) 93

Branchlets stout, dark, subglabrous. Leaves
elliptic to oblong, 10-24(-28) by 5-12cm,
coriaceous, tough and stiff; base cuneate to obtuse;
apex acute, acuminate to caudate; nerves 8-15
pairs, very prominent below, seldom arcuate;
petiole 11/,-2'/,(—3'/,) cm, stout. Inflorescence up
to 15cm, compact, profusely branched up to
5(-8) times, not terminating in a dichasium, sub-
glabrous to puberulous; basal bracts up to 5 mm.
Sepals 4(—5). Petals 4. Stamens 4. Fruit ovoid to
elongate-ovoid, 2!/,-3'/, by 11/,-2 cm; persistent
disk prominent, bulging; sepals prominent.

Distr. Malesia: Sumatra (incl. Simalur I.), W
Java, Borneo, NE. Celebes, Moluccas (Ambon,
Ceram). Fig. 5.

Ecol. Common in primary lowland and mossy
forest, up to 1800 m. Fi. April—Aug., fr. June-
March.

Galls. This variety is very prone to fruit galls
and the largest ones in the genus (over 1!/, cm @)
occur here.

Vern. Sumatra: ahélat, awa ahélat uding, awa
énti, awa simangurach, tutun simangurah, Simalur
I.; bung, médang tima, tanah, Bencoolen. Borneo:
médang aima. Moluccas: soya.

Note. This variety is noted for the generally
large dimensions of leaves and inflorescence,
though there is a reduction in size from Borneo to
Moluccas.

e. var. Clarkeana (KING) DANSER, Blumea 1 (1934)

62; MATTHEW, Blumea 23 (1976) 72. — M.
clarkeana KING, incl. var. macrophylla Kinc —
M. korthalsiana WANGERIN var. macrophylla

WANGERIN — Vitex eg lab od ELMER — M. prem-
noides HALL. f. — M. trichotoma BL. var. tenuis
DANSER.

Branchlets slender, grey, subglabrous. Leaves
oblong to elliptic-oblong, 5—12(-18) by 2—4(-8'/,)
cm, coriaceous; base cuneate to obtuse; apex acute
to acuminate; nerves 5-7 pairs, seldom arcuate;
petiole 1-1'/,cm, slender. Inflorescence up to
10cm, compact, branched 3(—4) times, often ter-
minating in a dichasium. Sepals 4. Petals 4.
Stamens 4. Fruit ovoid, 1*/,-2 by */.-1 cm; per-
sistent disk inconspicuous; sepals prominent.

Distr. Peninsular Thailand (Pattani) and
Malesia: Sumatra, Banka, Malay Peninsula,
Borneo, Philippines (Mindanao). Fig. 5.

Ecol. Primary forest, from low altitude to
1100m. Fl. Jan.—Aug., fr. July—Febr. Galls occur on
stem and fruit.

Vern. Philippines: /amog.

4. Mastixia eugenioides MATTHEW, Blumea 23
(1976) 73.

Tree up to 30 m; d.b.h. up to 30 cm; branchlets
stout, opposite, glabrous. Leaves opposite, elliptic
to oblong-elliptic, 4-12 by 2-5'/,cm, thick-
coriaceous, glabrous; base cuneate; apex acumi-
nate to caudate; nerves 5-7 pairs, prominent
beneath, with intermediary ones and distinct veins;
petiole 1'/,—2'/, cm, stout. /nflorescence up to 8 cm,
rather stout and compact, glabrous, up to 4 times
branched, at times terminating in a dichasium;
branches ‘of the first order opposite; higher order
bracts triangular, under 3mm; lower bracts
lanceolate, up to 5mm, all glabrous. Submature
flower bud 2mm @. Sepals 4, broader than long,

94

FLORA MALESIANA

[ser. I, vol. 8?

thick, glabrous. Petals 4, thin, glabrous outside.
Stamens 4. Ovary glabrous. Fruit (unripe) ovoid,
2'/, by 1'/,cm; persistent disk and sepals incon-
spicuous.
Distr.
Fig. 4.
Ecol. Primary (often Dipterocarp) lowland
forests, up to 400 m. Fl. July—Aug., fr. Sept.
Notes. Leaf scars conspicuous; inflorescence
notably erecto-patent when young, spreading later.
The inflorescence and flowers somewhat resemble
those of M. rostrata ssp. rostrata, but the stout
branchlets with strictly opposite leaves and stout
petiole, prominent intermediary veins, and fruits of
different shape with thick pericarp, make this
species quite distinct.
All the 9 collections are from a restricted area.

Malesia: Borneo (Sarawak, Brunei).

5. Mastixia rostrata BL. Mus. Bot. 1 (1850) 258;
Mia. Fl. Ind. Bat. 1, 1 (1856) 773, (1858) 1095;
K. & V. Bijdr. 5 (1900) 92; WANGERIN, Pfi. Reich
Heft 414 (1910) 22; Koorp. Atlas 1 (1913) t. 191;
DANsER, Blumea 1 (1934) 52; Burk. Dict. (1935)
1428; Doct.v.LEEUWEN, Ned. Kruidk. Arch. 51
(1941) 207; Back. & BAKH. f. Fl. Java 2 (1965)
159; MevER, Bot. News Bull. Sandakan 8 (1976)
65; MATTHEW, Blumea 23 (1976) 73. — M. jung-
huhniana Mia. FI. Ind. Bat. 1, 1 (1856) 772. —
M. margarethae WANGERIN in Fedde, Rep. 4 (1907)
335; Pfil. Reich Heft 41* (1910) 21. — M. cuspidata
BL. var. margarethae HALL. f. Beih. Bot. Centralbl.
34, 2 (1916) 41. — M. caudatifolia MerR. Pl. Elm.
Born. (1929) 233.

Tree up to 30 m; d.b.h. up to 50 cm; branchlets
slender, (sub)opposite or scattered, glabrous.
Leaves (sub)opposite or scattered, elliptic to oblong-
elliptic, 4-10 by 2-5cm, chartaceous to sub-
coriaceous, glabrous; base cuneate; apex caudate
over 1 cm; nerves 4-6 pairs, prominulous beneath;
veins obscure; petiole 1—2'/, cm, slender. Inflores-
cence up to 6cm, slender, compact or lax, sub-
glabrous, up to 4 times branched, branches of the
first order (sub)opposite or scattered; bracts
triangular, under 3mm, glabrous. Submature
flower bud 1-2!/,mm @. Sepals 4, broader than
long, thin, glabrous. Petals 4, glabrous outside.
Stamens 4. Ovary glabrous. Fruit ovoid to oblong,
1+/,-3 by 3/,-1 cm, persistent disk prominent or
inconspicuous, sepals inconspicuous.

Distr. Malesia: Sumatra, Banka, Java, Borneo,
Lesser Sunda Is. (Sumbawa, Flores). Fig. 4.

KEY TO THE SUBSPECIES

1. Submature flower bud 2!/, mm @. Inflorescence
compact, branches of the first crder (sub)-
opposite. Leaves (sub)opposite. Galls absent

a. ssp. rostrata

1. Submature flower bud 1 mm @. Inflorescence
lax, branches of the first order scattered. Leaves
scattered. Galls frequent. . b. ssp. caudatifolia

a. ssp. rostrata — M. junghuhniana Mia.

Branchlets (sub)opposite. Leaves (sub)opposite,
less often scattered ; petiole 11/,-2 cm. Inflorescence
compact, generally under 4 cm, glabrous; primary
branches (sub)opposite. Submature flower bud
2//,mm @.

Distr. Malesia: West & Central Java, Lesser
Sunda Is. (Sumbawa, Flores). Fig. 4.

Ecol. Moist forest, from low altitude up to
1400 m. Fl. March, very fragrant, fr. May-July.
Galls absent. Ripe fruit dark blue.

Vern. Java: daun kaju téndjo, daun kitadjas,
huru gading, kilburoy, kiléjas, kiténdjo, kitindjo,
lalakina, tjangkar. Flores: bumis, rau, tapadeke.

Notes. Specimens from the Lesser Sunda Is.
have larger leaves than those from Java.

The reference in KANJILAL & Das (Fl. Assam 2,
1938, 371) to this species seems erroneous, as it
does not occur on the Asian continent. If the state-
ment “‘stamens 3”’ is correct, the plant can even not
belong to Mastixia.

b. ssp. caudatifolia (MERR.) MATTHEW, Blumea 23
(1976) 74. — M. margarethae WANGERIN — M.
caudatifolia MERR. — Fig. 3f (galls).

Branchlets scattered. Leaves scattered; petiole
up to 1'/, cm. Inflorescence very lax, up to 6 cm,
puberulous at the nodes; primary branches
scattered. Submature flower bud 1 mm @.

Distr. Malesia: northern half of Sumatra,
Banka, Borneo. Fig. 4.

Ecol. Primary forest, from the lowland up to
1600 m. F/. June—Oct., fr. Aug.—March. Globose to
elongate galls are common, specially those on
fruits. Sometimes they resemble a legume and can
be up to 31/, cm long.

Vern. Borneo: patoli entelit, Iban lang.

Note. It is almost impossible to separate sterile
materials of ssp. caudatifolia from those of M. cus-
pidata, though their flowers are entirely different.

2. Series Alternae

MATTHEW, Blumea 23 (1976) 75.

Inflorescence branches of the first order scat-
tered; branchlets and leaves scattered; nodes
terete; fruit generally ellipsoid or oblong.

Distr. Ceylon and continental Asia; through
Seti but absent in New Guinea and Solomon
S.

6. Mastixia macrocarpa MATTHEW, Blumea 23
(1976) 75.

Tree up to 21 m; d.b.h. up to 20 cm; branchlets
stout, scattered, woolly. Leaves scattered, elliptic-
oblong to oblong, 13-30 by 51/,-15 cm, subcoria-
ceous, villous, especially below; base cuneate, at
times slightly oblique; apex acuminate; nerves
7-10 pairs, prominent below, villous; veins promi-
nulous, puberulous to villous; petiole 4-7 cm,
stout, woolly. Inflorescence up to 9cm, stout,
woolly, branched up to 4 times; branches of the
first order scattered ; bracts triangular to lanceolate,
up to 8 mm, densely woolly. Submature flower bud
4mm @. Sepals 5, broader than long, villous.
Petals 5, velvety outside. Stamens 5. Ovary densely
villous. Fruit oblong-ovoid, 4-4'/, by 2 cm; per-
sistent disk inconspicuous, sepals prominent, up to
5 mm.

Distr. Malesia: Borneo (Sarawak), Philippines
(Luzon); 2 collections. Fig. 6.

Ecol. Lowland forest. Fi. Oct., fr. June. Sticky
resin on the branches; fruits very pale green.

Note. Leaves and fruits are the largest known in
the genus; lenticels up to 3 by 1 mm; leaf scars up

1977]

CORNACEAE (Matthew)

2

to 4 by 4mm; peduncles of terminal cymes up to
6mm; pedicels 2mm; sepals 2 mm broad at the
base; filaments 3 mm; anthers 1'/, mm; receptacle
3-31/, by 2mm; style 4mm; stigma 5-lobed,
appearing bifid.

®@ M. mecrocarpa Matthew
O m. glauca matthew
@ M. cuspidate Slume

Fig. 6. Localities of three species of Mastixia.

7. Mastixia glauca MATTHEW, Blumea 23 (1976)
76

Tree up to 15m; branchlets stout, scattered,
glabrous. Leaves scattered, obovate, 7-16 by
4'/,-8'/, cm, thick-coriaceous, glaucous and waxy
below, glabrous; base obtuse; apex apiculate;
nerves 4-5 pairs, with intermediary ones, all
obscure; veins obscure; petiole 2-3'!/, cm, stout,
glabrous. Inflorescence up to 5 cm, rather stout and
compact, subglabrous to sparsely puberulous,
branched up to 5 times; branches of the first order
scattered; bracts triangular, under 3mm. Sub-
mature flower bud 3 mm @. Sepals 4(-5), as long
as broad, sparsely puberulous. Petals 4(—5), thick,
appressed-hairy outside. Stamens 4(-5). Ovary
sparsely puberulous.

Distr. Malesia: Borneo (Sarawak: Mt Santu-
bong), 3 collections Fig. 6.

Ecol. Lowland forest. F/. April-May.

Note. Tender bark of branchlets yellowish;
inflorescence clearly broader than long; 5-merous
flowers only occasional.

8. Mastixia tetrapetala Merr. Philip. J. Sc. 13
(1918) Bot. 42; En. Philip. 3 (1923) 242; MATTHEW,
Blumea 23 (1976) 76, f. 5 (map). — M. pachyphylla
Mere. Philip. J. Sc. 13 (1918) Bot. 325; En. Philip.
3 (1923) 241. — M. crassifolia MeRR. Philip. J. Sc.
26 (1925) 486.

Tree up to 8(-15)m; branchlets very stout,
scattered to subopposite, subglabrous. Leaves
scattered to subopposite, crowded at apices of
branchlets, obovate to oblanceolate, 5-15 by
2-7 cm, thick coriaceous, glabrous; base cuneate;
apex acute to acuminate; nerves 6—8(-12) pairs,
usually prominent below; veins prominulous
below; petiole 1-2'/,cm, stout, glabrous. IJn-

escence up to 3-(6) cm, stout, very compact,

berulous to villous, 2(—3) times branched;
lee of the first order scattered; higher order
bracts triangular, under 3 mm; lower ones lanceo-
late up to 5 mm, puberulous. Submature flower

bud 5 mm @. Sepals 4, broader than long, glabrous
to appressed-hairy. Petals 4, thick, glabrous to
appressed-hairy. Stamens 4. Ovary glabrous to
appressed-hairy. Fruit ellipsoid, 2-3 by 1%/,—
11/, cm; persistent disk and sepals inconspicuous.

Distr. Malesia: Philippines (Luzon, Catan-
duanes), 7 collections.

Ecol. Primary forest, from low altitude up to
2300 m. Fi. Febr.—March, fr. Sept.—Febr.

Notes. Branchlets stout with conspicuous leaf
scars and fibrous bark. Phyllotaxis tends to be
obscured owing to congestion of parts: leaves are
generally crowded towards the apices of branch-
lets. Inflorescence branches do not always elongate
as in other species.

The species is quite distinct and stands rather
isolated from others in the stoutness of parts, the
large, 4-merous flowers, and the large, ellipsoid
fruits.

Two other species of MERRILL, M. pachyphylla
and M. crassifolia are considered conspecific with
M. tetrapetala. There are indeed certain differences:
leaves of M. crassifolia generally have 8-12 nerves
per side prominent below, and massive fruits.
M. pachyphylla has (sub)opposite leaves and pri-
mary inflorescence branches. However, when
examined together, it is seen that both M. pachy-
phylla and M. crassifolia are extreme variations of
M. tetrapetala.

9. Mastixia cuspidata BL. Mus. Bot. 1 (1850) 256;
Mia. FI. Ind. Bat. 1, 1 (1856) 772; HALL. f. Beih.
Bot. Centralbl. 34, 2 (1916) 41; DANseR, Blumea 1
(1934) 55, excl. var. margarethae; MATTHEW,
Blumea 23 (1976) 79. — M. pentandra BL. var.
cuspidata Miq. Fl. Ind. Bat. 1, 1 (1858) 1095;
WANGERIN, Pfi. Reich Heft 414 (1910) 26. — M.
bracteata CLARKE, FI. Br. Ind. 2 (1879) 746; KING,
J. As. Soc. Beng. 71, ii (1902) 73; WANGERIN,
Pfl. Reich Heft 41* (1910) 26, f. 1 G-K, N-O;
DANSER, Blumea 1 (1934) 54.

Tree up to 24 m; d.b.h. up to 40 cm; branchlets
very slender, scattered, subglabrous. Leaves
scattered, obovate, elliptic or oblong, 4-12(—16) by
2-4(—6) cm, subcoriaceous, glabrous; base cuneate;
apex abruptly cuspidate (over 1 cm), oblique;
nerves 4 (or 5) pairs, arcuate, imp above;
veins obscure; petiole 1—1'/, cm, slender. Inflores-
cence up to 4cm, rather slender, subglabrous to
puberulous, 2(—3) times branched; branches of the
first order scattered; higher order bracts subulate,
lower ones foliaceous, over 10 mm, passing into
foliage leaves. Submature flower bud 3 mm @.
Sepals 5, broader than long, subglabrous. Petals
5, thick, densely appressed-hairy outside. Stamens
5. Ovary densely appressed silky-hairy. Fruit
oblong, 1'/,-2 by °/,-l1.cm; persistent disk and
sepals inconspicuous.

Distr. Malesia: Sumatra, Banka, Malay
Pensinula, Borneo. Fig. 6

Ecol. Primary and secondary forests, from low
altitude up to 900 m.

Vern. Sumatra: bébung, kundur. Banka: mén-
kapas. Malay Peninsula: dadaru. Borneo: biansu-
gunong, Sarawak.

10. Mastixia BL. Bijdr. (1826) 654; DC.
Prod. 4 (1830) 275; BL. Mus. Bot. 1 (1850) 256;
Mig. Fl. Ind. Bat. 1, 1 (1856) 771, (1858) 1095;

96 FLORA MALESIANA

K. & V. Bijdr. 5 (1900) 88; Merr. Philip. J. Sc. 1
(1906) Suppl. 111; DANserR, Blumea 1 (1934) 49;
BACK. & BAKH. f. Fl. Java 2 (1965) 159; MATTHEW,
Blumea 23 (1976) 80, f. 5 (map), 6. — M. arborea
[non (WIGHT) BEDD.] CLARKE, FI. Br. Ind. 2 (1879)
745, p.p.; KANJILAL & Das, Fl. Assam 2 (1938)
370; HUNDLEY & CuitT, Trees Shr. Burma ed. 3
(1961) 119. — M. cambodiana Pierre, Fl. Coch.
(1892) t. 260 B; WANGERIN, Pfl. Reich Heft 41
(1910) 29; EvrarD, FI. Gén. I.-C. 2 (1923) 1195. —
M. scortechinii Kinc, J. As. Soc. Beng. 71, ii
(1902) 73; WANGERIN, Pfi. Reich Heft 41* (1910)
27, f. 1 L-M; DANseER, Blumea 1 (1934) 56. — M.
Philippinensis WANGERIN in Fedde, Rep. 10 (1912)
273; Merr. En. Philip. 3 (1923) 241; CHAo,
Taiwania 5 (1954) 94, 99, f. 37; Lt & CHAO, Quart.
J. Taiwan Mus. 7 (1954) 124, f. 19. — M. parvi-
folia HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 41;
DANSER, Blumea 1 (1934) 51. — M. subcaudata
Me_rr. Philip. J. Sc. 13 (1918) 43; En. Philip. 3
(1923) 242. — M. megacarpa RIDL. FI. Mal. Pen. 1
(1922) 891. — M. chinensis MERR. Sunyatsenia 3
(1937) 256; Li, Taiwania 1 (1938) 94. — M. alterni-
folia Merr. & CHUN, Sunyatsenia 5 (1940) 153. —
M. poilanei TarpigEvu, Fl. Camb. Laos & Vietnam
8 (1968) 16. — Fig. 3c—e (galls).

Tree up to 37 m; d.b.h. up to 75 cm; branchlets
slender or stout, scattered, subglabrous to puberu-
lous. Leaves scattered, obovate, elliptic to oblong-
elliptic, 4-16 by 2-8cm, chartaceous to thick-
coriaceous, subglabrous: base cuneate to attenuate;
apex acute or acuminate to caudate; nerves 4~7(-9)
pairs, distinct below; veins distinct below; petiole
stout or slender, 1-4 cm. Inflorescence up to 8 cm,
slender or stout, subglabrous to densely appressed-
hairy, up to 3(-4) times branched; branches of the
first order scattered; bracts either all triangular,
under 3 mm, or lower ones lanceolate, up to 15
mm; basal inflorescence axes of the first order
subtended by ordinary bracts or by leaves.
Submature flower bud up to 3!/, mm @. Sepals 4 or
5, broader than long or as long as broad, thick,
puberulous, to appressed-hairy. Petals 4 or 5, thick,
glabrous to appressed-hairy. Stamens 4-5. Ovary
puberulous to appressed-hairy. Fruit ovoid to
oblong, 2!/,—31/, by 1-11/, cm; persistent disk con-
spicuous or not; sepals inconspicuous.

Distr. Continental SE. Asia (NE. India,
Bhutan, Burma, Thailand, Tonkin, S. Yunnan)
and throughout Malesia; not yet recorded from
the Lesser Sunda Is. and New Guinea.

KEY TO THE SUBSPECIES

1. Flowers basically 4-merous. Inflorescence stout,
rusty-puberulous. Corolla glabrous outside.
Leaves obovate, coriaceous; apex acute; base

b. ssp. moluccana

1. Flowers basically 5-merous.

pairs; veins distinct beneath. Fruit ovoid.
3. Basal bracts lanceolate, up to 15 mm. Fruit
larger than 3 by 1'/,cm . a. ssp. pentandra
3. All bracts triangular, under 3 mm. Fruit up to
Dae Oy ein os co -< CSSD. CHINeNSIS
2. Leaves up to 4-12 by 11/,-5 cm; nerves up to 6
pairs; veins obscure beneath. Fruit oblong.

{[ser. I, vol. 8?

4. Length of sepals up to half as long as wide.
Leaves chartaceous to subcoriaceous. Fruit
2'/,-3 cm long : . d. ssp. philippinensis

4. Length of sepals almost as long as wide.
Leaves coriaceous to thick-coriaceous. Bracts
uniformly triangular. Fruit 1°/,-2cm long

e. ssp. scortechinii

a. ssp. pentandra.

Tree up to 34 m; branchlets stout. Leaves elliptic
to oblong-elliptic, 8-16 by 4-8 cm, coriaceous;
base cuneate; apex acuminate; nerves 6~-7(-9)
pairs; veins distinct below; petiole stout, 2-4 cm.
Inflorescence up to 8 cm, stout, densely appressed-
hairy; basal bracts lanceolate, up to 15 mm. Sepals
5, broader than long. Petals 5, appressed-hairy
outside. Stamens 5. Fruit ovoid, 3—-31/, by 11/,-11/,
cm.

Distr. Malesia: West & East Java.

Ecol. In humid, mixed forest, 400-500 m. Fi.
July—Dec.

Vern. Java: Auru lilin, tenjau.

b. ssp. moluccana MATTHEW, Blumea 23 (1976) 81.

Tree up to 15 m; d.b.h. 20 cm; branchlets stout.
Leaves obovate, 8-15 by 3-8 cm, coriaceous; base
attenuate; apex acute; nerves 5-6 pairs; veins
distinct below; petiole stout, 2!/,-3 cm. Inflores-
cence up to 5cm, stout, rusty puberulous; basal
bracts up to 4 mm. Sepals 4(—5), broader than long.
Petals 4(—5), glabrous outside. Stamens 4(—5).
Fruit (immature) ovoid, 11/, by 3/4 cm.

Distr. Malesia: Moluccas (Morotai).

Ecol. Mixed rain-forest, up to 1000 m. Fl. May.
Once a leaf-gall was noted.

Notes. The basal pair of lateral inflorescence
branches often occur in the axils of normal leaves,
a tendency noted in ssp. philippinensis. Flowers
relatively large, yellowish to greenish; corolla
dome-shaped (in bud). Calyx margin wavy; petals
4 by 3 mm; filaments 3 mm; anther 1 mm; style
11/, mm. The only fruit seen is detached and im-
mature.

The arrangement of the primary inflorescence
branches is at times obscure.

The soft, coriaceous texture of the leaves, dark
above, and pale below, the stout inflorescence with
rusty indumentum and the few, large, 4-merous
flowers with glabrous dome-shaped corolla (in
bud) distinguish this subspecies from the others.
It is yet only known from Morotai I.

c. ssp. chinensis (MERR.) MATTHEW, Blumea 23
(1976) 83. — M. chinensis MERR. — Fig. 3¢ (galls).

Tree up to 20m; branchlets stout. Leaves
elliptic to elliptic-oblanceolate, 8-20 by 4-8 cm,
coriaceous; base attenuate; apex acute; nerves 6-8
pairs; veins distinct below; petiole stout, 1%/,-
2'/, cm. Inflorescence up to 8 cm, subglabrous to
appressed-hairy; all bracts uniform, under 3 mm.
Sepals 5, broader than long. Petals 5, appressed-
hairy outside. Stamens 5. Fruit oblong, 2-2'/, by
1 cm.

Distr. NE. India, Bhutan, N. Burma, Thailand,
S. China (Yunnan), Tonkin; in Malesia: Malay
Peninsula (Kedah, once).

Ecol. Mixed forests up to 1900 m. Fl. May-
June, fr. Aug.—May. In India galls and domatia
occur.

1977]

d. ssp. philippinensis (WANGERIN) MATTHEW,
Blumea 23 (1976) 85.— M. philippinensis WANGERIN
—WM. subcaudata MERR. — Fig. 3d-e (galls).

Tree up to 22'/,m; brachlets slender. Leaves
obovate to elliptic, 4-12 by 1'/,-5 cm, chartaceous
to subcoriaceous; base attenuate; apex acuminate;
nerves 4-6 pairs; veins obscure beneath; petiole
slender, 1-2 cm. Inflorescence up to 7 cm, slender,
subglabrous to puberulous; all bracts triangular,
under 3 mm. Sepals (4-)5, broader than long.
Petals (4-)5, glabrous to appressed-hairy outside.
Stamens (4-)5. Fruit oblong, 2!/,-3 by 1*/,—1"/, cm.

Distr. Malesia: throughout the Philippines.

Ecol. In forests, from low altitude up to 1350 m.
Fl. May-Sept., fr. Jan.Dec. Galls are rather
frequent on stem, leaf and fruit.

Note. Ssp. philippinensis is distinguished from
ssp. scortechinii in the generally smaller height and
smaller and thinner leaves, more slender inflores-
cence axis, the lower 1 or 2 inflorescence axis (axes)
a the axil(s) of normal leaves, and the oblong
Tuits.

e. ssp. scortechinii (KING) MATTHEW, Blumea 23
(1976) 86. — M. scortechinii KING — M. mega-
carpa RipL. — M. parvifolia HALL. f.

Tree up to 37m; branchlets stout. Leaves
obovate to oblong, 4-12(-15) by 3—5(—6) cm, thick-
coriaceous; base cuneate to attenuate; apex acute
to acuminate; nerves 4-6 pairs; veins obscure
beneath; petiole stout, 11/,-2'/, cm. Inflorescence
up to 8 cm, stout, puberulous to villous; all bracts
triangular, under 3 mm. Sepals (4-)5, as long as
broad. Petals (4~-)5. Stamens (4-)5. Fruit oblong,
2!/,—-2!/, by 13/,-2 cm.

Distr. Thailand (once); in Malesia: S. Sumatra,
Banka, Malay Peninsula, Borneo, Celebes.

Ecol. Primary forests, from low altitude up to

CORNACEAE (Matthew) 97

2400 m. Fi. fr. Jan.—Dec. Inflorences and fruit galls
occur.

Vern. Malaya: médang pisang. Banka: mén-
kapas. Borneo: kaju wulu, médang surugan.

Note. Ssp. scortechinii is distinguished from
ssp. pentandra by the generally obovate and smaller
leaves, less stout inflorescence, uniformly short
bracts and oblong fruits with thick pericarp.

Excluded

Mastixia cuneata BL. Mus. Bot. Lugd. Bat. 1
(1850) 257 = Notaphoebe umbelliflora (Bu.) BL.
Cf. HALL. f. Beih. Bot. Centralbl. 34, 2 (1916) 42;
DANSER, Blumea | (1934) 68.

Mastixia gracilis KiNG, J. As. Soc. Beng. 74, ii
(1902) 73; WANGERIN, Pfi. Reich Heft 41+ (1910)
28; DANsER, Blumea 1 (1934) 68; MATTHEW,
Blumea 23 (1976) 90 = Vaccinium bancanum Mia.
var. tenuinervium J. J. S. (Ericaceae), according to
the type number Wray 1528 mentioned by
SLEUMER, Blumea 11 (1961) 76. — Ed.

Mastixia heterophylla BL. Mus. Bot. Lugd. Bat.
1 (1850) 257; WANGERIN, Pfl. Reich Heft 414 (1910)
28; DaANsER, Blumea 1 (1934) 69; MATTHEW,
Blumea 23 (1976) 90. — HALLItER f. suggested this
to be Gomphandra capitulata BEcc., but this was
questioned by SLEUMER, Blumea 17 (1969) 193.
According to us this sterile sheet (L 901, 169-350)
collected by PRAETORIUS in Palembang, is not a
Mastixia but we cannot give a proper identification.

Mastixia tetrandra (THW.) CLARKE. — DANSER,
Blumea 1 (1934) 56, referred two Sumatran speci-
mens to this species, which is hitherto only found in
Ceylon and the Andaman Is. One of these is
sterile and the other is in bud; they can equally well
be referred to M. rostrata ssp. rostrata, and their
identification remains doubtful. Cf MATTHEW,
Blumea 23 (1976) 77.

Excluded

Cornus caudata Hassk. & ZOLL., nom. illeg., C. ilicifolia (BL.) HAssK. & ZOLL., C. serrulata (BL.)
Hassk. & ZOLv., and C. stricta ZoLL. & Mor. are all combinations or names made by HAssKARL and
ZOLLINGER based on Polyosma BL., because of their opinion that this genus of the Saxifragaceae would be
synonymous with Cornus L. Cf. HAssKARL, Cat. Hort. Bog. (1844) 168 and ZoLLINGER, Natuur- &

Geneesk. Arch. Neérl. Ind. 2 (1845) 10.

ONAGRACEAE (P. H. Raven, St. Louis)!

Annual or perennial herbs (in Mal.), occasionally somewhat woody near the base,
sometimes aquatic. Leaves spiral or opposite. Stipules absent or reduced, deltoid.
Flowers mostly 4-merous, rarely 5-merous (in Mal.), solitary or arranged in a
terminal racemose inflorescence, subtended by (often reduced) leaves or bracts.
Bracteoles absent or 2 at the base of the ovary. Floral tube short or absent. Sepals
erect, persistent. Petals caducous, contorted in aestivation, white, pink or yellow,
sometimes emarginate. Stamens 4, 5, 8, or 10, in 2 whorls, rarely with an inter-
mediate number, epipetalous ones sometimes shorter. Anthers usually versatile,
sometimes seemingly basifixed by reduction: pollen single or in tetrads. Ovary
inferior, (in Mal.) 4- or 5-celled and with c~ ovules; summit of the ovary (disk) flat
to conical (in Mal.), sometimes with depressed nectaries surrounding the bases of
the epipetalous stamens. Style simple; stigma capitate, clavate or globose, often
4-lobed. Ovules with axial placentation, |-pluriseriate. Fruit (in Mal.) a mostly long
and slender loculicidal or irregularly rupturing capsule. Seeds rounded or elongate,
in Ludwigia sometimes embedded in powdery or surrounded by cork-like endocarp
tissue, in Epilobium with a chalazal plume of trichomes (coma); endosperm absent;
embryo straight.

Distribution. About 17 genera and more than 600 spp. in tropical and temperate regions, with a dis-
tinct centre of diversity on the northern hemisphere in the New World, in Malesia two native genera which
are both almost ubiquist.

Ecology. Ludwigia is largely confined to the hot lowland and hills usually in wet or damp localities,
Epilobium is confined to the higher mountain regions.

Dispersal. Epilobium spp. are manifestly wind dispersed by virtue of their coma. Ludwigia spp. depend
on dispersal by water and possibly incidental exozoic dispersal by water birds; in Ludwigia hyssopifolia
there are two kinds of seed, one of which is enveloped by a corky tissue derived from the endocarp,
enhancing their buoyancy.

Pollination. Almost all of the Malesian species are self-pollinated, shedding pollen directly on the
stigma at or before anthesis and rarely visited by insects. In Ludwigia peruviana, introduced in the Old
World, the anthers are extrorse and shed pollen away from the stigma; thus outcrossing is predominant.
Some outcrossing probably also occurs in the relatively large-flowered L. adscendens and L. octovalvis,
which are known to be visited by insects, and in Epilobium detznerianum, in some populations of which the
stigma is even held above the anthers. In our area, Here (Dansk Bot. Ark. 5, 1927, 18) reported Melipona
sp., a bee, visiting the flowers of Ludwigia peruviana (as Jussieua peruviana), and Bombus rufipes at the
flowers of the locally naturalized Oenothera stricta (as O. lamarckiana) and Fuchsia magellanica (as F.
coccinea). The Melipona bees were not observed to contact the anthers or stigma of the large-flowered
Ludwigia, but would certainly do so in visiting smaller-flowered species. All of the Malesian species are
genetically self-compatible.

Morphology & Anatomy. The Onagraceae are distinctive in their monosporic, 4-nucleate,
‘Oenothera type’ embryo sac development; in the nearly universal presence of viscin threads among the
pollen; and in the loose construction of their pollen exine. Most species of Epilobieae and about half of
Ludwigia shed their mature pollen in tetrads; these include all Malesian species except L. adscendens and
L. hyssopifolia, in which the pollen is shed singly. Intraxylary phloem occurs throughout the family
adjacent to primary xylem, and interxylary phloem (included phloem) is found in many genera but not in
Ludwigia (CarLquist, Ann. Mo. Bot. Gard. 62, 1975, 386); in these features Onagraceae resemble many
other Myrtales.

The stomata are surrounded by three or more subsidiary cells, sometimes resembling those of Cruciferae.

All Onagraceae have an inferior ovary and a floral tube, which is prolonged beyond the ovary in all
except Ludwigia and Epilobium sect. Chamaenerion.

In several species of Ludwigia half-submerged parts of the stem are covered by a whitish aerenchyma;
in L. adscendens short roots at the nodes are transformed into inflated, elongate aerophores enhancing
floating on water.

Chromosomes. All species of Epilobium sect. Epilobium, a taxon that includes all Malesian species,
which have been examined have had a gametic chromosome number of n = 18. Species of Ludwigia have
a gametic chromosome number of m = 8 and multiples. These genera differ from most others in Onagra-

(1) With co-operation of the General Editor in framing the manuscript. ’
The author gratefully acknowledges the support of the U.S. National Science Foundation to the studies
of Onagraceae.

(98)

1977] ONAGRACEAE (Raven) 99

ceae in having small chromosomes that are heteropycnotic and dark-staining throughout the mitotic
cycle. Naturally occurring interchange heterozygotes, abundant in the tribe Onagreae, are not known to
occur in either group. The original basic chromosome number of the family is x = 11, as in Fuchsia,
Circaea, and others.

Hybridization. Hybrids are rare between the recognized species of Ludwigia. In Epilobium sect.
Epilobium they are occasional where two or more entities come together, but their occurrence is limited by
the predominant autogamy or cleistogamy of most species and to some extent by ecological differentiation
also. A wide range of fertilities is characteristic of these hybrids, as explained in detail in our monograph
of the Australasian species (RAVEN, D.S.I.R. New Zeal. Bull. 216, 1976), and cytoplasmic differences
sometimes occur. Two of the Malesian species occur together in N. Luzon, and a few individuals suggest
hybridization; four occur together in the mountains of New Guinea, with hybrids probably occasional
but poorly studied so far. All species of Epilobium found in Malesia have the same chromosome arrange-
ment that is predominant in Eurasia, from where they doubtless came.

Chemotaxonomy. Raphides, needle-like crystals of calcium oxalate, are ubiquitous in the vegetative
parts of Onagraceae. The few reports of alkaloids are doubtful and seem to indicate rather the presence of
secondary amines. Ellagic acid occurs. Among the flavonoids reported from the family, flavonols based
quercetin are ubiquitous, whereas kaempferol and more highly oxygenated types based on myricetin are
frequent. The anthocyanins include predominantly malvidin and cyanidin derivatives, with the latter
predominant in the rose-purple petals of Epilobium. The yellow petals of most species of Ludwigia are
colored by carotenoids, with the chalcone isosalipurposide forming a non-ultraviolet-reflective centre in
many species, including L. peruviana.

Uses. Only some species of Ludwigia are mentioned to be in use for minor medicinal purposes; see
under Ludwigia spp.

KEY TO THE GENERA

1. Capsule loculicidally dehiscent with 4 valves; axis persistent. Seeds comose. Floral tube present.
Petals 4, white, pinkish or red. Stamens 8. Stem-base without aerenchyma. . . . . 2. Epilobium
1. Capsule irregularly dehiscent; axis not persistent. Seed not comose. Floral tube absent. Petals yellow,
if white or creamy then flowers 5-merous and floating aquatic plant adorned with short spongy aeren-
chyma-roots at the nodes. Stamens 4-10. Stem-base not rarely covered by aerenchyma. 1. Ludwigia

1. LUDWIGIA

Linn£, Gen. Pl. (1754) 55; Sp. Pl. 1 (1753) 118; Munz, Bull. Torr. Bot. Cl. 71
(1944) 152; HaRA, J. Jap. Bot. 28 (1953) 289; A. & R. FERNANDES, Garcia de Orta 5
(1957) 109; RAVEN, Reinwardtia 6 (1963) 327. — Jussiaea LINNE, Gen. PI. (1754)
183; Sp. Pl. 1 (1753) 388; BAcK. Trop. Natuur 3 (1914) 59; Fawcett, J. Bot. 64
(1926) 10; MuNz, Darwiniana 4 (1952) 179. — Nematopyxis Mia. FI. Ind. Bat. 1, 1
(1855) 600. — Fig. 1, 4, 5.

Slender herbs, erect or creeping and rooting at the nodes, to large shrubs.
Underwater parts often swollen and spongy or bearing inflated white spongy aero-
phores. Leaves alternate or opposite, mostly entire. Stipules absent or reduced,
deltoid. Flowers borne singly, clustered, or arranged in an inflorescence. Brac-
teoles lacking or conspicuous, usually two, at or near the base of the ovary. Floral
tube absent. Sepals 3-7, persistent after anthesis. Petals as many as the sepals or
absent, caducous, yellow or white, with contorted aestivation. Stamens as many as
or twice as many as the sepals, or flowers very rarely with an intermediate number
of stamens; anthers usually versatile but sometimes apparently basifixed by reduc-
tion. Pollen shed in tetrads or singly. Disk (summit of the ovary) flat to conical,
often with depressed nectaries surrounding the bases of the epipetalous stamens.
Stigma hemispherical or capitate, the upper 1'/,-*/, receptive, often lobed, the
number of lobes corresponding to the number of locules. Ovary with a number of
cells equal to the number of sepals, very rarely more; placentation axial; ovules
pluriseriate or uniseriate in each cell, in one species uniseriate below, pluriseriate
above; if uniseriate, the seeds sometimes embedded in powdery or woody endocarp

100 FLORA MALESIANA [ser. I, yokes

from which they detach easily or with difficulty. Capsule irregularly dehiscent, or by
a terminal pore, or by flaps separating from the valve-like top. Seeds rounded or
elongate, the raphe usually easily visible and in some sections equal or nearly equal
in size to the body of the seed.

Distr. According to my synopsis (Reinwardtia 6, 1963, 329) 75 spp., all over the world; in Malesia 6
spp., one of which is certainly introduced.

Ecol. One aquatic and the other species mostly in swampy or damp places, often in rice-fields, from the
lowland up to c. 2100 m, mostly below 1000 m. Flowers last only one day.

Taxon. I have divided the genus into 17 sections, the largest of which (sect. Myrtocarpus) is neotropical.
They are often shrubby with large, 4- or 5-merous flowers, dimerous stamens, prominently 4- or 5-ribbed
capsules, free seeds and pollen grains shed in tetrads. They appear phylogenetically central in the genus.
In Malesia this section is represented by an introduced weed, L. peruviana. Close to this section are one
African (sect. Africana) and one American section (sect. Macrocarpon) with terete capsules. Following
these is a series of small Old World sections which have the stamens reduced to one whorl; in one African
section flowers are 3-merous. L. hyssopifolia forms a monotypic section unique in having two kinds of
seeds, those in the lower part of the capsule uniseriate and embedded in the endocarp, those in the upper
part pluriseriate and free, while pollen grains are single, Other sections, not represented in Malesia, have
all of the seeds loosely embedded in powdery endocarp. The structure of the seed is important in the dis-
crimination of sections.

The second major line of the genus consists of species in which the seeds are embedded in coherent
chucks of woody endocarp which render the capsule a tough unit from which it is difficult to separate the
seeds. The two sections belonging to this line have basically 5-merous flowers and pollen shed singly.
Through the disentangling of these relationships it appears that the number of stamens is not decisive for
dividing the genus into two genera as this would go across relationships and lead to heterogeneous assem-
blages of species.

Each Malesian species belongs to a different section and being so small in number it seems not useful to
give descriptions of these sections; I refer to my revision (1963).

The cradle of the genus is probably South America with an important secondary centre of evolution in
Africa. It is one of the most primitive genera in the family.

Note. It has appeared that seeds retain viability in the herbarium in unpoisoned, not too old speci-
mens; flowering plants can thus be raised from fruiting herbarium specimens.

KEY TO THE SPECIES

1. Stamens twice as many as sepals.
2. Seeds pluriseriate, free (not embedded in endocarp).
3. Plant subglabrous to appressed-pubescent. Capsule terete. Petals 5-17 by 4-17 mm. Style 1.5-
3.5 mm. Raphe equal in diameter to the body of theseed. . ......... 2. L. octovalvis
3. Coarse, strongly pubescent or villous plant. Capsule strongly 4-angled, villous, with flat sides.
Petals 15-24 by 16-26 mm. Style 1 mm long. Raphe not more than '/, the diameter of the body of the
Sr MERE eee | Oc eee. dea ame soy = whe: « SIS yay! acngssl zyciai uSGuis: 41 Me, ie Ase Ae 1. L. peruviana
2. Seeds at least below uniseriate and embedded in endocarp.
4. Seeds in the c. 1/, upper part of the capsule pluriseriate and free. Sepals 4. Petals 2-3 mm long
_ 5. L. hyssopifolia
4. Seeds all uniseriate in each cell of the capsule and embedded in endocarp. Sepals 5—7 (rarely 4). Petals
4.5-23 mm long.
5. Aquatic, with floating branches forming erect clusters of spongy, spindle-shaped aerophores
(aerenchyma). Petals white or creamy, with yellow atthe base. . ...... 6. L. adscendens
5. Plant not forming such aerophores on the decumbent branches. Petals bright golden-yellow with a
darker spot at the base. New Zealand, Australia, Pacific Is., Formosa, China, and Japan, also in the
New World, might possibly be found in East Malesia. Cf. Aston, Aquat. Pl. Austr. (1973) 144, f. 55
L. peploides (H.B.K.) RAVEN
1. Stamens as many as sepals, very rarely more in some flowers.
6. Seeds pluriseriate in each cell of the capsule. Petals elliptical, 1-3 by 0.7-2 mm. Capsule terete. Seeds

ira egy Oa NI) oe 8. | hab 5a so beet te, Patera late tetas dace 3. L. perennis
6. Seeds uniseriate in each cell of the capsule. Petals narrow spathulate, 1.3-2.2 by 0.4-0.9 mm. Capsule
Slightly 4-anpied. Seeds'0:5—0)6 by: O:3Simm- 20) ee ee eo BO 4. L. prostrata

1. Ludwigia peruviana (L.) Hara, J. Jap. Bot. 28 Steen. Fl. Sch. Indon. (1949) 305. — Oenothera
(1953) 293; RAvEN, Reinwardtia 6 (1963) 345, hirta LInNé, Sp. Pl. ed. 2, 1 (1762) 491. — Jussieua
map 14. — Jussiaea peruviana LINNE, Sp. Pl. 1 hirta (L.) Sw. Obs. Bot. (1791) 142, non LAmMK,
(1753) 388 ; Back. Trop. Natuur 3 (1914) 61; Onkr. 1789; Back. Ann. Jard. Bot. Btzg Suppl. 3 (1909)
Suiker. (1930) 470, Atlas t.445; ALstoNin Trimen, 406. — Jussiaea speciosa RiDL. J. Bot. 59 (1921)
FI. Ceyl. 6 (1931); Munz, Darwiniana 4 (1942) 131; 259; Fl. Mal. Pen. 1 (1922) 828.

1977]

ONAGRACEAE (Raven)

101

Shrub 0.5-3 m, entirely villous, the hairs often
multicellular, especially in the inflorescence; long
inflated aerophores arising from submerged,
buried roots. Leaves lanceolate to broadly lanceo-
late, 4-12 by 0.3—-1.5 cm, narrowly cuneate at base,
apex acute to acuminate; nerves 12-22 pairs; sub-
marginal vein not prominent; petiole 3-12 mm.
Flowers in upper leaf axils. Bracteoles lacking or up
to 7 mm long, subulate. Sepals 4 or 5, lanceolate,
irregularly serrulate, 10-18 by 4-8 mn, villous.
Petals bright yellow, veiny, suborbicular, 15-24 by
16-26 mm, shallowly emarginate, with a claw 1-3
mm. Stamens 8 or 10, subequal; filaments 2-3.5
mm; anthers 3-4.5 mm long, extrorse and not
shedding pollen directly on the stigma at anthesis.
Pollen shed in tetrads. Disk elevated 1-2 mm, with
a depressed densely white-hairy nectary around the
base of each epipetalous stamen. Style c. 1 mm;
stigma broadly elongate-hemispherical, 2-3 mm
high. Capsule villous, 1.2-3 by 0.6—-1 cm, light
yellowish brown with 4 prominent dark brown
ribs, 4-angled, thin-walled, readily and irregularly
loculicidal; pedicel 2-4.5 cm. Seeds pluriseriate in
each cell, free, light brown, finely striate and cellu-
lar pitted, obovoid, 0.6—-0.8 mm long; raphe */, to
1/, the width of the body.

Gametic chromosome number (Old World
populations), n = 40.

Distr. Native of the New World, from the SE.
United States throughout South America, intro-
duced and naturalized in Malesia since the 2nd
half of the last century, collected in Malaya,
Sumatra, Java (common in West), but obviously
still absent from many areas.

Ecol. Along ditches and in moist places, mostly
in the lowland but ascending to c. 1400m. Fi.
Jan.—Dec.

Vern. Banka: pitjanket, M; Java: tjatjabéan, S,
lombokan, J.

Note. BAckKer (Onkr. Suiker. 1930, 470)
observed that in inundated situation the plant
produces aerophores which are emitted by shallow,
horizontal roots; they are erect but their tips
usually float on the water.

2. Ludwigia octovalvis (JAcQ.) RAVEN, Kew Bull.
15 (1962) 476; Reinwardtia 6 (1963) 356, maps
19-20, incl. ssp. brevisepala (BRENAN) RAVEN ef
ssp. sessiliflora (MicH.) RAVEN; HENRY & PRIT-
CHARD, Bot. Div. Lae, Bot. Bull. 7 (1973) 132,
fig.; Everist, Pois. Pl. Austr. (1974) 393. —
Jussiaea suffruticosa LINNE, Sp. Pl. 1 (1753) 388;
Bru. Fl. Austr. 3 (1867) 307; F.v.M. Descr. Not.
Pap. Pl. 4 (1876) 60; CLARKE, FI. Br. Ind. 2 (1879)
587; Koorp. Exk. Fl. Java 2 (1912) 703; Rip.
Trans. Linn. Soc. Bot. II, 9 (1916) 57, incl. var.
hirta Ripv.; GaGn. Fl. Gén. L.-C. 2 (1921) 986;
C. T. Wuite, Proc. R. Soc. Queensl. 34 (1922) 48;
Back. & BAKH. f. Fl. Java 1 (1963) 261. — Oeno-
thera octovalvis JACQ. En. Syst. Pl. (1760) 19. —
Jussiaea pubescens LINNE, Sp. Pl. ed. 2, 1 (1762)
555. — L. perennis (non L.) Burm. /. Fl. Ind. (1768)
37. — Jussiaea angustifolia LAMK, Encycl. 3 (1789)
331. — Jussiaea villosa LAMK, I.c.; RipL. Fl. Mal.
Pen. 1 (1922) 828. — Jussieua octovalvis (JACQ.)
Sw. Obs. Bot. (1791) 142. — Jussiaea angustifolia
Bi. Bijdr. (1826) 1132, non Lamxk, 1789; BAcK.
Trop. Natuur 3 (1914) 62. — Jussiaea blumeana
DC. Prod. 3 (1828) 55. — Jussiaea burmannii DC.

l.c. 57. — Jussiaea exaltata Roxs. (Hort. Beng.
1814, 33, nomen) Fl. Ind. ed. Carey 2 (1832) 401. —
Jussiaea costata Pr. Epim. Bot. (1849) 217. —
Jussiaea junghuhniana Mia. FI. Ind. Bat. 1, 1 (1855)
627; VAL. Bull. Dép. Agric. Ind. Néerl. 10 (1907)
41. — Jussiaea erecta (non L.) RIDL. J. Bot. 59
(1921) 258; Fl. Mal. Pen. 1 (1922) 827, incl. var.
exaltata (Roxs.) Ripi.; Back. Onkr. Suiker.
(1930) 470, Atlas t. 446; STEEN. Arch. Hydrobiol.
Suppl. 10 (1932) 314. — L. pubescens (L.) HARA,
J. Jap. Bot. 28 (1953) 293. — Fig. 1.

Usually robust, well-branched herb, sometimes
woody at the base, up to 4 m, subglabrous or with
sparse or dense appressed or spreading pubescence.
Leaves lanceolate or narrowly lanceolate, to
narrowly ovate, or subovate, 2-14 by 0.5-4 cm,
narrowly to broadly cuneate at the base and
attenuate at apex; nerves 11-20 pairs, sub-
marginal vein well developed; petiole up to 1 cm.
Bracteoles reduced or to 1 mm long. Sepals 4, ovate
or lanceolate, 6-15 by 1-7.5 mm. Petals yellow,
broadly obovate or cuneate, emarginate, 17 by
2-17 mm. Stamens 8, epipetalous ones shortest;
filaments 1-4 mm long; anthers 0.5-4 mm long,
extrorse but soon crumbling and shedding pollen
directly on the stigma. Pollen shed in tetrads. Disk
slightly raised, with a white-hairy sunken nectary
surrounding the base of each epipetalous stamen.
Style 1.5-3.5 mm; stigma subglobose, shallowly
4-lobed, 1.2-3 mm @. Capsule thin-walled, 1.7-
4.5cm by 2-8mm, terete, pale brown with 8
darker ribs, readily and irregularly loculicidal;
pedicel up to 10mm. Seeds pluriseriate in each
cell, free, brown, rounded, 0.6-0.75 mm long,
0.5-0.7 mm wide including the inflated raphe
which is equal in size to the body of the seed and
evenly transversely ridged.

Gametic chromosome numbers, n = 16, 24.

Distr. Throughout the tropics of the world,
between c. 32° N and 30° S.

Ecol. Mostly in humid places, damp grassland,
rice-fields, along ditches and water-courses, in
swamps, lakes and pools, drains, sandy or silty
floodbanks, gravelly riverbeds, on floating islands
in lakes, on floating logs in lagoons, sago swamps,
mountain peat swamps with sedges, also in old
native gardens and coconut plantings, from the
lowland up to c. 1000 m, in Java and Celebes up to
1400 m, in New Guinea up to 2100 m. Fi. Jan.—
Dec.

Several collectors mention that it propagates by
runners and that old leaves turn reddish. The lower
part of the stem is at times coated by aerenchyma.
In inundated condition aerophores are produced;
see under |. L. peruviana.

DOocTERS VAN LEEUWEN (Zoocecidia, 1926, 427
f. 808; Ned. Kruidk. Arch. 51, 1941, 204) recorded
fruit galled by beetles and aphid galls on the ter-
minal leaves of the branches.

Uses. In Java minor medicinal qualities are
ascribed to this species, amongst others against
sprew. Rumpuius, who described it under the name
herba vitiiginum (Herb. Amb. 6, p. 49) did not
mention uses (Heyne, Nutt. Pl. 1927, 1206).
BurkiLL (Dict. 1935, 1274) reported that the
mucilaginous leaves, after which the plant is called
‘lakomayer’ = ‘water Vitis’, are used for poulticing
in a variety of complaints; it has also been recorded
as used for headaches, orchitis, glands in the neck,

102 FLORA MALESIANA [ser. I, vol. 8?

i Me: oom

ay

\}

Fig. 1. Ludwigia octovalvis (JACQ.) RAVEN. a. Habit in flower, b. withered fruits, both x #/, (a after
BACKER, 1940, b JUNGHUHN CCXxII).

1977]

diarrhoea, dysentery, nervous diseases, and as a
vermifuge. In WILKINSON’s Dictionary it is said
that a kind of tea is made from the leaves. Also in
India medicinal properties are ascribed to it.

Vern. Malaya: buyang samalam, lakom ayér,
pujang malam, yénlidah, M; Sumatra: urang aring
itém, Simalur I.; Java: gagabusan, tjatjabéan, S,
kalaménja, Md, salah njowo, J; Lesser Sunda Is.:
pangambo, E. Sumbawa; Philippines: tayilakton,
Tag., balansuit, Mag., malapako, tubong-talapan,
Bik., pachar-pachar, Sul., palangdisin, Ig., talang-
kau, Ilk., halangot, naudyawa tubig, If.; Celebes:
kélétélé téngén, Tonsaw. dial., Minahassa;
Moluccas: daun panu, Ambon; New Guinea: ewo,
onarenare, Kapauko lang., kampur, Sakaj bivouac,
Merauke, pfauhanu, Kutubu, mayenke, Orne lang.,
Kaiye, rowijetwi, Enga lang., Yogos, togorarah,
Wapi lang., Marok, rama-rama, Matapaili lang.

Note. In my revision (1963) I distinguished three
subspecies, more or less geographically defined,
which I wish to withdraw here.

3. Ludwigia perennis LINNg, Sp. Pl. 1 (1753) 119;
A. & R. FERNANDES, Garcia de Orta 5 (1957) 114,
475; RAVEN, Reinwardtia 6 (1963) 367, map 21. —
L. parviflora Roxs. (Hort. Beng. 1814, 11, nomen)
Fl. Ind. 1 (1820) 440; Brn. Fl. Austr. 3 (1867) 307;
Kurz, J. As. Soc. Beng, 46, ii (1877) 91; CLARKE,
Fl. Br. Ind. 2 (1879) 588; TRIMEN, FI. Ceyl. 2
(1894) 234; Rip-. Fl. Mal. Pen. 1 (1922) 828; BAck.
Onkr. Suiker. (1930) 471, Atlas t. 447; BAck. &
Baku. f. Fl. Java 1 (1963) 261. — L. lythroides
BL. Bijdr. (1826) 1134. — L. gracilis Mig. Fl. Ind.
Bat. 1, 1 (1855) 629. — Jussiaea perennis BRENAN,
Kew Bull. 8 (1953) 163.

Annual herb up to | m, subglabrous or minutely
puberulent on younger parts. Leaves narrowly
elliptic to lanceolate, 1-11 by 0.3—-2.7 cm, narrowly
cuneate at base, apex subacute; nerves 6-12 pairs;
submarginal vein weakly developed; petiole
2-15mm, winged. Sepals 4, rarely 5, deltoid,
(1.3-)2-3.5 by (0.5-)0.7-1.8 mm, glabrous or
minutely puberulent. Petals yellow, elliptic, 1-3 by
0.7-2 mm. Stamens usually 4 or 5, rarely more;
filaments 0.3-0.7 mm; anthers 0.5-0.7 by 0.5-
0.7 mm, shedding pollen directly on the stigma at
anthesis. Pollen shed in tetrads. Disk slightly
elevated, glabrous. Style 0.7—1.5 mm; stigma glo-
bose, 0.4-0.5 mm @. Capsule thin-walled, glabrous
or puberulent, 3-16 (-19) mm long, terete, pale
brown, readily and irregularly loculicidal; capsule
sessile or on a pedicel up to 6 mm, often + nodding.
Seeds pluriseriate in each cell, free, brown with
fine brown lines, ellipsoid-rounded, 0.3-0.5 by
0.2-0.25 mm; raphe very narrow and inconspi-
cuous.

Gametic chromosome number, 1 = 8.

Distr. Tropical Africa (from Senegal, Lake
Chad, and the Sudan south to E. Congo and N.
Natal), Madagascar, through continental SE. Asia
(Ceylon to S. China, Hongkong, and Hainan), and
throughout Malesia (except Borneoand Moluccas)
to tropical Australia and New Caledonia. Fig. 2.

Ecol. Sunny, humid or marshy situations
fallow rice-fields, sawah dikes, along ditches, rivers
and water-courses, dry riverbeds, sugarcane fields
on heavy clay, damp places in Eucalypt savannah
—" aero from the lowland to c. 500 m. Fi.

an.-—Dec.

ONAGRACEAE (Raven)

103

In Java and Madura I. decidedly preferring
regions subject to a dry monsoon.

Fig. 2. Range of Ludwigia perennis L. (after RAVEN,
196

4. Ludwigia prostrata Roxs. (Hort. Beng. 1814, 11,
nomen) Fl. Ind. 1 (1820) 441; Bru. FI. Austr. 3
(1866) 308; CLARKE, Fl. Br. Ind. 2 (1879) 588;
TRIMEN, FI. Ceyl. 2 (1894) 234; Merr. FI. Manila
(1912) 355; Rpt. Fl. Mal. Pen. 1 (1922) 829;
Me_rr. En. Philip. 3 (1923) 220; Back. Onkr.
Suiker. (1930) 472, Atlas t. 448; Back. & BAKH.
f. Fl. Java 1 (1963) 261; RAvEN, Reinwardtia 6
(1963) 374, map 23. — L. fruticulosa BL. Bijdr.
(1826) 1133. — L. leucorhiza BL. I.c. — Nema-
topyxis pusilla Mia. FI. Ind. Bat. 1, 1 (1855) 630. —
Nematopyxis prostrata Mig. l.c. — Nematopyxis
fruticulosa Mia. l.c.

Annual herb 0.1-0.6m, subglabrous, often
reddish-tinged. Leaves elliptic or narrowly elliptic,
1-13 by 0.3-2.7 cm, glabrous or with a few minute
hairs along the veins, narrowly cuneate at the base,
apex acute; submarginal vein weakly developed;
petioles 4-25 mm, distinct. Sepals 4, deltoid, 1.3-
2.5 by 0.7-1.1 mm, glabrous. Petals yellow, narrow-
ly spatulate, 1.3-2.2 by 0.4-0.9mm. Stamens 4;
filaments 0.8-1.2 mm; anthers 0.4-0.5 mm wide,
broader than long, closely appressed to the stigma
and shedding pollen directly on it at anthesis.
Pollen shed in tetrads. Disk slightly elevated,
glabrous. Style c. 1 mm; stigma globose, c. 0.5 mm,
the upper half receptive. Capsule thin-walled,
glabrous, 12-22 by 0.8-I1 mm, +4-angled, pale
brown, readily and irregularly loculicidal, the
seeds showing plainly as indentations in the walls
at maturity. Seeds uniseriate in each cell, free, pale
brown, speckled or striped transversely with narrow
darker brown stripes, plump, ovoid, apiculate at
one end, 0.5-0.6 by 0.3 mm; raphe narrow, linear.

Gametic chromosome number, 7 = 8.

Distr. Tropical SE. Asia (Ceylon and S. Deccan
to N. India, Assam, S. China and Andamans); in
Malesia: Malay Peninsula, Java, Lesser Sunda Is.
(Timor), Borneo, and Philippines (Palawan,
Luzon, Negros, Mindanao). The single record from
Australia (N. Queensland: Mossman R.) recorded
by C. T. WuiTe (Proc. R. Soc. Queensl. 50, 1939,
78) is a misidentification of L. hyssopifolia. On the
anon L. prostrata is in Malesia a rare species.

ig. 3.

Ecol. Paddies, fallow and planted, by ditches,
along riverbanks, in swampy places, rather rare,
from lowland up to c. 800 m. Fl. Jan.-Oct.

VERN. Philippines: alubihud, P.Bis.

Fig. 3. Range of Ludwigia prostrata Roxs. (after
RAVEN, 1963).

5. Ludwigia hyssopifolia (G.DON) ExELL, Garcia
de Orta 5 (1957) 471; A. & R. FERNANDES, /.c. 471,
474; RAVEN, Reinwardtia 6 (1963) 385, map 30;
HENTY & PRITCHARD, Bot. Div. Lae, Bot. Bull.
7 (1973) 131, fig. — Jussiaea linifolia VAHL, Eclog.
Am. 2 (1798) 32, non L. linifolia Por. 1813; BACK.
Trop. Natuur 2 (1913) 20, fig.; ibid. 3 (1914) 61;
Onkr. Suiker. (1930) 469, Atlas t. 444; STEEN.
Arch. Hydrobiol. Suppl. 10 (1932) 314; Back. &
BAKH. f. Fl. Java 1 (1963) 260. — Jussiaea hyssopi-
folia G.Don, Gen. Syst. 2 (1832) 693. — Jussiaea
suffruticosa (non L.) RIDL. J. Bot. 59 (1921) 257;
Fl. Mal. Pen. 1 (1922) 827. — L. prostrata (non
Roxs.) C. T. Wuite, Proc. R. Soc. Queensl. 50
(1939) 78. — Fig. 4.

Annual herb 5 cm to 3 m, often becoming woody
at the base; young growth and inflorescence
minutely puberulent; elongate aerophores arising
from buried submerged roots. Leaves lanceolate,
1-9 by 0.2-3 cm, narrowly cuneate at the base,
apex acuminate; nerves 11-17 pairs; submarginal
vein not prominent; petiole 2.5-18 mm. Sepals 4,
lanceolate, 2-4 by 0.7-1.2 mm, finely puberulent,
3-nerved. Petals yellow, fading orange-yellow,
elliptic, 2-3 by 1-2 mm. Stamens 8, pale greenish
yellow, epipetalous ones shorter; filaments of
episepalous stamens 1-2 mm, those of epipetalous
ones 0.5-1 mm; anthers 0.4-0.6 mm wide, 0.2-
0.3mm high, shedding pollen directly onto the
stigma at anthesis. Pollen grains shed singly. Disk
+ elevated, with a depressed ciliate nectary sur-
rounding the base of each epipetalous stamen.
Style pale greenish yellow, 1-1.5 mm; stigma de-
pressed-globose, c. 0.6-1.2mm @, 0.5-0.8 mm
high, shallowly 4-lobed, the upper portion recep-
tive. Capsule relatively thin-walled, finely puberu-
lent, 1.5-3cm by 1-1.2 mm, subterete, enlarged
in the upper ?/, to 4/3, subsessile. Lower seeds uni-
seriate in each cell of the capsule, nearly vertical,
brown, oblong, 0.7-0.85mm long, each firmly
embedded in a cube of relatively hard endocarp;
raphe '/, the diameter of the body. Seeds in upper
inflated portion of the capsule multiseriate, free,
ovoid, 0.35-0.5 mm long, paler brown than the
lower seeds and with a narrower raphe. Lower part
of capsule at first marked by distinct bumps
corresponding to the position of the uniseriate
seeds, but as the endocarp hardens and swells, the
capsule becomes smooth.

FLORA MALESIANA

[ser. I, vol. 82

Gametic chromosome number, n = 8.

Distr. Tropical Africa (Dakar to Lake Chad,
S. Sudan and S. Congo), continental SE. Asia
(Ceylon to Hainan), throughout Malesia to
Micronesia and N. Australia (Cape York Penin-
sula and Arnhem Land).

It is difficult to fix the native country of this now
widely spread palaeotropical weed which has no
close relatives. It might have been introduced in
tropical Africa where it is relatively local and con-
fined to the west, but it was collected in Sao Tomé
as early as 1822.

Ecol. A very common weed of pools, along
drains, shallow ditches, water and river edges, in
paddies and humid, fallow garden land, on waste
land, fallow sawahs, and in clearings, both on clay
and humid white sand (Borneo), in pools in Euca-
lypt savannah (Wetar I.), on Mt Kelud (Central
Java) as a pioneer on old volcanic mudstreams
(lahars), from the plains up to c. 1000 m. Fi. Jan.-
Dec;

BACKER (Trop. Natuur 2, 1913, 133) described in
detail the biology of this species, of which the
flowers open at 7 a.m. Seeds are gradually released
by decay of the pericarp of which the vessels remain
persistent. To his surprise both kinds of seed were
buoyant for 16 days, after which they sank. On the
17th day he observed one buoyant seedling, but
unfortunately no mention was made from which
kind of seed; his experiment was through inter-
ference discontinued. He made also notes on the
aerophores emitted by the shallow roots as occur
also in L. peruviana and other species.

DOCTERS VAN LEEUWEN (Zoocecidia, 1926, 428;
Ned. Kruidk. Arch. 51, 1941, 204) reported galled
fruits caused by beetles.

Leaves are often attacked by a blue beetle:
Graptodera cyanea (BACKER, Trop. Natuur 2, 1913,
132).

Uses. HEYNE (Nutt. Pl. 1927, 1206) noted that
in N. Celebes it is used for poulticing pimples.
According to BuRKILL (Dict. 1935, 1273) it is in
Malaya generally stocked by Chinese herbalists,
but its use is not clear; it was once recorded that
an infusion of the root is swallowed by Malays for
syphilis. In the Philippines the whole plant is used
for black dye (QUISUMBING, Medic. Pl. Philip.
1951, 676).

Vern. Sumatra: méligai, M, Banka; Java:
(djukut) anggéréman, mainang, tjatjabéan, S;
Philippines: pasau-na-hapai, sila sila, Tag., ilam-
num-wiliyan, Mag., kakaggin diloba, If., barigaud,
Bik., manakatud, Ik., talang-duron, Pamp.,
tohod-téhod, Bik.; Celebes: kayu ragi, Manado.

Note. The dimorphous seeds are very unusual
and it would be most interesting to have informa-
tion on the properties of the two seed types with
respect to germination.

6. Ludwigia adscendens (L.) Hara, J. Jap. Bot. 28
(1953) 290; A. & R. FERNANDES, Garcia de Orta 5
(1957) 475; RAVEN, Reinwardtia 6 (1963) 387, maps
31, 33; Aston, Aquat. Pl. Austr. (1973) 142; HENTy
& PRITCHARD, Bot. Div. Lae, Bot. Bull. 7 (1973)
130, fig. — Jussiaea repens LINNE, Sp. Pl. 1 (1753)
388, non L. repens Forst. 1771; F.v.M. Descr. Not.
Pap. Pl. 4 (1876) 60; CLARKE, FI. Br. Ind. 2 (1879)
587; O. K. Rev. Gen. Pl. 1 (1891) 251, incl. var.
pilosa O. K. et var. glaberrima O. K.; TRIMEN, FI.

ONAGRACEAE (Raven) 105

1977]

(

u

N

a i TN
SV LZ a

Y] AS 7 f
any
"TS

J

re

iN

x "/, (a after

Fig. 4. Ludwigia hyssopifolia (G. Don) Exe . a. Habit, in flower, b. withered fruits, both

Backer, 1940, b VerBoom 9).

106

Ceyl. 2 (1894) 233; Koorp. Exk. Fl. Java 2 (1912)
703; BACK. Trop. Natuur 3 (1914) 56, 60, f. 1-5;
RIDL. Fl. Mal. Pen. 1 (1922) 827; GAGNn. Fl. Gén.
I.-C. 2 (1925) 987; Hocur. Candollea 3 (1925) 479,
incl. ssp. glabrata Hassv. f. albiflora Hocnr.;
HARTSEMA, Flora (Allg. Bot. Z). n.s. 22 (1927) 242,
t. 3; Back. Onkr. Suiker. (1930) 469, Atlas, t. 443;
STEEN. Arch. Hydrobiol. Suppl. 10 (1932) 314,
f. 62; Fl. Sch. Indon. (1949) 305; Back. & BAKH.
f. Fl. Java 1 (1963) 260. — Jussiaea adscendens
LinnE, Mantissa 1 (1767) 69. — Jussiaea fluviatilis
BL. Bijdr. (1826) 1132. — Fig. 5.

Herb with prostrate or ascending stems, rooting
at the nodes, with conspicuous white, erect,
spindle-shaped, mucronate aerophores arising in
clusters at the nodes of the floating stems and from
the roots, the more or less erect stems to 60 cm;
floating stems to 4m; plants normally glabrous,
but the branches growing on dry ground densely

FLORA MALESIANA

[ser. I, vol. 8?

villous and rarely flowering. Leaves broadly
oblong-elliptical, 0.4-7 by 0.7-4cm, narrowly
cuneate at base, apex acute or obtuse; nerves 6-13
pairs; submarginal vein not prominent; petioles
long. Flowers in upper leaf axils. Bracteoles
present near base of capsule, deltoid, c. 1.2 by 1.3-
1.5mm. Sepals 5, deltoid-acuminate, 5-11 by
2-3.2 mm, glabrous or villous. Petals creamy white,
yellow at base, obovate, rounded at apex, 9-18 by
6-10 mm. Stamens 10, epipetalous ones slightly
shorter; filaments white, 2.5-4mm; anthers
1.2-1.8mm long, apparently shedding pollen
directly on the stigma at anthesis. Pollen grains
shed singly. Disk + elevated, with a depressed
white-hairy nectary surrounding the base of each
epipetalous stamen. Style white, 4-8 mm, densely
long-hairy in lower half; stigma globose, green,
1.5-2 mm @, 1-1.2 mm high, the upper 2/; recep-
tive. Capsule glabrous or villous, 1.2-2.7 cm by

Che: “a

y
i

SANY
FD
>
Zan

Fig. 5. Ludwigia adscendens (L.) HARA, X 2/; (after BACKER, 1940).

1977]

ONAGRACEAE (Raven)

107

344 mn, light brown, with 10 conspicuous darker
brown ribs, terete, the seeds evident between the
ribs as bumps c. 1.5mm apart; capsule thick-
walled, very tardily and irregularly dehiscent;
pedicel 2.5—5.5 cm. Seeds uniseriate in each cell of
the capsule, pale brown, 1.1-1.3mm long, +
vertical, firmly embedded in coherent cubes of
woody endocarp 1.2—1.5 mm high, 1—-1.2 mm thick,
the endocarp firmly fused to the capsule wall.

Gametic chromosome numbers, 7 = 8, 16.

Distr. Continental Asia (from Ceylon to S.
China), throughout Malesia, in Australia one
locality in W. Arnhem Land. Fig. 6.

Ecol. Freshwater pools, ditches, swamps, fallow
and wet paddies, very common, from the lowland
up to c. 1600 m. Fi. Jan.—Dec.

Fig. 6. Range of Ludwigia adscendens (L.) HARA
(after RAVEN, 1963).

BACKER (Trop. Natuur 3, 1914, 56) depicted and
described in detail the biology. The root system
consists of three kinds, long + unbranched anchor
roots, shorter much-branched feeding roots, and
the erect spongy aerophores. BACKER cut the latter,
but the plant remained (only very slightly less)
buoyant. After pollination the pedicel bends down
and the fruit ripens in the water (as in several other
aquatics); the fruit decays and releases the cork-
winged seeds which are buoyant.

On desiccated muddy soils a never-flowering
terrestrial form often occurs, marked by very small
crowded leaves and a stronger pubescence.

Uses. Malays in Perak use it for poulticing inskin
complaints (BURKILL, Dict. 1935, 1273). Batak
people use this (and other aquatics, like also do the
Chinese) to feed pigs, and it is recorded to be eaten
as salad in Indo-China. QUISUMBING (Medic. Pl.
Philip. 1951, 677) reported that it is used in a
decoction as an astringent for dysentery.

Vern. Water primrose, E; Sumatra: buang
buang, Toba-Batak; Java: pangeor, M, ganggéng
landeuj, kambang peutjit, ruba silah, (rumput)
kologa, S, krangkong, kréma, patjar banju, tapak
doro, J; Philippines: sigang-dagat, Tag., gabi-gabi,
Mag., tabagan, If.; New Guinea: agidahano,
Kutubu lang.

Note. L. adscendens appears to be allied more
closely to the American L. helminthorrhiza (MART.)
Hara than to any Old World species. Together
with the mostly yellow-flowered African L.
Stolonifera (GUILL. & PERR.) RAVEN, these three
are the only species that regularly produce clusters
of erect inflated aerophores at the floating nodes,
although other species have descending root-like
aerophores at these nodes and may have long,
spongy aerophores from the submerged under-
ground parts.

Excluded or Doubtful
ae i tenella BurM. f. Fl. Ind. (1768) 103, t. 34,
2.

There is no unanimity of opinion about the
identity of BURMAN’s plant which he said to have
come from Java. MERRILL (Philip. J. Sc. 19, 1921,
369) suggested it to be a form of L. octovalvis which
I doubt in view of the specific epithet and the
species with which it is being contrasted. ALSTON (in
Trimen, Fl. Ceyl. 6, 1931, 130) took it up for
L. hyssopifolia, and this suggestion seems the most
plausible. From BuRMAN’s description, which
contains almost certainly errors, it cannot be
identified. Unfortunately, or perhaps fortunately,
the type at G could not be found.

Ludwigia erigata LINNE, Mantissa 1 (1767) 40. —
L. triflora Desr. in Lamk, Encycl. 3 (1792) 615,
nom. illeg. subs., belongs to the Rubiaceae.

Ludwigia trifolia Buro. f. Fl. Ind. (1768) 36;
Houtt. Nat. Hist. 2, 7 (1777) 344 is according to
MERRILL (J. Arn. Arb. 19, 1939, 368) Oldenlandia
biflora L. (Rubiaceae).

2. EPILOBIUM

Linné£, Gen. Pl. ed. 5 (1754) 164; Sp. Pl. 1 (1753) 347; HAUSSKNECHT, Monogr.
Epilob. (1884); RAVEN, Bull. Br. Mus. Nat. Hist. Bot. 2 (1962) 325, 13 maps, pl. 33-
39; Brockig, New Zeal. J. Bot. 4 (1966) 366, 2 fig.; Bothalia 9 (1967) 309, 7 fig.;
RAVEN, Blumea 15 (1967) 269, 7 fig.; BRockir, New Zeal. J. Bot. 8 (1970) 94;
Raven, D.S.I.R. New Zeal. Bull. 216 (1976) 321 pp., 158 fig. — Fig. 7.

Perennial herbs, often flowering in the first year, occasionally somewhat woody
near the base. Leaves (in Mal.) opposite below, spirally arranged above. Flowers in
the axils of the often greatly reduced upper leaves. Floral tube short (in Mal.) or
essentially absent. Sepals 4, erect. Petals 4, white, pink, or purple, emarginate

108 FLORA MALESIANA [ser. I, vol. 8?
en

VII\ VI BASS
PY) A Ha VO aw. .
“s

7. SVE
(Pu ere we

Oka LL SSeS

AUVN
\- }

Via

PX

Fig. 7. Epilobium hooglandii RAVEN. a. Habit, nat. size, b. node, x 10, c. seed with base of coma, x 30. —
E. detznerianum SCHLTR ex Diets. d. Node, x 10. — E. prostratum Wars. e. Seed, x 30 (a—c HOOGLAND
& PULLEN 5540, d HOOGLAND & PULLEN 5687, e WOMERSLEY c.s. 6103). After Raven, 1967.

1977] ONAGRACEAE (Raven) 109

(in Mal.). Stamens 8, in 2 whorls, the epipetalous ones shorter. Stigma (in Mal.)
clavate or rarely globose, usually surrounded by the shedding anthers at maturity.
Ovary 4-locular, the ovules very numerous. Fruit a long, slender, loculicidal cap-
sule. Seeds very numerous, small, with a chalazal plume of trichomes (coma).

Distr. About 200 spp., well-represented in temperate regions, mostly on the northern hemisphere, with
the greatest centre of morphological diversity in the western U.S.A., at relatively high latitudes and alti-
tudes; in the tropics confined to the mountains; in Malesia: rare and local, in W. Central Sumatra, E. Java
(Mt Tengger), Lesser Sunda Is. (Lombok, Sumbawa, Timor), Philippines (N. Luzon, Panay), SW.
Central Celebes (Latimodjong Range), Moluccas (Central Ceram), and New Guinea.

Ecol. Characteristic of open, disturbed habitats or grassland or the alpine zone, not normally found in
primary forest.

Dispersal takes easily place by wind as the light seeds are provided with a silky coma.

It is still rather surprising that, whereas open habitats are very common in the Malesian mountains,
with their numerous volcanoes and alpine habitats, the number of localities is so very restricted. It must
probably be considered that though dispersal may be easy, the coma only acts efficiently in dry air and this
is in the tropics with their frequent rain and cloud formation and nightly fogs during most of the year a
rather rare climatic situation. In this respect the west-monsoon, blowing from Asia southeastwards (Nov.—
March) is most unsuitable, as these winds are laden with moisture. The east-monsoon, blowing from
Australia northeastwards (May-Sept.) is in this respect definitely more favourable as this is a dry wind.
This wind regime is a consequence of the situation of the Asian and Australian continents.

Local dispersal by water might also occur, as the lowest localities in New Guinea are all along stream-
sides, but this may also be due to the downward air-stream over rushing water.

For hybridization see under the family.

Genesis. The species occurring in Sumatra (1 sp.) and in Luzon (2 spp.) also occur in continental SE.
Asia, the one in E. Java and the Lesser Sunda Is. (E. hirtigerum) also occurs in Australia, Tasmania, and
New Zealand. The four remaining species are all endemic in East Malesia, 3 being confined to New Guinea
and 1 also occurring in Central Celebes and Ceram. They are closely related to species found in Australia
and New Zealand, from which they, and E. hirtigerum, were probably derived via the east-monsoon dis-
cussed above.

I have argued that the Australasian species ultimately show affinity to those of continental Asia and
their ancestors must have reached Australia across the tropical mountains of Malesia. My assumption is
that this southeastward penetration of the genus occurred in the Pliocene.

KEY TO THE SPECIES

1. Buds and flowers erect, the inflorescence erect or somewhat drooping.
2. Stems pubescent only along conspicuously elevated lines running down from the margins of the
DEES. Meee GLY, fe Ow, E. wallichianum
2. Stems pubescent all around.
3. Upper leaves strongly reduced. Inflorescence slightly nodding (Luzon).
ween waves ovate, 0:4-1;5. cnt wide. i) 22. arte ie Lee ea oe 1. E. brevifolium
4, Leaves very narrowly elliptic to linear, 0.1-O.5Scm wide... .... . 2. E. platystigmatosum
3. Upper leaves not notably reduced. Inflorescence erect (Java, Lesser Sunda Is., New Guinea).
5. Plants finely glandular-pubescent; leaves usually ericoid in appearance; petals steerer ae ;
. E. keysser

og Maenwe? se | ot bhp We Sa Se el Kae Oe Bee ee Ga

5. Plants densely strigulose; leaves broader, not ericoid; petals white or very pale pink

4. E. hirtigerum
1. Buds and flowers drooping, becoming erect in fruit.

6. Stems with elevated glabrous lines running from the center of each petiole and elevated pubescent lines
from their angles; petals 7-14 mm long 6. E. detznerianum

6. Stems without elevated lines, uniformly pubescent all around; petals 2.5-8.5 mm long.
7. Petals 6-8.5 mm long; seeds 1-1.4mmlong. .......... 7. E. hooglandii
7. Petals 2.5—5(-6) mm long; seeds 0.7-0.9mm long. . . . . 2-1 ee eee 8. E. prostratum

efi tol .t? g

1. Epilobium brevifolium D. Don, Prod. Fl. Nepal.
(1825) 222; RAveN, Bull. Br. Mus, Nat. Hist. Bot. 2
(1962) 361.

ssp. trichoneurum (HAUSSKNECHT) RAveN, Bull.
Br. Mus. Nat. Hist. Bot. 2 (1962) 362. — E. tricho-
neurum HAUSSKNECHT, Oest. Bot. Z. 29 (1879) 54;
Monogr. Epilob. (1884) 208; H. LéveiLvé, Ic. Gen.
Epilob. (1910) t. 84. — E. philippinense C.B. Ros.
Philip. J. Sc. 3 (1908) Bot. 209; Merr. En. Philip. 3
(1923) 220, excl. syn.

Robust, perennial herb, 15-40 cm, the under-
ground parts not scaly; plants strigulose, stems
pubescent all around. Leaves mostly opposite,
alternate in and near the inflorescence, the upper
ones reduced, strigulose along the veins and
margins especially below, the nerves evident, ovate,
1-3 by 0.4-1.5cm, serrate; petiole 1-2 mm. In-
florescence slightly nodding. Flowers erect, borne
in the axils of reduced upper leaves. Floral tube
1.82.4 mm across, |-1.3 mm deep. Sepals 2.5-5.5
by 1-1.3 mm. Petals obovate, 4.5-8 by 3-5.5 mm,

110

rose purple, the notch 1-1.5 mm deep. Anthers
1-1.2 mm long; filaments of the longer stamens
1.2-2.5 mm, those of the shorter 1-1.5 mm. Style
2.5-4 mm. Stigma clavate, 1.8-2.3 mm high, c.
1 mm thick, surrounded by the anthers at anthesis.
Capsule 3.5-7cm long, on a pedicel 0-1 cm.
Seeds 0.9-1.2 by 0.4-0.5 mm, papillose, blackish
brown, not beaked, obovoid, blunt at both ends,
the coma 5—7 mm long, white.

Gametic chromosome number, n = 18.

Distr. Continental SE. Asia (SE. Tibet:
Chumbi Valley, to Assam, Burma, and W. China);
in Malesia: Philippines (Mountain Province of
N. Luzon).

Ecol. Wet open slopes in the pine region, along
streams and by springs, 1400-2100 m. Fi. July, Oct.

Note. E. brevifolium ssp. trichoneurum is one of
three subspecies of a species which ranges from
Himachal Pradesh in the Western Himalaya east-
wards throughout the Himalaya and southern
China to Formosa, northern Luzon, northern
Vietnam, and northern Burma. In northern Luzon,
it occurs together with the other species found in
the Philippines, E. platystigmatosum, and one plant
of the collection CLEMENS 16385, suggests hybridi-
zation between these two entities, which are usually
widely distinct morphologically, although doubt-
less more closely related to one another than to
other Malesian species.

2. Epilobium platystigmatosum C. B. ROBINSON,
Philip. J. Sc. 3 (1908) Bot. 210; MerrR. En. Philip. 3
(1923) 221; RAVEN, Blumea 15 (1967) 272. —
E. cephalostigma var. linearifolium HisauT1, J. Jap.
Bot. 14 (1938) 143, f. 3. — E. formosanum MaAsum.
Trans. Nat. Hist. Soc. Formosa 29 (1939) 62;
OnwIl, Fl. Japan (1965) 657. — E. sohayakiense
Korpz. Act. Phytotax. Geobot. 8 (1939) 61.

Slender, perennial herb, 15—40 cm tall, the under-
ground parts not scaly; plants strigulose, densely so
in the inflorescence, the stems pubescent all around.
Leaves mostly opposite, alternate in and near the
inflorescence, the upper ones reduced, strigulose
along the veins and margins especially below, the
nerves evident, very narrowly elliptic or linear,
1-3(-4) by 0.1-0.5 cm, weakly and sparsely ser-
rulate; petiole 1-4mm. Inflorescence slightly
nodding. Flowers erect, borne in the axils of reduced
upper leaves. Floral tube c. 1 mm across, c. 0.8 mm
deep. Sepals 3-4 by 1.2-1.6 mm. Petals narrowly
obovate, 3.3-4.5 by c. 2mm, white or pale pink,
the notch c. 1 mm deep. Anthers 0.2-0.3 mm long;
filaments of the longer stamens c. 1.8 mm, those of
the shorter c. 1.2mm. Style c. 2mm. Stigma
broadly clavate, c. 0.8 mm high, c. 0.4 mm thick,
surrounded by the anthers at anthesis. Capsule
glabrescent, 3-5 cm long, on a pedicel 0-1.8 cm.
Seeds 0.8-1 by 0.3-0.4 mm, papillose, not beaked,
obovoid, blunt at both ends, the coma 4-6 mm
long, white.

Gametic chromosome number, n = 18.

Distr. Japan, China (Hupeh, Kiangsu), For-
mosa, and Malesia: Philippines (N. Luzon:
Benguet Prov.; Panay: BS 31439).

Ecol. Chiefly along small streams and about
cliffs, 1200-2400 m. F/. April-June.

Note. I can find no difference between the species
generally known as E. formosanum and the sup-
posed Philippine endemic populations of E. philip-

FLORA MALESIANA

{[ser. I, vol. 8?

pinense. This species is apparently not common on
the Asian mainland.

3. Epilobium wallichianum HAUSSKNECHT, Oéest.
Bot. Z. 29 (1879) 54; RAVEN, Bull. Br. Mus. Nat.
Hist. Bot. 2 (1962) 365; Blumea 15 (1967) 272. —
E. nepalense HAUSSKNECHT, Oest. Bot. Z. 29 (1879)
53, p.p.; Monogr. Epilob. (1884) 218, p.p.; H.
LEVEILLE, Ic. Gen. Epilob. (1910) t. 120. —
E. duclouxii H. LEvEILLE in Fedde, Rep. 6 (1908)
110; Ic. Gen. Epilob. (1910) t. 144. — E. sarmen-
taceum (non HAUSSKNECHT) BUNNEMEJER, Trop.
Natuur 10 (1921) 57, f. 9.

Erect perennial herb 15-SOcm, from a long
rhizomatous base from which leafy shoots arise;
plants strigulose, more densely so above, with
elevated pubescent lines running down from the
sides of the petioles, stems thick and hollow. Leaves
opposite in lower half of the plant, alternate
above, the margin and nerves densely strigulose,
narrowly ovate or lanceolate, subacute at the apex,
obtuse at the base, sharply and densely serrulate,
1.5—4 by 0.5—1.5 cm; petiole short but distinct, up
to 2 mm. Inflorescence densely strigulose with an
admixture of glandular trichomes, somewhat
nodding in bud. Floral tube c. 2 mm across, c.
0.8mm deep. Sepals c. 5 by c. 2 mm, apiculate.
Petals obovate, c. 8 by 3.5-4 mm, pale violet, the
notch shallow, c. 1 mm deep. Anthers 1.3-1.5 mm
long; filaments of the longer stamens c. 2 mm,
those of the shorter c. 1 mm. Style 3-3.5 mm.
Stigma globose, c. 1.5 mm thick, surrounded by
the anthers at anthesis. Capsule densely strigulose
with an admixture of erect, glandular trichomes,
erect, 5-9.5 cm long, on a pedicel 1-2 cm. Seeds
1.2 by 0.6 mm, coarsely papillose, dark brown, the
ei 6-7 mm long, white, tinged with brown at the

ase.

Distr. Continental SE. Asia (W. Nepal to
Yunnan, south to the Khasya & Naga Hills), in
Malesia: Central W. Sumatra (Mt Kerintji), one
collection.

Ecol. Along river-bank, c. 2000 m.

4. Epilobium hirtigerum A. CUNN. Ann. Mag. Nat.
Hist. 3 (1839) 33; HAUSSKNECHT, Monogr. Epilob.
(1884) 291; H. LEveILLE, Ic. Gen. Epilob. (1910)
t. 18; ALLAN, Fl. New Zeal. 1. (1961) 279; Bur-
BIDGE & Gray, Fl. A.C.T. (1970) 276; WiL.is
Handb. Pl. Vict. 2 (1972) 464; RaveN, D.S.I.R.
New Zeal. Bull. 216 (1976) 141, f. 63-64, 65-66
(maps). — E. brasiliense HAUSSKNECHT, Oéest.
Bot. Z. 29 (1879) 119. — E. sarmentaceum (non
HAUSSKNECHT) Koorp. Exk. FI. Java 2 (1912) 704;
Back. & BAKH. f. FI. Java 1 (1963) 262. —
E. cinereum (non A. RicH.) RAVEN, Blumea 15
(1967) 273, pro specim. mal.

Robust, erect, clumped perennial herb, 15-100
cm, the underground parts not scaly; plants densely
strigulose, the stems pubescent all around. Lower-
most /eaves opposite, the rest alternate, densely
strigulose, especially below and along the veins,
nerves evident, narrowly lanceolate, 0.6-3 by
0.2-0.5 cm, coarsely serrate, subsessile. Inflores-
cence erect. Flowers erect, borne in the axils of
upper leaves. Floral tube c. 1 mm across, c. 1 mm
deep. Sepals 3—5.5 by 0.8-1.6 mm. Petals obovate,
3-5 by 2-3.2 mm, white or very pale pink, the
notch c. 1mm deep. Anthers 0.7-1 mm long;

1977]

ONAGRACEAE (Raven)

111

filaments of the longer stamens 2-2.8 mm, those of
the shorter 1-1.8mm. Style 1-3mm. Stigma
clavate, 1.5-2.7 mm _ high, 1-1.5 mm thick, sur-
rounded by the anthers of anthesis. Capsule
3-5(-6)cm long, on a pedicel 0-2cm. Seeds
0.9-1.2mm long, 0.35-0.45mm thick, finely
papillose, brown, not beaked, obovoid, blunt at
both ends, the coma 5-8 mm long, white.

Gametic chromosome number, 7 = 18.

Distr. Very widely distributed, South America
(Argentina, Uruguay, Brazil: Santa Catarina),
New Zealand, Australia, and Malesia: Lesser
Sunda Is. (Timor, Sumbawa, Lombok) and East
Java (Mt Tengger).

Ecol. Moist places, grasslands; in E. Java the
only locality is near the single small well on the
otherwise dry and barren extinct volcanic cone of
Mt Widodaren on Mt Tengger caldera, at 2100 m.
In Lombok in Casuarina forest; in Timor in
Eucalypt savannah; 1800-2200 m. F/. Oct.—Jan.

Note. Variable species. The Malesian specimens
belong to a highly autogamous Australian form
with small white flowers which occurs in swampy
places throughout the lowlands of Australia (from
Queensland to temperate W. Australia and Tas-
mania) and is predominant in New Zealand. I have
assumed that it came there from Australia and has
spread again from New Zealand to South America,
either by natural dispersal or by man.

5. Epilobium keysseri DreLs, Bot. Jahrb. 62 (1929)
486; HooGL. Blumea Suppl. 4 (1958) 223; Borc-
MANN, Z. Bot. 52 (1964) 124, 143; RAVEN, Blumea
15 (1967) 274, f. 1 (map).

Clumped erect perennial herb 12-60 cm, often
ericoid in aspect, + woody at the base, the under-
ground parts not scaly; plants finely glandular-
pubescent. Lowermost /eaves opposite, the rest
alternate, coriaceous, subglabrous, the nerves
obscure, margin revolute, narrowly elliptic to
elliptic, 0.5-1.3 by 0.1-0.2(-0.4) cm, with a few
coarse teeth on each side, subsessile. Inflorescence
erect. Flowers erect, borne in axils of upper leaves.
Floral tube 1-2 mm across, 0.7—1 mm deep. Sepals
3-5.6 by 1-2.5 mm. Petals obovate, 4.5—-8 by 2.8-4
mm, purplish rose, the notch 1.5-2mm deep.
Anthers 0.7-1 mm long; filaments of the longer
stamens 1.8—2.5 mm, those of the shorter 0.5-1.5
mm. Style 2.5-4mm. Stigma clavate, 1.5-2 mm
high, 0.5—0.7 mm thick, surrounded by the anthers
at anthesis. Capsule 3-6 cm long, glabrescent, on a
pedicel 1-2.5cm. Seeds 0.9-1.2 by 0.4-0.6 mm,
papillose, brown, with a short pellucid beak, the
coma 5-7 mm long, white.

Gametic chromosome number, 1 = 18.

Distr. Malesia: New Guinea (Mt Wilhelmina
in West, common on many summits in East), many
collections. Fig. 8.

Ecol. Subalpine and alpine meadows and grass-
lands, open shrubberies, swampy treefern grass-
land, occasionally epiphytic on treefern trunks,
forest glades, often in succession after ground-fires,
(1600-)1950—3800 m. F/. (Jan.—)June—Aug.(—Dec.).

Vern. Aingum, Tomba, gonema, Chimbu,
yandepai, Enga lang., Wabag, papal, Enga lang.,
Poio, tamitan, Mendi lang., Giluwe.

Notes. A distinctive species but clearly belong-
ing to the Australasian assemblage.

Most plants, with their narrow leaves, are dis-

Fig. 8. Range of Epilobium keysseri Diets (after
RAVEN, 1967).

tinctly ericoid in appearance, and thus parallel
representatives of many other typically non-ericoid
groups that occur in alpine and subalpine regions
of New Guinea.

6. Epilobium detznerianum SCHLTR ex DIELs, Bot.
Jahrb. 62 (1929) 485; RAVEN, Blumea 15 (1967) 277,
f. 3, 5 (map). — E. papuanum RIDL. var. alpestre
RiDL. Trans. Linn. Soc. Bot. II, 9 (1916) 58. —
E. papuanum (non RID.) HooGL. Blumea Suppl. 4
(1958) 228. — Fig. 7d.

Clumped perennial herb 3-15 cm, the under-
ground stems not scaly; plants mostly glabrous,
with elevated, strigulose lines running down from
the margins of the petioles and glabrous ridges
running down from back of petioles. Leaves mostly
opposite, alternate in the inflorescence, coriaceous,
nerves obscure, broadly elliptic or ovate, obtuse at
apex and base, entire, 0.3-1 by 0.2-0.7 cm;
petiole 1-1.5mm, short but distinct. Flowers
nodding, the ovaries erect, borne in the axils of
upper leaves. Floral tube 1.5-3 mm across and
about as deep. Sepals 4.5-7 by 1-2.5 mm. Petals
obovate, 7-14 by 3-6 mm, bright purplish rose,
the notch c. 2 mm deep. Anthers 0.8-1.2 mm long;
filaments of the longer stamens 4—5.5 mm, those of
the shorter 3.3-4.5 mm. Style 5.5-8 mm. Stigma
broadly clavate, 1.3-1.7 mm high, c. 1 mm thick,
surrounded by or held just above the anthers at
anthesis. Capsule erect, subglabrous, 4-5 cm long,
on a pedicel 2.5-8.5cm. Seeds (0.9-)1-1.5 by
0.5-0.7 mm, not beaked, finely papillose, pale
brown, the coma c. 8 mm long.

Gametic chromosome number, 7 = 18.

Distr. Malesia: New Guinea (Mts Carstensz &
Wilhelmina in West, Telefomin, Mts Sarawaket &
Wilhelm in East). Fig. 9.

Fig. 9. Range of Epilobium detznerianum SCHLTR
ex Dies (after RAVEN, 1967).

112

FLORA MALESIANA

[ser. I, vol. 8?

Ecol. Subalpine and alpine grasslands and
ridges, open slopes and bogs, earth screes, sub-
alpine forest and its grassy edges, rock clefts, cliff
crevices in alpine thickets, on dripping slate land-
slides, more rarely on sandy or gravelly gully
beds; 3000-4500 m. Fi. (Jan.—Febr.—)May-—Sept.
(—Dec.).

Note. An attractive alpine species reaching the
highest elevations in the genus in New Guinea. It is
usually more condensed in habit than E. hooglandii
and differs at once from that species in the glabrous
ridge decurrent from the center of each petiole and
the elevated pubescent lines decurrent from the
edges of the petiole. It grows sympatrically with
E. hooglandii and E. keysseri, but no intermediates
have been observed; intermediates with the former
species would be very difficult to detect.

7. Epilobium hooglandii RAVEN, Blumea 15 (1976)
278, f. 2,6 (map). — E. pedunculare (non A. CUNN.)
F.v.M. Trans. R. Soc. Vict. I, 2 (1889) 7. — E.
detznerianum (non SCHLTR ex DiELS) HOoc_.
Blumea Suppl. 4 (1958) 228. — Fig. 7a—c.

Caespitose perennial herb with decumbent
branches, the erect portions 10—25(—45) cm long;
plants glandular pubescent along elevated lines
running down from the margins of the petioles
below, more densely and uniformly so above, and
with an admixture of strigulose pubescence in the
inflorescence. Leaves mostly opposite, alternate in
the inflorescence, subcoriaceous, nerves + visible
in dried material, broadly elliptic to ovate, acute or
obtuse at apex and base, entire or with a few teeth
on the margins, 0.5-1.3 by 0.3-0.9 cm; petiole
1-3 mm, short but distinct. Flowers nodding, the
Ovaries erect, borne in the axils of upper leaves.
Floral tube 1.4-2mm across, 1-1.2mm deep.
Sepals 3-4 by 1-1.6 mm. Petals 6-8.5 by 2.5—4.5
mm, pink to purplish rose, the notch c. 1.5 mm
deep. Anthers 0.7-1 mm long; filaments of the
longer stamens 3—6 mm, those of the shorter 2-4
mm. Style 2.5-6 mm. Stigma clavate, 2—2.5 mm
long, 1-1.2 mm thick, surrounded by the anthers
at anthesis. Capsule erect, glabrescent, 5-8 cm long,
on a pedicel 4-12 cm. Seeds 1-1.4 by 0.3-0.45 mm,
finely papillose, brown; coma 5-8 mm long, white.

Gametic chromosome number, n = 18.

Distr. Malesia: New Guinea (Mt Wilhelmina
in West, many localities in East).

Ecol. Subalpine and alpine meadows, fire-
induced alpine grassland, near boulders, in alpine
shrubbery, on peaty grassland, near waterfalls, in
secondary forest on limestone cliffs, on stream-
bank gravel, stony creek beds, along forest tracks,
between grass tussocks in old lake basin, in pen-
dent masses on rocks, occasionally in moist,

eG areas; 2000-4150m. FI. May—Nov.
—Jan.).
Vern. Dirimpia, Chimbu, Masul, nonami,

Mairi, Mondo.
Note. Differs from the closely related E.
detznerianum by its evenly pubescent stems, and

from E. prostratum by large flowers and seeds. It
grows sympatrically with the other three New
Guinean species. Occasional collections are inter-
mediate with E. prostratum and obviously result
from incidental hybridization. However, the two
species keep in general amply distinct and are as
well differentiated as most recognized species of
the genus.

8. Epilobium prostratum Wars. Bot. Jahrb. 16
(1893) 15, 23; RAVEN, Blumea 15 (1967) 280, f. 4,
7 (map). — E. papuanum Rip-. Trans. Linn. Soc.
Bot. II, 9 (1916) 57. — Fig. 7e.

Similar to E. hooglandii, but differing as follows:
habit lax, spreading. Leaves 0.4-0.8 by 0.2-0.4 cm.
Floral tube 0.8-1.1 mm across, 0.6-0.9 mm deep.
Sepals 1.5-3 by 0.6-1.4 mm. Petals 2.5—-5(-6) by
1.3-2.5 mm, very pale pink to purplish rose, the
notch c. 1mm deep. Anthers 0.5-0.7 mm long;
filaments 1.8-3.5 mm. Stigma clavate, 0.8—1.5 mm
long, 0.50.9 mm thick, surrounded by the anthers
at anthesis. Capsule 3.5—-5 cm long, on a pedicel
5-10 cm. Seeds 0.7-0.9 by c. 0.3-0.4 mm, finely
papillose, brown, the coma 5-8 mm long.

Gametic chromosome number, n = 18.

Distr. Malesia: SW. Central Celebes (Latimod-
jong Mts), Moluccas (Ceram), and throughout
New Guinea. Fig. 10.

Fig. 10. Range of Epilobium prostratum WARB.

Ecol. Moist open places, especially on rocky
alluvium bordering streams, stream-banks across
treefern grassland, along river-bank in limestone
scree, on fallen trees on open ridge, rarely in
montane forest, sometimes colonizing landslips;
(1200-)1900-3400 m, in Celebes and Ceram at
c. 2750-3400 m. Fl. Jan.—Dec.

WARBURG’s type specimen was found by HELL-
WIG at an exceptional low altitude of 1200 mina
streambed.

Vern. New Guinea: kimbil, Enga lang., Poio,
dirimpia, Chimbu, Masul, kokorabadi, Mandi lang.

Notes. Closely similar to E. hooglandii, but
readily distinguished for the most part by smaller
flowers and seeds. Occasionally intermediate
specimens (obviously hybrids) are found where the
two occur together.

In New Guinea it descends to lower altitudes and
has correspondingly a much wider range.

Introduced
Fuchsia boliviana CARR. Rev. Hort. (1876) 150; as F. corymbiflora (non R. & P.) BACK. & BAKH. f. FI.

Java 1 (1963) 264.

Locally cultivated and perhaps established in the mountains of West and East Java (Mts Malabar and

Tengger) between 1500 and 2000 m.

1977] ONAGRACEAE (Raven) 113

Easily distinguished from the following species by its drooping inflorescences of bright red flowers with
a floral tube 5-6 (instead of less than 1) cm long.

Fuchsia magellanica LAMK var. gracilis (NICH.) BAILEY: BACK. & BAKH. f. Fl. Java 1 (1963) 264; as
F. coccinea (non SOL. ex AITON) CuRT. — BUNNEMEWER, Trop. Natuur 10 (1921) 56, fig.; Wisse, ibid. 11
(1922) 480, fig.; Hocur. Candollea 2 (1925) 480; BAck. & BAKH. f. Fl. Java 1 (1963) 264.

This species, native of temperate South America, has repeatedly been reported to occur cultivated but
also naturalized in anthropogenous places and on volcanic ash on the mountains of West, Central, and
East Java (Mts Patuha, Malabar, Diéng, Sindoro, and Tengger) between 1000 and 2100 m, and in the
mountains of Sumatra (Karo Lands).

Oenothera stricta LEDEB. ex Link, En. Hort. Berol. 1 (1822) 377, ssp. stricta; as O. erythrosepala (non
BorsAs) BACK. & BAKH. f. Fl. Java 1 (1963) 262. — Oenothera sp. Doct.v.LEEUWEN, Verh. Kon. Ak. Wet.
A’dam II, 31 (1933) 191.

Docrers VAN LEEUWEN /.c. introduced this species from seed he collected in Hawaii and sowed on the
summit of Mt Pangrango in West Java, at 3000 m altitude, in 1921. It is maintaining itself there and,
although self-pollinating, is visited by Bombus rufipes (HEIDE, Dansk Bot. Ark. 5, 1927, 18) and doubtless
by nocturnal insects as well. It is a native of temperate South America, widely cultivated and naturalized.

It may be distinguished from all other species of the family in Malesia by its combination of a long
floral tube and yellow petals; the flowers open at sunset.

Cultivated

Representatives of several genera — Clarkia and Gaura among them — are cultivated, mainly in the
mountains. BACKER & BAKHUIZEN VAN DEN BRINK f., Fl. Java 1 (1963) gave an account of these.

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

Trees, shrubs, lianas, very rarely herbaceous (extra-Mal.); twigs often lenticellate
and nodes with gland fields; spines very rare (extra-Mal.). Stipules absent. Leaves
simple or mostly compound (digitate or impari-1—4-pinnate), (in Mal.) decussate,
rarely in whorls of 3-4, often provided with glands underneath, in the New World
often provided with terminal tendrils, rarely scattered or in pseudo-whorls (extra-
Mal.); domatia sometimes present (fig. 8b, 23h). Inflorescences bracteate, cymose,
but not rarely thyrses contracted to racemiform or racemose inflorescences, or
even reduced to solitary flowers (extra-Mal.), terminal, axillary or from the old
wood. Pedicels mostly with 1-2 bracteoles. Flowers usually very showy, rather large,
bisexual, articulate with the pedicel or not. Calyx connate, closed in bud and later
(not rarely irregularly) splitting into lobes, or cupular, or spathaceous, or lobed
from the beginning and with equal or unequal, valvate lobes, developing earlier
than the corolla, often glandular outside and inside with water and slime producing
glands and hydathodes, persistent or circumscissile caducous along an abscission
line. Corolla sympetalous, campanulate, tubular, funnel- or salver-shaped, mostly
zygomorphic, lobes equal or unequal, valvate or imbricate in bud, tube often with a
narrow cylindrical (constricted) lower part (basal tube) and a widened upper part
(upper tube). Stamens 5 almost equal, or mostly 4 didynamous, the 5th sterile,
rudimentary, adnate to the corolla tube, mostly inserted at the rim of the basal
tube and not rarely (glandular) hairy at the insertion, more rarely inserted higher
up. Anthers basifixed, 2-celled, rarely one cell barren or 1-celled, introrse, dehiscing
lengthwise, usually the anthers connivent in pairs; anther cells often free and diver-
gent, connective not rarely produced. Disk intrastaminal, mostly annular, rarely
absent. Ovary superior, 2-celled, rarely 1- or 4-celled (extra-Mal.); style filiform,
stigma usually 2-lipped, sensitive. Ovules (in Mal.) in each cell on the septum in
two or more rows of 3-04, mostly on 2 placentas. Capsule 2-valved, either loculicid
with the septum perpendicular to the valves — sometimes provided with an addi-
tional transverse false septum — or septicid with the septum parallel with the valves,
or (extra-Mal.) an indehiscent, 1-celled, soft or hard-shelled, pulpy berry. Seeds in
each cell attached to the dissepiment in one or more rows, inserted transverse to
axis of fruit, anatropous, mostly on both sides with hyaline wings; embryo exal-
buminous, the cotyledons mostly notched, sometimes on both sides. Germination
always epigeal. .

Distribution. About 120 genera and some 650 spp., mainly in the tropics and subtropics,
roughly between 40° N and 30-35° S, very few in the warm-temperate zone; in Malesia: 14
native genera of which 2 are endemic, viz Hieris in Penang and Lamiodendron in Papuasia.
Among the remaining 12 one occurs through the Old World (Dolichandrone), 7 are shared with
continental SE. Asia (two of which extend also to Africa and Madagascar: Fernandoa, Stereo-
spermum) and 4 with Australia and Melanesia; the latter occur in Malesia only in the east except
Deplanchea which ranges westward to Sumatra.

In the family tropical Asia and Africa share a few genera (Markhamia, Fernandoa, Stereosper-
mum, and Dolichandrone), but Africa and America share only one, viz Tecoma. This latter
affinity goes further, though very disjunct via Campsidium (Chile) and Campsis (N. America and
E. Asia) to Tecomanthe-Pandorea-Neosepicaea (Moluccas to Three Kings Is. and E. Australia).
Otherwise there appear to be only two other transoceanic ties, viz tribe Crescentieae which is
shared by Africa and the Americas, and the genus Catalpa which occurs in E. Asia and the
Caribbean area.

As GENTRY (Brittonia 25, 1973, 227-230) has shown, the average number of species per genus

(114)

1977] BIGNONIACEAE (van Steenis) 115

is only 5, which is very small in comparison with many other families, but can only partly be
explained by a possibly small generic concept. There are quite a number of monotypic genera
(in Malesia 5), but they are well defined in many characters and stand very apart.

This, and the worldwide distribution of the family, and the disjunctions in ranges, definitely
point to relict survival and ancient origin, onwards of which period the three tribes have under-
gone a separate, independent development on the continents, mainly leading to differentiation in
Indo-Australia and in the New World, with the greatest abundance in the latter. Unfortunately,
the fossil evidence (only Tertiary) is meagre and untrustworthy (SCHUMANN in E. & P. Nat. Pfl.
Fam. 4, 3b, 1894, 208), both to macrofossils and to pollen.

Ecology. Within the family there is a fairly wide coverage of habitats and there are quite a few
which are confined to arid conditions (Rhigozum and Catophractes in S. Africa, Tecomella in
Arabia, Phyllarthron bernierianum in Madagascar, Dolichandrone filiformis, D. heterophylla and
the linear-leaved drought form of Pandorea pandorana in Australia). A few are rheophytic
(Astianthus and Chilopsis in the Americas). A few are warm-temperate (Catalpa) or subtropical-
alpine (U/ncarvillea incl. Amphicome in the Sino-Himalayas and Argylia and Campsidium in the
Andes). One species is bound to the mangrove (Dolichandrone spathacea in Indo-Melanesia).

The majority, however, belongs to the tropical forest, mostly the everwet type, but a fair num-
ber in the seasonal type, below c. 2000 m; only Tecomanthe ascending to c. 3100m in New
Guinea.

In Malesia most are evergreen, but Oroxylum indicum (fig. 5) and Dolichandrone spathacea can
stand leafless in the dry season for many months. Fernandoa macroloba is also deciduous, as are
the species of Stereospermum.

Habit. The majority of the Malesian genera are small or large trees, the only climbing genera
being Nyctocalos, Hieris, Tecomanthe, Pandorea, and Neosepicaea; in subalpine heathland
Tecomanithe may be forced to creep on other vegetation.

Most trees are of medium size, but species of Stereospermum, Fernandoa, Pajanelia, and also
Radermachera gigantea may attain quite good dimensions. Oroxylum is a short-lived nomad tree.

Dominance. Almost all species occur scattered in the forest and several are very rare indeed.
An exception is Dolichandrone spathacea, bound to the swampy, brackish inner mangrove,
which according to CORNER (Wayside Trees, 1940, 164) is in the North of the Malay Peninsula,
in Perlis, a feature of the country, flanking roads and standing as an upright poplar in the rice-
fields. Fig. 16. In Great Natuna I. (NW. off Sarawak) I found Pajanelia longifolia locally very
common in coastal forest, but this was probably encouraged by devastation. In secondary forests,
on earth slides, abandoned fields, and on fresh volcanic ash Radermachera glandulosa and R.
gigantea may be frequent in the pioneer upgrowth in Java (also on Krakatao), but this high
frequency is local; KOoRDERS mentioned it for G. Telemojo and Pringombo in Central Java. Even
Oroxylum indicum which is a nomad plant bound to secondary growths is always found in only
a few specimens. Lamiodendron magnificum was once mentioned by Brass as forming a com-
munity on a gravel bank behind the beach, in Normanby I., but this tree is extremely rare other-
wise.

Flower biology. In many species (like in several Gesneriaceae, Verbenaceae, Solanaceae) the
calyx develops much earlier than the corolla and is closed in bud. Inside of the calyx with water
and slime producing glands and hydathodes in which the corolla develops. These so-called
waterbuds are very characteristic, especially in such large-flowered species as Spathodea campanu-
lata, the tulip tree, which derives a Dutch and Malay name from this feature (spuitjesboom,
panchot) which is enjoyed by children to play with. It is one of the few biological phenomena
which are entirely confined to the tropics, as far as I am aware.

Treus (Ann. Jard. Bot. Btzg 8, 1889, 38-46, t. 13-15) made an anatomical study of the glands
inside the calyx of Spathodea campanulata and on his instigation GREsHOFF examined the exudate,
dissolved organic and inorganic substances, which appeared similar to those of leaf-hydathodes.
Koorpers (ibid. 14, 1897, 354-469, t. 21-27) extended this subject with research on some other
genera of Bignoniaceae (Parmentiera, Kigelia, Crescentia, Fernandoa adenophylla, Radermachera
gigantea) and some other plants, confirming TrReus’s results.

Pollination. Flower-shape, -colour, -position, and -scent are very different in the mostly showy
flowers of Bignoniaceae, and the syndromes attract different visitors.

116 FLORA MALESIANA (ser. I, vol. 8?

Bats are frequently visiting species of certain genera, another phenomenon restricted to tropi-
cal plants. According to FAEGRI & VAN DER PuL (Principles Pollination Ecology, ed. 2, 1971,
154) the attraction syndrome is: nocturnal anthesis, whitish, creamish or drab greenish or dark
purple colour, stale or sour, unpleasant smell reminiscent of fermentation at night, large quantity
of nectar and pollen in large anthers, large-mouthed and coarse flowers on strong stalks sticking
out of the foliage or cauliflorous to flagelliflorous flowers, thus coming into easy reach for landing.
This is found in Malesia in several cultivated genera (Kigelia, Crescentia, Parmentiera, Mark-
hamia, etc.) but occurs also in the native Fernandoa adenophylla, Pajanelia, and Oroxylum.
Fig. 7. Notwithstanding the many papers and records of observation — corollas show claw marks
after these visits — it is not proved to my satisfaction that visits of bats are compulsory for polli-
nation cq. fertilisation, experimenting being in this field deplorably meagre. My doubt is streng-
thened by observations by HARRIS & BAKER in Ghana where Kigelia is native (J. West Afr. Sc.
Assoc. 4, 1958, 28) and can set fruit in absence of bats; they observed also frequent visits by
sphingids but they doubt effective pollination by these.

Birds, humming birds and sun-birds, frequently visit certain species, the attraction syndrome
being: tubed, vividly coloured (orange, scarlet), diurnal, mostly odorless, nectar-producing
tubular flowers. Here also many observations are made, e.g. in Tecoma (Tecomaria). To this class
belong in Malesia some species of Radermachera (R. ramiflora), Neosepicaea, Tecomanthe, and it
can be expected for Deplanchea. Also the cultivated Spathodea campanulata is frequented by
birds (cf. BEUMEE, Trop. Natuur 14, 1925, 28-30), notably kutilans and ?djalaks, at Bogor; they
severely damage the corolla. Here again the question whether bird-visits are compulsory for
pollination cg. fertilisation is inadequately supported by experiments. Caution is necessary to
conclude to the necessity of cross-pollination, as e.g. HUNTER (Rec. Auckl. Inst. Mus. 6, 1967,
169-170, t. 24) recorded that in Tecomanthe speciosa, of which cuttings of a single plant led to its
cultivation, fertilisation — that is selfing — could be effected by hand-pollination, but later also
naturally by bees, although far from its native habitat.

Moths. A few species have the moth-attraction syndrome for flower visitors, which implies:
nocturnal, very fragrant flowers with abundant honey, in mostly pale or white, long-tubed or
salver-shaped corollas. Fig. 1, 8, 15. This is found in species of Nyctocalos, Dolichandrone, Hieris,
Millingtonia, and some species of Radermachera (e.g. the Chinese R. sinica, R. pentandra, and
R. peninsularis). Probably long-tongued sphingids (hawk-moths) visit these flowers.

Bees and butterflies. Possibly bees visit flowers of species not belonging to the three categories
mentioned above.

The ecological role of the many sorts of extra-floral nectaries in unexplained.

It is a fact that in general fruit setting is scarce in Bignoniaceae; in several the fruit was only
occasionally found long after the plant had been described in flower. In Malesia the fruit is still
unknown of Hieris and Lamiodendron. Even after abundant flowering fruit production is often
very low with 1 or 2 fruits in each inflorescence, except in Radermachera glandulosa and Tecoma
stans. With all these flower visit devices one would expect otherwise.

Dispersal. Bignoniaceae occur throughout the tropics and several are still found in the sub-
tropics of the whole world. One might ascribe this to their having winged seed (except Crescentieae
and a few other exceptions), but against expectations they are almost absent from oceanic
islands. Bignoniaceae occur all along the coasts of the West Pacific, notably in New Guinea and
in Australia species of Tecomanthe and Pandorea are not rare, but the only occurrence in the West
Pacific islands is a common Australian Pandorea in New Caledonia and Lord Howe lI., anda
peculiar Tecomanthe in a single locality of the Three Kings Is., the northernmost territory of New
Zealand.

Obviously wind dispersal has not been as effective as one would expect.

Dispersal by seawater is common in Dolichandrone spathacea, a back-mangrove species,
ranging from the western Deccan Peninsula to North Luzon, south to Timor and southeastwards
to New Caledonia; the range is almost continuous without gaps. Fig. 14. It is most peculiar,
however, that so far it has never been found in the mangroves of northern Australia. Its seeds have
thickish corky wings instead of flimsy wings as usual in most members of the family (except the
fleshy indehiscent fruits of the Crescentieae and a few other exceptions as e.g. Pauldopia) and are
most excellently adapted to be dispersed by seawater.

1977] BIGNONIACEAE (van Steenis) 117

Seedlings. These are very uniform in all tribes of the family; in the embryo the foliaceous
cotyledons are flat in one plane, mostly emarginate at both ends, hypocotyle and rootlet are
small. Germination is epigeal by stretching of the hypocotyle. The first leaves are mostly simple,
as in most compound-leaved families; they are often dentate. In a very few genera with thick seeds
there may develop — possibly by intrusion of the testa — a false septum in the seed, in Malesia
notably in Stereospermum and here also the cotyledons are folded. The only exception is men-
tioned by LusBock (Contr. Knowl. Seedlings 2, 1892, 334, fig. 569) for ‘“Bignonia insignis’ with
fleshy connate cotyledons and hypogeal germination; the name is evasive and at Kew the identity
could not be traced; presumably the record rests on an error.

Literature: E. BureEAu, Monogr. Bign. (1864); J. LusBock, Contr. Knowl. Seedlings 2 (1892)
332-345, fig. 569-575; R. S. Troup, Silvic. Indian Trees 2 (1921) 684-693; J. A. Duke, Ann. Mo.
Bot. Gard. 52 (1965) 349, pl. 20; G. DE LA MENSBRUGE, La germination et les plantules essences
arbres Céte d'Ivoire; Techn. For. Trop. Nogent-sur-Marne, Paris (1966) 332-333; D. BURGER,
Seedlings (1972) 52-54; C. S. SCHOPMEIJER, Seeds Woody PI. U.S., Agric. Handb. 450 (1974) 260,
281, 321.

Juvenile plants of Pandorea pandorana show leaves very different from the mature foliage, in
having many jugae and being coarsely dentate. Tecoma filicifolia NICHOLS. was based on such
material. This led also to a serious misinterpretation of Tecoma leptophylla Bu., from New
Guinea, of which the juvenile leaves (fig. 37b) are Pandorea pandorana but the flowers belong to
Neosepicaea.

Taxonomy. Since the basic work on the systematy by E. BuREAU (Monographie, 1864), the
treatment of the family in Flora Brasiliensis by BUREAU & SCHUMANN (1896-97), and the treat-
ment of SCHUMANN in the Pflanzenfamilien (1895) the traditional subdivision of the family in 5
tribes has proved satisfactory. Crescentieae with 1-celled berries occur in Africa and the Americas,
two other monogeneric tribes are South American, while the bulk of the family belongs to Big-
nonieae and Tecomeae, of which the latter are about balanced as to number of genera in the Old
and New World, but Bignonieae are predominantly American. These two tribes are largely dis-
tinguished on the dehiscence of the fruit, loculicid in Bignonieae and septicid in Tecomeae.

In passing it may be remarked that Gentry (PI. Syst. Evol. p. 126, 255) recently advocated that
Crescentieae of the neo-tropics and of Africa-Madagascar are of separate descent and would
represent two parallel evolutionary lineages; this suggestion is more based on geographic argu-
ment and evolutionary hypotheses than on morphological arguments.

The delimitation against other families of Sympetalae is well-defined, but there are a few genera,
notably Wightia and Paulownia, which are sometimes referred to Bignoniaceae, though FENZL
(Denkschr. K. Bay. Bot. Ges. Regensburg 3, 1841, 227-230), BUREAU, SCHUMANN, VON WETT-
STEIN, and other specialists referred them to Scrophulariaceae. A survey of opinions I gave in my
paper on Wightia (Bull. Bot. Gard. Btzg III, 18, 1949, 214-216), in which I excluded it from
Bignoniaceae. Even recently Paulownia is sometimes casually treated as Bignoniaceous (e.g.
SCHOPMEVER, Agric. Handb. U.S.A. 450, 1974, 527), although the embryo is embedded in
endosperm; furthermore the stigma is different from that in Bignoniaceae, the anthers have no
prolonged connective, there is no rudimentary stamen and the seeds are provided with several
wings and seem to be laterally attached, not transverse as in Bignoniaceae. For Wightia I tabulated
(1.c.) the relation to both families. Its seeds have no endosperm, but the absence of a staminode,
the structure of the stigma, the central placenta and the absence of a produced connective on the
anthers point distinctly to Scrophulariaceae. The seed is quite differently attached as compared
with Bignoniaceae, viz laterally and the wing surrounds the entire seed. Its wood has two kinds
of medullary rays, narrow and broad ones, a character which, at least in Malesian Bignoniaceae,
is absent.

Though the capsule in Wightia is septicid and in Paulownia loculicid, both genera have the
same kind of axile placentation, in which the thickened placenta becomes detached from the
valves as a subquadrangular seed-cake, showing their close affinity, completely differing from the
situation in Bignoniaceae.

According to SURYAKANTA (J. Palyn. 9, 1973, 73) the pollen of both genera differs from that in
Bignoniaceae and resembles that of Scrophulariaceae.

NAKAI (J. Jap. Bot. 24, 1949, 13) accommodated Paulownia in Paulowniaceae, probably in-

118 FLORA MALESIANA [ser. I, vol. 8?

duced mostly by its arboreous habit and fruit; they certainly merit to be placed in a separate
tribe or subtribe of Scrophulariaceae. We regard nowadays the arboreous habit as primitive in
herbaceous families and we might conclude that they are ancient relicts from a period when
Bignoniaceae and Scrophulariaceae had a common matrix.

Also in South America there are two woody genera of the Scrophulariaceae which were at
times referred to Bignoniaceae, viz Schlegelia (syn. Dermatocalyx) and Gibsoniothamnus, accord-
ing to GENTRY (Fieldiana, Bot. 34, 1971, 55; Ann. Mo. Bot. Gard. 61, 1974, 533-537); see also
LEINFELLNER (Oest. Bot. Z. 121, 1973, 13-22). They are (hemi-?) epiphytic shrubs or lianas, a
similar habit as in Wightia.

Genetics. Chromosomes. DARLINGTON & WyLIiE (Chrom. Atlas, 1955) and Moore (ed.)
(Regn. Veg. 90, 1973) gave for 26 genera x = 20 (2n = 40) and they belong to Tecomeae,
Bignonieae and Crescentieae, both from the palaeo- or neotropics. There is one higher number
x = 22 (Amphilophium, South America, Niedzwedzkia = Incarvillea) and several lower ones:
Pandorea, and some doubtful countings in Tecoma x = 19, Tecomanthe dendrophila 2n = 36
(Christine BRIGHTON in litt.), Jacaranda x = 18, Tecoma capensis x = 17, Oroxylum, Milling-
tonia, Argylia (from South America) x = 15, Spathodea x = 13, and Incarvillea x = 11. In
supplement indices Campsis is also given as 16 and Oroxylum as 14.

I have scanned the numbers of Scrophulariaceae, Gesneriaceae and Verbenaceae, but can find
no reliable ties, Bignoniaceae being obviously more homogeneous than those.

The number given for Paulownia, 2n = 40, x = 10, might as well fit Bignoniaceae as Scrophu-
lariaceae.

Hybridisation. Not many species hybrids are known to me, but those known are interesting,
as there are at least two between species of East Asia and SE. North America which are now very
disjunct after the Pleistocene Ice Age; it is not impossible that they formed part of more con-
tinuous populations in the warmer Pliocene via Beringia. This idea is supported by the fact that
in both cases the hybrids are fertile.

E. C. SmitH (J. Arn. Arb. 22, 1941, 219) reported on Catalpa ovata DON x C. bignonioides
WALT. (= x C. syringifolia Sims). Haploid all have 20 chromosomes (SAx, J. Arn. Arb. 14, 1933,
274).

Then there is x Campsis tagliabuana (VIVIAN1) REHDER, a hybrid between the Chinese C.
grandiflora (THUNB.) K. Scu. (C. chinensis (LAMK) Voss.) and C. radicans (L.) SEEM. which pro-
duces fertile progeny (cf. STEARN, Bot. Mag. 169, 1953, t. 198).

The third one is also bi-continental, Tecoma smithii W. WATSON (Gard. Chron. 14, 1893, 649,
fig. 104; E. Smitu, ibid. 16, 1894, 64; cf. also Gartenflora 44, 1895, 51, fig. 14). This is a reputed
hybrid, which E. SMITH made at Adelaide, in 1882, between 7. velutina (a hairy variety of T. stans)
and T. capensis. It was propagated by cuttings, but it produced seed and its offspring of seedlings
diverged in size and flower colour. Curiously SPRAGUE, in a succinct note (Fl. Cap. 4, 2, 1904,
448) reduced it to T. alata DC., without referring to its hybrid nature.

Anatomy. Wood. Of the Malesian Bignoniaceae only a small proportion of tree species is
known wood-anatomically; the climbing and scandent species are fully unexplored in this
respect. Anomalous structure has, however, been described for several genera outside Malesia.
As far as known, the Malesian tree genera are wood-anatomically rather homogeneous: with
simple, rarely also reticulate, perforations to the vessels, homogeneous rays, mainly paratracheal,
aliform or confluent parenchyma, and fibres with simple to minutely bordered pits. Except for its
unusually narrow rays, Dolichandrone spathacea from the mangrove swamps does not differ
appreciably from the inland genera in its wood structure.

Leaves. Very poorly known for the Malesian representatives. Diversity of stomatal type and
indumentum (non-glandular and glandular hairs in a variety of forms) certainly deserves detailed
comprehensive studies, which will probably yield important additional taxonomic characters.

References: General surveys: SOLEREDER, Syst. Anat. Dicot. Stuttgart (1899, 1908); METCALFE
& CHALK, Anat. Dicot. Oxford (1950). — Wood: JaNssonius, Mikrographie des Holzes 4
(1926) 721-753 (Dolichandrone, Oroxylum, Stereospermum); PANSHIN, Philip. J. Sc. 48 (1932)
143-205 (Dolichandrone); DrescuH, Mal. For. Rec. 15 (1941) 50 (Deplanchea, Dolichandrone,
Pajanelia, Stereospermum); JANSSONIUS, Blumea 6 (1950) 450-452 (affinities); SEBASTINE, J.
Ind. Bot. Soc. 34 (1955) 299-306 (Pajanelia); JuTTE, Nova Guinea 10 (1959) 242 (Deplanchea). —

1977] BIGNONIACEAE (van Steenis) 119

Leaves: SIEBERT, Ann. Mo. Bot. Gard. 35 (1948) 123-136 (glands); PALIWAL, Flora 159 (1970)
124-132 (stomata). — P. BAAs.

Pollen morphology. Bignoniaceae have a long history of pollenmorphological study,
starting with the pioneer studies of H. MOHL (1835). The first author to present a detailed pollen-
morphological survey of the family, drawing attention to the taxonomical significance of the
pollen characters, was URBAN (1916). He concluded that (1) any attempt to base the main sub-
division of the family on pollen characters would group together taxonomically unrelated genera,
(2) for generic delimitation pollen had limited value. Later studies by several authors have
confirmed this (cf. BUURMAN, in the press).

Inaperturate, tricolpate, stephanocolpate and pericolpate apertural types occur in the family.

In a few genera tetrads are found.

Size ranges between 25 um in Astianthus antisyphilitica and 100 um in Nyctocalos cuspidata,
shape varies between suboblate and subprolate.

Remarkable is the amount of variation which may occur within genera or even intraspecifically.
In Stereospermum inaperturate, tricolpate and perisyncolpate pollen is found, while in Anemo-
paegma longepetiolatum inaperturate, stephanocolpate, pericolpate grains as well as tetrads
occur. In such cases sculpture affords more constant characters.

The tricolpate type is dominant in the family and is found in all four tribes. It rarely shows well
developed equatorial endoapertures. Instead, characteristically ruptured aperture membranes
are present, especially in Tecomeae. Operculate colpi occur in Argylia. A subdivision of the tri-
colpate type is possible on sculpture, which mostly varies between perforate and reticulate.

In some genera very complex pollen grains are present, such as those of Nyctocalos (fig. 2).

The tricolpate pollen grains in Bignoniaceae resemble those in Scrophulariaceae and Myopora-
ceae. The similarities with Pedaliaceae, stressed by ERDTMAN (1952) refer to a rather specialized
pollen type and may not reflect close affinity.

References: MOHL, Ann. Sc. Nat. II, 3 (1835) 304-346; URBAN, Ber. Deut. Bot. Ges. 3 + (1916)
728-758; ERDTMAN, Pollen morphology and plant taxonomy. Angiosperms. Almqvist &
Wiksell, Stockholm (1952) 73-74; BUURMAN, Contribution to the pollenmorphology of the
Bignoniaceae, with special reference to the tricolpate type (in the press). — J. MULLER.

Chemotaxonomy. Bignoniaceae share a number of biochemical tendencies with
Verbenaceae, Labiatae, Scrophulariaceae and with several other families of WETTSTEIN’s Tubi-
florae. Most of their outstanding chemical characters were already mentioned and discussed in
my ‘Chemotaxonomie der Pflanzen’ vol. 3 (1964) 268-281, 645-646, to which the reader is
referred. Much phytochemical information, however, became available only in more recent time.
Recent results confirm the trends already apparent in 1963; they are summarized in the following
pages. Chemical characters of Bignoniaceae may ultimately prove to be very useful in tracing
inter- and intrafamiliar relationships.

(1) Most members seem to produce and accumulate iridoid glucosides (formerly often called
pseudoindicans). Since a long time Bignoniaceae are known to contain labile glycosidic bitter
principles. Such a compound was isolated in 1888 from the bark and fruits of Catalpa bignonioides
WALTER and called catalpin (name changed later to catalposide). The structure of catalposide was
definitely established in 1962; it is an aucubin-type (C,-aglucone) ester glucoside and one of the
first pseudoindicans for which clearcut structural and biogenetic relationships with iridodial
and nepentalactone were demonstrated (hence the name iridoid glucosides for a presently very
large group of constituents of dicotyledonous plants). Catalposide (tastes bitter) is an ester of
p-hydroxybenzoic acid with catalpol (= 7,8-epoxy-aucubin). Catalpol and catalposide occur in
all species of Catalpa (leaves, stems, fruits) and catalpol (= catalpinoside) was also isolated from
barks of Paulownia tomentosa Steup. and P. fargesii FRANCH. where it occurs together with
syringin (V. PLouvier, C. R. Ac. Sc. Paris 272D, 1971, 1443). Probably catalpol and catalposide
occur in many more members of the family. In most recent times some related glucosides were
isolated from Bignoniaceae. Vanilloyl-catalpol (= amphicoside) is a constituent of Amphicome
emodi LinvL, and veratroyl-catalpol occurs in Tecomella undulata Seem. 5-Hydroxycatalpol
(= macfadyenoside) was isolated from Macfadyena cynanchoides MORONG. All iridoid glucosides
mentioned hitherto have structures based on the aucubin-derivative catalpol. The first non-
aucubin-type glucoside described from Bignoniaceae is tecomoside with a C,o-aglucone; it was

120 FLORA MALESIANA [ser. I, vol. 8?

isolated from Tecoma capensis LINDL. (A. BIANCO et al. Gazz. Chim. Ital. 105, 1975, 195). It is to
be expected that much more iridoid glucosides will be detected in the family in future.

(2) Some Bignoniaceae produce alkaloids. So far only pyridine-type and piperidine-type
alkaloids with an iridoid Cy 9- or rarely Cy-skeleton were identified definitely in species belonging
to this family. This fact strengthens the belief that the tendency to produce iridoid compounds is
a very important character of Bignoniaceae. Thusfar simple iridoid alkaloids were described for
species of Campsis (boschniakine), /ncarvillea (plantagonine, indicain), Tecoma (tecomanine,
tecostidine, tecostanine, boschniakine, 4-noractinidine and several derivatives of skytanthine).
The basic constituents of Amphicome (now reduced to Incarvillea), Newbouldia and other genera
may belong to the same group of alkaloids.

A recent review of the chemistry, distribution and systematic meaning of all presently known
main groups of iridoid plant constitutents was published by S. ROSENDAL JENSEN et al. (Bot.
Notis. 128, 1975, 148-180).

(3) Many Bignoniaceae synthesize naphthaquinones and corresponding anthraquinones by
prenylation of o-succinylbenzoic acid. This pathway to quinonoid naphthalene- and anthracene-
type secondary metabolites is presently known from taxa belonging to Rubiaceae, Verbenaceae,
Scrophulariaceae, Bignoniaceae and possibly Acanthaceae and Gesneriaceae. In roots, woods
and barks of Bignoniaceae lapachol, lapachonone, «- and B-lapachone and dehydro-«-lapachone
occur frequently. These monomeric naphthaquinonoid compounds are often accompanied and
sometimes replaced by more complex dimeric constituents like tectol, guayacanine and guayine
and by corresponding anthraquinones such as tectoquinone and 2-methyl-3-hydroxyanthra-
quinone. Woods which contain appreciable amounts of these quinonoid compounds are more or
less resistant to marine borers, white ants and Fungi. At the same time such woods may be the
causes of skin irritations and of allergic skin diseases in man. Lapachol- and tectoquine-type
substances are presently known from species of the genera Catalpa, Heterophragma, Kigelia,
Paratecoma, Phyllarthron, Stereospermum, Tabebuia, Tecoma, Tecomella, and Zeyhera. R. H.
THOMSON has reviewed the chemistry and distribution of quinones and related compounds in his
book ‘Naturally occurring quinones’ (2nd ed. 1971). The phthalide catalpalactone from the wood
of Catalpa bignonioides WALTER and C. ovata G. Don arises from the same pathway as lapachol
and its congeners (H. INouyE et a/. Chem. Pharm. Bull. Tokyo 23, 1975, 384, 392, 2523). On the
other hand it should be stressed that the red-coloured naphthaquinones of Boraginaceae (e.g.
alkannin) which are structurally very similar to lapachol are produced along a totally different
biosynthetic pathway (cf. E. LEISTNER, Chinoide Farbstoffe, Ber. Deut. Bot. Ges. 88, 1975,
163-178).

(4) The “‘tannins’”’ mentioned for many Bignoniaceae in the older phytochemical literature
(e.g. DEKKER, 1913) seem to be glycosides and esters of o-diphenolic compounds. Orobanchin
(= verbascoside)-type ester glycosides were definitely demonstrated to occur in species of
Campsis, Catalpa, Eccremocarpus and Pandorea. A review of this type of polyphenolic plant
constituents which simulate true tannins in some respects is to be found in my ‘Chemotaxonomie
der Pflanzen’ vol. 5 (1969) 250-252. Orobanchin yields a molecule of caffeic acid, 3,4-dihydroxy-
phenylethanol, glucose and rhamnose each. Just as in most other families of Sympetalae true
tannins are replaced in Bignoniaceae by more or less complex esters and glycosides of o-diphenolic
cinnamic acid derivatives. Moreover, simple esters of caffeic acid and biosynthetically related
derivatives of cinnamic and benzoic acids are present in large amounts in many Bignoniaceae.
The recent investigations of V. B. PANDEY and B. DasGupTA with the bark of Tecomella undulata
SEEM. (veratroylglucose = tecomin: Experientia 26, 1970, 1187) and of M. SUGUMARAN et al.
with leaves of Tecoma stans H.B.K. (16 aromatic acids: Ind. J. Exper. Biol. 13, 1975, 93) exemplify
this trend. p-Hydroxybenzoic acid is present as an ester in all species producing catalposide;
probably this phenolic acid is rather ubiquitous in the family. The presence of appreciable
amounts of hydroquinone (in living cells as the glucoside arbutin?) in leaves of Jacaranda
mimosaefolia D. Don (S. SANKARA SUBRAMANIAN ef al. Phytochemistry 12, 1973, 220) might be
connected with a strong tendency to produce and accumulate p-hydroxybenzoic acid; if this is
actually the case hydroquinone (and arbutin?) may be detected in much more Bignoniaceae in
future. Jacaranone, a quinonoid compound which exhibits antitumor and cytotoxic activity was
recently isolated from leaves and twigs of Jacaranda caucana PITTIER (M. OGurRa et al. Lloydia

1977] BIGNONIACEAE (van Steenis) 121

39, 1976, 255); it seems to be derived from tyrosine and is chemically very similar to the Cornus
quinol glucoside (= cornoside) which is also present in leaves of Digitalis purpurea (Bot. Notis.
128, 1975, 174).

(5) According to J. B. HARBORNE (Phytochemistry 6, 1967, 1643) leaf flavonoid patterns of
Bignoniaceae are close to those of Acanthaceae, Gesneriaceae, Labiatae and Scrophulariaceae.
Features which support such a statement are the replacement of flavonols by flavones in many
species, the relatively frequent occurrence of flavones with an unsubstituted B-ring (e.g. chryson,
baicalein), of 6-hydroxylation of chrysin (baicalein), apigenin (scutellarein) and luteolin (6-
hydroxyluteolin) and of 0-methylation of flavones. The latter trend is illustrated by Zeyhera
tuberculosa BUR. ex VERLOT which contains 5,6,7-trimethoxyflavone and 5,6,7,8-tetramethoxy-
flavone in leaves (J. P. KuTNEY & H. W. HANSSEN, Phytochemistry 10, 1971, 3298). The bitter
principle of the fruits of Sparattosperma vernicosum Bur. & K. SCH. was shown by J. P. KUTNEY
et al. (Phytochemistry 9, 1970, 1877) to be the 7-neohesperidoside of pinocembrin (= 2,3-
dihydrochryson).

(6) Free triterpenic acids occur in appreciable amounts in leaf waxes of many families of
Tubiflorae (especially Verbenaceae, Labiatae and Plantaginaceae) and reJated orders. It is of
interest in this respect that ursolic acid was isolated in recent time from leaves of Bignonia
diversifolia H.B.K., Campsis radicans SEEM., Catalpa bignonioides WALTER, Heterophragma
quadriloculare K. Scu., Jacaranda mimosaefolia D. Don (not definitely identified) and Paulownia
tomentosa STEUD. The bark of Jacaranda mimosaefolia yielded lupenon.

(7) Many members of Verbenaceae, Labiatae, Scrophulariaceae and Plantaginaceae replaced
starch by stachyose-type oligosaccharides as storage carbohydrates. The same trend seems to
exist in Bignoniaceae. Large amounts of stachyose occur in species of Catalpa (roots, wood, bark),
Newbouldia laevis SEEM. (roots) and Paulownia tomentosa (stem).

(8) Most representatives of Tubiflorae produce starch-free seeds which are rich in proteins and
oils. The seed oils are often characterized by a high degree of unsaturation. In this respect
Bignoniaceae conform to the rule. Their seeds generally contain 20-35% of oil. In some taxa
oleic and (or) linolic and (or) linolenic acid are the only major fatty acids of the seed oils (e.g.
species of Crescentia, Niedzwedzkia = Incarvillea, Paulownia and Stereospermum). In other taxa
the ‘normal’ fatty acids are accompanied or replaced by large amounts of unusual fatty acids
such as conjugated trienoic acids (species of Catalpa, Chilopsis, Jacaranda), Cy -keto-acids
(Cuspidaria pterocarpa DC.), octadeca-trans-3,cis-9,cis-12,cis-15-tetraenoic acid (Tecoma stans
H.B.K.) or hexadec-9-enoic and octadec-11l-enoic acid (Doxantha unguis-cati Miers). M. J.
CHISHOLM and C. Y. Hopkins discussed the chemistry of seed oils of 11 species representing 4
tribes (Canad. J. Chem. 43, 1965, 2566).

The preceding phytochemical picture places Bignoniaceae phytochemically very close to a
number of families of Tubiflorae, especially Verbenaceae, Labiatae and Scrophulariaceae. Still
other constituents are known from Bignoniaceae. Lack of acquaintance with their structures and
(or) with their distribution, however, does not yet allow a systematic evaluation. Saponins, which
seem to be rather widespread in the family but were never investigated in detail, belong to these
chemical characters. The same holds for a number of phenolic compounds isolated in recent
time, such as the lignans sesamin and paulownin from Paulownia tomentosa StEuD. and Phyl-
larthron comorense DC., the dilignol (a lignan-type compound) zeyherol from Zeyhera digitalis
HoeHNeE and the dihydroisocoumarins 6-methoxymellein, kigelin and 6-demethylkigelin from
Kigelia pinnata DC.

Concluding it may be stated that the intimate relationships between Bignoniaceae and Scrophu-
lariaceae which are indicated by genera like Catalpa and Paulownia (often placed in Scrophu-
lariaceae) are confirmed by phytochemistry. At the same time phytochemistry stresses a very
close coherence of a core of families of Tubiflorae; this core comprises Scrophulariales sensu
Cronouist (1968) and Lamiales sensu TAKHTAJAN (1969). — R. HEGNAUER.

Uses. There are no outstanding qualities marking Bignoniaceae as useful plants, otherwise
than ornamentals and these concern mostly the introduced species for which I refer to the special
key and account at the end. There are magnificent native species notably of Tecomanthe but they
have as yet not become in general use.

Good roadside trees are Millingtonia hortensis and Spathodea campanulata. A highly esteemed

122 FLORA MALESIANA [ser. I, vol. 8?

vegetable (/alab) with the Sundanese is Oroxylum indicum (flowers, buds, and very young pods).

For re-afforestation and holding terraces on slopes the pioneer qualities of species of Rader-
machera and Deplanchea might be useful.

The timber is in general not valuable and in nature not available in sufficient quantity. The soft
wood of Millingtonia hortensis was advertized as useful for tea-boxes. The only species yielding
sizeable timber of good quality are: Fernandoa macroloba, Pajanelia longifolia, Radermachera
gigantea, and the three species of Stereospermum, which all may be valuable for silviculture.

Terminology. The shape of the corolla has been defined as tubular (fig. 4b, 10e) in which
case there can be a distinction in a basal tube (on apex of which inside the stamens are inserted)
as in fig. 4b; funnel-shaped (fig. 23d, 26c), hypocrateriform or salver-shaped (fig. 1a, 8c, 15) or
infundibuliform (fig. 32a).

Notes. Since my thesis (Rec. Trav. Bot. Néerl. 24, 1927, 787-1049), here always cited as ‘Thesis
(1927)’, and subsequent revision in Bull. Jard. Bot. Btzg III, 10 (1928), I have remained always
much interested in this family and have published some revisions and many notes precursory to
the present treatment. I have to thank the late Mr. N. Y. SANDWITH (Kew) for namings of cul-
tivated species, and Dr. A. L. GENTRY (St. Louis) for recent information on them, Dr. H. HEINE
(Paris) for assistance in various matters, Mr. Michael GALorE (Lae) and Prof. E. J. H. CORNER
(Cambridge) for photographs, Miss Christine BRIGHTON (Jodrell Lab., Kew) for the first chromo-
some count in Tecomanthe, while I gratefully acknowledge precursory work performed by Mr.
J. C. DEN HARTOG on Tecomanthe and Pandorea in 1969/70 at the Rijksherbarium where he
worked as a graduate student.

KEY TO THE GENERA!
1. Erect trees or shrubs.
ee Peaved aitc. THOSUY I WHOLIS, 7 © oe See ss se es ea ee eee 6. Deplanchea
2. Leaves compound, almost always decussate.
3. Leaves 1-pinnate.
4. Leaf rachis with a sharp, keel-like ridge above. Leaflets 8-12 pairs. Capsules winged
12. Pajanelia

4. Leaf rachis not keeled above. Leaflets less than 6 pairs. Capsules not winged.

5. Corolla salver-shaped, white, nocturnal, the tube 12-18cm long, narrow-cylindric. Calyx
spathaceous to the base, 3-6 cm. Leaves nigrescent, with domatia. A flat pseudoseptum developed
parallel to the valves. Seeds rectangular, with corky wings as thick as the seed . 7. Dolichandrone

5. Corolla much shorter. No pseudoseptum. Seeds with hyaline wings. Leaves not nigrescent.

6. Mature leaflets serrate; crenate, or pmnatiid ~~ 5 ©." 2 <2 2". a>. ee 5. Tecoma
6. Mature leaflets entire.
7. Calyx regularly 5-lobed, rather thin, reticulately veined, short-hairy, 2-3 cm
11. Lamiodendron'
7. Calyx lobes unequal, often less than 5; calyx not thin and reticulate-veined.
8. Capsule rather flattened, with a flat septum, if terete with 10 ribs .. . . 10. Fernandoa
8. Capsule terete, with a terete septum, never with many ribs, sometimes one ridge on each
valve. No domatia in Mal. spp.
9. Seeds thick, in one row in each cell, each fitting in deep notch in the septum. Valves rather

ards Ovplesiince2 TOWS IN iGACH COM) ac ah as, aie eae ee ees 8. Stereospermum
9. Seeds thick, in many rows, the septum without notches. Valves thin. Ovules in many rows
Be cach cede 5 Lah I are ei Gs SRE ae 9. Radermachera

3. Leaves 2—3(—-4)-pinnate.
10. Flowers fleshy, very coarse: calyx 2-4 cm, corolla wide, dirty violet or reddish purple to liver
brown, 7-10 cm. Capsule flat, sword-shaped, 60-120cm. .......... 3. Oroxylum
10. Flowers not fleshy, much smaller or thinner, white or pink. Capsule smaller and of different shape.
11. Leaflets with domatia. Corolla white, salver-shaped, the tube 6-8 cm long, cylindric, 2 mm wide.
Capsule compressed parallel to the septum, the latter parallel to the valves; dchicoanead septicidal.
11. Leaflets without domatia. Corolla tube otherwise, wider. Capsule cylindric, with a terete corky
ane” septum attached perpendicular to the valves; dehiscence loculicidal. . . . . 9. Radermachera
. Lianas.

12. Corolla salver-shaped, with a narrow cylindric tube, 5-19 cm long. Capsule flat, large, with a
median lengthwise ridge. Septum parallel to the valves, dehiscence loculicid. . . . 1. Nyctocalos

12. Corolla not salver-shaped, tubular, infundibuliform or funnel-shaped.

(1) Of Lamiodendron and Hieris the fruit is still unknown.
The key given here is only to native and thoroughly naturalized species (only Tecoma stans).
A tentative key to the cultivated species is added in an appendix on page 180.

1977] BIGNONIACEAE (van Steenis) 123

13. Corolla tube + geniculate above the basal tube, upper tube slightly curved and flattened with a
prominent fold. Calyx c. '/, cm, below the very short lobes with a short spur-like tooth. Ovules
6-8 per cell, in + two rows. Leaflets 5, those of the lower pairsessile ...... 2. Hieris’

13. Corolla tube not geniculate, without a fold. Ovules ~ per cell in several rows. Capsule loculicid,
with boat-shaped valves.

14. Leaves digitate, with 3(—-5) leaflets. Corolla lobes valvate ......... 13. Neosepicaea
14. Leaves pinnate, with 1 or more pairs of leaflets. Corolla lobes imbricate.

15. Flowers in racemes, axillary, mostly on the old wood, rarely terminal (in a high-mountain sp.),
the rachis at the base usually with several crowded pairs of minute sterile bracts. Calyx large,
15-40 mm, distinctly lobed. Corolla large, mostly red, 5-12 cm long incl. the lobes, not bearded
in the mouth and upper part of the tube, but stuppose hairy at the insertion of the stamens (rim
of the basal tube). Corolla lobes usually deltoid, very narrowly overlapping. Anther-cells
eset bones. Vee . Se oo a ae 14. Tecomanthe

15. Flowers in thyrses, only occasionally depauperate into racemes, usually terminal, sometimes
axillary or on old wood. No sterile bracts at base of peduncle. Calyx small, mostly stunted or
very shortly lobed, 2-7 mm. Corolla incl. lobes up to c. 3'/, cm (in a single Australian species
with large lobes c. 5 cm), usually white or yellowish with red dots or streaks inside lobes and
mouth, often bearded in the mouth and upper part of the tube, sometimes with a hair-ring at the
insertion of the anthers. Corolla lobes widely overlapping, often roundish. Anther-cells 1*/,-
RE ey ae re Sof FT ae 15. Pandorea

Tribe 1. Bignonieae

B. & H. Gen. Pl. 2 (1876) 1027; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 209;
BUREAU, Fl. Bras. 8, 2 (1896) 16. — Eubignonieae ENDL. Gen. Pl. (1839) 712. —
Subtribe Eubignonieae DC. Rév. Bign., Bibl. Univ. Genéve (1838) 122; FENZL,
Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 262; DC. Prod. 9 (1845) 143.

Capsule septicid, the septum parallel to the valves. Frequently lianas, with
tendrils, mostly in the neotropics.

Note. Boyer (Hort. Maur. 218) and DC. (/.c.) included the two present tribes in
tribe Bignonieae.

1. NYCTOCALOS

T. & B. in Mig. J. Bot. Néerl. 1 (1862) 366; BUREAU, Mon. (1864) 52; Mia. Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 201; ibid. 3 (1867) 249; SEEM. J. Bot. 8 (1870) 147;
K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 219 (‘Nycticalos’); STEEN. Thesis
(1927) 805; Bull. Jard. Bot. Btzg III, 10 (1928) 178; Acta Bot. Neerl. 2 (1953) 306;
SANTISUK, Kew Bull. 28 (1973) 182. — Fig. 1-2.

Lianas, without tendrils. Leaves pinnately 3-foliolate (in one extra-Mal. sp.
I-pinnate with 5 leaflets). Leaflets herbaceous to chartaceous, entire, + elliptic,
acuminate, with a few scattered crateriform glands along and spaced from the
midrib underneath, above very fine punctate-pitted glandular. /nflorescence a short
lateral or terminal 8—12-flowered raceme. Pedicels bracteolate. Flowers nocturnal,
erect, fragrant; lobes widely imbricate. Calyx cup-shaped, truncate, with 5 horn-
like teeth, the latter with glands on both sides. Corolla almost actinomorphic,
salver-shaped, the long narrow basal tube dilated in the upper part, with 5 rounded
unequal or subequal lobes. Stamens inserted in the throat 4 with or without a rudi-
ment or 5, equal or 2 anterior ones sometimes longer, not exserted; anther-cells
divergent, versatile; filaments glabrous and no hairs near their insertion. Disk
annular, fleshy. Ovary with c~ rows of c ovules along the margins of the dissepi-
ment; style long, filiform. Capsule large, flat, stalked and acuminate but with

124 FLORA MALESIANA [ser. I, vol. 8?

v
4

—_

Fig. 1. Nyctocalos cuspidata (BL.) Miq. a. Habit, in flower, b. fruit, opened, showing seeds and dissepiment,
x 1/, (after MIQUEL, 1867)

parallel edges, valves with a central prominent rib; dissepiment thinnish, flat.
Seeds flimsy winged, roundish.

Distr. Three spp. in SE. Asia (Assam, Burma, Thailand, Yunnan) and West Malesia (Java, Borneo,
Celebes, and the Philippines). Fig. 3.

Ecol. Rare rain-forest lianas at low altitude.

Taxon. The genus stands isolated in the Old World flora. In many aspects the small New World genus
Tanaecium Sw. is very similar, but in this genus at least part of the leaflets carries a terminal tendril;
besides its fruit is not flat, but said to be cylindric or quadrangular with convex woody valves. Still I
believe it to be an ally in the New World. There is also a similar resemblance with the monotypic
South American genus Macranthisiphon BUREAU but that has 2-ranked ovules and a more elongate,
funnel-shaped corolla tube.

For a moment I thought that Nyctocalos pinnata STEEN. (from Yunnan, only known in fruit, /.c. 1953,
306) might belong to Hieris, but the very numerous seeds defeat this, as far as H. curtisii is concerned.
Hieris is, of course, the most intimate related genus, with the same punctate glands on the leaves and a
deceptively similar calyx; its pollen is quite different.

Too much importance has been ascribed to the structure of the androecium: 5 equal stamens to 4
didynamous; this varies as is explained under N. cuspidata. It led SEEMANN (/.c.) even to the inclusion of
the Australian Hausmannia jucunda into the genus, which belongs in fact to the Tecomeae with quite
different fruit and valvate corolla lobes.

Nomencl. The generic name is female, being derived from the Greek nux.

KEY TO THE SPECIES

1. Corolla tube c. 15-16 cm long, white. Calyx teeth horn-like, with a linear apex. Lateral petiolules
oe Se a ne ae een ee er ree ae Fy ea ee Se 1. N. cuspidata
1. Corolla tube c. 5-6 cm long, tinged palish yellow suffused with pinkish shade. Calyx teeth triangular,
acute.Lateral petiolules:2=3 mimi... «/i)4.. go ease See ee 2. N. brunfelsiiflora

1977]

1. Nyctocalos cuspidata (BL.) Mia. Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 249, t. 8B (‘cuspidatum’);
Merr. Philip. J. Sc. 1 (1906) Suppl. 237; C. B. Ros.
ibid. 6 (1911) Bot. 211; MeERR. En. Philip. 3 (1923)
443; STEEN. Thesis (1927) 813, incl. var. oblongum
STEEN.; Bull. Jard. Bot. Btzg III, 10 (1928) 180;
SANTISUK, Kew Bull. 28 (1973) 183. — Tecoma
cuspidata BL. Rumphia 4 (1849) 35. — N. macro-
siphon T. & B. Cat. Hort. Bog. (1856) 155, nomen.
—N. brunfelsiaeflorus (non T. & B.) Mia. Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 201, pro specim.
celeb. — N. thomsonii Hook. f. Bot. Mag. 93 (1867)
t. 5678; CLARKE, FI. Br. Ind. 4 (1884) 377; STEEN.
Thesis (1927) 809; Bull. Jard. Bot. Btzg III, 10
(1928) 180; SANTISUK, Kew Bull. 28 (1973) 183. —
Gelseminum cuspidatum O. K. Rev. Gen. Pl. 2
(1891) 479. — N. assamica Hook. f. ex. K.ScH. in
E. & P. Nat. Pfl. Fam. 4, 3b (1894) 221, nomen,
lapsus. — Fig. 1-2.

Leaflets elliptic, ovate, obovate, or narrow
oblong, rounded at base, acuminate to cuspidate at
apex, 6-11(—18) by 31/,-7(-10) cm; petiole 3-5 cm;
rachis 2!/,—31/,cm; petiolules !/,-1 cm. Pedicels
c.+/,-1 cm. Calyx c. 6mm. Corolla in bud pale
green, later creamy, the tube c. 15-19 cm; dilated
part c. 3-6cm; lobes rounded, c. 1'/,-2 cm.

Stamens 4, didynamous, with or without a filiform
rudimentary Sth one; anther-cells 6-10 mm; con-
nective with a filiform appendage 2-3 mm.
Capsule 16-24 by 33/,-47/,cm. Seeds (including
- Ped wings) rounded to obovate, 3-4 by
2!-3 cm.

Fig. 2. Nyctocalos cuspidata (BL.) MiQ. Pollen
grain, SEM x 500 (BS 10396).

Distr. SE. Asia (Assam: Mikir & Gowhatty
Hills) and Central Malesia: Philippines (Luzon,
Polillo, Palawan, Biliran, Mindanao, Basilan) and
Celebes (Manado, Kema, Bantaeng). The type
was said to have been collected in the Moluccas by
ZipPeL but this must be doubted. Fig. 3.

Ecol. Lowland rain-forests.

Notes. Hitherto importance was laid in keys on
the difference between WN. cuspidata and N.
thomsonii in that the former was described with 5
fertile stamens and the latter with 4 didynamous

BIGNONIACEAE (van Steenis)

125

stamens and a filiform rudiment. This was suspi-
cious as there were hardly any other differences. In
material on spirit of Celebes specimens cultivated
in Hort. Bog. I have now found flowers with
didynamous stamens whether or not accompanied
by a staminode. Furthermore, in RIEDEL s.n. from
Manado and in BS 10396 from Polillo there are 5
perfect stamens; but even here in one flower the
two anterior stamens were somewhat longer than
the others. There is thus variation in the degree of
tendency to zygomorphism. This is also visible in
the difference in size of the corolla lobes of which
one is mostly larger than the others. The taxono-
mical implication is the reduction of N. thomsonii.

In the Malesian specimens the calyx teeth appear
to be somewhat longer and more horn-like than in
the Assam specimens depicted by Hooker f.

2. Nyctocalos brunfelsiiflora T. & B. in Mig. J.
Bot. Néerl. 1 (1862) 367 (brunfelsiaeflorus’); Mia.
Choix (1863) t. vu; Ann. Mus. Bot. Lugd.-Bat. 1
(1864) 201, excl. syn. et specim. celeb.; ibid. 3
(1867) 248, t. 8A; STEEN. Thesis (1927) 811; Bull.
Jard. Bot. Btzg III, 10 (1928) 179 (‘brunfelsiiflorus’) ;
Acta Bot. Neerl. 2 (1953) 306; BAck. & BAKH.
f. Fl. Java 2 (1965) 536; SANTISUK, Kew Bull. 28
(1973) 183. — N. shanica MACGREGOR & W. W.
SmiTH, Rec. Bot. Surv. India 4 (1911) 280; STEEN.
Thesis (1927) 811; SANTISUK, Kew Bull. 28 (1973)
182, 183; Thai For. Bull. Bot. 8 (1974) 88.
Leaflets elliptic, oblong, or obovate, acuminate
to cuspidate, 7-13 by 3-6cm; petiole 4-7 cm;
rachis 11/,-3cm; petiolules 1-2mm. Racemes
5-10-flowered. Pedicels 1—-11/, cm. Calyx 5-6 mm,
suffused with reddish tinge in anthesis (ex coll.).
Corolla whitish afterwards yellowish, later apically
suffused with pink; tube 5—7 cm, the dilated part
some 2-2'/,cm; lobes rounded to obovate or
truncate, c. 2cm. Stamens (so far as known) 5.
fertile, equal or subequal. Capsule 10-13 by 3-4 cm,

Fig. 3. Range of the genus Nyctocalos T. & B.:
N. brunfelsiiflora T. & B. (triangles), N. cuspidata
(BL.) Mig. (dots), and N. pinnata Steen, (circle).

126 FLORA MALESIANA [ser. I, vol. 8?

NIf

ZS "J
tA) 4
ps

Y
ea

LE
= 2 a8

Fig. 4. Hieris curtisii (RIDL.) STEEN. a. Habit, x 4/2, b. corolla, nat. size, c. LS of ovary, disk and recep-
tacle, x 2!/,, d. calyx, x 2!/,,e. CS of ovary, x 7'/2, f. glands on ovary (HENDERSON S.n.).

1977]

BIGNONIACEAE (van Steenis)

127

Distr. SE. Asia (Upper Burma: S. Shan States;
Thailand: N. & SE.), in Malesia: Borneo (Sabah,
near Kudat; Sarawak, near Niah), SW. Java
(Wijnkoops Bay). Fig. 3.

Ecol. Lowland rain-forests, even in Burma
below 300 m.

Uses. TEYSMANN found it a beautiful ornamen-
tal; as far as nocturnally flowering plants can be.
He could easily propagate it by marcottes. VAN
HASSELT (in sched.) noted that in SW. Java crushed
leaves are rubbed against head and stomach aches.

Vern. Kakatjangan, S, SW. Java.

Notes. N. shanica was distinguished by having
glabrous anthers; these had in N. brunfelsiiflora
been described and depicted as hairy to the base.
This is, however, a lapsus: the base of the filaments
and tube in the vicinity of the insertion is only
dotted with small sessile granular glands. SANTISUK

(l.c.) said that the pollen would be different from
that of N. shanica, but I cannot accept this for
specific distinction. The colour of N. shanica was
described as white, but field data enumerated by
SANTISUK mention also creamy white, buds pur-
plish, and pale yellow flowers. Of N. brunfelsiiflora
MIQUEL mentioned them to be pale pinkish ‘tirant
légérement vers le jaune’, more purplish to anthesis.
BACKER said: corolla at first white, afterwards
yellowish; tube apically suffused with red. I do not
ascribe much importance to these faint colour
variances, especially nocturnal flowers often dis-
colour with age.

In comparing the scanty flowering material of
both species it seems that in N. shanica the widen-
ing of the corolla tube starts lower than in N. brun-
felsiiflora, but I can not accept this for specific
distinction.

2. HIERIS

STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 279, f. 13. — Fig. 4.
Slender woody twiner. Leaves 1-pinnate, with (1—)2(-3) pairs of entire leaflets.

Racemes (axillary or) terminal. Pedicels bracteolate. Flowers scattered. Calyx
cupular, articulated with the pedicel, short- or indistinctly 5-lobed, just below the
margin with 5 spur-like, upcurved teeth at the base of each lobe, each tooth with a
few glands on each side of its base. Corolla zygomorphic, the basal narrowed tube
c. 1/, of its length, + geniculate with the upper */, which is widened, flattened, and
gently curved; lobes subequal, suborbicular, finely capitate-glandular, papillose-
hairy outside at base. Stamens 4, didynamous, inserted at the end of the basal tube,
included, 5th rudimentary; anthers connivent in pairs, divaricate, connective
apiculate. Disk entire, pulvinate-annular, fleshy. Ovary ovate, subterete, with 2
grooves, microscopically glandular; ovules in + two rows of 3-4 in each cell.

Distr. Monotypic. Malesia: Malaysia: Penang I. (near village on north coast).

Taxon. Outstanding by the few ovules and the geniculate tube of the corolla. The structure of the
inflorescence seems to be racemose though flowers are articulated.

Notes. Unfortunately the fruit and seed of this most interesting plant is unknown; from the cross-
section of the ovary one might assume it to belong to tribe Bignonieae as the grooves of the ovary corres-
pond with the edges of the septum, while furthermore the calyx structure is a replica of that of Nyctocalos,
with which genus it seems closest related. In Nyctocalos, however, there is a very large number of ovules in
each cell, which are very few in the Penang plant, like in the South American genus Memora. By the
pinnate leaves Hieris shows resemblance with Nyctocalos pinnata STEEN. from Yunnan, which is unfor-
tunately only known in fruit, but has abundant seeds in each cell.

Mr J. MULLER told me (Febr. 1975) that the pollen of Hieris is not in the least resembling the showy
pattern of Nyctocalos.

1. Hieris curtisii (RipL.) Steen. Bull. Jard. Bot.
Btzg ILI, 10 (1928) 280, f. 13. — Tecoma curtisii
Ruiw-. J. As. Soc. Str. Br. 49 (1908) 26. — Pandorea
curtisii Ripc. Fl. Mal. Pen. 2 (1923) 553, f. 125;
Steen. Nova Guinea 14 (1927) 301; Thesis (1927)
846, f. 4(2). — Fig. 4.

Glabrous. Twigs terete, with very many small
lenticels ; nodes with glands and a dark transversal
line. Leaves (10—)15-20 cm pe petiole 2'/,-8 cm;
rachis c. 4-5 cm; petiolules of lower pair of leaflets
4-12 mm, of upper pair (0-)1-2 mm. Leaflets
herbaceous, slightly unequal-sided, ovate-oblong,
long-acuminate, 5-8 by 2~3 cm; nerves 4-5 pairs;

undersurface with scattered, small, rimmed-crateri-
form glands, upper surface with microscopical
pitted-punctate glands. Rachis rather densely
flowered, microscopically puberulous, 2-7 cm;
peduncle 2 cm, with barren bracts. Bracts linear,
2mm. Pedicels thin, 5-8 mm, with 1-2 minute
bracteoles in the lower half. Ca/yx 6 mm, purplish,
bluntly 5-ribbed, inside with dark red microscopical
capitate-glandular hairs. Corolla 4'/,-5 cm, tube
yellow, lobes whitish turning pale lilac, c. 1 cm @,
2 upper recurved, 3 lower erect; outside on transi-
tion of tube and lobes scattered, rather large,
brown red glands; basal tube 1'/,cm by 2mm,

128 FLORA MALESIANA [ser. I, vol. 8?

Distr. Malesia: Penang I., see above.

Ecol. Not well noted, 3 collections, all from
1898-1902; ff. June, July, Nov., and a cultivated
specimen in Hort. Sing., Lawn 0, in Febr.

near the insertion of the stamens scattered micro-
scopical red-tipped, capitate-glandular hairs as in
the calyx tube. Filaments glabrous, 12 and 14 mm;
rudiment 5mm, with reflexed apex; anther-cells
2!/,mm, connective appendage linear, 1 mm.
Ovary elliptic 1'/, by 1 mm; style 2 cm; stigmatic
lobes elliptic, 1'/, mm.

3. OROXYLUM

VENT. Dec. Gen. Nov. (1808) 8; K.ScuH. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 225
(‘Oroxylon’), I.c. 212 in clavi; STEEN. Thesis (1927) 816; Bull. Jard. Bot. Btzg III, 10
(1928) 181. — Calosanthes BL. Bijdr. (1826) 760; DC. Prod. 9 (1845) 177; BUREAU,
Mon. (1864) 45, t. 9. — Hippoxylon RaFin. Sylv. Tellur. (1838) 78, nom. illeg. —
Fig. 5, 7.

Glabrous tree, robust in all its parts. Leaves 2—3(—4)-pinnate, all nodes with in
sicco shrinking articulations; leaflets entire. Flowers very large, fetid, nocturnal, in
large terminal racemes (by exception in a thyrse). Calyx persistent, not articulated,
coriaceous, closed in bud, with a fine apical pore, later opening campanulate, +
entire. Corolla funnel-shaped, lobes 5, subequal, imbricate in bud. Stamens 5,
subequal, all fertile; anthers 2-celled, cells free, + parallel. Ovary with ~ rows of
ovules in both cells. Capsule flat, very large, sword-shaped, linear; dissepiment
flat, coriaceous. Seeds large in ~ rows; insertion linear, 1 cm wide.

Distr. Probably monotypic. From Ceylon, the Deccan and Himalayas through SE. Asia (also in S.
China: Yunnan, Kwangsi, Setchuan, Kweichou) and Malesia eastwards to the Philippines, Celebes, and
Timor. Fig. 6.

Ecol. Acharacteristic, short-lived nomad tree, nowhere gregarious, not in mature rain-forest but always
in openings, secondary growths and thickets, rather indifferent to climate (also in teak forest under
seasonal conditions) and soils, mostly below 1000 m, but in S. China up to 1375 m (HANDEL-MAZZETTI).

Taxon. A second species has been described, raised from seed, collected by A. HENRY in Yunnan, in
1889, in the Arnold Arboretum, and named O. flavum REHDER (in Sargent, Trees & Shrubs 1, 1904,
193, t. 92). REHDER discriminated this from O. indicum chiefly by the sulphur yellow colour of the nearly
symmetrical flowers, the plain not toothed or crisped corolla lobes, the splitting calyx, and the oblong
leaflets.

Several of these characters are not valid, especially if we take into consideration that REHDER’s plant
was an unbranched sapling of 3 m high. In such saplings the leaves are always somewhat longer and
thinner. The sulphur-yellow corolla is also rarely found in O. indicum from where I described it (1928) as
var. citrinum STEEN. on a cultivated specimen at Bogor so annotated by J. J.SmirH (C.H.B. XV.K.B.IX-
11). The calyx is indeed different from that in O. indicum, in being thinner and having 5 faint ribs, but it is
lobed by tearing, and this is also sometimes found in fruiting specimens of O. indicum. The corolla in
O. flavum is also regular and somewhat smaller than usual but an examination of the type showed an
exactly similar occurrence of hairs at the anther bases, the patelliform glands outside and the granular-
glandular hairs inside. Remains the plain, entire corolla lobes, and an other character figured by REHDER
but not mentioned by him, viz that the inflorescence is not a raceme, but a thyrse, with the lower stalks
5-flowered in double triads and the upper ones in simple triads, a situation never recorded or seen by me
in O. indicum. 1 cannot well account for these two differences, but they could be due to cultivation; in
on seperience tropical plants in hothouses often deviate from those in the wild, certainly in first-flowering
saplings.

1. Oroxylum indicum (L.) Kurz, Fl. Burma 2
(1877) 237; CLARKE, Fl. Br. Ind. 4 (1884) 378;
K. & V. Bijdr. Booms. 1 (1894) 66, Atlas 2 (1914)
t. 358; RIDL. Fl. Mal. Pen. 2 (1923) 548; Merr. En.
Philip. 3 (1923) 444; STEEN. Thesis (1927) 816; Bull.
Jard. Bot. Btzg III, 10 (1928) 181, incl. var. citrinum
STEEN. /.c. 184; OCHSE & BAKH. Ind. Groent. (1931)
77, f. 46; HAND.-Mazz. Symb. Sin. 7 (1936) 888;
CORNER, Ways. Trees (1940) 166, Atlas t. 29. —
Palega-pajaneli RHEEDE, Hort. Mal. 1 (1686) 77,

t. 43. — Bignonia indica var. « LINNE, Sp. Pl. (1753)
625; Roxs. Fl. Ind. ed. Carey 3 (1832) 110. —
Bignonia pentandra Lour. Fl. Coch. 2 (1790) 379.
—Bignonia tripinnata NORONHA, Verh. Bat. Gen. 5
(1790) art. 4, p. 8, nomen. — Spathodea indica
Pers. Syn. 2 (1807) 273. — Calosanthes indica BL.
Bijdr. (1826) 760; WiGut, Ic. Pl. 4 (1850) t. 1337-
1338; Mig. Fl. Ind. Bat. 2 (1858) 752; BUREAU,
Mon. (1864) 45, t. 9. — Bignonia quadripinnata
Bianco, FI. Filip. (1837) 499, ed. 3, t. 219. —

1977]

BIGNONIACEAE (van Steenis)

129

RRNA.

te
+o *
44
cy Sain

fe nd
“ne

4

Fig. 5. Oroxylum indicum (L.) Kurz, the ‘midnight horror’. Pole on left is a sapling that has flowered
and fruited and is temporarily leafless. Different branches of the same tree may be in leaf, flower or fruit
at the same time (Tg. Bukit, Sg. Sedili Ketchil, photogr. CorNER, June 1934),

Hippoxylon indicum RarFin. Sylv. Tellur. (1838)
78, nom. illeg. — Arthrophyllum ceylanicum Mia.
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27. — Arthro-
phyllum reticulatum BL. ex Mia. |.c., et corr. 318. —
Fig. 5, 7.

Smallish, glabrous, sparingly branched, semi-
deciduous tree, 6—-20(—27) m; trunk 10-40 cm @,
with grey bark and large leaf-scars; twigs thick, (as
the trunk at least at apex) pithy, later hollow,
lenticellate, as the leaf-rachis. Leaves tufted at
twig-ends, with a long petiole, '/,-2 m; leaflets long
petioled, ovate to oblong, acuminate, 4-11(—15) by
3-9 cm, cuneate, rounded or reniform at the trip-
linerved mostly oblique base, underneath distinctly

reticulate-veined, with some scattered gland fields
near the axils of the nerves and scattered, micros-
copical scales. Innovations of leaves and racemes
viscid. Racemes terminal, erect,'/,—1'/, m long, pith
of twig-apex, peduncle and rachis partitioned.
Pedicels long, with a few bracteoles in lower part,
2-4 cm. Calyx coriaceous, becoming almost woody
in fruit, containing water in bud, truncate or
irregularly shallow lobed by tearing, campanulate,
brown or dirty-violet, 2-4 by 1'/,-2 cm. Corolla
reddish purple to liver-brown to dirty violet out-
side, dirty yellowish to pinkish inside, with a foxy
stench, 7-10 cm long, the lobes subequal, in young
buds strongly folded into a massive apex, +

130

crisped or undulate-crenate, in flower patent to +
reflexed, outside with scattered patelliform glands,
the lobes inside with dense, almost sessile capitate
gland-hairs; basal tube wide, widened to base,
c. 1'/,cm. Stamens inserted in throat, their base
long hairy. Style 4-6 cm, dark violet as the sub-
entire, large disk. Capsule pendent, 45-120 by
6-10 cm, valves flat, almost woody, finally black.
Seeds incl. wings 5—9 by 2'/,—-4 cm.
Distr. As the genus. Fig. 6.

Fig. 6. Range of the genus Oroxylum VENT.

Ecol. As the genus. As a consequence of its
short-lived, short-sized nomad habit relatively rare
in tracts with largely high primary forest, e.g.
Borneo. In Malaya chiefly by villages and by rice-
fields (CORNER). Also not particularly common in
open but seasonally very dry tracts, such as the
Lesser Sunda Is. and in teak forest largely confined
to mixed forest stands. F/. Jan.—Dec., according to
Koorpbers in Java at the start of the dry season;
fr. July—May, the fruit remaining during the dry
season on often leafless stems.

CORNER (l.c.) gave a lively account of his obser-
vations on this grotesque treelet. He remarked:
“that each leaf develops as a unit and when it
withers it breaks up gradually in regular order from
the tip to the base: the leaflets fall off singly and
the main stalk and its side-stalks break up at the
joints: the bits accumulate round the base of the
trunk like a collection of limb-bones, so that we
may call it the ‘Broken Bones Plant’. The leaves
are crowded near the end of the stem or its
branches, and saplings, which remain unbranched
until after their first flowering at a height of some
15 ft., look like gigantic umbrellas. When the
saplings flower, the inflorescence develops from
the apical bud and therefore further upward growth
of the main stem is prevented. When the inflores-

FLORA MALESIANA

[ser. I, vol. 8?

cence has finished flowering, the leaves below it
fall off and the leafless stem is left as a pole with a
few sabre-like pods dangling from its extremity:
wherefore, we may call it the ‘Tree of Damocles’.
Then, after 3-4 weeks ina leafless state, one or more
lateral buds on the stem break out and grow into
side-branches which, in due course, flower, fruit,
shed their leaves and branch in their turn: and,
thus, the big trees are constructed sympodially
with open irregular crown and a few lanky ascend-
ing limbs. Each branch seems to flower independ-
ently of the others so that flowers, fruits and grow-
ing twigs may be found on the same tree.”

Flower biology. The flowers are nocturnal; on
each raceme 1-2 flowers open on one night.
According to CorRNER (i/.c.): “‘The corolla begins
to open about 10 p.m., when the tumid, wrinkled
lips part and the harsh odour escapes from them.
By midnight, the lurid mouth gapes widely and is
filled with stink. Before sunrise the corolla is
detached and slips off over the long style. The
flowers are pollinated by bats which are attracted
by the smell and, holding to the fleshy corolla
with the claws on their wings, thrust their noses
into its throat: scratches, as of bats, can be seen on
the fallen flowers of the ‘Midnight Horror’ next
morning.” Fig. 7.

Dispersal. The gauzy seeds slip out of the opened
pods and flit away on the breeze with the jerky
motion of a butterfly: so in noon-tide, we may call
the tree the ‘Midday Marvel’ (Corner, /.c.).

Uses. Popular with the Sundanese as a veget-
able (/alab), fresh young leaves and flowers; even
unripe capsule valves are eaten after being cooked
(HASSKARL).

In Bawean I. flowers are used against inflam-
mation of the eyes. The bitter bark is chewed in
Java for depurative purpose, especially after
delivery.

In Sarawak used for dyeing rattan of black shiny
baskets.

In West Java (Priangan) local people are con-
vinced that the tree is a protection of the house
against thieves, a superstition probably derived
from the sword-like shape of the capsules.

Vern. Midnight horror, E; Malaya: (beka)
kampong, blalai, blonglai (kaya), bulai kaju, kain,
merlai, poko bulai, Malacca; Sumatra: bolai,
Minangkabau, habreng, Atjeh, '(ka)kapung, M, :.
Sum., méngleo, Simalur I., abang-abang, Asahan;
Borneo: gimurai, Sarawak, Bidayan name; Java:
ki tongtorang, pongpor(r)ang, S, (kayu) lanang,
mungli, wongli, wungli, J, pedangan, Japara, délég,
kadjén djalér, kéok, padangan, raon, J (all once
noted), bunglo, punglo, Md; h. lema kaba, kowa,
Flores; Celebes: buli, Bantaeng, pohon padang,
Manado; karu kadang, kayu, Kutai; Philippines:
balilang-uak, pingka-pingka, p.-pinkahan, taghilau,
Tag., abang-abang, P.Bis., abong-abong, Bis.,
sakayan-bakus, Tagb., balay-uak, bunglui, Sul.,
baliuag, bungoi, C.Bis., banloi, Sub., barangau,
kamkampilan, Ilk., kampilan, Neg., maidbaid, Bik.

BIGNONIACEAE (van Steenis)

ee eeeUUUE a EEE ESS SEES UENSSE SEINE EERE ERS

Fig. 7. Oroxylum indicum (L.) Kurz. a. Top of raceme with two open flowers, * '/, b. bat arriving on a
flower, c. landed bat on a flower sucking honey (Old Bot. Garden, Univ. Malaya, Kuala Lumpur, photogr.
SoEPADMO, Sept. 1973, 9.30 p.m.).

132 FLORA MALESIANA [ser. I, vol. 8?

Fig. 8. Millingtonia hortensis L. f.a. Habit, x */2, b. detail of underside of leaflet, showing domatia, x 5, c.
flower, nat. size, d. anthers, one in CS, e. capsule, x 1/,, f. seed, nat. size (a after WALLICH, c-d after
BUREAU, b, e—~f SPANOGHE s.n. TIMOR).

1977] BIGNONIACEAE (van Steenis) 133

4. MILLINGTONIA

LINNE f. Suppl. (1781) 45, 291, non DONN, 1807, nec Roxs. 1820; K.Scu. in E. & P.
Nat. Pfi. Fam. 4, 3b (1894) 226, f. 89 j-k; STEEN. Thesis (1927) 825; Bull. Jard. Bot.
Btzg III, 10 (1928) 186. — Nevrilis RAFIN. Sylv. Tellur. (1838) 138, nom. illeg. —
Fig. 8.

Medium-sized evergreen or deciduous tree with corky bark. Leaves 2-3-pinnate,
with domatia. Thyrses lax, co-flowered, terminal. Flowers white, fragrant, noctur-
nal. Calyx small, truncate-campanulate, + 5-lobed, persistent. Corolla salver-
shaped, glabrous, with a very long, slender, basal tube at apex widening towards
the limb, limb at base short funnel-shaped, zygomorphic (+ 2-lipped), 5-lobed,
imbricate in bud. Stamens 4, didynamous, glabrous, inserted at the base of the
widened part of the tube (throat), shortly exserted, no staminode; anthers with
1 fertile cell, the other spur-like, barren; connective dorsal, swollen. Disk cup-
shaped, crenate. Capsule linear, compressed parallel to the septum, septicid-
dehiscent, valves flat. Seeds ~, thinly discoid, winged.

Distr. Monotypic. SE. Asia (India, Burma, Thailand, Indo-China, Yunnan) and Malesia: probably
native, in E. Java, Madura and Kangean Is., Lesser Sunda Is. (Bali, Sumbawa, Sumba, Flores, Timor),
and S. Celebes (SW. Peninsula; Muna I.), in many places also cultivated (Penang, Sumatra, Java, etc.) and
in dry areas running wild, may be wild also in N. Malaya (Perlis and Kedah, Corner, /.c.). Fig. 9.

Ecol. Lowland monsoon forest.

1. Millingtonia hortensis LiINNé f. Suppl. (1781)
291; Decne, Herb. Timor. (1835) 32; Span.
Linnaea 15 (1841) 326; Mia. Fl. Ind. Bat. 2 (1858)
753; BUREAU, Mon. (1864) 45, t. 8; F.-VILL. Nov.
App. (1880) 150, cult. Manila; CLARKE, FI. Br. Ind.
4 (1884) 377; K. & V. Booms. Java 1 (1894) 65;
STEEN. Thesis (1927) 826; Bull. Jard. Bot. Btzg III,
10 (1928) 187; CorNER, Ways. Trees (1940) 165;
Merr. J. Arn. Arb. 25 (1944) 316; BAcK. & BAKH.
f. Fi. Java 2 (1965) 234. — Bignonia suberosa
Roxs. Cor. Pl. 3 (1811) 11, t. 214, nom. illeg. — Big-
nonia cicutaria MART. Denkschr. K. Ak. Wiss.
Minchen 6, KI. Math. Phys. (1820) 153, t. D. —
M. dubiosa SPAN. in Hook. Comp. Bot. Mag. 1
(1835) 348, nomen. — Nevrilis suberosa RAFIN.
Sylv. Tellur. (1838) 138, nom. illeg. — Fig. 8.
Evergreen (or deciduous?) tree, 5-25m, to
30 cm @; bark corky, very rough, cracking; twigs
lenticellate. Mature leaves herbaceous, nearly
glabrous, 3—S-jugate, lower pairs pinnate, up to
1 m; leaflets ovate-lanceolate, acuminate, sinuate
or crenate, or entire, 2'/,-6 by 1'/,-3 cm; domatia
haired. Thyrse erect, 10-40cm, puberulous,
flowers fragrant, only few open at a time. Calyx
2-4 mm, teeth short, broad, obtuse, margin revo-

Fig. 9. Range of Millingtonia L. f.; delimitation in
Asia is slightly arbitrary.

lute. Corolla tube 6-8 cm by 2mm, widened to
mouth, limb 4-5cm @, lobes ovate, acute, out-
side with crateriform glands, c. 1'/, cm. Filaments
c. 10 and 14mm long; anthers 2mm, with a
small appendage at the base. Style to 8 cm long.
Stigmatic lobes ovate-acute, 1'/, mm. Capsule 30-
35 by 1'/,-1°/, cm. Seeds thin-discoid, 1'/,-3'/, by
1-1"/, cm including the wings.

Distr. As the genus. Fig. 9.

Ecol. A characteristic tree of regions subject to
annual drought (‘monsoon flora’), companion of
teak, fire-resistant by its thick corky bark and pro-

fuse capacity of suckering from roots, below 750 m
alt. In Timor common in Ziziphus stands (MEVER
Drees). F/. Jan.—Sept., mostly June.

Father ScumMurTz reported that it is in Flores not
deciduous; flowers appear at the end of the dry
season, before the first rains set in.

Uses. The soft, even-grained timber was some-
times advertized for tea-boxes but is not of high
quality. Tree sometimes used for parks or road-
sides, leaves as a poor substitute of opium in
cigarettes, sometimes received from the opium
factory in Java under the vernacular name gendje.

134

Vern. Indian cork tree, E, kurkboom, D,
kahombu, M (Sum.), amfiunan, sékar pétak, sékar
putih, J, karpoti, Kangean, kanongoh, Bali,
kétangar, Sumba, takah, Dawang lang., Timor, ai
katong inggar, takah, toka hau, Timor, ai kakassa,
Tetun lang., Port. Timor, katangka, Bug., Makas-

FLORA MALESIANA

[ser. I, vol. 8?

Notes. From Sumba the flowers have once
been noted to be red (IBoET 264), never confirmed.

MEER DreEES (Comm. For. Res. Inst. Bogor 33,
1951, 39) recorded that Millingtonia is deciduous
in the driest regions of Timor, but the scant field
notes do not confirm this.

sar, Kaulolo, Muna I.

Tribe 2. Tecomeae

ENDL. Gen. Pl. (1839) 711; FENZL, Denkschr. K. Bay. Bot. Ges. Regensburg 3
(1841) 261; B. & H. Gen. Pl. 2 (1876) 1029; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b
(1894) 209; BuREAU, FI. Bras. 8, 2 (1897) 300. — Subtribe Catalpeae DC. Rév.
Bign., Bibl. Univ. Genéve (1838) 123; Prod. 9 (1845) 203.

Capsule loculicid, the septum attached transverse to the valves. Mostly trees or
shrubs, rarely lianas, by exception with tendrils.

5. TECOMA

Juss. Gen. (1789) 139; REHDER, Mitt. Deut. Dendr. Ges. 22 (1913) 262; Britton,
Bull. Torr. Bot. Club 42 (1915) 372; URBAN in Fedde, Rep. 14 (1916) 313;
MELcuior, Ber. Deut. Bot. Ges. 59 (1941) 18-31. — Stenolobium D.Don, Edinb.
Phil. J. 9 (1823) 264; Seem. J. Bot. 1 (1863) 87; STEEN. Thesis (1927) 964; Bull.
Jard. Bot. Btzg III, 10 (1928) 217. — Tecomaria Spacu, Hist. Nat. Vég. 9 (1840)
137; FENZL, Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 266; SEEM. J. Bot. 1
(1863) 19-23; SPRAGUE, FI. Cap. 4, 2 (1904) 448; STEEN. Thesis (1927) 831; Bull.
Jard. Bot. Btzg III, 10 (1928) 193; BRUMMITT, Bull. Jard. Bot. Nat. Belg. 44 (1974)
421.

Erect or scrambling shrubs or small trees. No gland fields at the nodes. Leaves
l-pinnate, sometimes 1-jugate, or more rarely simple; leaf or leaflets incised or
serrate, densely microscopically glandular-punctate and with hairy domatia under-
neath. Pedicel with minute bracteoles. Flowers in terminal racemes or more often
raceme-like thyrses, yellow, orangish or scarlet. Calyx cupular or campanulate,
with 5, often apiculate deltoid lobes, glands scattered. Corolla tubular, with a short
basal tube, funnel-shaped, + straight or + curved, widened to the mouth, lobes
almost equal to unequal, imbricate in bud, minutely ciliate. Stamens 4, didyna-
mous, exserted or included; anther-cells divergent, often finally standing out trans-
versally, free or partly connate, sometimes hairy; Sth rudimentary. Disk cupular-
pulvinate to shallowly cup-shaped. Ovary narrow cylindric or oblong, compressed,
lepidote; ovules 2—-4-seriate in each cell. Capsule linear, -- compressed parallel to
septum; valves smooth. Seeds hyaline-winged all round, insertion punctiform.

Distr. Some dozen species in the New World, from extreme S. Arizona and S. Florida to northern
Argentina, especially in the Andes, and one species in southern Africa. Some species widely cultivated in
the tropics and subtropics and one of these locally naturalized in Malesia.

Taxon. I can see not sufficient reason to keep Tecomaria generically apart from Tecoma. It often is said
to differ by the exserted stamens and orange-red to scarlet flowers, but it has appeared that among the
many taxa of South American Tecoma (Stenolobium) there are taxa with exserted stamens and in some
the flowers are orangish. Inadvertently SEEMANN (J. Bot. 1, 1863, 19-23) also united them, but curiously
later distinguished Stenolobium (l.c. 87).

1977] BIGNONIACEAE (van Steenis) 135

According to SPRAGUE (FI. Cap. 4, 2, 1904, 448) there are only two valid characters, viz the number of
the rows of ovules in each cell (2 in Tecoma, 4 in Tecomaria) and the anthers. As to the first character, in a
dozen American genera this number varies, from 2-4, 2-6 and in Tabebuia even from 2-many; its
value seems therefore to be rather low. The second character holds: in American Tecoma the anther-cells
are completely free causing them in full anthesis to stand often perpendicular to the filament; in Tecomaria
they are connate in the upper 3rd or 4th part, so that they can not diverge so widely.

The intimate relationship between Tecoma and Tecomaria is emphasized by a reputed fertile hybrid,
x Tecoma smithii W. WATSON (see p. 118), between Tecoma velutina and Tecomaria capensis.

As the differences between the genera coincide with the geographical disjunction I am prepared to
9 at sectional level and refer Tecomaria to Tecoma sect. Tecomaria (SPACH) ENDL. Gen.

mt if

From southern Africa 3 spp. were described but F. Wuirte (For. Fl. N. Rhod. 1962, 380) and BRUMMITT
(Bull. Jard. Bot. Nat. Belg. 44, 1974, 419) distinguish only one.

In South America a thorough revision probably will also show reduction to fewer variable and raciated

species.

1. Tecoma stans (L.) H.B.K. Nov. Gen. Sp. 3
(1819) 144; DC. Prod. 9 (1845) 224; F.-VILL. Nov.
App. (1880) 151; MerR. Fl. Manila (1912) 428;
JOHNSTON, Proc. Cal. Ac. Sc. IV, 12, 2 (1924) 1166;
SANDWITH in Pulle, Fl. Surinam 4, 2 (1938)
79; CORNER, Ways. Trees (1940) 170, f. 44,
pl. 159; Back. & Baku. f. Fl. Java 2 (1965) 539;
GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 958,
f. 38, with full synonymy. — Bignonia stans
Linne, Sp. Pl. ed. 2 (1763) 871; Juss. Gen. (1789)
139; RECHINGER, Denkschr. K. Ak. Wiss. Wien 85
(1911) 356. — Stenolobium stans SEEM. Ann. Mag.
Nat. Hist. 10 (1862) 30; J. Bot. 1 (1863) 88, incl.
var. pinnata SEEM. type var.; BUREAU, Mon.
(1864) t. 13; MerRrR. En. Philip. 3 (1923) 444; STEEN.
Thesis (1927) 905; Bull. Jard. Bot. Btzg III, 10
(1928) 218.

Shrub, up to c. 1-4 m. Leaflets 1-3 pairs (cult.
sometimes 1-foliolate) lanceolate, acuminate,
serrate, glabrous, but often along midrib laxly
hairy, 3-10 by 1-4cm, cuneate at the base, no
proper petiolules; petiole 2-5cm. Racemes
glabrous, c. 5-15cm. Pedicels 5-10 mm. Calyx
campanulate, 5-7 mm, usually with some im-
pressed plate-shaped glands in middle part or
upper half, lobes short-ciliate. Corolla yellow,
3'/,-5 cm, limb up to 3'/,cm @. Stamens included,
anther-cells + pilose. Capsule acute, often lenti-
cellate, 10-22 by '/,-*/, cm. Seeds (incl. wings) 2 by
1/, cm, inserted in two rows on the margins of the
septum.

Distr. From Florida through Central and South
America to N. Argentina, widely cultivated in the
tropics, also in Malesia, and sometimes run wild,
naturalized e.g. in Tahiti, the Society Is. (Raiatea),
and the Marquesas (Nunuhiva), often together
with tree ferns and Gleichenia.

Notes. Vegetatively a rather variable species.
The normal form is with pinnate leaves, but some-
times there are 3-foliolate and even simple leaves
intermixed in one sheet. In Tahiti the specimens
have 5—6 pairs of leaflets. (This may be var. multi-
jugum R. E. Fries, Ark. Bot. 1, 1903, 401). In

exico a sheet had almost entire leaflets (Sumi-
CHRAST 1885). There is in America a form with

underneath woolly-hairy leaflets: 7. stans var.
velutina DC. (T. mollis H.B.K.), but the density of
the tomentum varies considerably in degree and I
am not very much in favour to recognize this; this
is also the opinion of STANDLEY (Trees, Shrubs
Mexico, 1926, 1319).

JOHNSTON (J. Arn. Arb. 21, 1940, 264) said that
in Mexico the normal-leaved form occurs in coastal
regions, the incised-leaved form ininland places and
the tomentose form south of these two, all three
replacing, suggesting subspecific segregation.

It is rather peculiar that, though the normal-
leaved form is widely cultivated throughout
Malesia, the only naturalized one is a fairly con-
stant form with deeply incised leaflets, which
seems to be rather rare in the Americas.

var. incisa G. Don, Gen. Syst. 4 (1838) 224;
J. K. MAHESWARI, Bull. Bot. Surv. India 3 (1961)
357. — T. incisa Sweet, Hort. Brit. ed. 1 (1827)
284, nomen. — T. stans var. apiifolium DC. Prod.
9 (1845) 224; Back. & BAKH. f. Fl. Java 2 (1965)
539. — Stenolobium stans var. apiifolium SEEM. J.
Bot. 1 (1863) 89; STEEN. Thesis (1927) 906; Bull.
Jard. Bot. Btzg III, 10 (1928) 218. — ? Stenolobium
incisum RosE & STANDLEY, Contr. U.S. Nat. Herb.
16 (1913) 174. — ? T. stans var. angustatum
REHDER, Mitt. Deut. Dendr. Ges. 24 (1915) 227.
— T. incisa (ROsE & STANDLEY) JOHNSTON, J. Arn.
Arb. 21 (1940) 264.

Leaflets (2—3—)4(—5) pairs, very coarsely toothed
to deeply pinnately incised, not rarely to the mid-
rib, making acute-triangular lobing, 5-10 by
1-—2'/, cm (incl. teeth).

Distr. Central America, widely cultivated in
the tropics, also in Malesia: naturalized in Timor,
Ternate, and SW. New Guinea (near Uta); also
naturalized in the Concan and N. Kanara (TALBOT).

Ecol. In Timor (Kupang and Baucau Plateau)
characteristic for red calcareous soils and lime-
stone, in the latter place gregarious in shrubberies,
flowering already at an early age; 5-500 m. FI.
mostly Aug.—Sept. (Oct.), fr. Oct.-Dec.

Vern. Yellow bells, E, Malaya; ai funan, Tétu
lang., E. Timor; dufa dufa, Ternate.

6. DEPLANCHEA

VIEILLARD, Bull. Soc. Bot. Normandie 7 (1862) 96; BUREAU, Bull. Soc. Bot. Fr. 9
(1862) 164; Beauvis. Gen. Montrouz. (1901) 90; STEEN. Thesis (1927) 906, f. 8-9;

136 FLORA MALESIANA [ser. I, vol. 8?

Bull. Jard. Bot. Btzg III, 10 (1928) 218, f. 2-3; Heine, Fl. Nouv.-Caléd. 7 (1976) 71,
f. 16-17. — Diplanthera BANKS & SOL. ex R.BR. Prod. (1810) 448, non THouaRs,
1806, nec SCHRANK, 1819; SCHEFFER, Nat. Tijd. N. I. 31 (1870) 332; B. & H. Gen.
Pl. 2 (1876) 1048; STEEN. Nova Guinea 14 (1927) 293. — Bulweria F.v.M. Fragm.
4 (1864) 147. — Fig. 10, 12-13.

Fig. 10. Deplanchea bancana (SCHEFFER) STEEN. a. Flower, b. fruit on thickened rachis, c. opened capsule,

seeds covering the septum, d. seed, all nat. size. — D. glabra (STEEN.) STEEN. e. Flower, nat. size, f. LS of

ovary, enlarged. — D. tetraphylla (R.BrR.) F.v.M. g. FLOwEr, nat. size (a—-d C.H.B. XIII-J—65, e-f GIELLERUP
583, g after VAN STEENIS, 1927).

Trees, with thick, pithy branches, robust in all parts. Leaves simple, in whorls of
3-4, + tufted to end of branches, entire, at base above with a few large crateriform
or saucer-shaped glands, underneath often fine-punctiform dotted and sometimes
with scattered larger crateriform glands; glabrous or with a yellow indument of
simple hairs. Thyrses terminal, erect, a thick short rachis with crowded, horizontal,
long-stalked triads or twice-forked triads. Flowers erect, showy, yellow, stalked.
Calyx on a 2-3 mm high, obconical, solid hypanthium, articulate with the pedicel,
closed in bud, with short lobes hairy at tip, in anthesis with 5 rather regular, acute
lobes or tearing into 2-5 unequal, irregular lobes, inside fine-glandular, outside
not rarely with few large crateriform glands, tip penicellate. Corolla imbricate in
bud, lobes ciliate, zygomorphous to degree, hardly with a distinct basal tube, tube
just or far exceeding the calyx. Stamens 4, didynamous, exserted, rarely a 5th
rudiment, + erect or recurved to one side, together with a style inserted shortly

1977] BIGNONIACEAE (van Steenis) 137

above the base of tube, base capitate-glandular hairy; filaments ribbon-shaped;
anther-cells free, wide-divergent. Disk annular, crenate. Ovary subsessile, glabrous,
2-celled, each cell with 2 closely placed placentas; style very long; stigma with 2
narrow lobes. Ovules o¥, in many rows. Capsule short-stalked, ellipsoid, with hard,
boat-shaped valves, erect; septum flattened, lens-shaped, thick. Seeds very many,
roundish, very thin hyaline-winged all around, punctate-inserted.

Distr. Probably 5 spp., 1 in West Malesia, 2 in New Guinea (of which | sp. also in N. Australia and the
other also in E. Borneo and Central Celebes), and 2 in New Caledonia. Fig. 11.

Ecol. Rain-forests with preference for light and secondary forest, kerangas forest, others in woodland
savannahs and invading grasslands, from sea-level to 1000 m.

Notes. The much increased collections gave a better understanding in specific delimitation and varia-
bility of characters, leading to reduction in the number of species. Especially the hairiness occurs to degree
and is occasionally deviating; in occasional specimens of D. bancana the calyx may possess dense long
hairs inside the calyx. For this reason I have reduced D. tubulosa STEEN. and D. coriacea STEEN. The
Australian D. hirsuta BAILEY I have reduced tentatively to D. tetraphylla; I believe it to be a juvenile form
which accounts for its sinuate leaf margin and occurrence of deviating phyllotaxis, decussate or whorls of 3.

For brevity’s sake the characters mentioned in the key are not repeated in the descriptions.

Specimens in fruit or in bud, or without corolla are difficult to identify.

Affinity. Deplanchea has no affinity to other Old World genera. BUREAU (Mon. 1864, 51) compared
it with the genus Delostoma from Andine South America with which it shares several characteristic
features: thick twigs, terminal inflorescences, simple leaves, and boat-shaped fruit valves. Delostoma
differs in having the valves said to be unequal, one flat, one boat-shaped, and further by triplinerved leaves,
a regular, dentate (sometimes ‘double’) stunted calyx, and pink or violet flowers.

KEY TO THE SPECIES

1. Corolla tubular, the tube + twice as long as the calyx, straight or slightly curved. Stamens and style
erect, + straight in anthesis. Leaves in whorls of 3, underneath almost always very laxly hairy on
midrib and nerves as is the petiole. Calyx lobes + equal, corolla lobes ditto : . 3. D. glabra

1. Corolla tube only for 2-5 mm exceeding the calyx, the limb distinctly zygomorphous, 2 lobes higher
connate, patent and longer than the others. Stamens and style patent-curved over this lobe or recurved.
Leaves in whorls of 3-4.

2. Branches of the thyrse triads, or flowers solitary. Pedicels 1'/,-2!/, cm long. Bud + cylindric in shape,
rarely pear-shaped, often with 5 faint ribs below the lobes. Calyx with a few to several large crateri-
form glands, the lobes + equal, in anthesis c. 1/,;-"/, as long as the tube. Corolla tube almost cylindric,
c. 10 by 5 mm. Stamens c. 3-3!/, cm. Leaves in whorls of 3 (by exception 4), hairy underneath or
glabrous bancana

2. Branches of the inflorescence often 2(—3) times forked. Pedicels !/,-2 cm. Buds pear- or spindle-shaped,
or obovoid, smooth. Calyx without crateriform glands, distinctly widened + campanulate in anthesis,
the lobes mostly unequally tearing, sometimes only 2 or 3, '/,-*/; as long as the tube in anthesis.
Corolla tube widened almost from the base, c. 1-1!/, cm high, 1'/, cm wide at the mouth in anthesis.
Stamens 4-4'/, cm. Leaves in whorls of (3—)4, always hairy underneath 2. D. tetraphylla

Se wae hoe Soe Se, See? eee VY ee es ee A ee ey a ee emer Nees 2 es ge Noe Oe We ee ee pe ee . D.

1 bancana (SCHEFFER) STEEN. Thesis

. Deplanchea orange, darker than corolla. Fruit 10-14 by 3'/, cm.
(1927) 921, incl. var. glabra STEEN. I.c. 923; Bull.

Seeds c. 3/,4cm @, incl. wings 3 by 2 cm.
Distr.

Jard. Bot. Btzg III, 10 (1928) 221, f. 2b, 3. —
Diplanthera bancana Scuerrer, Nat. Tijd. N. I. 31
(1870) 334; Hassk. Flora 53 (1870) 219; CLARKE,
Fl. Br. Ind. 4 (1884) 385; RipL. Fl. Mal. Pen. 2
(1923) 552. — D. coriacea Steen. Bull. Jard. Bot.
Btzg III, 10 (1928) 224, f. 2c-e. — Fig. 10a-d.
Small to large tree, 4-36 m; bole to 20 m; d.b.h.
15-150 cm, with small or larger buttresses; bark
finely fissured, flaky, wood soft, white. Leaves
chartaceous to coriaceous, obovate to elliptic, 9-34
by 5'/,-20 cm, apex rounded, rarely short-wide-
acuminate; base cuneate to cordate; glabrous to
yellow hairy in various degree, as is the thyrse;
petiole 3-6 cm. Peduncle 5-20 cm; rachis 2-5 cm;
primary lateral stalks of triads 2-4 cm. Calyx 12-
18 mm. Corolla tube inside at base densely capitate-
glandular hairy on insertion of stamens. Anthers

West Malesia: Sumatra (Palembang,
Asahan, Bencoolen, Indragiri, Tapanuli), Riouw
Is. (Karimon I., P. Temiang, P. Kedondong),
Malaya (also Penang I.), Banka (common),
Billiton, and Borneo. Fig. 11.

Ecol. In primary and secondary forests, in
Borneo not rare in heath forest, mostly on sandy
soils, podsols and wet kerangas, slopes of podsol
terraces, from sea-level up to 1000 m. F/. Jan.—Oct.

Vern. Méndjanbing, méngkubéng, méngkubung,
M (Banka), kayu chéndéru, Malacca, labu, Palem-
bang, méngkubong, mértapa, P. Temiang, kayu si
martim, baha, Batak, tui, M (P. Karimon, in error
with Dolichandrone?), éndjabiengien, Billiton.

Notes. A fairly variable plant. In addition to
the yellow-tomentose or velutinous haired typical
form as described by Scuerrer there occur glab-

FLORA MALESIANA

[ser. I, vol. 8?

=

Mh -. ; *! Ae, | |

e

Saeed :

< Xe
a aS e 24 fi
Fig. 11. Range of the genus Deplanchea VIEILLARD:
D. tetraphylla (R.BR.) F.v.M. (dots), D. glabra

(STEEN.) STEEN. (triangles), D. bancana (SCHEFFER)
STEEN. (line). In New Caledonia 2 endemic species.

rous forms, but in degree, with few hairs, or the
hairs still more reduced and confined to the lower
part of the petiolar groove, or only to the axillary
bud. There are also specimens of which the calyx is
inside long pubescent with 1-seriate hairs, notably
S$ 11989, 16427, 17591, 25411, SAN 32195,

ANDERSON 8398, HALLIER B 2507, VAN NIEL 4019, :

but they are hairy as the type or glabrous, or with
small glabrous leaves, rounded leaves or with short
acuminate tip. This is also the reason that I cannot
maintain D. coriacea, as the leaf-base varies from
cordate to rounded to cuneate, without correlation
with other sets of characters. Also the number of
crateriform glands varies and these are also found
in other specimens. I will not exclude the possibility
that in the field certain biotypes may be bound to
certain soil types, but I see no possibility to defini-
tions and formal recognition from the herbarium.

In ANDERSON 8398 from Sarawak the leaves are
in a whorl of 4.

Fruits are extremely scarce in the herbarium.

2. Deplanchea tetraphylla (R.BR.) F.v.M. Second
Syst. Cens. Austr. Pl. 1 (1889) 167; STEEN. Thesis
(1927) 916, incl. var. novoguineensis STEEN. l.c. 917;
Bull. Jard. Bot. Btzg III, 10 (1928) 220; Proc. R.
Soc. Queensl. 41 (1929) 55; Webbia 8 (1952) 435.
— Diplanthera tetraphylla R.Br. Prod. (1810) 449;
Bru. Fl. Austr. 4 (1869) 540; SCHEFFER, Nat. Tijd.
N. I. 31 (1870) 335; BANKS & SOL. Ill. Cook’s Voy.
2 (1901) 72, t. 229; BatLey, Queensl. Fl. (1901)
1137; Compr. Cat. Q. Pl. (1909) 368; WuiTE,
Proc. R. Soc. Queens}. 34 (1922) 52; ibid. 38 (1927)
259; LANE-PooLe, For. Res. (1925) 137; STEEN.
Nova Guinea 14 (1927) 293. — Bulweria nobilis-
sima F.v.M. Fragm. 4 (1864) 147. — D. bulwerii
F.v.M. ibid. 5 (1865) 72, (1866) 214. — ? Diplan-
thera hirsuta F. M. BAtLey, Bot. Bull. Dep. Agric.
Queensl. 14 (1896) 11; Queensl. Fl. (1901) 1137;
Compr. Cat. Q. Pl. (1909) 368. — Faradaya
chrysoclada K.Scu. & Laut. Nachtr. Fl. Schutzgeb.
(1905) 370; Beer & H. J. Lam, Blumea 2 (1936)
225, cf. LAM & MEEUSE, Blumea 3 (1938) 201. —
D. hirsuta (F. M. BAILEY) STEEN. Proc. R. Soc.
Queensl. 41 (1929) 56. — Fig. 10g, 12-13.

Tree, without buttresses, 4-25 m; d.b.h. 10 to
over 100cm; bole 1-17m; bark grey or grey-
brown, corky, furrowed and rectangular-fiaking;
wood pale straw-coloured. Leaves chartaceous to
coriaceous, usually obovate or oblong-obovate,

underneath yellow-velutinous, base somewhat
cuneate to stunted, exceptionally cordate, on the
base above with 1-7 cup-shaped large glands,
11-23(-60) by 7-14(-30) cm; petiole 2!/,-5 cm.
Peduncle 4-12 cm; rachis 3-9 cm; branches 2-71/,
cm; pedicels 1-2 cm. Calyx 12-14 mm. Fruit 5-11
by c. 2!/, cm. Seeds incl. the wings 2 by 1*/, cm.

Distr. NE. Queensland (incl. Thursday L.,
Fitzroy I.) and East Malesia: New Guinea and the
Aru Is. (Trangan and Wokam Is.). Fig. 11.

Ecol. Predominantly in the periodically dry
belts of New Guinea, also in gallery forest, very
rarely in rain-forest, almost confined to grassland
and wooded savannahs and associated with
Eucalyptus tereticornis (Central Distr.) or Mela-
leuca, but also in mixed savannahs (Antidesma,
Schefflera, palms, etc., at Merauke), not rarely
common, also a pioneer in fired areas, from sea-
level to c. 600 m, rarely at 1200 m (Mafulu). Fi.
May-Oct., fr. July—Oct., often flowers and fruits
together, but fruiting specimens very rare in the
herbarium. Dwarf specimens may in places flower
and fruit.

C. J. Srerers (Verkenningsrapport Berari
Komebwaller. Mimeo, Fak Fak, 1956, p. 6, 7, 10,
phot. 2) reported D. tetraphylla from sandy soils,
often inundated through an impervious subsoil in a
heathy forest of Melaleuca.

Field notes: style greenish yellow, filaments
yellow, anthers brown. According to VAN ROYEN
the flowers have a sourish-sweet scent and are
eaten by ‘luries’, lorikeets (at Merauke).

Uses. At Fak Fak the timber is used for prahus
by the Papuans.

Vern. Laargola, Trangan, Aru Is., kapul, M,
Merauke, bas, Sorong, Mooi lang., tembako d’ora,
Fak Fak, Ersania lang., pwan, Mumuni, Orokaiva
lang., pakawa, Maipa, Mekeo lang.

Notes. The phyllotaxis is not constant, several
specimens have whorls of 3, reminding of D.
bancana with which this species is closest related.
In 3m high saplings (DOCTERS VAN LEEUWEN
n. 38, cultivated in Hort. Bog. sub XVI.I.F.8) all
leaves were opposite.

Similarly saplings in Queensland may have
opposite leaves and, moreover, narrow oblong to
lanceolate leaves with wavy, even toothed margin
(L. S. SmitH 12382). Such plants were described
as D. hirsuta and may precociously flower; also
very small normal-leaved specimens may flower,
obviously at an early age, in New Guinea, possibly
stimulated by open, pyrogenous habitat. Such
specimens may also sucker.

3. Deplanchea glabra (STEEN.) STEEN. Thesis
(1927) 919, f. 8f, 1; Bull. Jard. Bot. Btzg Ili, 10
(1928) 225. — Diplanthera glabra STEEN. Nova
Guinea 14 (1927) 293. — D. tubulosa STEEN.
Thesis (1927) 926, f. 8g, k, m; Bull. Jard. Bot.
Btzg III, 10 (1928) 226. — Fig. 10e-f.

Tree, 11/,-22 m; d.b.h. 12-60 cm; bole 3-12m;
bark grey, scaly; mostly + glabrous in all its parts.
Leaves obovate-oblong to elliptic-oblong, coria-
ceous, usually very laxly haired on the midrib (and
nerves) below, very rarely yellow short-velutinous
on inflorescence, midrib, nerves and petiole, fine
dark-dotted beneath, 9-40 by 4'/,-25 cm, rarely
with some scattered larger glands; base rounded to

BIGNONIACEAE (van Steenis)

Fig. 12. Deplanchea tetraphylla (R.Br.) F.v.M. Inflorescence from above, capsules, partly opened, showing
dissepiment (fl. photogr. HOOGLAND 4249; fr. photogr. WomERSLEY, 1956, Sogeri).

140 FLORA MALESIANA [ser. I, vol. 8?

Fig. 13. Rather young tree of Deplanchea tetraphylla (R.BR.) F.v.M. in Bot. Garden Lae (photogr. M.
GALORE).

1977]

BIGNONIACEAE (van Steenis)

141

cuneate or subcordate, apex rounded; petiole
21/.-7 cm. Peduncle 3-7 cm; rachis 1-11 cm; triads
1/,-2 cm stalked; pedicels 4-15 mm. Bud spindle-
shaped to obovoid. Calyx glabrous, very rarely
haired on the mid-sepaline ribs, pustular towards
apex but the pustules hardly ever opening as
crateriform glands, 15-17 mm long, lobes 5 sub-
equal, 3-6 by 2'/.-5 mm, c. 21/,-3'/, times as
short as the tube. Corolla tube inside near the
stamens capitate-glandular hairy. Placentas in each
cell 2, nearly confluent. Capsule 6-9 by c.2-2'/, cm.
Seeds incl. wings 11/, by 1 cm.

Distr. Malesia rather common in North New
Guinea in the vicinity of Hollandia and Mt
Cyclops, also found twice in Central Celebes
(Malili, Tobela, Palopo) and once in E. Borneo
(Mt Nijapa, Kelai R., Berau). Fig. 11.

It is remarkable that the species is obviously
never found in other parts of New Guinea.

Ecol. Both in high forest and in savannah, but

flowering and fruiting already when small, 10-700
(—1000) m.

Field notes: pedicels red, filaments and style
yellowish green, anthers orange, glands on leaf-
base orange. No buttresses. Fi. fr. March—Oct.

Vern. Celebes: momo, Malili, kKalambutoh,
Toradja.

Notes. Although this species is usually almost
glabrous, except for some lax hairs on the midrib
beneath, KOSTERMANS & SOEGENG 444, from Hol-
landia, has very hairy inflorescences, midrib and
petioles, so that obviously the indumentum may
vary as it does in D. bancana.

Also the single Bornean specimen known so far
(KOSTERMANS 21491) is similarly hairy; it was found
on a mountain ridge at 1000 m alt.

The two specimens from Central Celebes I refer
to this species, although one is in fruit and the
other in bud, because: the calyx is + regularly
lobed, with a few pustules but without crateriform

more commonly in pyrogenous _ grassland
(Gleichenia-Ischaemum), aS a _ pioneer, often

glands and the small pod and seed do not match
D. bancana; both spp. have leaves in whorls of 3.

7. DOLICHANDRONE

(FENZL) SEEM. [Ann. Mag. Nat. Hist. III, 10 (1862) 31, nomen; J. Bot. 1 (1863) 226,
nomen] J. Bot. 8 (1870) 379, nom. cons.; K.SCH. in E. & P. Nat. Pfl. Fam. 4, 3b
(1894) 240, f. 92B-D; SpraGuE, Kew Bull. (1919) 303; STEEN. Thesis (1927) 928;
Bull. Jard. Bot. Btzg III, 10 (1928) 227. — Pongelia RaFIN. Sylv. Tellur. (1838) 78,
nom. rejic. — Dolichandra sect. Dolichandrone FENZL, Denkschr. Bay. Bot. Ges.
Regensb. 3 (1841) 265. — Fig. 15-16.

Trees with 1l-pinnate leaves (or scattered simple leaves, extra-Mal.), leaflets
entire (or serrulate, extra-Mal.). Flowers in few-flowered terminal racemes, salver-
shaped, white, fragrant, nocturnal. Calyx not articulate, closed in bud, later
spathaceous, caducous. Basal tube of corolla long, narrow-cylindric, upper part
inflated, lobes mostly crisped. Stamens 4, didynamous, 5th rudimentary, inserted at
the throat; anther-cells divergent. Disk annular. Ovules ~ in 4-6 rows, inserted
on 2 placentas in each cell, on the septum. Capsule elongate subcylindric to +
compressed, septum very narrow, false septum very broad and parallel with the
valves. Seeds hyaline-winged, in the Mal. sp. rectangular with thick corky wings.

Distr. Spp. 9, one in tropical E. Africa, 4 in tropical SE. Asia, 3 in tropical N. Australia, and one
ranging widely from Malabar through Indo-Malesia to New Caledonia. Fig. 14.

Ecol. Allinland species, except the wide-ranging D. spathacea which is a back-mangrove tree.

Typif. In the Code and in Ind. Gen. D. spathacea (L.f.) K.Scu. has been accepted as the type, following
SEEMANN (J. Bot. 1, 1863, 226). However, the lectotype must be chosen from the original materials
incorporated by Fenzt in Dolichandra sect. Dolichandrone, elevated by SEEMANN to generic rank. He
referred to Spathodea b of ENDLICHER and Spathodea R.Br. Under the first reference D. spathacea is not
represented, at most a Dolichandrone represented by Bignonia spathacea (non L.) sens. Roxsp. Corom.
t. 144, a wrong identification for D. falcata (WALL. ex DC.) Seem. (cf. SPRAGUE, Kew Bull. 1919, 308).
This then must be the type of the genus.

Taxon. The Australian species are in habit very different from the African and Indo-Malesian ones:
characteristic shrubs or small trees of xerophytic habit and xeromorphous structure, the leaves sometimes
obviously not decussate, coriaceous, sometimes entire, with fine parallel ascending veins, or leaflets even
needle-like. In addition I find the pods terete and the pseudoseptum not flat, but irregularly, corky swollen
with impressions of the seeds. According to URBAN (Ber. Deut. Bot. Ges. 34, 1916, 755) these species
would also be different in pollen from the other species. I have arranged them in subg. Coriaceae STEEN.
es 1927, 931, f. 10). They should probably be better arranged in a separate section, rather than in a
subgenus.

142

FLORA MALESIANA

[ser. I, vol. 8?

Fig. 14. Range of the genus Dolichandrone (FENZL) SEEM. and its species, the largest range being that of
D. spathacea (L. f.) K.ScH., a mangrove tree with buoyant seeds.

Three spp. are distinguished; D. filiformis (DC.) F.v.M. is a fairly constant one, with 3-5 filiform
leaflets, but D. heterophylla (R.BR.) F.v.M. is very variable, with simple to pinnate leaves (3-7 leaflets) also
varying in width, whereas D. alternifolia (R.BR.) SEEM. with ovate, simple leaflets shows a tendency to split
the leaf. Already SEEMANN (J. Bot. 8, 1870, 382) remarked that the latter two probably belong to one
variable species, which opinion I now tend to share; he accepted the epithet heterophylla.

1. Dolichandrone spathacea (L. f.) K.Scu. FI. Kais.
Wilh. Land (1889) 123; Merr. Fl. Manila (1912)
429; Int. Herb. Amb. (1917) 469; Sp. Blanc. (1918)
349; SPRAGUE, Kew Bull. (1919) 304; STEEN.
Thesis (1927) 937; Bull. Jard. Bot. Btzg III, 10
(1928) 227; C. T. Wuite, J. Arn. Arb. 10 (1929)
265; MerRR. Comm. Lour. (1935) 355; CORNER,
Ways. Trees (1940) 163, Atlas pl. 26-27; HEINE,
Fl. Nouv.-Caléd. 7 (1976) 81, pl. 18. — WNiir
Pongelion RHEEDE, Hort. Mal. 6 (1686) 53, t. 29. —
Lignum equinum RumpH. Herb. Amb. 3 (1750) 73,
t. 46. — Bignonia spathacea L. f. Suppl. (1781) 283;
REtTz. Obs. Bot. 5 (1788) 5; BLANco, FI. Filip.
(1837) 499. — Bignonia longissima Lour. FI.
Coch. (1790) 380, nom. illeg., non JAcQ. 1760. —
Bignonia javanica THUNB. Mus. Nat. Ac. Upps. 17
(1794) 150, nomen; Fl. Ceil. (1825) 7, nomen, cf.
STEEN. Blumea 6 (1950) 359. — Spathodea longi-
flora VENT. Choix (1803) 40; SPAN. Linnaea 15
(1841) 326. — Spathodea rheedii SPRENG. Syst. 2
(1825) 835, quoad syn.; WALL. Cat. (1832) n. 6516;
DC. Prod. 9 (1845) 206; Mia. FI. Ind. Bat. 2 (1858)
754. — Pongelia longiflora RaFin. Sylv. Tellur.
(1838) 79. — Bignonia longiflora WILLD. ex DC.
Prod. 9 (1845) 206. — Spathodea loureiriana DC.
l.c. 209. — Spathodea luzonica BLANCO, FI. Filip.
ed. 2 (1845) 350; ed. 3, 2 (1878) 284, t. 242. —
Spathodea diepenhorstii Miaq. Fl. Ind. Bat. 2 (1858)
754. — D. rheedii SEEM. J. Bot. 8 (1870) 380; Kurz,
Fl. Burma 2 (1877) 234; CLARKE, FI. Br. Ind. 4
(1884) 379; K. & V. Bijdr. (1894) 69; Gams -e, J.
As. Soc. Beng. 74, ii (1905) 377; Rip. Fl. Mal.
Pen. 2 (1923) 549. — D. longissima K.Scu. in E. &
P. Nat. Pfi. Fam. 4, 3b (1894) 240. — Fig. 15-16.
For fuller references see STEEN. (1928).

Evergreen, glabrous tree, 5-20 m; 10-40cm @;
wood soft, white. Leaves usually 3-4-jugate,
15-35 cm, stalked, in the herbarium nigrescent as
all other parts; young parts + viscid, young leaves
slightly pinkish in the field (CORNER); leaflets thin,
ovate-oblong to lanceolate, unequal-sided, entire,
long-tipped (in seedlings sometimes toothed), 6-16
by 3-7cm, underneath with hairy domatia.
Racemes 2-8-flowered. Rachis 2-3cm. Bracts
caducous. Bracteoles 0. Pedicels 2-4 cm. Flowers
not articulated. Calyx conical, coriaceous, usually
arcuate, beaked, circumscissile caducous, with
many microscopical glands and a field with large
crateriform glands at apex, 3—6(—8'/,) cm. Corolla
tube 12-18 cm long, the mouth 7-12 cm @; basal
tube 9-12 cm, gradually funnel-shaped expanded
above the throat for 4cm; lobes 5, broad, sub-
equal, frilled round the edge, with large glands,
2'/,-3 cm. Stamens not exserted. Style exserted.
Capsule flattened-cylindrical, + ribbed, straight or
+ arcuate, or twisted, tipped, 25-70 by 2-3 cm;
valves hard leathery, pseudoseptum flattish, hard
corky, c. 11/,-13/,cm wide. Seeds dark grey,
rectangular, in many rows, 12-18 by 68mm
including the thick corky wings; attachment a fine
line, 8-10 mm long.

Distr. From the coast of Malabar throughout
tropical SE. Asia and the whole of Malesia to New
Guinea, Micronesia (W. Carolines: Korror; Yap:
Tomil I.), the Solomons, the New Hebrides and
New Caledonia, not found in Australia and Poly-
nesia. Fig. 14.

Ecol. Confined to the back-mangrove and banks
of tidal rivers and estuaries. RIDLEY (Kew Bull.
1910, 203; J. As. Soc. Str. Br. 59, 1911, 40, 146)

BIGNONIACEAE (van Steenis)

143

Fig. 15. Dolichandrone spathacea (L. f.) K.Scu. Flowers and twigs in bud and fruit, magnification */;;
upper capsules opened (photogr. CORNER).

recorded it common in low-lying rice-fields near
Kanga village, Lower Siam, as the predominant
tree, which he ascribed as “‘relics of the time when
this whole country was a tidal swamp, gradually
filled up after the disappearance of the sea”. Other
seashore plants were also found in these paddy
fields, such as Euphorbia atoto. This inland occur-
rence is also stressed by Corner (/.c. 164) who
found it “frequently in coastal rice-fields; in Perlis
it is indeed a feature of the country; also in North
Kedah, as soon as one reaches Kodiang it attracts
attention, standing in the paddies as an upright
poplar and flanks the roads which lead to Kangar
and Singgora. Old tree trunks are massive and
fluted at the base, the crown tapering upward. The
old, opened twisted pods remain for a long time on
the tree.”

Brass found it very abundant in Daru I. (S. New
Guinea), while K. J. Wire found almost pure
stands in Umboi I. (Morobe Distr.) in swamps
behind the mangrove. He recorded it also from
freshwater swamps in the Markham Valley. At the
Bogor Botanic Gardens it is successfully cultivated
in freshwater.

The calyx is filled with water in bud. The very
young inflorescence and developed ovary is often
slightly glossy varnished in the herbarium, similarly
as is found in Radermachera, certainly by the exu-
date of glands which are found at the apex of the
calyx. Fl. fr. Jan.—Dec., flowers and fruits not rarely
found together. Koorpers (1894 /.c.) and HEYNE
(Nutt. Pl. 1371) say that in the dry season it may be
at times nearly leafless fruiting in Central and East
Java.

144

FLORA MALESIANA

[ser. I. vol. 8?

Fig. 16. Trees of Dolichandrone spathacea (L. f.) K.Scu. in the coastal rice-fields of Perlis where it is a
feature of the country (photogr. CORNER).

Pollination. Flowers open at dusk and drop
before sunrise; they must be pollinated by hawk-
moths with very long tongues to attain the honey.
In each inflorescence one flower is open at a time
(CORNER).

Dispersal. The corky seeds float readily and must
be dispersed by seawater. In this respect it is strange
that the species is not found in northern Australia
and Polynesia.

Uses. Of little use other than fire-wood; in
N. Borneo a collector deemed the wood useful for
making clogs and matches; in the Carolines (Koror
I.) leaves and fruit are said to be used as a sub-
stitute for betel leaves in chewing. HEYNE (Nutt. Pl.
1371) said that the wood is not durable, but light
and easy to work for small things in the house;
pieces of branches are sometimes used for floats of
fishing nets in East Java and the Karimon Djawa
Is.; in the Minahassa it is used for scabbards, in
Madura I. for masks for the topéng. In Madura a
cold concoction of the leaves is also used against
mouth sprew. RUMPHIUs said that in Ambon twigs
of lignum equinum (translation of kaju kuda) were
used for making hedges.

Vern. Malaya: poko kulo, tuj, M; Sumatra:
tuwé-éj, Atjeh, kudo-kudo uwi, Simalur, kuda kuda,
Pariaman, ki arak, Palembang; Java: kaju or ki
djaran, M, djarang, S, djaram, djaran pélok,
djaranan, kadjéng kapal, kaju pélok, kapal, J,
kadju djharan, kaju djaran binék, Md; Borneo:
kélaju, tuwi, Kutai, toi, tui, Brunei, Bajau lang.,
towi, Kedayan; Celebes: fojet, kaju pélumping,
sangi, tomana, Minahassa; Talaud Is.: sansarangi;
Alor: bombila; Ternate: djodjamé; Tidore: djamé;
Ambon: kaju kuda, kati kati; Philip.: tua, tui,
Tag., pata, Ilk., tafgas, Tagb., tanhas, C. Bis.,
tanghas, P.Bis., tewi, Mbo., tiwi, Tag., Bik., C.Bis. ;
New Guinea: rie, Holtekang, Wembie lang., pide,
pier, S. New Guinea, Asmat lang., aisumbu, Mani-
kiang lang., asember tiy, Oransbari, Hatam lang.,
daud, Cape Vogel, Wanigela; New Britain: /atiu,
W. Nakanai, favituviti, Gazelle Pen., Boava lang.;
Solomons: kwae kwaele, Guadalcanal, kwe kweale,
Malaita, kwe’ekwe’eali, Kolombangara, ririge,
Small Nggela, Kwara’ae lang.

Note. Especially leaflets of saplings may, at
times, show some serrulations on the margin and
may also be slightly hairy.

1977] BIGNONIACEAE (van Steenis) 145
hh EEE
8. STEREOSPERMUM

Cuam. Linnaea 7 (1832) 720; A.DC. Prod. 9 (1845) 210; B. & H. Gen. Pl. 2 (1875)
104, pro sect. Eustereospermum ; STEEN. Thesis (1927) 946; Bull. Jard. Bot. Btzg III,
10 (1928) 233; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 68. — Hieranthes RAFIN.
Sylv. Tellur. (1838) 79. — Dipterosperma HASSsK. Flora 25, 2 (1842) Beibl. 1, p. 28;
Cat. Hort. Bog. (1844) 152; Pl. Jav. Rar. (1848) 507. — Fig. 17, 19.

Fig. 17. Stereospermum personatum (HASSK.) CHATTERJEE. a. Habit, b. capsule, both x */,, c. seed, nat.
size (a after WicHT, Ic. 4, t. 1341, b-c BeuseKom & GEESINK 3662).

146 FLORA MALESIANA [ser. I, vol. 87

Rather tall, deciduous trees. Leaves 1-pinnate; leaflets a few pairs, diminishing
in size downwards, underneath usually with flat, dish- or cup-shaped glands or
glandular spots; no domatia (in Mal.). Thyrses well-branched, paniculiform, mostly
terminal, or on old wood. Flowers fragrant (in Mal.). Calyx usually short-lobed.
Corolla infundibuliform; basal tube mostly concealed in the calyx, upper part
usually funnel-shaped; mouth bilabiate, upper lip 2-, lower 3-cleft, lobes subequal,
rounded, crisped, toothed or laciniate. Stamens 4, didynamous, included, 5th
rudimentary; anthers glabrous, cells divergent. Disk cupular to annular. Ovary
cells each with 2 rows of many ovules. Capsule long linear, terete, mostly twisted,
usually 4-angular in section; septum thick, corky, terete, with alternating notches
to fit the thick seeds which appear in two rows; valves coriaceous. Seeds o, thick,
trigonous, wedge-shaped, with a cross-groove, on both sides thinly winged; coty-
ledons folded, 2-lobed, radicle straight.

Distr. Over a dozen spp., in tropical Africa and Madagascar, in SE. Asia as far east as Yunnan, in
Malesia: 2 spp. in Malaya, possibly also in Sumatra, and a doubtfully indigenous record of a third in
East Java. Fig. 18.

Ecol. Largely confined to regions subject to a seasonal climate, all in the lowlands, in everwet rain-
forest obviously deciduous and flowering after a dry spell.

Note. Besides the 3 Malesian spp. distinguished here, P. Dop mentioned S. cylindricum PIERRE from
Malaya (Fl. Gén. I.-C. 4, 1930, 582), but this must rest on an error as that species is only known from
Thailand and Indo-China (cf. SANTISUK, Thai For. Bull. Bot. 8, 1974, 22).

KEY TO THE SPECIES

1. Inflorescence (incl. flowers) viscid-hairy by patent capitate-glandular hairs. Leaves not glabrous.
eta tube gradually funnel-shaped widened, straight, Stamens glabrous at base. Capsule not
4-ridged.

2. Leaflets 0-5 mm stalked, base cuneate-attenuate. Capsule faintly 3-ridged on each valve, c. 15-18 mm

@, septum 8-13 mm @. Calyx campanulate, 5-7 mm. Corolla dull purple, yellow-streaked within,
c. 3cm (stretched); lobes crenulate; tube c. 13/,cm. Filaments towards the insertion with small
Seanlar glands. cot o Matas eee cee eee as ea eae Sea eee 2. S. chelonoides
2. Leafiets 5-11 mm stalked, rounded to subcordate at the base. Capsule only with 1 median ridge on
each valve, 8-12 mm @, septum 6-9 mm @. Calyx + cylindric, 8-12 mm. Corolla pale lilac, 6-7 cm;
lobes deeply fringed- -laciniate; tube c.4cm. Filaments glabrous ....... 1. S. fimbriatum

1. Inflorescence and leaves glabrous (or very rarely minutely puberulous). Corolla c. 3 cm, suddenly
widened and curved above the basal tube, yellowish, the limb with reddish veins and markings, lobes
crenulate. Throat and base of filaments densely hairy. Leafiets 5-15 mm stalked, long-acuminate.
Calyx campanulate, 4-5 mm. Capsule 4-ridged, 8-10 mm @; septum 4-5mm @ . -3. S. personatum

1. Stereospermum fimbriatum (WALL. ex G. Don)
A.DC. Prod. 9 (1845) 211; Kurz, Fl. Burma 2
(1877) 231; CLARKE, Fl. Br. Ind. 4 (1884) 383;
GaMBLE, Man. (1902) 516; J. As. Soc. Beng. 74, ii
(1905) 378; Rip. Fl. Mal. Pen. 2 (1923) 550;
STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 234;
Dop, Fl. Gén. I.-C. 4 (1930) 578; CoRNER, Ways.
Trees (1940) 172, pl. 33; CHATTERJEE, Bull. Bot.
Soc. Beng. 2 (1948) 69; SANTISUK, Thai For. Bull.
Bot. 8 (1974) 23. — Bignonia fimbriata WALL.
(Cat. (1832) n. 6500, nomen] ex G. Don, Gen.
Syst. 4 (1838) 221. — Fig. 19.

Very upright, deciduous tree, to 27-30m by
30-160 cm @; crown narrow, cylindrical, rather
: open; bark light grey, rough and flaky; young
Fig. 18. Range of the genus Stereospermum CHAM. _ leaves purple or violaceous. Leaves 30-75 cm, with
In Asia and Africa generalized, and numbers of rather sticky hairs, stalks yellowish; leaflets
species an approximation. Occurrence in Sumatra (2-)3(-4) pairs, rounded at the asymmetric base,

and Java doubtful. ovate-oblong, long-tipped, 8-16 by 3-7cm;

BIGNONIACEAE (van Steenis)

147

Fig. 19. Stereospermum fimbriatum (G.Don) A.DC.
Its slender habit; some trees deciduous; on the
churchyard in Malacca (photogr. CORNER).

petiolules 5-10 mm. Flowers in large spreading
viscid-pilose clusters, 8-30cm 2, on the bare
twigs or with the new leaves. Calyx tubular, with
5 very short pointed lobes, c. 8-12 mm. Corolla
dull white to pale pink or pale pinkish lilac, narrow
funnel-shaped, without a distinct basal tube, the
tube 4-5 cm long, the lobes beautifully long-fringed
(as a dainty night cap), c. 2 cm. Filaments glabrous,
inserted at c. 11-13 mm from the base. Capsule
more or less quadrangular, snake-like twisted,
35-60 cm by 8-12 mm; septum c. 5mm @. Seeds

c. 21/, cm long, 7mm wide, with rather thick wings.

Distr. Continental SE. Asia (Burma; Thailand:
Chiengmai to Peninsular Thailand); in Malesia:
Malay Peninsula (incl. Langkawi, Penang, and
Tioman Is.), possibly also in Sumatra.

The record from Sumatra rests on an un-
published, beautiful plate in a collection of
RAFFLES in the India Office Library & Records
(NHD 49/20), which might have been made in
Bencoolen, but according to Mr R. DEsMoND the
provenance is uncertain. It might be rare in
Sumatra, similarly as S. personatum.

Ecol. In the lowland and hill forests, in Burma
up to 1000 m, in Malaya in high forest and open
country: frequent in villages and belukar from
Malacca to Perlis and Kelantan, often on rocky
coasts and headlands. Fi. Febr.—June, fr. March-
Nov. “After the first spell of dry weather the leaves
are shed and the flowers appear on the bare
boughs in delicate clusters until the new foliage is
mature; in the early morning the corollas spin down
like snow-flakes and carpet the ground with pale
lilac blossom. There are many trees in the Christian
Cemetery at Malacca, and a fairer one for a grave-
yard would be hard to come by” (CORNER, /.c.).

Uses. A hard and durable fairly large timber
rather dark coloured, used for beams and posts and
said to be durable in the soil.

Roots and leaves are used medicinally for some
minor ailments: juice of leaves is dropped into the
ear for ear-ache; leaves pounded with lime are
applied to the skin for itch; a decoction of roots is
given as a protective medicine after childbirth
(BURKILL, Dict. 1935, 2082).

Vern. Malaya: chac(h)a(h), chachar, chechar,
chicha(r), M, lempoyan, beka(k) (BURKILL), snake
tree, E (CORNER).

2. Stereospermum chelonoides (L. f.) A.P.DC.
Bibl. Univ. Genéve II, 17 (1838) 125, pro comb.,
excl. ref. RHEEDE t. 26; A.DC. Prod. 9 (1845) 210,
pro basion.; HAINES, Kew Bull. (1922) 121, in text;
SANTISUK, Kew Bull. 28 (1973) 176; non auct. al. —
Bignonia chelonoides L. f. Suppl. (1781) 282, pro
typ., excl. ref. RHEEDE; non auct. al. — Bignonia
suaveolens Roxs. FI. Ind. ed. Carey 3 (1832) 104. —
Tecoma suaveolens G. Don, Gen. Syst. 4 (1838) 224.
— Hieranthes fragrans RaFIn. Sylv. Tellur. (1838)
79, nom. illeg. — S. suaveolens A.DC. Prod. 9
(1845) 211; WiGurt, Ic. 4 (1848) 9, t. 1342; Kurz,
Fl. Burma 2 (1877) 231; CLARKE, FI. Br. Ind. 4
(1884) 382; TrIMEN, FI. Ceyl. 3 (1895) 284;
GAMBLE, Man. (1902) 515; BRANDIS, Ind. Trees
(1906) 495; Haines, Fl. Bihar Orissa (1922) 656;
Beumée, Fl. Anal. Onderz. (1922) 33; GAMBLE,
Fl. Madras 2 (1924) 998; STEEN. Thesis (1927) 948,
incl. f, verticillatum STEN. l.c. 950; Bull. Jard. Bot.
Btzg III, 10 (1928) 236; Dorp, Fl. Gén. I.-C. 4
(1930) 588; CHATTERJEE, Bull. Bot. Soc. Beng. 2
(1948) 70; BACK. & BAKH. f. FI. Java 2 (1965) 540.

Deciduous tree, up to 30m, 80cm @; timber
dark, hard. Innovations viscid hairy. Leaves
opposite (rarely in whorls of 3), 30-50 by 15-25 cm;
leaflets 3—4 pairs, viscid-hirsute, glabrescent, rough
above and brittle when mature, ovate to obovate to
broadly oblong, acute to short-acuminate, entire
or fine-dentate, 523 by 3-10 cm; glandless or with
a few scattered spots; midrib finally puberulous

148

FLORA MALESIANA

[ser. I, vol. 8?

above, venation sparsely hirsute beneath; petiolules
thick, 2-3(-5) cm. Thyrse up to 25cm @, viscid-
pubescent with capitate-glandular hairs. Flowers
dull crimson to dull purple, yellow streaked within,
very fragrant. Calyx campanulate, viscid pubescent,
6-8 mm, shortly acutely 5-lobed. Corolla viscid-
hairy, 2!/,-3 cm long, the tube rather gradually
funnel-shaped, mouth long pubescent; lobes
subentire, + as long as the tube. Filaments in-
serted at 4-5 mm from the base, towards the inser-
tion with small sessile granular glands. Ovary
4-ribbed, sometimes sparsely glandular. Capsule
smooth or valves obscurely 3-ribbed, to 45 by
14/,-13/, cm; valves woody; septum 8-13 mm @.
Seeds 3/, by 3/4 cm.

Distr. Widely distributed in continental tropi-
cal SE. Asia, from Ceylon and the Deccan to
Assam and Burma, not yet reported for Thailand
and erroneously so for Indo-China (SANTISUK,
l.c.); in Malesia: very locally found in East Java,
but somewhat doubtful whether native.

BeuMEE /.c. recorded this tree for the first time
for East Java, where it was found locally in some
places in the (teak) forest districts S. Surabaya and
E. Tuban. He suggested that this occurrence would
fit the theory of a number of forest officers of early
import by Hindus of teak and some associated
trees (Butea monosperma, Schleichera oleosa, etc.)
and several other plants. I certainly agree that in
the Hindu period (roughly 800-1400 A.D.) plants
have come from India, especially those favoured
for sacred purposes; for example Cochlospermum
religiosum (L.) ALSTON, and others went to India
vice versa, as for example Santalum album L. (see
C. E. C. FiscHer, J. Bomb. Nat. Hist. Soc. 40,
1938, 458-467). The first is still only found near
Hindu temples in Bali and the latter is still spread-
ing in India. The disjunction between the localities
in East Java and India-Burma is in these cases
certainly caused by intentional dispersal by man in
historic time.

There are, however, a large number of other
plants showing this same disjunction, and all bound
to a seasonal climate, that is, subject to a distinct
annual period of drought. In a succinct analysis I
found these to belong to 4 classes (Hand. 8th Ned.
Ind. Natuurwet. Congr. Surabaya 1938, 1939,
408-409). Later I have further elaborated this
problem and tried to solve it (Reinwardtia 5, 1961,
420-429, maps 1-6). From this it appeared that the
ecological disjunction of the seasonal climate
between the colossal area it covers in SE. Asia
(south as far as Tenasserim) and a similar ecology
in Central & East Java and the Lesser Sunda Is. is
shared by a homologous plant-geographical dis-
junction of many hundreds of plants which do not
occur in everwet West Malesia, or only in very local
seasonal spots in Celebes and the Philippine
Islands. A fair number extend their range south-
eastwards to Australia. This proves that such pat-
terns are quite natural; I have assumed they
originated during the Pleistocene Glacial Period,
which created a temporary pathway for drought
plants between SE. Asia and Australia, to vanish
in the Late Holocene.

It could thus well be that also S. chelonoides does
occur in the native state in the East Javanese teak
forest. As a matter of fact no fruit has yet been
collected, although flowering was abundant. I

cannot subscribe to the opinion of BEUMEE that its
dispersal is here by vegetative means, because I
cannot well see by what vegetative means and
furthermore because it is difficult to see how it
would have maintained itself vegetatively in this
way for many centuries. On the other hand the
existence of a Javanese vernacular name is no
argument that it is native; experience tells us that
such names are often invented quickly. If it is
native, it remains curious that, though it is obvious-
ly of rare occurrence, it was only recently dis-
covered. It cannot be disproved, however, that its
seed was inadvertently introduced by the Forest
Service with teak seed from India or Burma.

Ecol. Seasonal forest and savannahs. FI. Sept.-
Oct. (India: April-June), fr. (Asia) Nov.—Dec.

Uses. GAMBLE, /.c. 516, said it is in SE. Asia
rather an important large tree by its durable timber
which is easy to work and good for building,
though the amount of heartwood is small. It also is
an excellent fire-wood and makes good charcoal.
The root and bark are used as a favourite tonic
native medicine. It also is important in sylviculture
for its very free seed reproduction; the fruit remains
long unopened on the tree and seed gets dispersed
at the very end of the hot season after the danger of
fire is nearly over, and can germinate with the first
rains. Even on exposed slopes and among grass its
good natural reproduction is noticeable.

BurRKILL (Dict. 1935, 2082) mentions that it
yields a gum of the tragacanth class. He also men-
tioned that it is referred to as a plant of magic in
Sanskritic India, patala, being the Sanskrit name,
of which modern vernaculars have been derived.
In this respect it is noteworthy that the Javanese
name ‘bedali’ is a name for Radermachera spp.
TRIMEN reported it in Ceylon planted near Buddhist
temples.

Vern. Djati tékén, kaju tékén, J.

Notes. In East Java one specimen had leaves in
whorls of three. Leaves of saplings and suckers may
show a serrate-dentate leaf margin. Such leaves are
sometimes also rather narrow and acuminate;
those of mature trees are broader and more wide
at apex.

Under S. personatum I referred to the lamentable
name change caused by erroneous interpretation of
the type of Bignonia chelonoides L. f. The first to
observe this was G. Don, /.c., who put ‘B. chelo-
noides Kon. under the synonyms of Tecoma
suaveolens. Then HAINES remarked that the type of
Bignonia chelonoides L. f. was currently named
S. chelonoides (Kew Bull. 1922, 121). I myself
(Thesis, 1927, 951) was of the same opinion. But
these observations were not evaluated nomen-
claturally until recently by CHATTERJEE and
SANTISUK.

3. Stereospermum personatum (HAssK.) CHATTERJEE,
Bull. Bot. Soc. Beng. 2 (1948) 70; SANTISUK, Kew
Bull. 28 (1973) 178; Thai For. Bull. Bot. 8 (1974)
26. — Padri RHEEDE, Hort. Mal. 6 (1736) 47, t. 26.
— Bignonia chelonoides (non L. f.) Rox. Fl. Ind.
ed. Carey 3 (1832) 106, p.p. — S. chelonoides [non
(L.f.) A.P.DC.] A.P.DC. Bibl. Univ. Genéve II, 17
(1838) 125, quoad ref. RHEEDE, t. 26; A.DC. Prod.
9 (1845) 210, p.p.; Kurz, Fl. Burma 2 (1877) 230;
CLARKE, FI. Br. Ind. 4 (1884) 382; TriMEN, FI.
Ceyl. 3 (1895) 283; Ripi. Fl. Mal. Pen. 2 (1923)

1977]

BIGNONIACEAE (van Steenis)

149

550; STEEN. Thesis (1927) 951; Bull. Jard. Bot.
Btzg III, 10 (1928) 237; Dorp. Fl. Gén. I.-C. 4
(1930) 579; CorNER, Ways. Trees (1940) 172, f. 43.
— Dipterosperma personatum Hassk. Flora 25, 2
(1842) Beibl. 1, p. 28; Cat. Hort. Bog. (1844) 152;
Pl. Jav. Rar. (1848) 507. — S. tetragonum A.DC.
Prod. 9 (1845) 210; Harnes, Fl. Bihar Orissa (1922)
655; Kew Bull. (1922) 121; GAMBLE, Fl. Madras 2
(1924) 998; HaNpD.-MaAzz. Symb. Sin. 7 (1936)
889. — Bignonia caudata A.DC. Prod. 9 (1845)
166. — S. hasskarlii Z. & M. ex ZOLL. Syst. Verz. 3
(1855) 54, nom. illeg., based on Dipterosperma
personatum Hassk.; Miq. Fl. Ind. Bat. 2 (1858)
756; Ann. Mus. Lugd. Bat. 1 (1864) 200. — S.
caudatum Mia. /.c. 200. — Fig. 17.

Deciduous, glabrous tree, up to 30 m, 75cm @;
bark pale pinkish grey becoming rather coarsely
fissured and flaky but not ridged. Young leaves
purplish or pinkish (CORNER). Leaves glabrous,
20—50 cm; leaflets 3-6 pairs, elliptic-oblong, gra-
dually tapering to the base, tip acuminate to cau-
date; 5-15 by 2'/,-6cm; underneath minutely
glandular-punctate, often with a few large flat
spot-glands (black in dry state); petiolules slender,
5-15 mm. Thyrses widely branched, paniculiform,
to 40 cm long. Flowers dingy yellow, cream within
with dark red stripes, in slender, erect, lengthening
thyrses 15—40 cm long, on the bare twigs with the
new leaves, c. 3 cm long, 1°/, cm wide at the limb.
Calyx campanulate, 6-8 mm, with 3-4 short acute
lobes, purple. Corolla with a narrow basal tube
4-5mm long, then campanulately widened and
curved, the bell-shaped part compressed with the
mouth closed and the underside grooved; tube
c. 13/,cm long; lobes crisped, the upper two re-
curved, the lower bearded at the mouth, ochre-
buff with brownish or purplish lines, pinkish
purple on the outside (CORNER). Filaments with a
dense hair tuft at the base. Capsule linear, + terete,
4-ribbed, curved or twisted, 8-45 cm by 8-10 mm;
septum 4-5mm @. Seeds 2 by */, cm, incl. wings

,cm.

Distr. Widely distributed from Ceylon through
entire continental tropical SE. Asia to Yunnan and
Indo-China; in Malesia: Malaya (very rare, only in
Penang, e.g. on Glugor Road, and Singapore),
possibly also in Sumatra.

Unfortunately there is no certainty about the
records in Indonesia: HASSKARL described his type
from trees cultivated in the Botanic Gardens at
Bogor; ZOLLINGER described S. hasskarlii also
from a cultivated tree in these gardens (ZOLLINGER

3069) but noted that it would have originated from
Bantam, West Java, adding the Sundanese verna-
cular ‘ki langir’; a duplicate of this number in
Paris is said by SANTISUK to have been annotated to
come from the Lampong Distr. in S. Sumatra,
which then probably is an error.

Then there is a collection said to have been col-
lected by KORTHALS with ‘Borneo’ printed labels.
This provenance is very unlikely, as the use of
these old labels has been proved to be often erratic.
These specimens may have come from West
Central Sumatra but may also have been collected
by KoRTHALS in the Bogor Botanic Gardens.
Plant-geographically the species might occur (or
have occurred) in Sumatra and West Java, but
probably as rare as in Malaya because of its
preference for seasonal forest conditions.

Ecol. Preferring lowland forests, up to 1000 m.
Fl. March-July (at Bogor Aug., Nov.—Febr.), fr.
Febr. (Asia), June (Malaya).

Uses. According to BurRKILL in India an im-
portant timber tree, especially in the northeast
where it is common; the hard grey wood is moder-
ately durable and easy to work, good for furniture,
used but less good for building; in Assam and E.
Bengal padri-wood is used for canoes and tea-
boxes. In S. India a cooling drink, from roots and
flowers, is given in fevers. The fragrant flowers are
offered in temples.

Nomencl. There has been a most unfortunate
confusion about the identity of Bignonia chelo-
noides L. f. (1781). This emanated from LINNE f.
who described it as a hairy plant (type herb.
Konig, in LINN), but added the reference to
Padri of RHEEDE, an other glabrous species with
long petiolules. This probably misled RoxBURGH
who applied LINNE’s epithet to the latter. This
interpretation was followed by almost all subse-
quent authors. In 1922 Harnes concluded that two
species were involved and he adopted for the
present one the name S. tetragonum DC. In 1948
CHATTERJEE replaced this by an older epithet of
HASSKARL.

Notes. As in several other members of this
family leaves from suckers and saplings may be
toothed or serrate at the margin; leaves of mature
trees are entire.

In a few continental Asiatic specimens a very
minute puberulous indument occurs on nerves
underneath in the inflorescence (KERR 1167,
KOSTERMANS 1056, LESCHENAULT 157).

9. RADERMACHERA

ZoLL. & Mor. in Zoll. Syst. Verz. 3 (1855) 53; BUREAU, Adansonia 2 (1861) 192,
t. 2; Mon. (1864) 50, t. 28; Mig. Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 250; SEEM.
J. Bot. 8 (1870) 145; Jackson, Ind. Kew. 2 (1895) 679 (‘Radermachia’); STEEN.
Thesis (1927) 953; Bull. Jard. Bot. Btzg III, 10 (1928) 238; Dop, Fl. Gén. I.-C. 4
(1930) 583; CHATTERJEE, Bull. Bot. Soc. Beng. 2 (1948) 71; SANTISUK, Thai For.
Bull. Bot. 8 (1974) 27; STEEN. Blumea 23 (1976) 121. — Lagaropyxis Mig. Ann.
Mus. Bot. Lugd.-Bat. | (1864) 198. — Mayodendron Kurz, Prel. Rep. For. Veget.
Pegu, App. D (1875) pl. | & 2; Fl. Burma 2 (1877) 232. — Stereospermum sect.

150 FLORA MALESIANA [ser. I, vol. 82

Fig. 20. Radermachera peninsularis STEEN. a. Leaflet, b. apex of thyrse, c. calyx and style, d. corolla, all

nat. size. — R. glandulosa (Bu.) Mig. e. Leaf base, underneath with dense glandfield, x 2, f. empty

capsule, with 2 valves and septum, x 1/3, g. section of septum, x 5, h. seed, x 2 (a—d LARSEN c.s. 31239,
e-h KOSTERMANS 6360A).

1977] BIGNONIACEAE (van Steenis) 151

Radermachera et Xylocarpaea B. & H. Gen. Pl. 2 (1876) 1047; CLarkg, FI. Br. Ind.
4 (1884) 383; K. & V. Bijdr. Booms. | (1894) 71. — Radermachera sect. Alatae
STEEN. Acta Bot. Neer]. 2 (1953) 307. — Fig. 20, 22.

Trees, rarely shrubs, evergreen (except 2 Indian and 2 Chinese spp.). Twigs and
floral rachis mostly distinctly lenticellate. Innovations sticky-lacquered. Leaves
2-3-pinnate; stalks and leaflets articulated; rachis sulcate; leaflets underneath
minutely but densely dotted, furthermore with gland-fields or scattered glands
underneath, mostly acuminate, very rarely laxly puberulous. 7hyrses terminal,
very rarely ramiflorous, in one sp. a raceme; bracts and bracteoles inconspicuous,
very rarely leafy; thyrses (and calyx) very rarely short-hairy. Flowers white, pink
with yellow streaks in mouth, more rarely greenish yellow, or golden yellow to
orange-red. Calyx closed in bud, rarely lobed from the beginning, splitting spatha-
ceously (not to the base) or mostly irregularly lobed, rarely stunted, often with
microscopical scale-like glands, moreover mostly with larger crateriform glands;
after anthesis almost always circumscissile-dehiscing at the base, along an abscis-
sion line, in one sp. persistent. Corolla either salver- or narrow funnel-shaped or
more commonly with a mostly short basal tube and often rather suddenly widened
into an upper tube; lobes mostly rounded unequal, not rarely ciliate. Stamens
didynamous with a Sth rudiment, but in one Chinese sp. 5 equal stamens, not
exserted; another-cells V-shaped divergent; filaments inserted at the apex of the
basal tube, except in two spp. capitate-glandular hairy at the insertion and in the
basal part, for the rest glabrous; connective produced. Ovary elongate often
minutely lepidote, glabrous, or tuberculate, never hairy; in both cells with several
rows of ovules; style filiform, mostly exceeding the anthers; stigma 2-lipped.
Capsule linear, terete, up to 75 cm; valves smooth, pergamentaceous, rarely +
woody, in one sp. tuberculate; septum terete, brittle, corky, but with shallow im-
pressions of the flat seeds, a narrow line on both sides testimony of attachment to the
middle of the valves. Seeds very «, flat, small, narrow, on both ends with a hyaline
wing.

Distr. About 15 spp. in Indo-Malesia, from the Deccan to S. China, Hainan, Formosa, and the southern
Ryu Kyus, most abundant in tropical SE. Asia; throughout Malesia, but not in the Moluccas proper and
New Guinea. Fig. 21.

Ecol. Lowland primary and secondary rain-forests, up to c. 1500 m, not rarely pioneering in disturbed
forest and on slopes.

Flower colour and corolla shape vary from pure white to orange-red and from hypocrateriform
(R. sinica (HANCE) HeEmsL. and R. frondosa CHUN & How) to tubular or campanulate. For the narrow-
tubular orange-flowered species pollination by birds can be expected (R. ramiflora) and for the pure
white, possibly nocturnal R. peninsularis hawk-moths may be the pollinating insects.

Ramiflory is found in Malesia in R. ramiflora; it occurs also occasionally in poor forms of R. pinnata
after leader-shoots have been damaged. In continental Asia it is also found in R. ignea (KURZ) STEEN.
and in R. hainanensis MERR.

Uses. Minor uses, see under the species.

Syst. Related to Stereospermum, which differs radically in having only two rows of ovules in each
cell, and thick trigonous seeds with folded cotyledons fitting in cavities of the septum; moreover, its leaves
are always 1-pinnate and most species are deciduous and prefer a seasonal climate.

Two small genera have recently been split off, both from SE. Asia, viz the monotypic Pauldopia STEEN.
(Acta Bot. Neerl. 18, 1969, 425) which has winged rachises, a curved corolla tube, wingless thickish seed,
a very thin septum, and a truncate calyx open in bud; and Barnettia SANTISUK (Kew Bull. 28, 1973, 172)
with 2 spp. in Thailand, with 1-pinnate leaves in pseudowhorls, a short-ellipsoid, -+- compressed capsule,
and tuberculate or immersed-glandular calyx and capsule and a cruciform septum; the latter genus
shows relationship with Heterophragma.

The closest ally of Radermachera is obviously the Afro-Asian genus Fernandoa (cf. Steen. Blumea 23,
1976, 133) from which it differs by the terete septum, the usually not ribbed or striate and thinner,

152 FLORA MALESIANA [ser. I, vol. 8?

narrower fruit valves, mostly 2-4-pinnate leaves, absence of domatia underneath the leaflets, and mostly
crateriform glands.

Notes. Specific delimitation, especially in Malesian material, proved difficult, as several species appear
to be variable and many were only known from the type, notably in the Philippine Islands. Degree of
pinnation is variable, often in one collection. Also the number and place of the larger glands on under-
surface of leaflets and on the calyx is liable to variation, as well as the flower colour and the way in which
the calyx splits. A critical scrutiny showed only few tangible characters and resulted into a severe reduc-
tion of species in Malesia.

KEY TO THE SPECIES

1. Calyx strongly lengthwise 5-6-ridged, 13/, cm, narrow, one side cleft down + halfway, 3-toothed at
apex. Leaves 1-pinnate, with 5 leaflets, coriaceous, very glossy above, with recurved edge, 7-14 by
3-4 cm. Corolla 4 cm, with narrow tube, slightly enlarged above; lobes c. 1 cm, obtuse.

6. R. coriacea

1. Calyx not lengthwise 5-6-ridged.

2. Corolla narrowly funnel-shaped, without distinction of basal and upper tube. Flowers in cauli-
florous racemes, with yellow tube and red limb. Leaves 2-pinnate. ..... . 1. R. ramiflora

2. Corolla with a cylindric basal tube, widened rather abruptly into an upper tube. Flowers in terminal
thyrses which are rarely raceme-like depauperated.

3. Calyx very short (3-5 mm), with a stunted rim, persistent. Leaves always 1-pinnate
2. R. glandulosa
3. Calyx longer, irregularly lobed, after anthesis circumscissile-caducous.
4. Corolla 2-3'/,(-4+/,) cm long (incl. lobes). Calyx 5-10(-13) mm long 3. R. pinnata

4. Corolla 5-6 cm long. Calyx 10-25 mm.

5. Filaments and inside of basal tube glabrous. Corolla tube outside towards apex and on lobes with
minute sessile glands. Leaves above puberulous on midrib and nerves, beneath with some scattered
glands. Corolla white or cream-coloured 4. R. insularis

5. Filaments and inside of basal tube near insertions capitate-glandular hairy. Corolla tube outside
towards apex mostly laxly and very short-hairy. Leaves glabrous above, beneath mostly with at
least a gland-field at base. Corolla mostly pinkish, sometimes white, with yellow streaks in the

mouth

1. Radermachera ramiflora STEEN. J. Bot. 72 (1934)
5; Blumea 23 (1976) 129.

Large tree, 24-30 m, stem 30-60 cm @; fluted at
base; bark grey, fissured. Innovations and racemes
lacquered, resinous sticky. Leaves (2—)3-pinnate, up
to 1 m, more or less crowded at the twig-ends; pin-
nae 4-6 pairs; leaflets elliptic-lanceolate, acuminate
at both ends, stalked, 31/,-8'/, by 11/,-31/,cm, under-
neath fine glandular-punctate and with scattered
small, shallow glands especially near the base, and
a few scattered flat glandular spots. Flowers thickly
set in closely placed ramiflorous pendent racemes
to 20 cm long, erect on curved pedicels 1-2 cm long
and with 3 bracteoles + halfway on an articulation.
Calyx in bud pear-shaped, closed, reddish green,
tubular, 2—3-lobed, eglandular outside, microsco-
pically glandular-papillose inside, c. 21/,—2'/, cm
long, 1 cm @ at mouth, lobes 4-6 mm, 1-2 split +
halfway. Corolla with yellow tube and red limb;
tube slightly curved, narrow salver-shaped, without
narrowed basal tube, 5-7 cm, the basal 11/,cm
densely pubescent with thick hairs, mouth 11/,-
2cm @; lobes rounded, subequal, + Icom @, +
papillose inside. Stamens (and style) reaching the
mouth, inserted halfway the tube, glabrous but
their adnate base lax glandular-papillose, thecae +
divergent, 4-5 mm; no produced connective. Disk
thick, annular-cup-shaped, faintly crenate. Ovary
ribbed, pistil c. 4cm, stigmatic lobes 2 mm,
very narrow. Capsule straight or twisted, 35-70 cm,
c. 5mm @; septum 2!/,-3mm @. Seeds 4-5 by
2'/, mm, the wings 6-7 mm.

Distr. Malesia: Sabah, Mt Kinabalu (Peni-
bukan, Dallas, Mesilau, Tenompok, Kota
Belud, Kp. Kiau I resthouse), not rare.

5. R. gigantea

Ecol. Rain-forest, also in disturbed forest on
hill side, 950-1500 m. Fl. Aug., Jan.—March, fr.
April, Dec.

Notes. A characteristic, isolated species: rami-
florous, flowers in racemes, shape of corolla.

Leaflets of suckers and saplings dentate. NOOTE-
BOOM & ABAN 1603 has only 2-pinnate leaves,
CLEMENS 30364 has them 2-3-pinnate.

Fig. 21. Range of the genus Radermachera Z. & M.

2. Radermachera glandulosa (BL.) Mia. Ann. Mus.
Bot. Lugd.-Bat. 3 (1867) 250; K. & G. J. As. Soc.
Beng. 74, ii (1905) 380; Koorp. Atlas Baum. 2
(1914) t. 356; STEEN. Thesis (1927) 965; Bull. Jard.
Bot. Btzg III, 10 (1928) 241; CorNeER, Ways. Trees
(1940) 168, f. 43; CHATTERJEE, Bull. Bot. Soc. Beng.

1977]

BIGNONIACEAE (van Steenis)

153

2 (1948) 74; STEEN. Acta Bot. Neerl. 2 (1953) 307;
Back. & BAKH. f. Fl. Java 2 (1965) 541; SANTISUK,
Thai For. Bull. Bot. 8 (1974) 27, f. 15; STEEN.
Blumea 23 (1976) 126. — Spathodea glandulosa
Bi. Bijdr. (1826) 762. — Bignonia porteriana
WALL. ex DC. Prod. 9 (1845) 165. — R. stricta
Z. & M. ex ZOLL. Syst. Verz. 3 (1855) 53; Rw . FI.
Mal. Pen. 2 (1923) 550. — Stereospermum glandu-
losum Mig. Suppl. (1860) 240, 565; CLarKE, FI.
Br. Ind. 4 (1884) 383; K. & V. Bijdr. Booms. 1
(1894) 74; KANyJILAL & Das, Fl. Assam 3 (1939)
404. — Lagaropyxis glandulosa Mia. Ann. Mus.
Bot. Lugd.-Bat. 1 (1864) 199. — Fig. 20e-h.

Small, glabrous, evergreen, crooked tree, up to
12m, 40 cm @; bark slightly pimply and peeling,
but not flaky or fissured, bitter; young leaves deep
purple, acrid. Leaves 1-pinnate; leaflets 2-5 pairs,
large, chartaceous, elliptic (mostly broad-),
rarely oblong-lanceolate, short-tipped, with a
conspicuous dark, dense gland-field at the oblique
base underneath (often bulging on upper surface),
10-30 by 5-17 cm. Thyrse narrow, 6-50 cm, gradu-
ally elongating, sometimes the upper part still
flowering while lower are in fruit. Calyx 3-5 mm,
cup-shaped, without abscission line, persistent,
truncate, purple, spotted with 5—7 purple glands in
a crescent. Corolla narrow, slender, tube narrow,
slightly curved, halfway rather gradually widening,
3(-4) cm, pinkish purple outside, white inside,
with a gland-field at outside of the ciliate lobes,
basal part of lower tube short capitate-glandular
hairy inside. Stamens hairy at insertion. Pods
hanging in bunches, straight, 15-30cm; valves
5-7 mm wide; septum 3-4 mm @. Seeds 10-16 by
11/,-2 mm.

Distr. Continental SE. Asia (Assam, Burma,
Thailand, Laos, China: Kwangsi, Kwantung) and
West Malesia: Malay Peninsula (also Penang),
Sumatra, Krakatao, West to East Java.

The two records from Borneo are erroneous: the
KoORTHALS specimens are mislocalized and BECCARI
811 (mentioned by MERRILL, En. Born. 1921, 525)
is a PS number. Both are from Sumatra.

Ecol. Primary and secondary forests and thick-
ets, frequently by streamsides, even rocky
Saraca-streams (CORNER), both under everwet
and under seasonal conditions (in Central and
East Java), from sea-level to 900 m, once reported
from 5000 ft (Cameroon Highlands). Fi. April,
July—Nov., fr. Jan.—Dec.

The fine seed would indicate easy dispersal, but
though the species is found in Penang and Krakatao
Is., it has never been collected in the Riau Is., not
in the islands west of Sumatra and also not in
those close to East Java (Madura I., Bali, and
Kangean) though it is found in Java as far east as
Bali Straits and R. gigantea extends east as far as
Alor. The glands at the leaflet basis are often black
from sooty moulds indicating actual glandular
excretion.

Uses. The timber is small and of negligeable
value.

Vern. Tuwi gadang, M, Minangkabau; kia bako,
ki hapit, ki langhit, kipahit, ki sakat, ki sikap, 8;
ambal, bangking, bangkongan, djelibru, godong
ambol, hambal, paras = gabret, klaju, lambal,
padali, pedali, pudang, J; sekar pote, Md.

3. Radermachera pinnata (BLANCO) Seem. J. Bot. 8

(1870) 147; Merr. Philip. J. Sc. 3 (1908) Bot. 336,
incl. var. glabra MERR. l.c.; Sp. Blanc. (1918) 350;
En. Philip. 3 (1923) 446; STEEN. Thesis (1927) 973;
Bull. Jard. Bot. Btzg IIT, 10 (1928) 248; Blumea 23
(1976) 129. — Millingtonia pinnata BLANCO, FI.
Filip. (1837) 501. — Méillingtonia quadripinnata
BEANGO; Ics: ed! 3,;, 2 (1878): 286, t. 252. — R:
banaibana BUREAU, Adansonia 2 (1861) 194. —
R. quadripinna Seem. J. Bot. 8 (1870) 147. — Stereo-
spermum pinnatum F.-VILL. Nov. App. (1880) 151;
Ro re, J. Linn. Soc. 23 (1884) 314; VipAL, Phan.
Cuming. Philip. (1885) 132; Rev. Pl. Vasc. Filip.
(1886) 203. — Stereospermum quadripinnatum
F.-VILL. Nov. App. (1880) 151; VmDAL, Syn. Atlas
(1883) 35, t. 73. — Stereospermum banaibanai
Ro.rFe, J. Linn. Soc. 23 (1884) 314. — Stereo-
spermum seemannii ROLFE, /.c. — R. acuminata
Merk. Philip. J. Sc. 3 (1908) Bot. 335; En. Philip. 3
(1923) 445; STEEN. Thesis (1927) 980. — R. mindo-
rensis MERR. Philip. J. Sc. 3 (1908) Bot. 338; En.
Philip. 3 (1923) 446; STEEN. Thesis (1927) 971. —
R. fenicis MERR. Philip. J. Sc. 3 (1908) Bot. 335,
434; En. Philip. 3 (1923) 446; STEEN. Thesis (1927)
979, incl. var. acuminata STEEN.; Jard. Bot. Btzg
III, 10 (1928) 261. — R. whitfordii MerR. Philip. J.
Sc. 7 (1912) 352; En. Philip. 3 (1923) 447; STEEN.
Thesis (1927) 963. — R. brachybotrys MERR.
Philip. J. Sc. 26 (1923) 489; E_mer, Leafl. Philip.
Bot. 10 (1939) 3809. — R. sorsogonensis ELMER ex
STEEN. Thesis (1927) 973; ELMer, Leafl. Philip.
Bot. 10 (1939) 3809. — R. elegans STEEN. Bull. Jard.
Bot. Btzg III, 10 (1928) 252, f. 8. — R. fenicis (non
MERR.) STEEN. /.c. 261, f. 11.

See for other synonyms under ssp. acuminata.

Distr. Malesia: Sumatra (also Simalur, Banka,
and Siberut Is.), Malaya, Borneo, Philippines,
Celebes (also Muna I.), and W. Moluccas (Sula
Is.: Taliabu).

Notes. It has appeared impossible to separate
the material into smaller species, as there is
gradual transition of the many populations,
especially in the Philippine islands, notably in
vegatative characters. R. brachybotrys is merely a
depauperate ridge facies. In one specimen 1- and
2-pinnate or biternate leaves are not seldom found
together. For brevity I have omitted from the
synonymy the many pro parte citations under
R. amoena, hypostictum and gigantea. Some speci-
mens from Borneo and Celebes show a tendency in
leaf-shape towards ssp. acuminata.

KEY TO THE SUBSPECIES

1. Leaflets usually chartaceous, elliptic, acuminate
to caudate; basal gland field mostly distinct, at
apex eglandular or with a few scattered glands

ssp. pinnata

1. Leaflets firmly chartaceous to coriaceous with
consequently less marked prominent venation
beneath, usually obovate to elliptic-obovate, at
apex mostly short- and blunt-tipped or blunt or
rounded; basal gland field dense, often one also

BEDDGKinn bhai’ Winn ae Ras ssp. acuminata

ssp. pinnata.

Tree to 20m, 15-40cm @. Leaves (1-)2-3-
pinnate, 25-50(—70) cm; leaflets elliptic-oblong,
acuminate, mostly at both ends, to caudate,
(3-)5—16 by (1'/,-)2-5(-8) cm, usually chartaceous,

154

FLORA MALESIANA

{ser. I, vol. 8?

io

Fig. 22. Radermachera pinnata (BLANCO) SEEM. ssp. acuminata (STEEN.) STEEN. Habit, in flower, Aug. 1970
(photogr. B. C. STONE).

with a basal gland field, apical gland field absent or
of scattered glands. Thyrses glabrous, sometimes
puberulous, (3—6—)15-60cm, in odd specimens
sometimes on the old wood. Calyx #/,-1(-11/,?) cm,
lobes unequal, (1—)2—4, glabrous, usually glandless.
Corolla pink to pale purplish, with yellow mark-
ings in the throat, 2—3'/,(—4"/,) cm, rarely slightly
lax short-hairy. Stamens glandular-hairy at their
insertion. Ovary and style glabrous, 12-17(-22)
mm. Capsule (6-)30-50cm; valves 4-6(—7) mm
wide; septum 2-2!/,mm @. Seeds (7-)13-15 by
2-3 mm.

Distr. Malesia: Philippines (the most common
species), Celebes (also in Muna I.), Moluccas
(Sula Is.: Taliabu).

Ecol. Lowland and montane primary and
secondary forests, on streamsides, up to 600 m.
Fl. Aug.—May, fr. Febr.—Nov.

Uses. Minor uses only, for carving and fuel.

Vern. Philippines: banai-bdnai, Tag., the com-
mon name, kalapuing, salai, tuing-huld, ulimbabon,
yabang-ydabang, Tag., banoi-banoi, Bag., ansohan,
badlan, Bis., labayanan, C.Bis., paling-udk, Bik.,
pagalayan, Bon., bunglai, Buk., atiatip, Ig.,
lanunisi, lasilak, Ybn., barangauan, I\k., bunlai,
Mbo,. banaibayan, paitan, pata, Pang., bani-bani,
Sbl., kutokong, Sub., hali-hdali, Sulu; Celebes:
ririh, Muna.

Note. In a few specimens some inflorescences
are ramiflorous, obviously due to damage of the
leader shoot.

ssp. acuminata (STEEN.) STEEN. Blumea 23 (1976)
129. — Spathodea lobbii T. & B. Nat. Tijd. N. I.

25 (1863) 413. — R. lobbii Mig. Ann. Mus. Bot.
Lugd.-Bat. 3 (1867) 250; SEEM. J. Bot. 8 (1870)
147; STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 243,
f. 5, incl. ssp. acuminata STEEN. l.c. 247, f. 6;
CORNER, Ways. Trees (1940) 168, f. 43 (‘lobbiana’);
SANTISUK, Thai For. Bull. Bot. 8 (1974) 29. —
R. amoena [non (WALL.) SEEM.] GAMBLE, J. As.
Soc. Beng. 74, ii (1905) 381; Rip. Fl. Mal. Pen. 2
(1920) 551. — R. corymbosa STEEN. Bull. Jard. Bot.
Btzg III, 10 (1928) 249, f. 7. — R. gigantea [non
(BL.) Miqg.] Burk. Dict. (1935). — Fig. 22.

Tree, 7-40 m, 60cm @. Leaves biternate or
2-pinnate; leaflets rather coriaceous, obovate to
obovate-elliptic, not or short- and blunt-tipped,
exceptionally acuminate, basal gland field well-
developed, apical one usually distinct, 8-15 by
31/,-71/, cm. Thyrse apical, fairly narrow, 15-25
cm. Calyx 1 cm, rarely to 11/, cm. Basal tube of
corolla rather suddenly widened.

Distr. S. Peninsular Thailand (Krabi, Pattani);
in Malesia: Sumatra (also in Banka; Mentawai
Is.: Siberut; Simalur I.), Malaya, Borneo, SW.
Philippines (Palawan, Culion).

Ecol. Primary and secondary forests, also in
open grasslands. In Malaya especially by streams
(CORNER), ON granite as well as on limestone; at
0-—400(-800) m. F/. May, July—March, fr. May-
Nov.

Taxon. Formerly often accepted as conspecific
(also by myself, 1927) with R. gigantea cq. amoena,
but certainly distinct.

Vern. Sumatra: kaju singamba, sindur langit,
sundur langit, Batak, kudo kudo pajo, Simalur,
mentu, tuih, tuwi(k), Banka; kapung suwi, kéka-

1977]

pung, Lampong; bunga pawang, seténgah burong,
tangkani, Malaya; Borneo: binutan, kudjuk langit,
Dajak-Kapuas; Culion: totancola, Tagb.

Note. I must admit that I have somewhat
hesitantly kept this apart as a subspecies from
true R. pinnata, from which it differs only in shape
and texture of leaflets and geographical range.
There are some specimens in Borneo which seem
transitional.

In the Philippines (Palawan and Culion) aber-
rant specimens are found with biternate leaves and
coriaceous leaflets with recurved margin and rather
prominent veins underneath. They were collected
in grasslands which may account for their habit.
However, in the same islands there are also larger-
flowered specimens which I have referred to R.
gigantea with similar habit. More field work in
these islands is needed to check my tentative
conclusions.

4. Radermachera peninsularis STEEN. Blumea 23
(1976) 128, f. la-d. — R. borii (non FiscHER)
SANTISUK, Thai For. Bull. Bot. 8 (1974) 30. —
Fig. 20a—-d.

Tree, 415m. Leaves 3-pinnate, 60-80 cm;
leaflets lanceolate-oblong, falcate-caudate, 5-8 by
2-3 cm, on both surfaces microscopically punctate
but only with a few scattered, very small ‘larger’
glands; midrib and main nerves (c. 4-5 pairs)
puberulous above (as in R. sinica). Inflorescences
terminal, similar to those in R. sinica; peduncle
firm, 15-35cm long, rachis 4-7 cm; full-grown
pedicels 2!/,-7 cm, halfway with 2 decussate linear
bracteoles c. 1'/,-3 cm long. Bracts long, linear,
exceeding the buds, the lowest up to 5cm long,
upper ones 2!/,-3 cm. Calyx campanulate, rather
wide and thickish, densely microscopically lepi-
dote, 2—2'/, by 1'/, cm, rather irregularly lobed for
1/,/, of its length, with 5 + distinct gland fields.
Corolla white or creamy, c. 6-7 cm long (incl. the
%entire lobes), with a rather wide (7/,cm) basal
tube c. 1'/, cm long concealed in the calyx on top of
which the filaments are inserted ; upper tube funnel-
shaped widened. Filaments c. 3cm, glabrous;
anthers 4'/,mm; connective appendage small.
Ovary glabrous. Capsule terete, tortuous, 60-70 cm
by 3-5 mm.

Distr. Peninsular Thailand (between Phangnga
and Krabi); in Malesia: Malay Peninsula (Cameron
Highlands), 2 collections.

Ecol. In Peninsular Thailand at 8°25’ N, 99°15’
E, in evergreen forest along a stream, on limestone,
at 50m altitude, in the Cameron Highlands at
1200 m in mixed rain-forest.

Notes. In habit deceptively like R. sinica, but
at once different by the shorter (6-7. cm), not
salver-shaped corolla and the much wider smooth
calyx. The corolla in R. sinica measures c. 8-12 cm
including the lobes, with a narrow, very gradually
widening tube. R. sinica also has the minute lax

bescent nerves and veins above and long narrow

SANTISUK /.c. referred the specimens to R. borii,
which I refer to R. sinica, a species ranging more
northerly in SE. Asia, viz from N. Assam, N.
Burma, and Tonkin to S. China, Hainan, Formosa,
and the southern Ryu Kyu Is.

5. Radermachera gigantea (BL.) Mig. Ann. Mus.

BIGNONIACEAE (van Steenis)

155

Bot. Lugd.-Bat. 3 (1867) 250; SEEM. J. Bot. 8
(1870) 146; Koorp. Atlas Baum. 2 (1914) t. 356
A-K:; Heyne, Nutt. Pl. (1927) 1371; STEEN. Thesis
(1927) 983, p.p., excl. syn. lobbii; Bull. Jard. Bot.
Btzg III, 10 (1928) 253, f. 9; Blumea 23 (1976) 126.
— Bignonia gigantea NORONA, Verh. Bat. Gen.
5 (1790) 70, nomen. — Spathodea gigantea BL.
Bijdr. (1826) 761; Mia. Fl. Ind. Bat. 2 (1858) 751.
— Bignonia amoena WALL. [Cat. (1832) n. 6512,
nomen] Pl. As. Rar. 2 (1831) 78, t. 183; LouDon,
Hort. Brit. (1830) 483, err. amara; G. Don, Gen.
Hist. 4 (1838) 222. — Bignonia oxyphylla DC.
Prod. 9 (1845) 169. — Stereospermum hypostictum
Mig. Sum. (1861) 565; CLARKE, Fl. Br. Ind. 4
(1884) 384, excl. syn. S. lobbii; K. & V. Bijdr.
Booms. 1 (1894) 72, excl. syn. lobbii; KANNLAL &
Das, Fl. Assam 3 (1939) 405. — Lagaropyxis
gigantea Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864)
198, incl. f. sumatrana et f. borneensis Miq. —
R. amoena SEEM. J. Bot. 8 (1870) 146; non GAMBLE,
J. As. Soc. Beng. 74, ii (1905) 381, quae est R. lobbii.
— R. elmeri Merr. Bull. Gov. Lab. Philip. 29
(1905) 48; Philip. J. Sc. 3 (1908) Bot. 334; En.
Philip. 3 (1923) 445; STEEN. Thesis (1927) 994. —
R. biternata MERR. Philip. J. Sc. 1 (1906) Suppl.
238; ibid. 3 (1908) Bot. 333; En. Philip. 3 (1923)
445; STEEN. Thesis (1927) 970. — R. palawanensis
Me_rr. Philip. J. Sc. 3 (1908) Bot. 336; STEEN.
Thesis (1927) 977. — R. elliptica MERR. Philip. J.
Sc. 3 (1908) Bot. 334; En. Philip. 3 (1923) 445;
STEEN. Thesis (1927) 964. — R. sibuyanensis ELMER,
Leafi. Philip. Bot. 4 (1912) 1485; Merr. En. Philip.
3 (1923) 447; STEEN. Thesis (1927) 992. — R. elmeri
var. fragrans ELMER, Leafl. Philip. Bot. 7 (1915)
2561; Merr. En. Philip. 3 (1923) 446. — R.
fragrans STEEN. Thesis (1927) 996. — R. punctata
ELMER ex STEEN. Thesis (1927) 982; ELmer, Leafi.
10 (1939) 3709. — R. borneensis STEEN. Bull. Jard.
Bot. Btzg III, 10 (1928) 258, f. 10.

Shrub or tree, (6—)20-40 m, up to 80 cm @; bark
and young leaves bitter. Leaves (1—)2(—3)-pinnate,
12-35(-80) cm, leaflets usually elliptic to oblong,
rarely somewhat obovate or lanceolate, shorter or
longer acuminate, 4-12(-15) by 2-6(-9) cm, at base
underneath mostly with a gland-field, at apex with
some scattered glands. Thyrses 8-40 cm, rather
open, terminal, ~©-flowered, glabrous. Flowers not
rarely fragrant. Calyx (1-)1'/,-2'/,cm long,
mostly 2-lobed, sometimes 1- or more-lobed ; glands
few or distinct. Corolla 5-6 cm long (incl. lobes),
pink or white, usually with yellow streaks in the
mouth, above the basal tube rather suddenly
widened, campanulate upper tube towards apex
almost always short-capitate glandular hairy, and
lobes ciliate. Filaments densely capitate-glandular
hairy at insertion. Style 2'/,-3 cm. Capsule 15-60
cm long, 5-8 mm @; septum 4mm @. Seeds 8-13
by 2-4 mm.

Distr. SE. Asia (Assam: Khasi & Jaintia Hills;
Burma: Manipur, Tavoy, WALLICH, CLARKE, /.c.,
not seen), and Malesia: Sumatra (incl. Billiton and
Banka), West to East Java (common in Central
and East Java), Lesser Sunda Is. (Bali, Sumbawa,
Flores, Alor), Bawean L., SE. Borneo, and Philip-
pines.

Ecol. Primary and secondary forests, also in
areas subject to a dry monsoon, and in East Java
in teak forest, very common on Mt Telemojo and
near Pringombo (Banjumas) (Koorpers, /.c.),

156 FLORA MALESIANA [ser. I, vol. 8?

Ve yaa \
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Breen Ne
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STRRSah ten era See

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=
Boss WENA

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wl
: : Wyn
Nantes y m
(eos wl
a, Pm

UTUrTTUTyY
a oe TT <I
f Mg Fe YQ ii oo 3s
MARS Ss ens, et Se

Fig. 23. Fernandoa macroloba (MiqQ.) STEEN. a. Habit of branchlet with young fruit, x 4/2, b. inflorescence

in bud, x 7/3, c. calyx, nat. size, d. corolla, x 3/,, e. CS of young capsule, nat. size, f. mature capsule,

x 1/2, g. seed, nat. size, h. detail of underside of leaflet, showing small domatia and flat gland, x 4, (a-e,
h C.H.B. XI-C-106a, fg DiEPENHORST 2353 HB, type).

a

se ne
ees

kz

1977]

BIGNONIACEAE (van Steenis)

157

from the lowland up to c. 1600 m. F/. Jan.—Dec.,
fr. Jan._Dec. Not rarely a pioneer in secondary
forest, cuttings sometimes used for strengthening
terracing on unstable slopes. Early flowering and
fruiting as a shrub.

Uses. Sometimes cultivated; also suitable for
re-afforestation. Timber is said to be strong and
durable, locally used for bridges and houses, but
not resistent against termites, and therefore of less
value for outdoor constructions; used for making
matches and matchboxes (HEYNE, /.c.).

Vern. Sumatra: tui (batu), M, Billiton,
Minangk., djamatan, djamaton, kaju angin, kaju
déling, radja matan, simaisaludang, sundér langit,
M, Karo-Batak, tangké, M, Atjeh, kulit bérilang,
M, Kroei, (ké)kapung tui, talas, M, Lampong;
West Java: ki padali, S; East Java: bédali (common
name), kédah, (ké)dali, kokok-kéjok, J, kaju raras,
karpotéh, potian, putian, sékar pétak, sékar pote,
Md; pédantén, Bali; atodjang, Alor; Borneo:
binutan, Dajak, Martapura, bunglai batu, b.
gunung; Philippines: Palawan: agtap, tantangan,
Tagb., sayo, Ig., barangau-a-nalabaga, pamaya-
bayen, Ik.

Notes. I have come to the conclusion that the
large-flowered species of the Philippines, R.
borneensis and R. gigantea, should be referred to
one variable taxon. Though there may be local
races they can not be properly distinguished, not
even on subspecific level, as their ‘characters’ fade
away; R. elliptica with 1-pinnate leaves goes via
biternate leaves to 2-pinnate leaves. R. sibuyanensis
has an almost glabrous corolla; R. fragrans has
fragrant flowers, but fragrancy is mentioned
frequently on labels, but also sometimes flowers are
said to be scentless. Flower colour is said to be
white or pink in the Philippines, to which mostly is
added the occurrence of yellow streaks in the
mouth. White flowers are only reported from
Borneo and the Philippines.

Especially a few Philippine specimens from Pala-
wan and Culion are aberrant; earlier collections
were described by MERRILL as R. palawanensis and
R. biternata. They were collected in grasslands and
differ by thicker, often biternate leaves, leaflets
with recurved margin and often distinctly arching
prominent veins on the undersurface. One number

(PNH 12319) has singularly slender tubular flowers
5 cm long. Other specimens with the same vegeta-
tive difference from normal R. gigantea have, how-
ever, smaller flowers (3—3!/, cm) and these I have
identified as R. pinnata ssp. acuminata. I must admit
that closer field work is needed to check my tenta-
tive conclusions.

R. amoena (WALL.) SEEM. was described from a
flowering shrub in Hort. Cale. which WALLICH
found worthy of an ornamental. WALLICH was
somewhat doubtful about its provenance, but
assumed that it was introduced by Mr C. TELFAIR
from Mauritius, where it must then also have been
cultivated. Though it is not known from Malaya
and Thailand, it does occur in Assam (Khasia &
Jaintia Hills, cf. KANNILAL & Das) and N. Burma
(Manipur: MEEBOLD 5169). These specimens I can-
not distinguish from R. gigantea. What KANJILAL
& Das mean in their key by ‘rusty-coloured cap-
sules’ is not clear to me; they mention the flowers to
be white.

Insufficiently known

6. Radermachera coriacea MERR. Philip. J. Sc. 3
(1908) Bot. 333; En. Philip. 3 (1923) 445; STEEN.
Thesis (1927) 961; Blumea 23 (1976) 131.

Leaves 1-pinnate, 20-30 cm; leaflets 5, coria-
ceous, oblong to elliptic-oblong, base acute, apex
blunt or obscurely blunt-tipped, margin recurved,
7-14 by 3-4cm, very glossy above; nerves 13
pairs. Panicles 15cm. Calyx 13/,cm, narrow,
strongly lengthwise 5—6-ridged, cleft on one side
halfway, 3-toothed at apex. Corolla 4 cm, the tube
rather narrow, slightly enlarged above, the lobes
obtuse c. 1 cm. Capsule 16 cm, valves S~7 mm wide
(ex descr.).

Distr. Malesia: Philippines: Central E. Luzon
(Tayabas Prov.: Baler), only known from the type
(MERRILL 1099); vern. name bibit parang.

Note. This is the only specific name in the genus
of which I could not trace the type, which may be
lost. A lengthwise ridged calyx I have not observed
in any species of the genus; the 1-pinnate leaves
and narrow leaflets seem rather characteristic. It
might possibly be an extreme form of R. pinnata
ssp. acuminata.

10. FERNANDOA

WELW. ex Seem. J. Bot. 3 (1865) 330, t. 37-38, sphalma Ferdinandia; ibid. 4 (1866)
123; ibid. 8 (1870) 280, sphalma Ferdinandoa; ibid. 9 (1871) 81; MILNE-REDHEAD,
Kew Bull. 3 (1948) 171; Heme, Adansonia 4 (1964) 467-470; Gentry, Ann. Mo,
Bot. Gard. 62 (1975) 480; STEEN. Blumea 23 (1976) 133. — Kigelianthe BAILL. Hist.
Pl. 10 (1891) 50. — Haplophragma P.Dop, Bull. Soc. Bot. Fr. 72 (1925) 889; STEEN.
Thesis (1927) 998; Bull. Jard. Bot. Btzg II, 10 (1928) 262. — Spathodeopsis
P.Dop, C. R. Ac. Sc. Paris 189 (1929) 1097. — Hexaneurocarpon P.Dop, l.c. —
Tisseranthodendron SILLANS, Bull. Soc. Bot. Fr. 98 (1951) 270-272, pl. — Fig. 23,
25.

Trees. Leaves |-pinnate; leaflets 2-5 pairs, diminishing in size downwards, ter-
minal one largest, beneath glabrous or tomentose with stellate hairs, with few

158 FLORA MALESIANA [ser. I, vol. 8?

scattered, larger, flat, spot-like glands and small hairy domatia in the nerve-axils.
Thyrses axillary or terminal, densely or laxly tomentose, or at least almost always
with (sometimes) small and appressed stellate hairs. Calyx tubular or campanulate,
unequally 2—S-lobed, with few to many warty or prominent glands in the upper
half. Corolla with a fairly large basal tube, rather suddenly campanulate-funnel-
shaped widened to the mouth; lobes undulate to crenate. Stamens 4, didynamous;
5th rudimentary; anthers divaricate. Disk annular. Ovary elongate, with 2 mar-
ginal placentas in each cell; ovules co. Capsule long, linear, terete, twisted or
straight, pendent, smooth or ribbed; septum flat, smooth, not rarely glossy,
thickish. Seeds numerous, rather rectangular, the wings rather narrow; insertion
punctiform.

Distr. About 4 spp. in tropical West Africa (Angola to Gabon) and East Africa (Tanganyika), 3 in
Madagascar, and 6 in Indo-Malesia (India to Indo-China); in Malesia 2 spp., in the Malay Peninsula and
N. Sumatra. Fig. 24.

Ecol. Tropical lowland forest.

Taxon. In Blumea /.c. I have given the reasons for uniting all these genera, by which the range of
Fernandoa is considerably extended and becomes similar to that of Stereospermum, Dolichandrone and
Markhamia. It differs from Markhamia and Dolichandrone in not having a false septum and a spathaceous
calyx; it differs from the continental Asian genus Barnettia in not having a cruciate septum, shortly
ellipsoid-oblong capsules and pseudo-whorled leaves. Its closest ally is Radermachera which has thinner,
linear-terete capsules which are not striate or ribbed and with thinner valves (except in R. sinica), and a
terete septum; moreover, the leaves in Radermachera are mostly 2—4-pinnate and the leaflets have no
domatia and another type of glands and never stellate hairs.

It is noteworthy that Fernandoa adenophylla shows in the aspect of flower size and shape, thick indu-
ment and especially in the leaves (leaflet texture and size diminishing downwards, often auricles) a
striking similarity in habit with Markhamia (e.g. M. cauda-felina), a genus which may have also ridged
pods in Africa: but Markhamia has a 4-angled septum and a spathaceous calyx! One gets the impression
that there are signs of reticulate affinity among the Afro-Asian genera, and possibly parallel evolution,
but it must be observed that several genera are obviously not unnatural in having distinct pollen types.

From its wide distribution Fernandoa might be supposed to be an ancient genus, from which Rader-
machera, Barnettia SANTISUK (cf. Kew Bull. 28, 1973, 172) and Stereospermum are derived specialisations.

Uses. All Indo-Malesian species seem to be good timber trees and of the Indo-Chinese species it is said
that the timber is good for all purposes and not attacked by termites. They may already flower and fruit
at an early age.

KEY TO THE SPECIES

1. Leaves hairy beneath. Inflorescence terminal, rusty tomentose, with stellate and branched hairs.
Calyx thick, campanulate, 2!/,-31/, cm, woolly-tomentose outside. Corolla thick, 6-7 cm, whitish to
yellowish. Capsule twisted, rusty-tomentose, 10-ribbed 1. F. adenophylla

1. Leaves glabrous except a few short hairs of the domatia. Inflorescences lateral, with appressed, fine,
partly stellate hairs. Calyx tubular, thin, c. 1-1!/,cm by 3 mm, fine appressed stellate-hairy. Corolla
thin, pink, short-hairy outside, c. 4-5 cm. Capsule straight or + twisted, glabrous, — ss

2. F. macrolo

1. Fernandoa adenophylla (G. Don) STEEN. Blumea
23 (1976) 135. — Bignonia adenophylla [WALL.
Cat. 6502] ex G.Don, Gen. Syst. 4 (1838) 221. —
Spathodea adenophylla DC. Prod. 9 (1845) 206;
Wiaur, Ill. 1 (1839) t. 160. — Heterophragma
adenophyllum SEEM. ex B. & H. Gen. Pl. 2 (1875)
1047; Kurz, Fl. Burma 2 (1877) 236; CLARKE, FI.
Br. Ind. 4 (1884) 381; Pratn, J. As. Soc. Beng.
60, ii (1891) 322; Koorp. Ann. Jard. Bot. Btzg 14
(1897) 417; GamBie, Man. (1902) 514; Ripv. FI.
Mal. Pen. 2 (1923) 551. — Haplophragma adeno-
phyllum (DC.) P. Dorp, Bull. Soc. Bot. Fr. 72
(1925) 890; STEEN. Thesis (1927) 1006, f. 13a,
14e-f; Bull. Jard. Bot. Btzg III, 10 (1928) 265;
GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 856, f. 14.

Tree, 4-20 m, not rarely poorly developed and

with irregular crown, 10-35cm @; innovations,
thyrse and calyx with dark rusty, multicellular
compactly branched hairs. Leaves 20-50 cm long;
leaflets 2-3(—4) pairs, subsessile, the lowest often
close to the base of the petiole and smallish and
orbicular auricle-like, the terminal leaflet largest,
mostly obovate to oblong to elliptic, obtuse to
acuminate, underneath with scattered crateriform
plate glands and glandular spots and with yellow-
ish-pubescent, stellate hairs (sometimes on short,
multicellular stalks), 7-24 by 4-19cm; petiole
0-8 cm, sulcate as is the rachis. Thyrse terminal,
stout, erect, lax-flowered, c. 20cm. Calyx cam-
panulate, c. 2!/,-31/, by 11/,-2 cm, inside sordid
white, thick, persistent; lobes '/.-1 cm, subequal,
entire. Corolla yellow-brown, white, yellowish

1977]

green (brown in sicco), outside woolly tomentose,
inside glabrous, basal tube c. 1'/,-2cm, upper
tube c. 3-4cm; the mouth c. 5cm @; lobes +
entire, 1—1*/, by 1*/,-2!/,cm @. Anthers + included,
cells + free. Ovary elongate, brown stellate hairy,
lcm, style 4cm, + exceeding longer stamens.
Capsule subterete, pendulous, twisted, rusty by
stellate hairs, 30-60 by 1'/,-2'/, cm @, the valves
with 5 strong prominent ribs; septum flat, shining,
corky, over 1 cm wide. Seeds variable in size in one
capsule, c. 2'/,—33/, by 3/,—1"/, cm incl. the smallish

gs.

Distr. Continental SE. Asia (Assam to Tenas-
serim and Chittagong, Burma, Thailand, Indo-
China; also in the Andaman and Coco Is.: PRAIN,
1891); in Malesia: only in the extreme northern
part of Malaya (Langkawi, Perlis: Chupeng;
Kedah: Alor Star; in 1882 also found on Bt.
Timah in Singapore I.).

Ecol. Mixed deciduous or evergreen monsoon
forest, also in bamboo forest often on limestone
and calcareous soils, but in Burma preferring
pervious siliceous soils (KuRz); 0-750m. FI.
May-Sept., fr. March-Sept.

KOSTERMANS (n. 1436) noted that in Thailand
the ‘white’ flowers open at night and drop the next
morning. Kurz /.c. stated it leaf-shedding in
Burma and flowering at the close of the cold
season. VAN DER Pu (Act. Bot. Neerl. 5, 1956,
ar stated that the nocturnal flowers are visited by

ts.

ht .
=o} ~\
j

BIGNONIACEAE (van Steenis)

ot S Ll Ee ALA

Fig. 24. Range of the genus Fernandoa Seem.
Figures above the hyphen indicate endemic species,
those below the hyphen non-endemic species for
each area or subarea. In Africa and SE. Asia the
delimitation is generalized.

. Fernandoa macroloba (MiQ.) STEEN. Blumea 23
i976) 136. — Spathodea sp. Teysm. Nat. Tijd.
N. L. 14 (1857) 345. — Spathodea macroloba Mia.
Sum. (1861) 565; Heyne, Nutt. Pl. ed. 1, 4 (1917)
167. — Heterophragma macrolobum BACK. ex
Heyne, Nutt. Pl. ed. 2 (1927) 1371. — Haplo-
phragma macrolobum (Miq.) STEEN. Thesis (1927)
1002, f. 13b, 14a—d; Bull. Jard. Bot. Btzg LUI, 10
(1928) 263, f. 12. — Fig. 23, 25.

Tall, deciduous tree, 15-40 m (bole 15-20 m),
20-70 cm @; no buttresses; larger twigs terete.
Leaves 25-40 cm; leaflets 3-5 pairs, a few mm
stalked to sessile, obovate- to elliptic-oblong,
glabrous, beneath with a few large spot-glands,
entire, abruptly acute-acuminate, cuneate at base,

159

the lowest pair often (2-4 cm) small and roundish,
14-18 by 4-7 cm; underneath some scattered dark
glandular spots and minute hairy domatia.
Thyrses rather short (2-8 cm), lateral, narrow and
dense, densely appressed pubescent; peduncle
short; pedicels c. '/,cm. Calyx tubular, less than
halfway with 2-3 incised, short, acute lobes,
1-1'/,cm by 3mm, appressed-pubescent by
appressed partly stellate hairs and a few warty
elevated glands in upper half, faintly 5-ribbed.
Corolla white or pale pinkish, of thin texture, out-
side minutely puberulous, 4-5 cm (incl. lobes);
basal tube c. 13/, cm long, upper one wide-funnel-
shaped, c. 1*/,cm long; lobes unequal obovate-
roundish, with crenulate margin, 11/,-13/, by
13/,-2'/, cm. Stamens 4, didynamous, inserted
halfway the basal tube. Disk annular, + puberu-
lous. Ovary lanceolate, densely yellow-tomentose,
4-angular, faintly 6-ribbed; ovules in 6 rows in
each cell; style delicate, 11/,-2cm, at base ap-
pressed-hairy. Capsule linear, terete, straight or
+ twisted, glabrous, 40-65 by 1-l'/,cm @;
valves thin-coriaceous, lenticellate; septum flat,
3/,-1 cm wide, shining, of corky texture. Seeds
2-2'/2 cm (incl. the very short hyaline wing) by
/,cm.

Distr. Malesia: northern half of Sumatra:
from the Res. Minangkabau and Pariaman north-
ward to Atjeh and even on P. Breuéh.

Fig. 25. Fernandoa macroloba (Miq.) STEEN. Twig
with flowers, nat. size al HuysmMans, Nov.
1956).

160 FLORA MALESIANA [ser. I, vol. 8?

d 2 Revel "SF

Fig. 26. Lamiodendron magnificum STEEN. a. End of twig, x 1/3, b. bud, nat. size, c. ramiflorous raceme,
d. terminal raceme, both x 1/3, e. LS section of ovary, x 3, f. CS of ditto, x 5 (BRAss 25543).

12, PAJANELIA

A.P.DC. Bibl. Univ. Genéve II, 17 (1838) 130, repr. p. 14; A-DC. Prod. 9 (1845)
227; BUREAU, Mon. (1864) 35, 50, t. 20; K.ScuH. in E. & P. Nat. Pfl. Fam. 4, 3b
(1895) 244. — Fig. 27, 29.

Sal aod

1977]

BIGNONIACEAE (van Steenis)

161

Ecol. Rain-forest, up to 350 m, according to
field labels in most localities rather common; see
e.g. TEYSMANN, /.c. Fl. Oct.—Dec., fr. Dec.—May.

Uses. An estimated timber tree, used for boats
etc. (HEYNE, /.c.).

Vern. Tuwé, Atjeh, tuhi, Karo-Batak, radja,
Batak, sungké (tjirit)), M, Padang Uplands,
sungkai rimbo, Minangkabau. The name sungkai
is also used in Palembang for another timber tree
with opposite pinnate leaves: Peronema canescens

Note. DEN BERGER & BIANCHI (Tectona 24,
1931, 894-903) noted that the vessels contain a
sulphur-yellow substance colouring red with alco-
hol: lapachol; a rare feature.

This tree is cultivated in Hort. Bog. sub n.
XI-C-106A and XI-H-27; unfortunately dupli-
cates of several other numbers (XI-I-10, 20 & 50)
have been distributed under this name also; they,
however, all belong to Dolichandrone serrulata
(WALL. ex DC.) SEEM.

(Verben.).

11. LAMIODENDRON

STEEN. Nova Guinea n.s. 8 (2) (1957) 381, f. 1. — Fig. 26.

Evergreen tree. Twigs thick, hard, lenticellate, nodes with pitted glands, inter-
nodes flattened. Leaves 1-pinnate; leaflets without domatia. Racemes or panicles
terminal or from the twigs, hairy and with numerous capitate-glandular hairs.
Thyrses terminal, sometimes reduced to short-racemiform. Flowers large, not
articulated with the pedicels. Pedicels bracteolate, often with abortive flowers in
the axil. Calyx halfway split, valvately 5-lobed already in bud, thin, veined. Corolla
campanulate, hardly zygomorphic, without basal tube, glabrous, glandular-punc-
tate; lobes 5, imbricate in bud, rounded, entire. Stamens 4 and a staminode,
inserted in the hairy throat. Ovary 4-sulcate, each cell with 2 placentas on the sep-
tum, ovules very many, in many rows; stigma with 2 oblong-spathulate lips.
Capsule (immature, 4'/, cm long) linear, shortly beaked, oval in CS with a flat
septum on which developed winged ovules; pericarp with 2 fine grooves in the
middle of the valves and obviously the capsule loculicid.

Distr. Monotypic. Malesia: E. New Guinea: Milne Bay Distr., d’Entrecasteaux Is. (Normanby I.) and

Louisiades (Rossel I.).

Ecol. Coastal lowland forest, rain-forest and sago swamps.
Taxon. Possibly related to Fernandoa, but lacking domatia and with small crateriform glands on the
leaflets beneath; the fruit structure may yield further criteria.

1. Lamiodendron magnificum STEEN. Nova Guinea
n.s. 8 (2) (1957) 381, f. 1; Blumea 18 (1970) 563. —

. 26.

Tree, 12-20m; flush purplish green. Leaves
4-5-jugate, glabrous, 30-50 cm; petiole (2'/,-)6-
10cm; petiolules 1-2 cm; leaflets firmly charta-
ceous, ovate-oblong, acuminate, oblique with
unequal base, 7-19 by 3'/,-9 cm; nerves 7-11 pairs;
beneath with scattered, small, pitted crateriform
glands. Thyrses glandular-hairy, 2-3 times
cymosely branched, up to 18cm, sometimes
seemingly from old wood reduced to racemiform
and rachis 1-3 cm. Bracts ovate-acute, 3-4 mm.
Pedicels 1'/,-2'/, cm, with 1-2 pairs of bracteoles
3-5 mm long. Calyx 2-3cm, with a few glands,
rather densely short capitate-glandular hairy;
lobes wide-ovate, mucronate, 1-1'/, by */,-1 cm.
Corolla showy, brilliant orange to apricot, dark

rple-red veined, with many small glands and the
obes (and tube) laxly capitate-glandular hairy,
8-10 by 4'/,-6 cm; lobes '/,-2 cm long, 3-3'/, cm
wide. Stamens 5-7 cm long, inserted near the base

of the corolla, at the base densely hairy; staminode
2 cm; anthers with free, divaricate cells, not versa-
tile, reflexed, 6mm long. Ovary quadrangular-
fusiform, 1 cm long, with large impressed glands,
moreover covered by dense, microscopical 1-celled
hairs and short-stalked peltate to capitate-glandu-
lar hairs; style 4-6cm; stigmatic lobes oblong-
acuminate, 4 mm.

Distr. Malesia; E. New Guinea (Milne Bay
Distr., Raba Raba Subdistr., Biniguni, May I track,
9°38’ S, 149°18’ E; Northern Distr., Popondetta
Subdistr., road to Gona-Arunda; near Wanigela;
near Alotau), Normanby and Rossel Is.

Ecol. In Normanby I. forming a community on
a gravel bank behind the beach fronting swamp
forest, near Wanigela in sago swamps, in Rossel I.
at 10m in a rain-forest on a stream bank, near
Biniguni in poor lowland rain-forest at 90 m, in
Popondetta in disturbed forest, near Alotau
CRUTTWELL Saw a Single tree.

Note. Worthy of introduction into cultivation
for the large showy flowers.

162 FLORA MALESIANA [ser. I, vol. 8?

Glabrous tree without buttresses, robust in all its parts. Leaves |-pinnate; rachis
and petiole sharp-keeled above. Thyrses coarse, erect, terminal; branches scattered,
1-3 times dichotomous. Calyx closed in bud, campanulate, coriaceous, with 5
irregular lobes, articulate ventricose-campanulate,

sey piad COCA
Wy Yl by

SSSS > SS
a
dl flsy
Vil

LCE

Wl

{(CUG Ue CULO
Y

“We

(Cec d(C (
PAPAL S nah hg fee
Mf

(CGC
Yyf 2. TD ha

Fig. 27. Pajanelia longifolia (WILLD.) K.SCH. a. Fragment of leaf, X */2, b. bud, c. flower, both nat. size,

d. capsule, e. ditto in CS, both x 1/,, f. seed, nat. size (a—c after WALLICH, Pl. As. Rar. 1, t. 95/96, e-f
Kerr 18547).

1977]

BIGNONIACEAE (van Steenis)

163

+ constricted above basal tube, zygomorphous; lobes 5, spreading to recurved,
nearly equal, 2 lobes connate halfway up, imbricate in bud. Stamens 4, inserted at
apex of lower tube, didynamous, subexserted, Sth rudimentary; filaments thick,
anther-cells divergent. Disk large, annular, fleshy. Ovary oblong-cylindric; ovules
in each cell o4, on 2 placentas, co-seriate. Capsule flat, obovate-lanceolate, tipped,
valves broadly winged along the margin. Seeds in several rows on the edges of the

septum, compressed, hyaline-winged.

Distr. Monotypic, from Malabar eastwards through SE. Asia (E. Bengal: Khasia, Sylhet; Burma:
Pegu, Travancore, Chittagong, Tenasserim; Thailand, Andaman Is.) to West Malesia: N. Sumatra (E.
Atjeh), Malaya (Perak, N. Kedah, ?Singapore, ?Penang), and the Natuna Is. (Sedanau, Bunguran,
Duperré) in the S. China Sea, NW. of Sarawak. Fig. 28.

F.-VILLAR (Nov. App. 1880, 150) recorded it erroneously from the Philippines.

Ecol. Lowland evergreen mixed forests, in the Andaman Is. also in deciduous forest.

I assume bats visit the probably nocturnal flowers, but there are no direct records.

1. Pajanelia longifolia (WILLD.) K.Scu. in E. & P.
Nat. Pfl. Fam. 4, 3b (1895) 244; STEEN. Bull. Jard.
Bot. Btzg III, 10 (1928) 267; ibid. 12 (1932) 164,
f. 2 (map). — Pajaneli RHEEDE, Hort. Mal. 1 (1678)
79, t. 44, sphalma in textu t. “45”. — Bignonia
indica var. B LINNE, Sp. Pl. (1753) 625. — Bignonia
longifolia WILLD. Sp. Pl. 3 (1800) 306. — Bignonia
pajanelia BucH. Hao. Trans. Linn. Soc. 13 (1821)
516, nom illeg., quoting WILLD. — Bignonia multi-
ius WALL. [Cat. 6503] Pl. As. Rar. 1 (1830) 81,

9 ; G. Don, Gen. Syst. 4 (1838) 221. —
B. multijt uga A.P.DC. Bibl. Univ. Genéve II, 17
(1838) 130: A.DC. Prod. 9 (1845) 227, excl. syn.
Lour. et Pers.; Mig. Fl. Ind. Bat. 2 (1858) 758;
BurEAU, Mon. (1864) 50, t. 20; Kurz, Prel. Rep.
For. Veget. Pegu (1875) App. A: xciii, App. B: 69;
Fl. Burma 2 (1877) 237 (‘Payanelia’); GAMBLE,
J. As. Soc. Beng. 74, ii (1905) 382; Rip. Fl. Mal.
Pen. 2 (1923) 549. — P. rheedii WiGur, Ic. 4 (1848)
t. 1343-4; Ill. 2 (1850) t. 161bis; Bepp. FI. Sylv. 3
(1872) clxix, t. 21-5; CLARKE, FI. Br. Ind. 4 (1884)
384; BRANDIS, Ind. Trees (1906) 494; PARKINSON,
For. Fl. Andam. (1923) 215. — P. bijuga, lapsus
mihi, in syn. Bull. Jard. Bot. Btzg III, 10 (1928)
267. — Fig. 27, 29.

Evergreen, small to large tree up to 30 m,1 m@,
changing leaves before flowering. Innovations
resinous. Twigs terete, thick, lenticellate. Leaves
l1-pinnate, 8-12-jugate, 40-120 cm; petiole thick,
to 15cm, rachis sharply keeled above, rounded
beneath; leaflets entire, oblique, ovate-oblong,
speed, chartaceous, 10-20 by 5-8cm, 3-5 mm
stalked; nerves anterior side 9-12, posterior
6-9 pairs; beneath with scattered or heaped
crateriform glands along the midrib. Thyrses up to
1m, coarse, rachis hollow, lcm @; pedicels
2'/,cm; bracteoles minute. Flowers with a soap
smell, se dy tle inside dull-purple shaded.
Calyx densely covered with microscopical
glands and besides with scattered, rimmed, dish-
shaped, large glands; splitting into 5 irregular,
crispately-crenulate, acute lobes, tube at base with
keel-like folds, persistent. Corolla 5—7'/, cm, thick,
+ constricted above base, pubescent except at
base; lobes imbricate in bud, in anthesis reflexed,
broad-obovate, crenulate and crispate; anthers
brown to nigrescent, cells divergent. Style lon
with a 2-lobed, clavate stigma. Capsule sti

30-45 by 5-9cm (incl. the 2-3cm wide, often
splitting wings), with a dorsal ridge, and a corky
margin on which the wings; septum quadrangular
in CS, contracted in the middle, corky. Seeds in
© rows on each margin of the septum, curved,
24/4-3 by 3/,-11/, cm (incl. the hyaline wings).

Distr. As the genus. Erroneously recorded for
the Philippines by F.-VILLar, Nov. App. (1880)
150. Rarely planted (Atjeh). Fig. 28.

Fig. 28. Range of the genus Pajanelia A.P.DC.

Ecol. Lowland primary and secondary rain-
forest, in the Natuna Is. common, with plenty of
seedlings, spared in coconut stands, elsewhere
scattered, riverbanks, etc., mostly coastal, from
sea-level to 100 m; in the Ghats recorded to 700 m
altitude. Fi. Jan.-April, Aug., fr. March-April,

ug.

Indian authors cite it as being evergreen, but in
the Natuna Is. I found it shortly deciduous. Kurz
found it in Burma an evergreen tree. He recorded it
from the Andaman Is. as ‘very abundant in the
leafless jungles’, that is: monsoon forest (Rep. Veg.
Andam., Is. 1870, 12, 43, 71). PARKINSON gave no
definite clue on the leaf change.

164

FLORA MALESIANA

{[ser. I, vol. 82

Fig. 29. Tree of Pajanelia longifolia (WILLD.) K.Scu.

showing the sparingly branched, + pachycaul

habit; behind is a limestone hill with dry evergreen
forest (photogr. CoRNER, 1935, Sg. Sedili).

The calyx contains water in bud, like other
coarse-flowered members (Spathodea, Oroxylum,
etc.).

Uses. GAMBLE (Manual Ind. Timb. ed. 1922,
517) said it is a good timber with close-grained
wood. PARKINSON /.c. found it common in the
Andaman Is. where it is used for canoes, planking,
and boat-building; timber smells like teak and
seems to withstand attacks of white ants. Kurz
(1870) recorded the largest tree with a trunk of
2m @. Inthe Natuna Is. it is estimated for building
boats.

In the latter islands a decoction of leaves is used
against fever; in Malaya a hot fermentation is
applied on the body for stomach disorders
(BURKILL, Dict. 1935, 1623).

Vern. Békak gunong, bongli, kaju bonglai, M,
Malaya, kaju sémua, Natuna Is., abeuéng laut,
Meulaboh, Atjeh.

Note. For obscure reasons BEDDOME (/.c.) found
this anomalous in the family, its flower reminded
of Jasmineae; but he also included Schrebera
(Oleaceae)!

1977] BIGNONIACEAE (van Steenis) 165

13. NEOSEPICAEA

DrELs, Bot. Jahrb. 57 (1922) 500, f. 1; STEEN. Thesis (1927) 899; Bull. Jard. Bot.
Btzg III, 10 (1928) 216; Nova Guinea n.s. 8 (1957) 173; ibid. Bot. n. 3 (1960) 15. —
Haussmannia F.v.M. Fragm. 4 (1864) 148, non Hausmannia DUNKER, 1846; K.Scu.
in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 223; STEEN. Thesis (1927) 901. — Nyctocalos
subg. Haussmannia SEEM. J. Bot. 8 (1870) 149. — Pandorea sect. Leptophyllae
STEEN. Thesis (1927), 841, in clav.; Bull. Jard. Bot. Btzg III, 10 (1928) 200. — Pan-
dorea sect. Grandiflores STEEN. Nova Guinea 14 (1927) 301, in clav., pro parte. —
Tecomanthe sect. Aurantiacae STEEN. Thesis (1927) 872, in clav.; Bull. Jard. Bot.
Btzg II, 10 (1928) 203. — Haussmannianthes STEEN. Proc. R. Soc. Queensl. 41
(1929) 50. — Fig. 30, 37c.

Large lianas. Twigs with a distinct gland-field on the nodes. Leaves digitately
compound; leaflets 3-5, sessile or short-stalked, only articulated at the base of the
petiolule, mostly unequal, terminal one largest, often with a metallic hue above
(s.s.), both faces with numerous microscopical glandlets, underneath besides with

Fig. 30. Neosepicaea viticoides Die.s. a. Habit, */,, b. LS of flower, showing also disk and staminode,

x 2,¢. CS of ovary, x 12. — N. leptophylla (BL.) Stren. d. Just opened capsule and one valve from inside,

x 1/,, €. two , * "/,, f. flower, x */, (a-c LEDERMANN 9809, after Diets, d-e Beccari PP 687, f
D. BERGMANN 261).

166 FLORA MALESIANA [ser. I, vol. 8?

few, flat, scattered, larger, round glands '/,—'/, mm @. Thyrses co-flowered, ter-
minal, axillary to ramiflorous. Calyx proportionally small, cupular to campanulate,
truncate and minutely toothed, or shallowly 5-lobed by tearing. Corolla short-
campanulate or more often + curved, narrow trumpet-shaped, glabrous, or out-
side at least the lobes puberulous-papillose ; lobes deltoid, valvate, tomentose on the
inner margin, lobes or tube inside sometimes with larger hairs; tube with a dense
ring of long hairs at the insertion of the stamens. Stamens 4, didynamous, exserted,
5th rudimentary; anther-cells divaricate. Disk cupular, enveloping the ovary base.
Ovary glabrous; style exserted; stigma 2-lamellate, 2-celled, with 2 placentas in
each cell and c ovules. Capsules stipitate, c. 10-20 cm long, narrow oblong, terete
to broad-ellipsoid in section, beaked; valves boat-shaped. Seeds co, thin-winged,
rectangular.

Distr. Queensland (1 sp.) and Malesia: New Guinea (3 spp.). Fig. 31.

Ecol. Rain-forests, from the lowland up to c. 2000 m.

Note. In the Campsis-alliance possibly closest related to Pandorea, different by the almost regular
flowers, the valvate corolla lobes, the exserted stamens and style, digitate leaves, and cupular disk clasping
the base of the ovary.

KEY TO THE SPECIES

1. Corolla brown, basal tube, upper tube and lobes all about '/, cm long; lobes inside bearded, tube on
one side so 1. N. viticoides
1. Corolla red or orange, at least 3 cm long, the tube many times as long as the lobes.
2 Gorolasorances | Oem longuncl: the lobese a ac . 2 <Ldathaies « . « 6 « Mee
2. Corolla red or pink, 2!/.-7 cm long incl. the lobes.

3. Corolla 5—7 cm long, puberulous to glabrous, lobes 1-2 cm long, outside sometimes with dark glands,
inside sometimes long-hairy. Thyrses axillary. Capsule 13-20 by 3 cm, with woody valves. Seeds
(incl. wings) 5 by 14/, cm 2. N. leptophylla

3. Corolla 2'/,-3 cm long, glabrous outside except occasionally the puberulous apex of the glandless
lobes !/, cm long, inside glabrous. Thyrses terminal and from the uppermost leaf-axils. Capsule
10-15 by 2!/, cm, the valves thin-coriaceous. Seeds (incl. wings) 3 by 1 cm. Queensland. Cf. STEEN.
Nova Guinea n.s. 8 (1957) 174 N. jucunda (F.v.M.) STEEN.

el ete eee ah elie Ae ets ile | 6P ets wi Ste 2s 6 erin ok ie 6 ele Om a ele Re

3. N. aurantiaca

ook ee is es. aii « See ek Jamese) eae ie (oh ve le) Aenate

Caw See aes «ae @ ee ee ph (ef

1. Neosepicaea viticoides DiELs, Bot. Jahrb. 57
(1922) 500, f. 1; S. Moore, J. Bot. 61 (1923) Suppl.
38; STEEN. Thesis (1927) 900; Bull. Jard. Bot.
Btzg III, 10 (1928) 217; J. Arn. Arb. 28 (1947) 423;
Nova Guinea n.s. 8 (1957) 174; ibid. Bot. n. 3
(1960) 15. — Fig. 30a-c.

High-climbing canopy liana. Leaflets (3-)4-5,
oblong-elliptic, acuminate, chartaceous, often
with a metallic hue above, sessile to 11/, cm stalked
by the tapering base; 8-17 by 3-71/, cm; petiole
3-10 cm. Thyrses terminal and in uppermost leaf-
axils, 5-20 cm long. Pedicels c. 1/,—'/, cm. Calyx
cupular, 3-5 by 3-5 mm, shallow-lobed, glandless.
Mature corolla 2'/,-3 cm, outside papillose, basal
tube about as long as the widened upper tube and
as the lobes, brown, with dark streaks inside; lobes
long-hairy within, 3-4 mm; tube with a long-hairy
zone inside to the insertion of the stamens.

No fruit has as yet been collected, and it has only
once been found in West New Guinea.

Distr. Malesia: East New Guinea (18 collec-
tions), in W. one.

Ecol. Rain-forest, not rare, from sea-level to
c. 1500 m. Fl. Jan.—Oct.

Notes. A homogeneous species, but the flower
colour is variously defined probably in part due to
the age of the flowers: dull ochre, dull yellow with
maroon markings, brownish olive, dull red or
brick-purple, brown and purple within; petals
velvet red. Young leaves are purple.

Fig. 31. Range of the genus Neosepicaea DIELS; in
New Guinea 2 (?3) and in Queensland 1 endemic
species.

2. Neosepicaea leptophylla (BL.) STEEN. Nova
Guinea, Bot. n. 3 (1960) 15. — Tecoma leptophylla
BL. Rumphia 4 (1849) 35, quoad flor.; Mus. Bot. 1
(1849) 27; Mia. Fl. Ind. Bat. 2 (1858) 758; Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 197; K.ScH. in

1977]

BIGNONIACEAE (van Steenis)

167

K.Sch. & Laut. Fl. Schutzgeb. (1900) 540. —
Gelseminum leptophyllum O.K. Rev. Gen. Pl. 2
(1891) 480. — Pandorea leptophylla Boer. Hand.
2 (1899) 600; Diets, Bot. Jahrb. 57 (1922) 449;
STEEN. Nova Guinea 14 (1927) 301, t. 33; Thesis
(1927) 843; Bull. Jard. Bot. Btzg III, 10 (1928) 200;
Proc. R. Soc. Queens]. 41 (1929) 46, 56. — N.
superba STEEN. Nova Guinea n.s. 8 (1957) 173;
ibid. Bot. n. 3 (1960) 15. — Fig. 30d-f.

Large liana, up to 20m, 3cm @. Leaflets 3-5,
oblong elliptic, acuminate, chartaceous to coria-
ceous, often with a metallic hue above, base
cuneate in degree, from sessile to not rarely
narrowed to a pseudo-petiolule to 3cm long;
6-18 by 3-llcm; petiole 5-12cm. Thyrses
obviously axillary, or on old wood, 5-30 cm long.
Pedicels */,-1*/, cm. Ca/yx cupular, not appressed
to corolla, 4-7 by 6-7 mm, margin + entire to
shallowly lobed, glabrous, often with a few glands.
Mature corolla 5—7 cm (incl. the lobes c. 1-2 by
1/,—/,cm), tube 2!/,cm wide at mouth, slightly
curved, outside glabrous or papillose (except at
base), purplish (see notes), lobes outside papillose
or glabrous, with few to several dark glands or
glandless, inside sometimes with long hairs; tube
inside glabrous or with a few or a line of long hairs.
Capsule 13-20 by 2'/, cm, terete or oval in section,
the valves woody. Seeds (incl. the wings) 4-5 by
11/, cm.

Distr. Malesia: New Guinea.

Ecol. Rain-forest from the lowland to c. 2000 m.
Fl. March—May, July—Nov.

Vern. Je-up, Kebar (once noted).

Notes. Fruits have only once been collected by
Beccari (Arfak Mts).

All inflorescences hitherto observed are axillary,
a single one is from old wood.

The flower colour seems to vary, and to change.
Vink (BW 11404) noted buds: calyx green, tube
pale green, lobes purple; submature flower base of
tube yellow, further orange-red, inside yellow,
orange-red veined. PULLEN (7729) noted on mature
flowers: pale purple, lobes recurved, inside white
with purple streaks. Others say merely flowers pale

violet ; PLEYTE (887) noted yellow. As in N. jucunda
the lobes obviously are refiexed in mature flowers.

I have reduced here N. superba which was dis-
tinguished by the papillose-puberulous corolla,
dark glands on the outside of the lobes, and long
hairs within extending in a narrow line into the
tube. Three collections have this well expressed
(BRASS 23829, PAYMANS 53 and PULLEN 7729).
However, there are other specimens defeating these
characters: SATAKE 834 and PLEYTE 887 have no
hairs in the tube and hardly any glands on the lobes;
BeEccarI s.n. has a puberulous corolla, but no long
hairs; BERGMANN 226 has an occasional hair in the
tube and an occasional gland on the lobes, but a
glabrous corolla; BERGMANN 261 has the long hairs
inside, but a glabrous corolla without glands and
thus comes nearest to BLUME’s type.

The Papuan species is close to N. jucunda from
Queensland. This has a much smaller, glabrous
corolla except occasionally the puberulous tip of
the lobes, the latter constantly much smaller; also
the thyrses are generally terminal whereas they
always seem lateral in N. leptophylla; its fruit is
smaller and the valves much thinner.

3. Neosepicaea aurantiaca (DIELS) STEEN. Blumea
15 (1967) 298. — Tecomanthe aurantiaca DIELS,
Bot. Jahrb. 57 (1922) 497; STEEN. Nova Guinea 14
(1927) 296; Thesis (1927) 874, 834, f. 3d; Bull.
Jard. Bot. Btzg III, 10 (1928) 204. — Fig. 37c.

Leaflets 3, coriaceous; petiole 6-7 cm; blade
oblong, acuminate, cuneate at base, 10-15 by
4'/,-6 cm; nerves 8-10 pairs. Thyrses axillary;
peduncle 8cm. Calyx 5-8 by 8-9 mm. Corolla
orange, c. 10 cm long, 2!/,—23/, cm wide at apex;
lobes inside puberulous, 11/,-2!/, cm.

Distr. Malesia: East New Guinea (Etappenberg,
2 haa Distr.: LEDERMANN 9561, Bt), once found at

Om.

Note. The cupular calyx and thyrsoid inflores-
cence stamp this as a Neosepicaea. It might turn
out to be an exceptionally large-flowered form of
N. leptophylla.

14. TECOMANTHE

BAILL. Hist. Pl. 10 (1891) 41; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 230;
Boer. Handl. 2 (1899) 590; Diets, Bot. Jahrb. 57 (1922) 496; STEEN. Nova
Guinea 14 (1927) 294; Thesis (1927) 864; Bull. Jard. Bot. Btzg III, 10 (1928) 201,
incl. sect. Dendrophilae, Volubiles et Saxosae STEEN. I.c. 205, 208, 210; Pac. PI.
Areas | (1963) 288, map. — Campana RuMPH. ex Post & O.K. Lexicon (1904) 95,
nom. inval. — Pandorea sect. Grandiflores STEEN. Nova Guinea 14 (1927) 301, pro
parte. — Fig. 32-36.

Small to large lianas, climbing or creeping (in mountain heaths). Glands on
twig-nodes small. Leaves 1-pinnate, 1—7-jugate; leaflets entire or toothed, under-
neath very finely punctate-glandular. Racemes short, pendent, from efoliate nodes
on old wood, very rarely axillary or terminal; peduncle short, with some crowded,
small, sterile bracts at base, in fruit thickening like a brachyblast; rachis short (up
to c. 7(-13) cm); flowers opposite, in the axil of a small, narrow fugacious bract.

[ser. I, vol. 8?

FLORA MALESIANA

168

1977] BIGNONIACEAE (van Steenis) 169

Pedicels with 2 small, narrow bracteoles. Calyx closed in bud, persistent, c. 11/,-4
em, with 5 fairly large, deltoid + equal lobes short-hairy along the margin,
rarely split on one side. Corolla infundibuliform, the basal tube gradually widening
upwards, + straight, mostly pink, c. 5—12 cm long, inside near the insertion of the
stamens stuppose or lax-hairy and sometimes with capitate-glandular papillae;
limb mostly slightly zygomorphous, very rarely distinctly zygomorphous; lobes in
bud narrowly imbricate, mostly deltoid. Stamens didynamous, mostly included,
Sth rudimentary; anther-cells almost free, (in Mal.) c. 4 mm and almost always
divaricate. Disk thick, annular. Ovary glabrous, in each of the 2 cells with c» rows
of ovules attached on 2 placentas on the dissepiment; stigma long, filiform, with 2
spathulate stigmas. Capsule linear-terete or flattened, stipitate and beaked, with
2 coriaceous or almost woody, smooth, wide or very narrow boat-shaped valves.
Seeds o, orbicular, with fairly large, thin-membranous wings.

Distr. Species 5, 1 in the Three Kings Is. at the N. tip of New Zealand, 1 in East Queensland, the others
in Malesia: Moluccas (Ternate, Halmaheira, Ambon, Ceram, Aru Is.), throughout New Guinea (incl.
Misool, Biak, Jappen, New Britain and Woodlark I., Trobriands), and ?Solomons (Bougainville). Fig. 34.

Ecol. Primary and secondary rain-forest, mossy forest and mountain heaths, from sea-level up to
3100 m.

One collector noted honey in the flowers. It is likely that the diurnal flowers are bird-pollinated, but
there are no records.

Tecomanthe speciosa was derived from cuttings of the single plant found in nature. It has in cultivation
produced flowers and can obviously propagate by self-pollination. The corolla is sometimes (in the
herbarium) already + open before full maturity. In one case rather long, unbranched, pendent, aerial
roots were observed emitted from a node. In juvenile specimens leaves tend to be more toothed than in
mature foliage.

Taxon. As I stated before Tecomanthe belongs with Pandorea to a distinct circum-Pacific affinity of
lianas, including Campsis in East Asia and North America and Campsidium in Chile, all sharing a similar
shape of flower and fruit. The South African genus Podranea is more remote and with its inflated calyx
and linear capsules possibly more allied to Tecoma. The four genera can be keyed out as follows:

1. Flowers in racemes, opposite. Evergreen, not climbing with roots. Calyx with well-developed lobes,
1-4cm long. Corolla with a hair-ring (lax or stuppose, sometimes replaced in part by capitate-
glandular papillae) near the insertion of the stamens.

Racemes almost always on the old wood, rarely axillary or terminal. Peduncle at the base with
crowded sterile bracts, rachis up to 7 cm, pendent, flowers close together. Calyx large, 1/,—4 cm.
Corolla 6-12 cm, tube not contracted below the slightly or distinctly zygomorphous limb; lobes
deltoid, in bud narrowly imbricate. Anthers mostly inciuded, cells almost always divaricate. Valves
of the capsule widely or narrowly boat-shaped; endocarp not removable .... . Tecomanthe
2. Racemes terminal on leafy twigs. Peduncle at base without bracts; flower pairs spaced. Calyx
c. 1 cm. Corolla c. 3'/, cm long, tube contracted below the limb; lobes rounded. Stamens + exserted;
anther-cells free but parallel. Capsule narrowly elliptic-oblong, with removable papery endocarp.
NE itis TW Macys eg. i's de) Seed? hth) a Tk Wi eI RIS Ce Ny | my Campsidium

1. Flowers paniculate in terminal thyrses, very rarely depauperate in racemes but then the calyx much

smaller than 1 cm. Corolla lobes widely imbricating. Anther-cells divaricate.
3. Deciduous, climbing with roots. Leaflets 5 pairs, distinctly sharply serrate. Calyx large, c. 1'/,-3 cm,
with large lobes. Corolla inside glabrous, large, c. 5'/,-8cm ...........004 Campsis
3. Evergreen, not climbing with roots, rarely erect (in arid country). Leaflets 1-7 pairs, not distinctly
sharply serrate in mature specimens. Calyx small, stunted or short-lobed, 2!/;-8 mm. Corolla smaller,
1'/,-3'/,(—5S) cm, almost always with a hair-ring near the insertion of the stamens and long hairs one-
en Cisne Minn tomer pat Of Cie Fi ey ds ck we ee 0 8 peatrw late Pandorea

Uses. Almost all species are ornamental and can be cultivated in tropical and subtropical countries;

they are not hardy. Propagation by cuttings or seed.

KEY TO THE SPECIES AND SUBSPECIES

1. Corolla cream-coloured, woolly tomentose in the upper half, 6-8 cm long, the limb very zygomorphic.
Stamens exserted, anthers c. 10 mm, with parallel cells. Calyx often split on one side to the base, the

Fig. 32. Tecomanthe dendrophila (BL.) K.Scu. a. Habit, '/,, b. pistil, c. anther, both enlarged, d. capsule,
* "/,, — T. ternatensis Steen. ¢. Leaf, f-g. inflorescence in bud and flower, all x */, h. CS of ovary (a-c
after BLuMe, d LAE 58656, e-h BeGuin 1201).

FLORA MALESIANA [ser. I, vol. 8?

Fig. 33. Tecomanthe dendrophila (BL.) K.Scu. in the mossy forest on Mt Cycloop, N. New Guinea, at
1200 m (photogr. VAN ROYEN).

1977] BIGNONIACEAE (van Steenis) 171

lobes very unequal. Leaflets 5, orbicular-elliptic, apex broadly rounded to slightly notched, 8-18 by

5-11 cm, fleshy, coriaceous when dry. Capsule terete, pointed at both ends, c. 16 by 2'/; cm; valves

thick, almost woody. Seeds 3-4 by 1!/, cm (incl. wings). Cf. HUNTER, Rec. Auckl. Inst. Mus. 5 (1958) 41,

pl. 6-7; Hunt, Bot. Mag. 179 (1972) t. 618. Three Kings Is. (New Zealand) T. speciosa Out.

1. Corolla at most puberulous in the upper half of the lobes, not very zygomorphic. Anther-cells 3-4 mm
long, divaricate. Calyx never split on one side to the base, lobes equal or unequal. Leaflets acute to
acuminate.
2. Uniseriate hairs near the insertion of the stamens very lax, few or almost absent. Rachis of raceme
glabrous. Corolla 4"/,-7 cm long.

3. Leaflets in 3-4 pairs, c. 31/,-6 by 1'/,-3 cm, about twice as long as wide, herbaceous, veins between
the main nerves usually distinct; lower lateral petiolules 0-3 mm. Calyx c. 11/, cm long (incl. lobes).
Corolla whitish, later pink tinged, 4'/,-6 cm, the lobes 2-5 by 10-15 mm, with dark dots in trans-
parent view. Stamens as long as the style, + exserted. No capitate-glandular papillae near the inser-
tions of the stamens 2. T. ternatensis

3. Leaflets in 2 pairs (4 or 5), c. 3-8'/, by 11/,-4 cm, at least twice as long as wide, often narrower,
obviously rather fleshy, at base rounded to truncate, often oblique, veins between the 4-5 main
nerves hardly visible; lower lateral petiolules 4-9 mm, longer than upper ones. Calyx 2'/,-3 cm long
(incl. lobes). Corolla tube pale, limb pink to rosy- purplish, tube marked with purplish lines inside,
4*/,-7 cm, lobes ovate-triangular, (?/,-)1—-1°/, by (@/s-)1"/.-1?/, cm, without dark dots. Stamens +
shorter than the style, not exserted. Near the insertions of the stamens mainly capitate-glandular
papillae and no or few uniseriate hairs. Capsule 5'/, by 2 cm. Cf. C. T. WHITE, Queensl. Nat. 4 (1920)
100, f.; STEEN. Proc. R. Soc. Queensl. 41 (1929) 49. Queensland. .... T. hillii (F.v.M.) STEEN.

2. Hairs near the insertion of the stamens in a stuppose ring. Corolla 6-13 cm. Rachis of inflorescence
often puberulous.

4. Leaves 1-2-jugate; leaflets fairly large, averagely 5-10 by 2'/,-5 cm, mostly herbaceous, usually
entire, occasionally with a few coarse teeth to apex; rachis not winged. Lateral pedicels 2-8 mm.
Racemes finally rich-flowered (6-20), on the old wood of coarse lianas. Capsule broad-elliptic in
section, c. 17—22(—30) cm long, c. 3(—33/,) cm wide and come the valves wide-boat-shaped, hard,
almost ‘woody. Seeds including wing 2!/,—31/, by 1*/,-1/.¢ 1 (ee Ie dendrophila

4. Leaves 2-7-jugate; leaflets small, averagely 1'/,-4(—8) by a 13/,(-33/,) cm, herbaceous to coriaceous,
the margin almost always toothed; the sulcate rachis (very) narrow-winged ; lateral pedicels 0-2 mm.
Racemes pauciflorous (2-6), lateral or terminal, on small, slender lianas. Capsule flat and com-
pressed, 8-14 cm long, the valves very much compressed- boat-shaped, coriaceous. Seeds including
wings c. 11/,-2 by 1-11/, cm.

5. Leaves 3-6(—7)-jugate, 4-jugate leaves always present. Leaflets thick-coriaceous to chartaceous
(or even herbaceous), elliptic with usually short, acute apex, brittle in the herbarium, the nerves
and midrib usually impressed above, never strongly prominent. Calyx 11/,-3 cm. Corolla 6-10 cm.
Staminode usually less than 1 cm long 3. T. volubilis

8: rpc 2(-3)-jugate, 4- and more-jugate leaves absent. Leaf-apex acute to cuspidate. Staminode

7-3 cm.
6. Leaflets coriaceous, tough when mature, not easily breakable, sharply toothed, with nerves and
veins conspicuously prominent on both sides. Calyx (13/,—)2-3'/, cm. Corolla 7-12 cm.
3b. T. volubilis ssp. tenax
6. Leaves coriaceous to herbaceous, brittle in the herbarium, toothed but not sharply so; midrib
and veins flat or slightly sulcate above, somewhat prominent beneath. Calyx 11/,-2'/, cm. Corolla
Dea). SPasOOA, Sass 2 294i0L . bya. we. We 0h om 3a. T. volubilis ssp. silvicola

1. Tecomanthe dendrophila (BL.) K.Scu. in K.Sch.
& Laut. Fl. Schutzgeb. (1900) 539; Recu. Denk-
schr. K. Ak. Wiss. M.-N. KI. Wien 89 (1913) 603;
Diets, Bot. Jahrb. 57 (1922) 496; STEEN. Nova
Guinea 14 (1927) 297; Thesis (1927) 880; Bull.
Jard. Bot. Btzg III, 10 (1928) 206; Laur. Bot.
Jahrb. 62 (1928) 292; Loruian, J. R. Hort. Soc. 83
(1958) 295; Sykes, Stud. Cult. Pl. N.Z. 1 (1966) 43,
f. 18; Herkvorts, Fl. Trop. Climb. (1976) 73, f. 97.
— Campana rubra RumpH. Herb. Amb. (1755)
Auct. 42; an Pandorea?, Merr. Int. Rumph. Herb.
Amb. (1917) 469. — Tecoma dendrophila BL.
Rumpnia 4 (1849) 35, et Dendrophila trifoliata BL.
sub. t. 190; Mus. Bot. 1 (1849) 25; Mig. FI. Ind.
Bat. 2 (1858) 757; Ann. Mus. Bot. Lugd.-Bat. 1
(1864) 197; K.Scu. Bot. Jahrb. 9 (1887) 218; FI.
Kais. Wilh. Land (1889) 123; Wars. Bot. Jahrb. 13
(1891) 418; F.v.M. Descr. Not. 9 (1890) 64. —
Tecoma amboinensis BL. Rumphia 4 (1849) 35;
Mus. Bot. 1 (1849) 26; Mio. Fl. Ind. Bat. 2 (1858)
757. — Campsis dendrophila Seem. J. Bot. 5 (1867)

373. — Campsis amboinensis SEEM. l|.c. 374. — T.
bureavii BAILL. Hist. Pl. 10 (1891) 41; K.Scn. in
E. & P. Nat. Pfi. Fam. 4, 3b (1894) 230. — Gelse-
minum amboinense et dendrophilum O.K. Rev. Gen.
Pl. 2 (1891) 479. — Pandorea dendrophila BorRv.
Handl. 2 (1899) 600. — Pandorea amboinensis
Boer. l.c. — T. gloriosa S. Moore, J. Bot. 61
(1923) Suppl. 38; Steen. Nova Guinea 14 (1927)
299; Thesis (1927) 888; Bull. Jard. Bot. Btzg III,
10 (1928) 210. — 7. venusta S. Moore, J. Bot. 61
(1923) Suppl. 38; Steen. Nova Guinea 14 (1927)
298; Thesis (1927) 897, f. 5b, incl. var. parviflora
Sreen.; Bull. Jard. Bot. Btzg III, 10 (1928) 216;
Hunt, Bot. Mag. 180 (1975) t. 693; HerKLots, FI.
Trop. Climb. (1976) 71, f. 96, col. pl. 7. — T. ellip-
tica Steen. Nova Guinea 14 (1927) 296, t. 34D;
Thesis (1927) 876, f. Sa; Bull. Jard. Bot. Btzg III,
10 (1928) 205. — T. acutifolia Steen. Nova Guinea
14 (1927) 297; Thesis (1927) 879; Bull. Jard. Bot.
Btzg II, 10 (1928) 206. — T. amboinensis STEEN.
Nova Guinea 14 (1927) 298; Thesis (1927) 890;

172

FLORA MALESIANA

[ser. I, vol. 8?

Bull. Jard. Bot. Btzg III, 10 (1928) 211. — T.
gjellerupii STEEN. Nova Guinea 14 (1927) 298;
Thesis (1927) 896; Bull. Jard. Bot. Btzg III, 10
(1928) 215. — Fig. 32a-d, 33.

A tall liana, up to 20(—30) m. Leaves 1-2-jugate;
leaflets ovate to elliptic or oblong-lanceolate,
herbaceous to chartaceous, entire or with a few
coarse teeth to the top, apex rather blunt to acumi-
nate, (3—)5—13 by (1'/,-)2!/.-7 cm; nerves flat above
or slightly impressed, prominent beneath; rachis
not winged; lateral petiolules 2-8 mm. Racemes on
the old wood, the rachis c. 1/,-7(-13) cm, with
usually 6-20 densely set flowers; pedicels 1-2 cm.
Calyx herbaceous to + coriaceous, (11/4,—)1'/,-4 cm
long, for */,—'/, incised, greenish tinged red to
purple-brown, the lobes triangular, blunt to
cuspidate, 8-15 by 5-10 mm, midrib prominent or
not. Corolla 7-11cm long including the broad
triangular acutish to blunt lobes 3/,-1"/, by 1-2. cm,
the tube pink, rosa or pale carmine, the lobes
creamy to yellowish, sometimes streaked with
purple lines, or pink all over, inside near the inser-
tion of the stamens stuppose-hairy. Anthers c.
4mm long, divaricate. Capsule almost cylindric,
stiped and beaked, 17—22(-30) by 3-33/, by 3 cm,
with hard, almost woody, boat-shaped valves.
Seeds including the thin wing 21/,-3'/, by 12/,-
YE an.

Distr. Malesia: Moluccas (Ambon, Ceram, Aru
Is.), New Guinea (throughout, and incl. Misool,
Jappen, Biak, Woodlark I. & New Britain), and
?Solomons (Bougainville), 120 collections.

MILLAR & VANDENBERG collected this species
(NGF 48505) at Arawa Plantation, Kieta Sub-
distr., Bougainville, cultivated in a garden ‘from a
native vine’. I feel not certain that it is native in
Bougainville; it might have been introduced from
Papua.

Ecol. In swampy or dry rain-forests, sometimes
riverine forest, once on limestone, from sea-level
up to c. 1500 m. Fl. April—Dec., fr. June—Nov.

Vern. Asee, Maibrat lang., fiyo, Wapi lang.,
Marok; Sepik: gwimbipuk, gwoimbipok, Waskuk,
sanie, Ambuti, ilei, Wagu, yakomenga, Narak &
Ganja, Mt Hagen.

Notes. Through the great increase in collections
it has appeared impossible to maintain several
formerly described species. The characters of the
leaves, calyx and corolla show transient, not
correlated variation. Though the number of herba-
rium collections in which 1- and 2-jugate leaves
occur together is restricted, they do occur on one
plant in cultivation and in the forest according to
collectors. The calyx shows a great variation in
size and degree of incision. Puberulous hairiness
may occur on the rachis, pedicels, the calyx, the
midrib and nerves beneath, the apical part of the
corolla-lobes, and on the leaf-rachis. Aberrations
are sometimes found in individual specimens: a
multi-lobed, wide calyx in BW 11242; a very thin,
tortuous, 20 cm long rachis of a lax raceme in NGF
11866 and BW 13352; once a leaf with 6 and 7
leaflets; once long unbranched roots produced
from a node of the old wood (JANOWSKI 427); an
axillary raceme (BRASS 28745).

Seemingly open flowers measure sometimes only
5 cm, but I assume this to be caused in drying of
immature flowers and tardy growth.

L

Fig. 34. Range of the genus Tecomanthe BAILL.

Figures above the hyphen indicate endemic species,

those below the hyphen non-endemic species. The

Australian T. hillii (F.v.M.) STEEN. occupies two
areas.

2. Tecomanthe ternatensis STEEN. Thesis (1927)
893; Bull. Jard. Bot. Btzg III, 10 (1928) 214, f. 1. —
Fig. 32e-h.

Large liana, up to 30 m or more; stem to arm-
thick. Leaves 3—4-jugate; leaflets elliptic, herba-
ceous, base + rounded, apex short-acute, with
some teeth towards the apex, c. 3!/,—7 by 11/,-3 cm;
veins between the nerves usually distinct; lower
lateral petiolules 0-3 mm. Racemes on the old
wood, 2-6 cm long, densely rich-flowered. Pedicels
c. 1 cm. Calyx pale green, 11/,-2 cm long, the lobes
deltoid, 7 by 5mm. Corolla whitish, later tinged
pink, 5-6 cm long including lobes, lobes wide and
short, c. 2-5 by 10-15 mm, with dark spots in
transparent view; tube near the insertions of the
stamens with very few hairs and no capitate-glandu-
lar papillae. Stamens as long as the style, +
exserted.

Distr. Malesia: Moluccas (Ternate, Halma-
heira) and NW. New Guinea (Biak I.), 5 collections.

Ecol. Primary and secondary forest, in Biak on
coralline limestone, in the Moluccas at 500-600 m.
Fl. Sept.-Nov., April.

Note. The Biak specimens are rather poor but
clearly belong to this species.

3. Tecomanthe volubilis Gipps, Arfak (1917) 179;
Diets, Bot. Jahrb. 57 (1922) 498; STEEN. Nova
Guinea 14 (1927) 299; Thesis (1927) 885; Bull.
Jard. Bot. Btzg III, 10 (1928) 209. — T. nitida
STEEN. Nova Guinea 14 (1927) 299, t. 33; Thesis
(1927) 887; Bull. Jard. Bot. Btzg III, 10 (1928) 209.
— T. arfaki STEEN. Nova Guinea 14 (1927) 300,
t. 34B; Thesis (1927) 884, f. Sd, j; Bull. Jard. Bot.
Btzg III, 10 (1928) 208. — Fig. 35a—c.

Small, slender climber, 2-5 m. Leaves 3-6(-—7)-
jugate, with 4-jugate leaves always present;
leaflets mostly glossy on both sides, dark green
above, pale beneath, mostly coriaceous, brittle in
the herbarium, ovate, obovate to elliptic, rarely
lanceolate, base usually cuneate, apex acute, rarely
blunt, margin in exposed places recurved, usually

1977]

BIGNONIACEAE (van Steenis)

173

with 1-5 pairs of bluntish teeth, 3/,—21/, by +/.-
11/, cm, usually sessile but lateral pedicels up to
2mm, nerves above usually impressed, beneath
usually prominent, often nigrescent in sicco.
Racemes axillary or terminal, rachis !/,-2!/, cm;
flowers 1-3 pairs, pedicels 3/,-11/, cm, both often

Fig. 35. Tecomanthe volubilis Gisss. a. Habit, b.
unripe capsule, c. leaf. — T. volubilis ssp. silvicola

Steen. d. Leaf. — 7. volubilis ssp. tenax STEEN. e.

Leaf, underside, f. leaflet, upper surface. All x '/,

(a VAN Roven & Steumer 7417, b Brass 9052,

c BW 3050, d ee 12904, e-~f KALKMAN
).

lax-puberulous. Calyx green suffused with red,
coriaceous to herbaceous, 11/,-3 cm, tube as long
as or up to 2 times as long as the acute triangular
lobes. Corolla pink to carmine, inside creamy,
sometimes streaked red inside, 6-8(—10) cm includ-
ing the triangular, acute lobes 1'/,-2!/, by 1/,-
3cm. Staminode usually less than 1 cm. Capsule
11-14 by 34cm, compressed; valves coriaceous.
Seeds c. 11/,cm @.

Distr. Malesia: New Guinea (Mts Arfak, incl.
Nettoti, Tamrau; Wissel Lakes, Wichmann,
Carstensz, Lake Habbema, Star Mts, Telefomin,
Bosavi), 20 collections.

Ecol. Mossy thickets and heaths, often burned,
ridges in high forest, open scrub and secondary
forest, sometimes on peaty soil or on limestone,
1250-3100 m. Fil. Sept.-Febr. (once April), /r.
Aug., Nov., Jan., April.

Vern. Basengga, Hattam lang., Arfak, daibuda,
débébuda, Kapauku lang., Wissel Lakes.

Note. At lower altitude and more shaded locali-
ties leaflets tend to be larger and less rigid and
coriaceous, with less impressed nerves above and
flat leaf margin.

3a. ssp. silvicola STEEN. nov. ssp. — T. saxosa
DrEts, Bot. Jahrb. 57 (1922) 498; STEEN. Nova
Guinea 14 (1927) 889, f. 5c; Thesis (1927) 297;
Bull. Jard. Bot. Btzg III, 10 (1928) 211. — 7.
cyclopensis STEEN. Nova Guinea 14 (1927) 298,
t. 34A; Thesis (1927) 895; Bull. Jard. Bot. Btzg III,
10 (1928) 214. — T. nitida (non STEEN.) HERKLOTS,
Fl. Trop. Climb. (1976) 73, f. 98. — Fig. 35d.

Differt a T. volubili foliolis 2(—3)-jugatis, apice
acutis vel cuspidatis; staminodiis plerumque 1+*/,-
3 cm longis. — Typus: VAN ROYEN & SLEUMER 7117
(L), NW. New Guinea, Vogelkop Peninsula,
Tamrau Mts, 1350 m.

Slender liana. Leaflets 2(-3) pairs, mostly her-
baceous to chartaceous, usually elliptic to ovate-
oblong, or lanceolate-oblong, entire but mostly
with several pairs of teeth, base cuneate to rounded,
apex acute, lateral leaflets 13/,—5(—8) by 3/,—21/,(—3)
cm, terminal one longest; pedicels +/,-6 mm.
Racemes axillary or on old wood. Calyx usually
herbaceous, 11/,—3(—3*/,) cm, halfway incised or +
less. Corolla 5—7(-9-11) cm, including the lobes.
Capsule flat, 8-12 by 2-3cm. Seeds 1*/,-2 by
1-1/, cm.

Distr. Malesia: New Guinea (throughout, but
far more common in East New Guinea, common in
Morobe Distr.), 30 collections.

Ecol. In the understorey of Nothofagus-
Araucaria and Castanopsis forest, more rarely in
moss forest and on ridges, mostly in fairly tall
forest, (1000—)1500—2400(—3000) m (once found at
80 m between Hollandia and Sentani). F/. Jan.—
Dec., fr. March, June, Oct.

Note. I regard this montane forest plant to
represent a race of 7. volubilis; it is rather variable
in foliage, some specimens looking transitional to
low altitude specimens of ssp. volubilis.

3b. ssp. tenax STEEN. nov. ssp. — Fig. 35e-f.

Differt a foliolis 2(-3)-jugatis, coriaceis, venis
nervisque utrinque perspicue prominentibus. —
Typus: LAE 60706, leg. Crorr et al., E. Papua,
S. slopes of Mt. Giluwe, 6°7’ S, 143°55’ E, fl. fr.
25~-xu1-1973 (L, iso in LAB).

174

Smallish liana, 2-10 m; branchlets puberulous
to subglabrous. Leaflets 2-jugate (very rarely 1- or
3-jugate), dark green above, pale green beneath,
glossy on both sides, ovate to lanceolate, coria-
ceous, not nigrescent, 2—5'/,(-8) by 1-2(-—4) cm,
terminal one largest, very tough, margin especially
towards apex with sharp teeth. Racemes axillary,
rarely terminal, lax, with (1—-)2-3 pairs of flowers;

FLORA MALESIANA

[ser. I, vol. 8?

bordering glades, 1800-3000 m. Fi. June, Sept.—
Jan. (once in May), fr. (twice) July, Dec.

Vern. Tserki, Enga lang., taugurapu, Tari,
igidumbroki, Mendi lang.

Note. The material is very homogeneous in 25
collections seen and though fertile characters with
ssp. silvicola are overlapping it can easily be recog-
nized vegetatively without any transitions.

rachis and pedicels + Jlax-puberulous; rachis
1-5 cm; pedicels 11/,-21/, cm. Calyx mostly thin,
2'/,-31/, cm, tube 11/,—-2(-3) times as long as the
lobes; reddish white to green with pink tinge; lobes
triangular, mucronate. Corolla pink to crimson
(once noted white), the lobes yellowish to white,
streaked red within, 7-12 cm long including the
triangular lobes c. 1/, by 1*/,cm. Capsule 8-14
by 2-3 cm (septum 2!/,cm stalked). Seeds c. 1*/,
by 1 cm including wing.

Distr. Malesia: New Guinea: Papua (E., S. &
W. Highlands, largely c. 6° S and 143-144° E), but
also in West New Guinea (Bernard Camp, Iden-
burg R.). Fig. 36.

Ecol. Mountain forest, often mossy, often with
Nothofagus dominating, sometimes in shrubs

Fig. 36. Localities of Tecomanthe volubilis Gipss
ssp. tenax STEEN.

15. PANDOREA

SPACH, Hist. Vég. 9 (1840) 136; Mon. (1864) 49; K.Scu. in E. & P. Nat. Pfi. Fam. 4,
3b (1894) 230; STEEN. Nova Guinea 14 (1927) 301, incl. sect. Parviflores STEEN., in
clay.; Thesis (1927) 294; Proc. R. Soc. Queensl. 41 (1929) 43. — Tecoma sect. Pan-
dorea ENDL. Gen. PI. (1839) 711; DC. Prod. 9 (1845) 225. — Tecomanthe sect. Mon-
tanae STEEN. Bull Jard. Bot. Btzg III, 10 (1928) 204. — Fig. 37, 39.

Lianas, only exceptionally (in arid countries) erect. Glands on twig-nodes small.
Leaves (1—)2-4(—7)-jugate; leaflets with microscopical glands and not rarely with
few to many larger crateriform scattered glands underneath; petiole in some species
with some ventral large glands near the base. Thyrses terminal, sometimes addi-
tional partial axillary thyrses in the upper leaf-axils; peduncle without sterile
bracts at the base; depauperate thyrses may appear occasionally as racemes.
Calyx closed in bud, small (less than 7 mm), cup-shaped to campanulate, stunted
or very shallowly lobed, sometimes tearing. Corolla generally small, the tube
cylindric or infundibuliform, incl. lobes at most 5 cm long, limb usually zygomor-
phous, the lobes small or large, in bud widely imbricating; throat and ventral side
of the tube mostly long-hairy and often with a hair-ring near the insertion of the
anthers. Stamens didynamous, almost always inserted; anther-cells divaricate,
c. 1*/,-2 mm long; 5th rudimentary. Disk annular or + pulvinous. Ovary elongate,
each cell with 2 placentas and many ovules. Capsule stipitate, + beaked, rather
thick, terete or + flattened; valves widely boat-shaped, firmly coriaceous; dis-
sepiment flat, oblong, thickish, with marginal seed-scars. Seeds many, roundish,
thin-winged.

Distr. Six spp., Central, N. & E. Australia, Tasmania, Lord Howe I., New Caledonia, Solomons
(Bougainville), and East Malesia: New Guinea (New Britain included), Moluccas, and the Lesser Sunda
Islands (Lombok, Flores, Timor). Fig. 38.

Ecol. In Malesia in rain-forest, from sea-level to 2450 m.

1977]

BIGNONIACEAE (van Steenis)

175

KEY TO THE SPECIES

1. Corolla tube glabrous outside, usually rather wide, without hair-ring inside near the insertion of the
stamens. Base of filaments glandular and also the ovary with similar sessile glands. Calyx cupular,
c. 2-3 mm. Venation not prominent above. Filaments inserted close to the base of the tube, which is

not narrowed

1. P. pandorana

1. Corolla tube (except at the very base) outside puberulous. Base of filaments without sessile glands.

Venation above mostly prominent.

2. Flowers large, 4-5 cm long (incl. lobes), white with crimson throat, the tube c. twice as long as the
rounded lobes. Leaves 2-jugate, the entire leaflets blunt at apex, nerves and veins not distinctly

prominent above. Queensland and New South Wales

P. jasminoides (LINDL.) K.ScH.

2. Flowers much smaller, at most 3!/, cm, the tube 4—6 times as long as the lobes; lobes less than c. 1 cm
diameter. Leaflets acute, nerves and veins usually distinctly prominent above, mostly toothed towards

the apex.

3. Corolla 12-15 mm long, narrow-cylindric, the tube c. 6 times as long as the lobes. No hair-ring at
the staminal base. Calyx 3-5 mm. Petiole above at base with one or a few large glands; leaf-rachis

narrowly winged.

4. Leaflets entire, 3—4-jugate, 5—12 by 2-5 cm. Corolla c. 12-13 mm, tube + curved, c. 2-3 mm wide,

no beard in the mouth and tube, lobes c. 2 mm. Ovary orbicular, with sessile glands. Venation on
upper surface of leaflets raised, but not fine-tessellate. Flowers cream-coloured, lobes and throat
pink-shaded. Queensland. Cf. STEEN. Proc. R. Soc. Queensl. 41 (1929) 46, f. 1.

P. baileyana (MAID. & R. T. BAKER) STEEN.

. Leaflets apically toothed, (1—)2—3-jugate, 6—81/, by 3'/,-4 cm. Corolla tube straight, c. 20 mm long,

c. 5mm wide, mouth and upper part of tube inside bearded, lobes c. 3-4 mm. Ovary obconical-
oblong, eglandular. Venation on upper surface of leaflets raised, fine-tessellate. (Corolla tube
wee, tne lobes Whitey. <2 <<) 3.6) 2. ee eee ee ee ee ec ee 2. P. stenantha

3. Corolla 20-35 mm long, the tube c. 4 times as long as the lobes, the mouth and upper part of the

tube inside bearded. Calyx c. 6-7!/, mm. Petiole without glands; rachis narrowly winged; leaflets
dentate in upper part, venation not fine-tessellate raised above.

5. Corolla straight, tubular, 20-25 mm long, with yellow tube, the lobes white to pale red or streaked

red, inside with a distinct hair-ring near the insertion of the stamens. Ovary + conical, eglandular.
Pedicels slender, 1-2 cm 3. P. montana

. Corolla c. 30-35 mm long, white with pale yellow mouth, infundibuliform, with a fairly narrow

lower part of the tube, widened apically, inside at the insertion of the stamens with a few hairs.

Ovary ellipsoid, with sessile glands. Pedicels c. 5mm. Queensland. Cf. Steen. J. Arn. Arb. 12

ney eee, a eT eS eee

1. Pandorea pandorana (ANDR.) STEEN. Bull. Jard.
Bull. Btzg III, 10 (1928) 198; Proc. R. Soc. Queens}.
41 (1929) 43; J. H. Wiis, Handb. Pl. Vict. 2
(1972) 578; BEADLE, EvANS & CAROLIN, FI.
Sydney Reg. ed. 2 (1972) 502; Herk ots, Fl. Trop.
Climb. (1976) 69, f. 91. — Bignonia pandorana
ANpDR. Bot. Rep. 2 (1800) t. 86. — Bignonia
pandorea Vent. Jard. Malm. (1803) t. 43. —
Bignonia pandorae Sims, Bot. Mag. 22 (1805) t. 865.
— Tecoma australis R.Br. Prod. (1810) 471; DC.
Prod. 9 (1845) 225, incl. var. meonantha (LINK)
DC.; Bra. FI. Austr. 4 (1869) 537; BAILEy,
Queensl. Fl. 4 (1901) 1134, incl. var. pandorea
(VENT.) BAILEY (= var. typ. , var. meonantha et
var. linearis BAILEY, I.c. pl. 45. — Bignonia australis
Arr. Hort. Kew. ed. 2, 4 (1814) 34. — Bignonia
meonantha Link, En. Berol. 2 (1822) 130. —

Tecoma meonantha Sweet, Hort. Brit. (1827) 284; .

G. Don, Syst. 4 (1838) 224; Harris, Wild Fl. Austr.
(1938) 151, pl. 6 (as T. australis). — Tecoma
diversifolia G. Don, Syst. 4 (1838) 225; DC. Prod. 9
tea) 225. — P. australis Spacn, Hist. Nat. Vég. 9
1840) 136; K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b
1894) 230; Diets, Bot. Jahrb. 57 (1922) 498;
. Nova Guinea 14 (1927) 302; Thesis
(1927) 859, incl. ssp. pandorea Steen. |.c. 861, ssp.
meonantha Steen. lc. 862, et ssp. linearis STEEN. l.c.
863; Domin Bibl. Bot. 22 (1929) 1153, incl. var.
oxleyi, nom. illeg., et var. meonantha, l.c. 1154. —
Tecoma floribunda CUNN. ex DC. Prod. 9 (1845)
225. — Tecoma oxleyi CuNN. ex DC. L.c.; J. M.
Brack, Trans. R. Soc. S. Austr. 39 (1915) 836;

P. nervosa STEEN.

Wuite & FRANCIS, Proc. R. Soc. Queensl. 37
(1926) 166; Harris, Wild Fl. Austr. (1938) 151,
pl. 39. — Tecoma ochroxantha KTH & BoucueE, Ind.
Sem. Hort. Berol. (1847) 12, sec. Bro. 1869. —
Tecoma leptophylla Bt. Rumphia 4 (1849) 35;
STEEN. Nova Guinea 14 (1927) 301, t. 33, pro parte,
pro fol. sol. — Tecoma austro-caledonica BUREAU,
Bull. Soc. Bot. Fr. 9 (1862) 163; Marp. Proc. Linn.
Soc. N.S.W. 39 (1914) 382; Compton, J. Linn.
Soc. Bot. 45 (1921) 373. — Tecoma ceramensis
T. & B. Nat. Tijd. N. I. 25 (1863) 412; Mig. Ann.
Mus. Bot. Lugd.-Bat. 1 (1864) 197, t. 5, incl. var.
elliptica Mig. — P. austro-caledonica Stem. Gard.
Chron. (1870) 1085; BAILL. Hist. Pl. 10 (1891) 40;
K.Scu. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 230;
GUILLAUMIN, Fl. Nouv. Cal. (1948) 317 (‘austro-
caledonicum’); Heine, Fl. Nouv.-Caléd. 7 (1976)

. 20. — Campsidium filicifolium BUuL,
Wholesale List New, Beaut. & Rare PI. (1874) fig.;
Cat., ex JOHNSON & Hoaa, J. Hort. 51 (1874) 366;
A. VAN GEERT, Cat. n. 74 (1874): T. Moore, FI. &
Pom. (1874) 280. — Tecoma filicifolium NICHOLSON,
Dict. Gard. 4 (1887) 13; cf. STEEN. Blumea 15
(1967) 146. — Gelseminum pandorea et ochroxan-
thum O.K. Rev. Gen. Pl. 2 (1891) 480. — P. cera-
mensis BAILL. Hist. Pl. 10 (1891) 40; K.Scu. in
BE. & P. Nat. Pfl. Fam. 4, 3b (1894) 230 (‘ceramica’);
Steen. Nova Guinea 14 (1927) 302; Thesis (1927)
852. — Tecoma pandorana ‘SKEELS, U.S. Dep.
Agric. Bur. Pl. Ind. Bull. 282 (1913) 62. — P,
acutifolia Sreen. Nova Guinea 14 (1927) 303,
t. 34C; Thesis (1927) 855; Bull. Jard. Bot. Btzg UL,

[ser. I, vol. 82

FLORA MALESIANA

176

plants, both
New Guinea,

ge of juvenile
b ZIPPEL s.n.

/2(a after BLUME,

c LEDERMANN 9561).

1

Owers and fruit, b. folia

(ANDR.) STEEN. a. Habit, fl
aca (DIELS) STEEN. c. Flower, x

pandorana

— Neosepicaea auranti

Fig. 37. Pandorea

Xena.

1977]

10 (1928) 196. — P. poincillantha STEEN. Nova
Guinea 14 (1927) 302; Thesis (1927) 857, incl. var.
fragrans STEEN. — Tecoma doratoxylon J. M.
BLACK, Trans. Proc. R. Soc. S. Austr. 51 (1927)
383; GARDNER, En. Pl. Austr. Occ. (1930) 118. —
P. doratoxylon J. M. BLACK, Trans. Proc. R. Soc.
S. Austr. 61 (1937) 248; Fl. S. Austr. pt 4 (1957)
773, f. 1106. — Campsis pandorana STEEN. FI.
Males. I, 4 (1948) xxi. — Fig. 37a-b.

Malesian specimens: Often large liana, 20-30 m.

Leaves 2-4(-6)-jugate; leaflets usually ovate-
elliptic to oblong, mostly entire and acuminate,
3-10 by 1'/,-6cm, glabrous, underneath with
few or many scattered large crateriform glands,
nerves not prominent above; midrib sulcate;
lateral petiolules 0-10 mm; petiole without glands
at the base above. Thyrses terminal, lateral or
from old wood, c. (1—)5—20 cm, glabrous, rarely
puberulous. Pedicels c. '/,-1'/, cm, rarely longer.
Calyx cupular, stunted or short-lobed, thin, c.
2-3 mm. Corolla 1-2(-3)cm long including the
lobes, mostly rather inflated-tubular, the tube
mostly + twice as long as the lobes, glabrous
outside, lobes mostly densely papillose-puberulous,
mouth and tube inside bearded on the ventral side,
light yellow, the zygomorphous limb and tube
inside streaked or mottled red or purple dotted,
without a hair-ring near the base of the stamens
and no proper basal tube. Stamens included, at
their base glandular-papillose, inserted very near
the base of the tube. Ovary glandular-papillose.
Capsule acute, c. (5—)9-12 by (13/,-)2'/,-3 by
2-2'/,cm; valves coriaceous; dissepiment rather
thick, 6'/,-8 by 17/,-2 cm, the seed scars marginal.
Seeds c. 2'/,-3 by 1*/,-2 cm including the hyaline
wings.
Distr. Central, N. & E. Australia, Tasmania,
Lord Howe I., New Caledonia, N. Solomons
(Bougainville), and East Malesia: New Guinea
(incl. New Britain), Moluccas (Morotai, Halma-
heira, Ambon, Ceram, Key Is.), and Lesser Sunda
Is. (Lombok, Flores, Timor); 65 collections.

Though Tecoma filicifolium, a juvenile form, was
said to have come from Fiji, I have shown (1967,
Lc.) that this hailed from New Caledonia.

Sims claimed that Loppices nurseries had
received Bignonia pandorae from Norfolk L,
but this seems to rest either on an erroneous localis-
ation or on a cultivated source.

Ecol. In Malesia in primary and secondary rain-
forest, from sea-level up to c. 2000(—2400) m,
getting distinctly scarcer upwards of 1350 m. Fi.
Jan.—Dec., fr. Oct.-Dec.

Taxon. A quite well recognizable species in
spite of a fair degree of variability. This is in part
ontogenetic, the juvenile form having narrow,
many-jugate, crenate, small leaflets; these are
sometimes still found on odd twigs of mature-
foliaged plants.

main variation is in Australia in the leaves,

the rain-forest (type variety) form having ovate to
elliptic 2-jugate leaflets, whereas in drier places
jugate leaves occur with lanceolate leaflets
(descri as T. meonantha), while in still more
arid places the 2-6-jugate leaves have almost linear
leaflets (described as T. oxleyi and T. aig ded
The latter form may at times be scrambling, bushy
or even erect (spearwood bush) and carry racemose
inflorescences. Though the typical representatives

BIGNONIACEAE (van Steenis)

177

of these three forms are distinct, they are con-
nected by a clear series of specimens with inter-
mediary characters, which already induced BAILEY
to say that he named the three forms as varieties
merely for convenience. Whether these forms are
genetically different taxa (races) or merely pheno-
typic forms can only be established by experiments.
BatLey added that flowers of the type variety
would emit a strongly disagreeable odour while
T. meonantha would have fragrant flowers. This
matter must be solved by field botanists.

In Malesia the 3-4-jugate leaves are often
narrower, sessile, and also more toothed than the
2-jugate ones; it looks like a matter of lingering
neoteny.

At higher altitudes, 1300-2000 m, leaflets tend
to be more coriaceous. In SCHMUTZ 3178 the veins
are by exception prominent above.

The New Caledonian form seems to be a local
race with small flowers and small roundish, dentate
leaves.

In rain-forest the leaflets are usually entire, or
with a few coarse teeth towards the apex, and
usually they have underneath a fair number of
crater-like larger glands, in Australian specimens
these are scarce or absent.

Lateral petiolules are usually short (2-5 mm),
but in HyLAND 5092 they measure 1!/, cm.

la. ssp. timorensis STEEN. nov. ssp.

Differt a speciminibus malayanis floribus com-
parate magnis (2-3 cm longis, lobis incl.), stamini-
bus atque stylo exsertis, ceterum ore atque tubo
floris barbis longis destitutis. — Typus: C. W. Kooy
363 (L), pr. Temef, S. Central Timor, fi. 18—vn—
1966, c. 800 m.

Leaflets 2-4-jugate, without crateriform glands
beneath. Calyx 24mm, stunted, minutely

5-mucronulate. Corolla 2-3 cm long incl. lobes.
Mouth and tube not bearded inside, the puberulous
papillae from the lobes extending in the tube in a
lax way. Stamens and style exserted.

Distr. Malesia; Lesser Sunda Is. (Timor); 4
collections.

Fig. 38. Range of the genus Pandorea Spacu.
Figures above the hyphen indicate endemic species,
those below the hyphen non-endemic species.

178

FLORA MALESIANA

[ser. I, vol. 8?

Ecol. On limestone (once) and along ravine in
mountain Eucalypt forest, c. 700-1000 m. FI.
March, May, July.

Vern. Tufe, Dawan lang., non amisu, Niki-Niki,
non fuléh, Mt Mutis.

Notes. Though undoubtedly P. pandorana, the
Timor race deviates within the species by lacking
the usual beard in the mouth and tube of the
corolla and in the genus by exserted stamens and
style. It is remarkable that the specimens from
Lombok and Flores do not belong to this sub-
species but agree with the Moluccan specimens.

2. Pandorea stenantha D1E.s, Bot. Jahrb. 57 (1922)
498; STEEN. Nova Guinea 14 (1927) 302; Thesis
(1927) 850; Bull. Jard. Bot. Btzg III, 10 (1928)
197. — Fig. 39e-f.

Large liana (stem to 2!/, cm @). Leaves (1-)2-3-
jugate, the leaflets coriaceous, ovate-oblong, acute,
towards apex toothed, venation raised on both
sides, above fine-tessellate, 6-8'/, by 31/.-4cm;
rachis narrowly winged; petiolules 0-8 mm;
petiole near the base above with one or few large,
sometimes raised glands, sometimes also one at the
articulation of the rachis. Thyrses axillary and

LX
its

xy
7s
[ikke

U

N)

Nxt

Fig. 39. Pandorea montana (Digs) STEEN. a. Leaf,

b. leaflet, x 1/., c. reticulate venation above, d.

inflorescence, x 14/,. — P. stenantha DIELS. e.

Inflorescence, x '/,, f. tessellate venation of leaflet

above (a, c NGF 13905, 6 LEDERMANN 9916, d

NGF 21251, e SOEGENG 360, f VAN ROYEN &
SLEUMER 5816).

terminal or on old wood, 10-15cm, fine-
puberulous. Pedicels 4-10 mm. Calyx 3-5 mm,
campanulate, with 5 short, broad-deltoid lobes.
Corolla tube yellow, the lobes white (ex typ.),
tube c. 20mm long and c. 5mm wide, outside
puberulous-papillose, the mouth and upper part
of the tube bearded, lobes c. 3-4 mm; no hair-ring
at the insertion of the stamens. Stamens included.
Disk cupular. Ovary flattened, obconical-oblong,
glandless.

Distr. Malesia; New Guinea (Sepik Distr. : April
R.; Mt Cyclops; SE. Irian: Ingembit) ; 3 collections.

Ecol. Rain-forest, 125-800 m. Fl. June, Nov.
Flowers once noted to be fragrant.

Notes. Apparently a rare species; available
flower material unfortunately rather inadequate.
VAN RoyEN noted: flowers white at base of tube
pale purple; SOEGENG: tube dirty yellow, lobes
lilac. The petiolar glands were not mentioned by
Drs in his brief description, but I noted them on
the type (1927).

3. Pandorea montana (DIELS) STEEN. nov. comb. —
Tecomanthe montana DiELs, Bot. Jahrb. 57 (1922)
497; STEEN. Nova Guinea 14 (1927) 296; Thesis
(1927) 875, f. 3c; Bull. Jard. Bot. Btzg III, 10
(1928) 204. — Fig. 39a-d.

Slender liana, to finger-thick. Leaves 3-4(-5)-
jugate, leaflets chartaceous to coriaceous, elliptic-
oblong, acute, towards apex toothed or shallowly
crenate, venation raised on both sides, but not
fine-tessellate above, c. 2-5!/, by 1-21/, cm, rachis
narrowly winged; petiole without basal glands;
petiolules almost absent. Thyrses axillary and
terminal, lax, almost glabrous. Pedicels filiform,
1-2cm. Calyx c. 7mm, with 5 short, broad-
triangular teeth. Corolla + infundibuliform,

0-25 mm long, the tube narrowed to the base,
straight, limb zygomorphous, c. 8-10 mm wide at
the mouth, the lobes 24mm, bearded in the
mouth and upper part of the tube, the lobes white
to pale red or streaked red, the tube light brown
or yellow flushed red outside, yellow inside, with a
distinct stuppose hair-ring near the insertion of the
filaments. Stamens included. Disk cupular. Ovary
+ conical, eglandular. Capsule flattened, 4-8'/,
by 11/,-3 by 1/.-1 cm, sessile to stiped and short-
beaked; valves hard, almost coriaceous to woody,
narrow boat-shaped ; dissepiment thick-coriaceous,
3-10 by 14/,-21/, cm. Seeds 11/,-23/, by 1-2 cm.

Distr. Malesia: East New Guinea (Sepik:
Hunstein Mts: Lordberg, Hunsteinspitze, 1000-
1350 m, 4 coll.), loc. class.; Morobe Distr.: Mt
Kaindi, 7°25’, 146°45’ E, 12 coll.; Mt Giluwe,
6°10’, 144° E.

Ecol. Mountain rain-forest, often mossy,
forest edges and regrowths, 1000-1350, 2100-
2450 m. Fi. Febr., May-June, Aug.—Nov., /r.
Aug.—Sept., Dec.

Notes. The thyrsoid inflorescence and flower-
size point to congenerity with Pandorea; luckily I
had made in 1927 a small drawing of the type and
observed several flower details on the original
material not mentioned by Drets in his brief
description. Unfortunately almost all duplicates at
L lack flowers, by careless distribution. The flower
colour noted on labels varies rather considerably :
tube cream, yellow, golden brown or red outside,
lobes white, cream, pale purple, streaked red.

1977] BIGNONIACEAE (van Steenis) 179

Doubtful & Excluded

In this list are combined all names of Malesian and SE. Asian plant names which are excluded or of
which the identity is uncertain or of which I have not seen the types

Bignonia albida Bu. Verh. Bat. Gen. 9 (1823) 195; StEupb. Nomencl. 2 (1841) 204 = Aeschynanthus
albidus (BL.) STEUD. (Gesneriaceae).

Bignonia angustifolia BL. Verh. Bat. Gen. 9 (1823) 194; Cat. Hort. Bog. (1823) 82 = Aeschynanthus
angustifolius (BL.) STEUD. (Gesneriaceae).

Bignonia comosa Roxs. (Hort. Beng. (1814) 95, nom. semi-nudum] FI. Ind. ed. Carey 3 (1832) 103; DC.
Prod. 9 (1845) 144; Mig. Fl. Ind. Bat. 2 (1858) 751. — Spathodea comosa G. Don, Gen. Syst. 4 (1838)
222, said to come from the Moluccas, is according to the type (Herb. Martius, in BR) Clerodendron
lanuginosum BL. 1825 (Verbenaceae). The detached fruit on the sheet was not described and belongs to
some SE. Asian Bignoniaceae.

Bignonia compressa LAMK, Encycl. 1 (1785) 424; G. Don, Gen. Syst. 4 (1838) 220, said to come from
the East Indies, is according to PERRIER DE LA BATHIE from Madagascar (Fl. Madag. 178, 1938, 59)
(Colea decora DC. Prod. 9, 1845, 241) = Rhodocolea racemosa H. PERRIER.

Bignonia fraxinoides PERROTTET, Mém. Soc. Linn. Paris 3 (1824) 102, nom. semi-nudum, said to grow in
East = aig is cf. STEEN. Blumea 15 (1967) 146 probably not from Java,; the name ought to be discarded
entirely.

Bignonia glauca WALL. Cat. 6506, nomen, non DECNE 1844. The type at Kew, a sterile specimen, was
annotated “‘perhaps not Bignoniaceae”’ by C. B. CLARKE. It has recently further been annotated “prob.
Meliaceae”’ by ALAN RADCLIFFE-SMITH.

Bignonia hirsuta LAMK, Encycl. 1 (1785) 422; WILLD. Sp. Pl. 3 (1802) 299; G. Don, Gen. Syst. 4 (1838)
225. — Tecoma hirsuta DC. Prod. 9 (1845) 173, 222.

Said to have come from “‘l’Indes”’; leaves opposite, digitate, with 5 leaflets, stalked and with petiolules;
leaflets oblong, cuneate at base, emarginate at apex, downy, slightly pubescent beneath. Flowers small,
curved, short reddish-yellow hairy. Calyx truncate, with 4 minute teeth. Stamens 4, exserted.

Vitex (Verb.) might be involved, but all its Indo-Malesian species have acuminate leaves and in Vitex
the corolla is not curved and if hairy the corolla is greyish. It might possibly be an American plant,
Tabebuia, or allied to that.

LAMARCK described the specimen from herb. Jussieu. Dr. H. HEINE made elaborate but unfortunately
unsuccessful attempts in herb. Jussieu and Lamarck to locate the specimen in the Paris Herbarium; there
is also no trace of it at Geneva; DE CANDOLLE did not see any material.

Bignonia laeta WALL. Cat. 6505A, cf. CLARKE, Fl. Br. Ind. 4 (1884) 376, is according to SPRAGUE,
Kew Bull. (1919) 306 = Dolichandrone serrulata SEEM.

Bignonia longiflora ReEINW. msc. ex DE VriesE, Pl. Ind. Bat. Or. (1856) 9, nomen = Aeschynanthus
longiflorus (BL.) DC. (Gesneriaceae).

ignonia macrostachya WALL. [Cat. 6504, nomen] ex G. Don, Gen. Syst. 4 (1838) 221; DC. Prod. 9
(1845) 166. Mr. R. K. Brummitt, Kew, kindly remarked on this (in litt. 15—vm1—1975) that ““WALLICH
8504 consists of two rather long inflorescences in bud only (one corolla almost open) and a fairly stout
piece of stem, and bears no leaves, open flowers or fruits. It has been annotated ‘Bignonia macrostachya
WALL. (and of G. Don & DC.)’ by C. B. CLARKE, but has no more recent identification. It seems to me to
be fairly clearly referable to Pajanelia longifolia (WiLLD.) K.ScH. My opinion seems to be supported by a
specimen laid away in the main herbarium under this species in a red folder (though it is not obvious what
it is supposed to be a type of), collected in Khasiya by GrirrirH and labelled ‘Bignonia macrostachya
WALL. Cat. 6504 & — rostrata WALL. Herb. 6503A’. The specimen WALLICH 6503A does indeed also
seem to be this species. The citation of WALLICH 6505 by G. Don in validating the name B. macrostachya
is presumably an error for 6504.

Bignonia moluccana DC. Prod. 9 (1845) 144; Mia. Fl. Ind. Bat. 2 (1858) 751. — B. discolor A. Ricn.
Sert As Astrol. (1834) xxix, non R.Br. 1814, said to come from the Moluccas. The description would tally
with Gmelina asiatica Lour. (Verbenaceae), but a sheet with an original label (in P) was identified by
E. Bureau as Bignonia capreolata L. which does not agree with the description. Confusion with labels and
specimens must have taken place and the name should be discarded. Cf. Sreen. Blumea 15 (1967) 146.

Bignonia purpurea THUNB. FI. Jav. (1825) 15, nomen. Of unknown identity, not mentioned by JUEL.

Bignonia ramiflora Decne, Nouv. Ann. Mus. Paris 3 (1834) 381, repr. Herb. Timor. Descr. (1835) 53. —
? Bignonia colei G. Don, Gen. Syst. 4 (1838) 221. — Colea ramiflora DC. Prod. 9 (1845) 241; Mia. Fl. Ind.
Bat. 2 (1858) 759. — Colea colei M. L. Green, Stand. Sp. Nom. Cons. (1926) 55-63; STEEN. Bull. Jard.
Bot. Btzg III, 10 (1928) 277, excl. syn. alter. — Colea timorensis in sched., in syn., ex PERRIER.

This rests on a mislocalized specimen of Porvre in herb. Jussieu from Madagascar, and is according to
PERRIER DE LA BATHIe, Ann. Mus. Col. Marseille 46 (1938) 43 = Rhodocolea racemosa H. PERRIER.

Bignonia ternatea REINW. ex DE Vriese, Reinwardt’s Reize (1858) 495, 644, nomen = Dichrotrichum
ternateum REINW. ex DE VRIESE.

Bignoniacea incerta: ZOLL. Syst. Verz. Heft 3 (1855) 53-54, based on ZOLLINGER 2214 = Wightia
borneensis Hoox. f. ssp. ottolanderi (KOORD.) STEEN. (Scrophulariaceae).

Colea aberrans BAiL. Bull. Soc. Linn. Paris 1 (1889) 687 rests on a specimen said to have been collected
by Potvre in Timor, but came from Madagascar according to PERRIER DE LA BATHE, Ann. Mus, Col.
Marseille 46 (1938) 28, and = Rhodocolea racemosa Perrier var. humblotiana H. Perrier.

Dolichandrone falcata (WALL. ex DC.) Seem.: F.-ViILL. Nov. App. (1880) 151. According to MERRILL,
En. Philip. 3 (1923) 445, obviously an erroneous record from the Philippines of this Asian species.

180 FLORA MALESIANA [ser. I, vol. 8?

Dolichandrone tulipifera Bru. in B. & H. Gen. Pl. 2 (1876) 1046; F.-ViILL. Nov. App. (1880) 151; MerR.
En. Philip. 3 (1923) 445. This is an erroneous, non-existing combination for Spathodea tulipifera G. DON
which was used by F.-VILLar for the tulip tree, Spathodea campanulata, which he saw cultivated at Manila.

Hadongia eberhardtii GAGN. Not. Syst. 14 (1950) 30, from Indo-China, is according to VIDAL, Bull. Soc.
Bot. Fr. 106 (1959) 352 a cultivated specimen of Citharexylum spinosum L. (Verbenaceae).

Hausmannia mollis K.Scu. ex STEEN. in sched.; Thesis (1927) 902; Hausemannia mollis F.v.M. sphalm.
Ind. Kew. Suppl. 1 (1906) 16 (Hausemannia mollis K.Scu.) = Archidendron molle (K.ScH.) DE Wit

(Leguminosae).

Markhamia cauda-felina (HANCE) CRAIB: SPRAGUE, Kew Bull. (1919) 310. — Dolichandrone sp. CERON,
Cat. Pl. Herb. Fl. For. Filip. (1892) 127. Now considered to be Markhamia stipulata (WALL.) SEEM. var.
cauda-felina (HANCE) SANTISUK, cf. Thai For. Bull. Bot. 8 (1974) 15.

The collection on which this was based is VIDAL 3398 (K), from Montufar, Albay Prov., Luzon. MERRILL
noted (J. Arn. Arb. 35, 1954, 154) that it was possibly occasionally introduced for forestry purposes by
VIDAL from S. China. As no later collections were ever made its cultivation seems to have been ephemeral.

Stereospermum cylindricum PIERRE ex P. Dop, Fl. Gén. I.-C. 4 (1930) 581, a species from Indo-China and
Thailand, was mentioned by Dop, /.c. 582 to occur in Malaya, but on what evidence is unclear. I found no
sheets in Paris to corroborate this.

Tripinna tripinnata Lour. Fl. Coch. (1790) 391. — Tripinnaria cochinchinensis Pers. Syn. 2 (1807) 173. —
Tripinnaria asiatica SPRENG. Syst. 2 (1825) 842, taken for a Bignoniacea by several authors. According to
MERRILL, Comm. Lour. (1935) = Vitex tripinnata (LourR.) MERR. (Verbenaceae).

CULTIVATED BIGNONIACEAE

There are quite a number of Bignoniaceae cultivated in Malesia; they stem from all parts of the
tropics. Frequently they hardly set any seed. Tecoma stans does so profusely and this has led to
its naturalization. Jacarandas also set seed but did not naturalize.

As Bignoniaceae are often very showy plants and are largely tropical there is no end to their
introduction. Therefore the survey given below may be or at least become incomplete.

It should also be remembered that cultivated plants are often neglected by botanical explorers
and are mostly scantily represent in herbaria.

These introduced species have mostly not been critically studied by me, but it is assumed that
their names are correct. I acknowledge with great thanks the loyal collaboration of the late Mr.
N. J. SANDWITH (Kew) who formerly named at my request certain introduced species, and of
Dr. A. H. Gentry (Missouri Botanical Garden, St. Louis) who was so kind as to check this
appendix.

Some papers or appendices are dedicated solely to cultivated Bignoniaceae or have taken them
up and often give keys and illustrations:

BACKER, C. A. & BAKHUIZEN VAN DEN BRINK Jr, R. C. 1965. Flora of Java 2: 534-542.
CHATTERJEE, D. 1948. A review of Bignoniaceae of India and Burma. Bull. Bot. Soc. Beng. 2:

75-79.

Faris, H. A. 1959. Las plantas cultivadas de la Republica Argentina. Bignoniacas. Inst. Bot.

Agr. 10, fasc. 173: 57 pp., 25 fig.

GentTRY, A. H. 1973. Ann. Mo. Bot. Gard. 60: Flora of Panama, part IX, fam. 172: 781-977, 41
fig.

HEINE, H. 1976. Flore de Nouvelle-Calédonie 7: 91-93.

HERKLotTS, G. 1976. Flowering tropical climbers: 63-74, fig. 80-101, col. pl. 5-7.

Ho.ttum, R. E. 1941. The Bignonia family in Malayan gardens. M.A.H.A. Mag. 11: 3-11.

SANTISUK, T. 1974. Bignoniaceae. Thai For. Bull. Bot. 8: 1-46.

Sykes, W. R. 1966. Studies of cultivated plants in New Zealand. 1. Bignoniaceae. New Zeal.

D.S.I.R. Inf. ser. 54: 63 pp., 25 fig.

ARTIFICIAL KEY TO CULTIVATED BIGNONIACEAE IN MALESIA

1. Climbing plants.
2. Leaves at least 2-jugate. Tendrils absent.
3. Stamens exserted. Corolla narrow-tubular, scarlet or sulphur-yellow .... . Tecoma capensis
3. Stamens included. Corolla not narrow-tubular, lavendar or white streaked with carmine.
4. Calyx truncate, at most 6 mm, including minute teeth, not inflated.
5. Corolla c. 4-5 cm long, white with crimson throat, the tube puberulous outside. Calyx c. 6 mm.
Pandorea jasminoides

1977] BIGNONIACEAE (van Steenis) 181

5. Corolla (1-)2-3 cm long, pale yellow, lobes purple dotted or streaked, the tube glabrous outside.
Peboxt 2 —s nt PP ted es. Sse d. 5 CIEE ee Baye . . . Pandorea pandorana
4. Calyx campanulate, inflated, white, distinctly 5-lobed, 11/, cm long . . . Podranea ricasoliana
2. Leaves 1-jugate. Tendrils in a number of leaves present.
6. Corolla lobes valvate; tube narrow, without a distinct basal tube. Stamens exserted. Flowers
Beontrorange. ls teks oes. oS. 0 eee . . . . . Pyrostegia venusta
6. Corolla lobes imbricate.
7. Calyx spathaceous, thin, 3 cm long. Corolla very large, rose-purple, (S—)7—9 cm. Disk absent.
Phryganocydia corymbosa
7. Calyx regular, not spathaceous. Disk present.
8. Inflorescence, calyx and outside of corolla tube densely hairy.
9. Calyx 5-6 mm long, 6-8 mm wide at the mouth. Inflorescence pauciflorous, almost a raceme,

with 2-4 pairs of opposite flowers. Tendrils branched . . . . . Pithecoctenium cynanchoides
9. Calyx 3-4 mm long, 2'/,-4 mm wide at the mouth. Thyrse rich-flowered, the flowers in triad
Sees, Fears Unbranched 6). . «(2 . {Rae eal Arrabidaea mollissima

8. Inflorescence, calyx and outside of corolla tube glabrous.

10. Tendrils undivided. Leaves obovate with long-cuneate base, the two basal nerves straight,
running up over halfway the blade, in the narrow angle with the midrib a large dark coloured
piandiheld-Pseudostipules'present’ ..: . (Gente Ties oe ea Saritaea magnifica

10. Tendrils 3-parted. Leaves not obovate, and no such gland field in the narrow angle at the base.

11. Corolla yellow, outside of lobes glabrous or lepidote. Plant without garlic odor.
12. Tendril with 3 claws. Pseudostipules scale-like. Calyx thin, broadly campanulate
Macfadyena unguis-cati
12. Tendril 3-fid. Pseudostipules foliaceous, 5~7(—15) mm in diameter. Calyx coriaceous, cupular.
Anemopaegma chamberlaynii
11. Corolla pale mauve to pale purple, outside of lobes puberulous. Plant smelling of garlic.
Pachyptera hymenaea
1. Erect shrubs or trees.
13. Leaves in scattered fascicles. Fruit indehiscent, hard-shelled with fleshy pulp.
MOPPCHUESISUUDIC 2 5 wk ce 8 se ee in oe ee en Crescentia cujete
Saereaves S-r1Oliate, petiole'winged) <.ysc:") .. AM Lae eee ee Crescentia alata
13. Leaves decussate or in whorls. Fruit dehiscent or fleshy and without a hard shell.
15. Leaves simple.
16. Leaves elliptic on a long, slender petiole. Thyrses terminal. Capsule linear with long-hairy, linear

Penerarned tree 2. 60). Le CGR). ees Bs ks ee eh va Catalpa longissima
16. Leaves small, obovate, tapering into the base; no proper petiole. Flowers ramiflorous. Fruit an
oblong berry. Twig nodes witha pairofthormns ........... Parmentiera aculeata

15. Leaves compound.
17. Leaflets 3, sessile, articulated on top of a winged petiole. Fruit fleshy.
18. Branchlets usually with 2 short, ascending thorns at most nodes. Fruit costate and curved, less
er eee long ed3 used ee are anes... Ot) AO cei Se oe Parmentiera aculeata
18. Branchlets unarmed. Fruit neither curved nor costate, usually more than 30 cm long, candle-like.
Parmentiera cereifera
17. Leaflets 5 or more.
19. Leaves digitately compound. Leaflets 5.
20. Leaflets stellate-hairy beneath. Flowers yellow. Calyx rusty stellate-tomentose
Tabebuia chrysantha
20. Leaflets lepidote, otherwise glabrous. Flowers pink. Calyx lepidote.
21. Leaflets acuminate or sharply acute. Inflorescence usually many-flowered. Mature capsule
midore,than:22 cm long: \.:;. aR. oe eG. Beal Sa eee Tabebuia rosea
21. Leaflets obtuse. Inflorescence few-flowered. Mature capsule usually less than 15 cm long
Tabebuia pallida
19. Leaves pinnate.
22. Leaves 2-pinnate.
23. Corolla infundibuliform, lilac. Capsule broad-ellipsoid, with woody valves. Staminode longer
than the stamens, glandular-pubescent, especially at apex.
24. Corolla glabrous, distinctly sigmoid. Leaflets acute, with oblique base.
Jacaranda obtusifolia ssp. rhombifolia
24. Corolla densely short-hairy, not sigmoid. Leaflets cuspidate, base hardly oblique
Jacaranda mimosifolia
23. Corolla salver-shaped, white, the tube 6-8 cm long, 2 mm wide. Capsule linear. Staminode
BOSSE IE PrP «DPE HE aay OS Millingtonia hortensis
22. Leaves 1-pinnate.
25. Calyx coarse, 2-7 cm long. Corolla wide-campanulate, coarse and large.
26. Calyx irregularly lobed, 2-3 cm. Flowers inside dark red, nocturnal, in long, pendent racemes.

Berry massive, sausage-shaped hs AOE Grist Peri? emis ark « Kigelia africana
26. Calyx spathaceous, 4-7 cm. Flowers orange-red, in erect terminal thyrses, diurnal. Capsule
dehiscent; with: winged seeds pauii at) «FOR WETR ey hs . . . Spathodea campanulata

25. Calyx short 5-lobed, 5-6 mm, Flowers not coarse.

182

FLORA MALESIANA

{[ser. I, vol. 8?

27. Corolla infundibuliform, yellow, with included stamens. Leaflets lanceolate, serrate, some-

times deeply incised

tin aie ees) 43:4... ee ey Ee Tecoma stans

27. Corolla narrow-tubular, usually orange red, sometimes sulphur-yellow, with exserted stamens.

Leaflets ovate, dentate

Anemopaegma chamberlaynii (Sims) Bur. & K.Scu.
Fl. Bras. 8, 2 (1896) 128; HerKLots, Fl. Trop.
Climb. (1976) 65, f. 82. — Bignonia chamberlaynii
Sims, Bot. Mag. (1820) t. 2148. — Bignonia
scandens VELL. Fl. Flum. 6 (1825) 232, t. 22. —A.
scandens MELLO ex K.Scu. in E. & P. Nat. Pfi.
Fam. 4, 3b (1894) 215; BACK. & BAKH. f- Fl. Java 2
(1965) 536.

Glabrous. Pseudostipules foliaceous, ovate to +
orbicular, 5-7(-15) mm. Leaflets ovate-oblong, to
lanceolate-oblong, acute, 5-14 by 21/,-5'/, cm.
Tendrils 3-fid. Flowers in 2-8-flowered axillary
racemes. Calyx campanulate, truncate, 7-8 mm.
Corolla 4-5 cm, pale yellow.

Distr. Brazil, introduced in East Java as an
ornamental, at Malang and Kali Baru (Besuki),
250-600 m; also seen from Rangoon, Burma
(DICKASON 6660). All Asian material has smallish
ovate leaflets, 5-6 by 2!/,-31/,cm and smallish
acute pseudostipules 5-7 mm long; in America
both can obtain much larger size.

Arrabidaea mollissima (H.B.K.) Bur. & K.Scu.
Fl. Bras. 8, 2 (1896) 46; SEIBERT, Carnegie Ins.,
Wash. 522 (1940) 406; DUGAND, Caldasia 3 (1945)
255:

Leaves patent, lax hairy, the longer hairs mostly
gland-tipped; leaflets ovate, acuminate, 4-12'/, by
21/,-7!/,cm. Flowers pink to rose, 3'/,-5 cm,
mouth whitish with yellow spot.

Distr. Mexico and Central America to Colum-
bia and Venezuela; cultivated in the vicinity of
Manila.

Catalpa longissima (JAcQ.) Dum. Cours. Bot. Cult.
2 (1802) 190; Sims, Bot. Mag. (1808) t. 1094;
SANDwITH, Rec. Trav. Bot. Néerl. 34 (1937) 228;
Apams, Fl. Pl. Jamaica (1972) 669; LiTTLe Jr.,
Trees Puerto Rico & Virgin Is. 2 (1974) 890, f. 700.
— Bignonia longissima Jacq. En. Pl. Carib. (1760)
25. — Macrocatalpa longissima BRITTON, J. N.Y.
Bot. Gard. 19 (1918) 8.

Tree up to 30 m, occasionally deciduous. Petioles
slender ; blades ovate-lanceolate, acute, (3—)5-11 by
(12/,-)2-4 cm. Flowers in small panicles, white,
pinkish on the lobes, yellow in mouth with purple
markings, 21/,-3 cm long. Calyx 2-cleft. Capsule
35-75 cm by 4mm.

Distr. Jamaica, Hispaniola, Martinique, intro-
duced for ornamental or forestry purposes in other
tropics, e.g. in the Marianas (Guam, Saipan) and
the Philippines (Luzon: Lamao For. Res.).

Vern. French oak, Haitian oak, Mast-wood,
Yoke-wood, E.

Crescentia alata H.B.K. Nov. Gen. Sp. 3 (1819)
158; F.-Vitt. Nov. App. (1880) 151; VIDAL,
Sinopsis Atlas (1883) 35, t. 73, f. C; MeERR. FI.
Manila (1912) 430; Sp. Blanc. (1918) 350; En.
Philip. 3 (1923) 447; STANDLEy, Trees Shrubs Mex.
(1926) 1324; Back. & Baku. f. Fl. Java 2 (1965)
542; GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 829.
— C. trifolia BLANCO, FI. Filip. (1837) 489; DC.

Tecoma capensis

Prod. 9 (1845) 247; BLANco, FI. Filip. ed. 3. 2
(1878) 271, t. 327. — Otophora paradoxa BL.
Rumphia 3 (1847) 146; Mig. Fl. Ind. Bat. 1, 2
(1859) 560 (L). — Parmentiera alata Miers,
Trans. Linn. Soc. Bot. 26 (1870) 166.

A crooked tree, 5-14 m, to 25cm @. Leaves in
scattered fascicles on the twigs, with a winged
petiole, 21/,-11 cm; leaflets brittle, lanceolate-
obspathulate, sessile, 1-—4'/,cm. Flowers 1-2
rami- and cauliflorous; calyx 2-lobed to the base;
corolla brownish with brown-purple venation,
rank-scented, with a transversal fold, 6 cm.
Fruit with a hard shell, + globular, 5-10 cm @.

Distr. Mexico to Costa Rica, from Mexico
early introduced by the Spaniards via Guam in the
Philippines, also in Java (rare) and Rabaul (New
Britain).

Vern. Philippines: cruz-cruzan, Tag., hoja cruz,
Spanish.

The pulp is in Mexico sometimes used as a
medicine; the shells are less in demand than those
of Crescentia cujete, but used for the same pur-
poses. No fruits seen from Malesia.

Crescentia cujete LINNE, Sp. Pl. (1753) 626;
STANDLEY, Trees Shrubs Mex. (1926) 1324; STEEN.
Thesis (1927) 1010; Bull. Jard. Bot. Btzg III, 10
(1928) 274; SerBeRT, Carnegie Inst. Wash. 522
(1940) 383; Back. & Baku. f. FI. Java 2 (1965)
542; GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 831.
— C. ovata Burm. f. FI. Ind. (1768) 132, nom.
semi-nudum (G).

Crooked tree to 10 m, 30 cm @. Leaves in scat-
tered bundles on the rough twigs, obspathulate,
sometimes short-acuminate, without petiole, up to
26 by 7!/, cm. Flowers solitary or in pairs on the
twigs, of a musty odor; calyx 2-lobed to the base;
corolla 4-7cm, dirty white or pale greenish,
purplish veined, finally turning dull purplish, the
wide tube with a transversal fold. Fruit broad-
ellipsoid to globular, 13—20 by up to 30 cm, indehis-
cent, with a hard shell.

Distr. Central America, very widely and early
distributed in the American and other tropics,
throughout Malesia, in the lowland, grown in
lawns, parks and used for hedges.

Vern. Tabu kaju, S. Sum., bila, Djakarta,
sékopal, sikadel, J, bila radja, Kangean, bila
bilanda, Makas., calabassa, Moluccas, bér(e)nuk,
S, bua no, Ternate, buwano, Halmaheira, Calabash
tree, E.

The pulp is sometimes used for medicinal
purpose and the hard shells are commonly in use
for drinking cups, vessels, and carving.

The flowers are bat-pollinated. Cf. Porscu, Oest.
Bot. Z. 80 (1931) 31-44, t. 9-10.

I saw the type of C. ovata Burm. f. which was
described from Java and which was in Index
Kewensis reduced to C. cucurbitina L. (now
Amphitecna latifolia (MILL.) GENTRY); the type
consists of 3 leaves and a single damaged flower;
the sheet carries no name, but the note ‘Kalbas,
4 stam., 1 pistile’.

1977]

Jacaranda mimosifolia D. Don, Bot. Reg. 8 (1822)
t. 631. — J. ovalifolia R.Br. Bot. Mag. 49 (1822)
t. 2337. — J. acutifolia (non H. & B.) auct.; STEEN.
Bull. Jard. Bot. Btzg III, 10 (1928) 270; Back. &
Baku. f. Fl. Java 2 (1965) 239.

Distr. Tropical America, Bolivia to NW.
Argentina. Seen from W. Java, Hawaii, S. Africa,
Madagascar, Congo. Perhaps not distinct from the
Peruvian J. acutifolia H.B.K.

Suitable as a roadside tree and in parks. Flowers
often when leaves are shed.

The names of Don and BRown were published
on the same day on material from the same source;
a unique nomenclatural case!

Jacaranda obtusifolia H.B.K. ssp. rhombifolia
(G. F. W. Meer) GENTRY, Mem. N.Y. Bot.
Gard., in the press. — J. rhombifolia G. F. W.
Mewer, Fl. Esseq. (1818) 213. — J. filicifolia
D. Don, Edinb. Phil. J. 9 (1823) 266; STEEN.
Bull. Jard. Bot. Btzg III, 10 (1928) 269; HoL_trum,
M.A.H.A. Mag. 3 (1933) 188, fig.; BACK. & BAKH.
f. Fl. Java 2 (1965) 239.

Distr. Northern South America, Venezuela to
Guiana. Seen from Malaya, Java (W. Java, also
Semarang and Malang), and Borneo (Sandakan,
Kuching).

Good roadside tree, and for large gardens and
parks.

Kigelia africana (LAMK) BTH. in Hook. Niger FI.
(1849) 463; SpraGugE, FI. Trop. Afr. 4, 2 (1906)
536; Merr. Fl. Manila (1912) 430; En. Philip. 3
(1923) 444; H. Herne in Hutch. & Dalz. Fl. W.
Trop. Afr. ed. 2, 2 (1963) 385; MERxM. & SCHREIB.
Prod. Fl. SW. Afr. fam. 128 (1967) 3. — Bignonia
africana LAMK, Encycl. 1 (1785) 424. — Crescentia
pinnata Jacq. Coll. 3 (1789) 203, t. 18. — Tecoma
africana G. Don, Gen. Syst. 4 (1838) 224. — K.
pinnata DC. Prod. 9 (1845) 247; KoeNs, Trop.
Natuur 1 (1912) 167, 6 f. — K. aethiopica (FENZL)
Decne in Deless. Ic. Sel. Pl. 5 (1845) 39, t. 93;
Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 275;
Back. & BAKH. f. Fl. Java 2 (1965) 542. — See for
further synonyms HEINE, vide supra.

Widely branched tree, to 20 m. Leaves decussate
or in whorls of 3-4, up to 50 cm; leaflets oblong,
entire or serrate distally, glabrous or hairy on
nerves, to 20 by 6 cm. Flowers nocturnal, coarse, in
terminal, pendent, narrow panicles up to 2 m long;
calyx greenish, 2-3 cm, 2-lipped; corolla outside
yellowish, veined, inside dark wine-red, 5-10 cm;
basal tube as long as the calyx or longer. Berry
sausage-like, 25-50 by 7'/,-15 cm, often on still
flowering panicles.

Distr. Africa, widely cultivated in other tropics
as an ornamental tree in parks and along roads, in
Malesia not rare.

Herne /.c. concluded that the genus consists of
one species only as the species distinguished by
Srapr are racial and grading.

The calyx contains much watery slime in bud
(cf. Koorp. Ann. Jard. Bot. Btzg 14, 1897, 407-
411). The flowers are in Malesia invariably visited
by bats. Cf Faircui_p, Trop. Gard. Bull. July
1968, 5; Harris & Baker, The Nigerian Field 40:
151-158; J. West Afr. Sc. Assoc. 4 (1958) 25-30;
Koens, Trop. Natuur 1 (1912) 167, f. 1-6;
McCann, J. Bomb. Nat. Hist. Soc. (1931) 467-471,

BIGNONIACEAE (van Steenis)

183

3 f. They are, however, also frequented by hawk-
moths and Harris & BAKER //.cc. concluded that
bats are not essential pollinating visitors.

Macfadyena unguis-cati (L.) A. GENTRY, Brittonia
25 (1973) 236; Ann. Mo. Bot. Gard. 60 (1973) 874.
— Bignonia unguis-cati LINNE, Sp. Pl. (1753) 623.
— Doxantha unguis-cati MIERS em. REHDER, Mitt.
Deut. Dendr. Ges. (1913) 262; HerKLOoTs, FI.
Trop. Climb. (1976) 66, f. 87. — Bignonia tweediana
LINDL. Bot. Reg. 26 (1840) t. 45, non GrisEB.;
BuysMaN, Flora 107 (1915) 361, cult. in Java. — ?
M. dentata Bur. & K.Scu. FI. Bras. 8, 2 (1897) 291;
STEEN. Bull. Jard. Bot. Btzg III, 10 (1928) 186;
Back. & BAKH. f. Fl. Java 2 (1965) 538.

Leaves very variable, those of juvenile plant
appressed to substratum, very small; later to 5 by
3'/, cm, ovate, dentate, but in other forms elliptic-
oblong and hardly dentate. Calyx irregularly
lobed to spathaceous or subspathaceous. Corolla
4'/,-8 cm.

Distr. Mexico to Brazil and N. Argentina, some-
times cultivated in Malesia.

Specific characters in this genus seem to be vague;
those used to distinguish M. uncata (ANDR.)
SPRAGUE & SANDW. from M. unguis-cati by GENTRY
(Ann. /.c. 871) he declared himself (Brittonia
l.c. 236) as inconstant. The one specimen I have
seen from Java agrees with the plate of M. dentata
but for the non-spathaceous calyx. Shape of calyx,
margin of leaves and length of corolla seem to be
very variable in this species which has already a very
large synonymy (GENTRY, Ann. /.c. 871). Does not
fruit in Java; propagated by suckers and cuttings.

Millingtonia hortensis L. f. Suppl. (1781) 291. See
for a full treatment p. 133.

Distr. SE. Asia, probably also native in the
Lesser Sunda Islands.

Suitable as a tree for roadsides and parks.

Pachyptera hymenaea (DC.) GENTRY, Brittonia 25
(1973) 236; Ann. Mo. Bot. Gard. 60 (1973) 888,
with full synonymy.

Glabrous liana. Vegetative parts smelling of
garlic, Pseudostipules bract-like, 1'/, mm. Leaflets
triplinerved, ovate-oblong, short-acuminate, 31/,—
7 by 1'/,-3 cm; venation prominent on both sides.
Flowers in short axillary racemes. Calyx tube
4-5 mm, with minute prominent pustular glands.
Basal tube of corolla twice as long as calyx, in all
3-4 cm. Anthers glabrous. Fruit flattened, 16-22 by
1'/,-1'/,cm; valves with a central rib. Seeds
1-1'/, by 3-3'/, cm, incl. the membranous wing.

Distr. Acommon liana ranging from Mexico to
Brazil of tropical dry forest, sporadically also in
moist forest, introduced in the Philippines (Manila,
Mindanao) and E. Java (Surabaya).

Another species may also be cultivated which is
in flower not easy to distinguish from P. hymenaea,
viz P. alliacea (LAMK) GENTRY; this latter species
a a short oblong fruit with thick, corky wingless

Ss.

The genus Pseudocalymna Samp. & KUHLM. has
been reduced to Pachyptera A.DC. by GENTRY
(Lc.).

Pandorea jasminoides (LINDL.) K.Scu. in E. & P.
Nat. Pfl. Fam. 4, 3b (1894) 230; Steen. Thesis

184

FLORA MALESIANA

[ser. I, vol. 8?

(1927) 847; Bull. Jard. Bot. Btzg III, 10 (1928) 195;
Proc. R. Soc. Queens]. 41 (1929) 48; BAcK. &
BAKH. f. Fl. Java 2 (1965) 538; Herktorts, FI.
Trop. Climb. (1976) 69, f. 90. — Tecoma jasmi-
noides LINDL. Bot. Reg. (1939) t. 2002.

Fairly tall climber; leaflets 4-7(-9), lanceolate,
blunt, 2!/,-5 by 1-2cm. Corolla white streaked
with carmine in the mouth, 4-5 cm long, short
hairy. ;

Distr. NE. Australia, not rarely cultivated in
the tropics and also subtropics (Mediterranean,
N. New Zealand), in Malesia rarely cultivated
(e.g. West Java), in the temperate zone in green-
houses.

Propagated by cuttings; seeds very rare.

Pandorea pandorana (ANDR.) STEEN. Bull. Jard.
Bot. Btzg III, 10 (1928) 198; Back. & Baku. f.
Fl. Java 2 (1965) 538. For full references see p. 176.

Distr. East Australia, New Caledonia, Lord
Howe I., Solomons (Bougainville), New Guinea,
Moluccas, and Lesser Sunda Islands.

In Malesia an unfrequent ornamental, but culti-
vated here and there through the tropics and sub-
tropics (also in the Mediterranean and N. New
Zealand).

Propagated by tjankoks (marcotting) and cut-
tings.

Parmentiera aculeata (H.B.K.) SEEM. Bot. Voy.
Herald (1854) 183; SEIBERT, Carnegie Inst. Wash.
522 (1940) 385; Gentry, Ann. Mo. Bot. Gard. 60
(1973) 899. — Crescentia edulis Desv. J. Bot. 4
(1814) 112. — Crescentia aculeata H.B.K. Nov.
Gen. Sp. 3 (1819) 158. — P. edulis DC. Prod. 9
(1845) 244 (heterotypic with Crescentia edulis
DEsv.).

Tree 7-8 m. Leaves articulated with very hard,
thorny extension of the nodal bark, the latter
remaining a permanent, ascending, sharp thorn;
leaflets elliptic, narrowed at both ends, 2-5 by
1-3cm; axillary fascicled leaves mostly simple.
Flowers 1-several together, terminal, axillary or on
branches or stem. Calyx spathaceous, 3-5 cm;
corolla white, 6-7 cm. Berry pendent, cylindric,
curved, costate with thick ribs, 8-17 cm by over
3cm 2.

Distr. S. Mexico to northern Central America,
cultivated elsewhere in the tropics; in Malesia rare:
W. Java, Luzon, also seen from Cairns (N.
Queensland).

Young sterile offshoots have no thorns and
possess coarsely dentate leaflets.

Parmentiera cereifera SEEM. in Hook. J. Bot. &
Kew Gard. Misc. 3 (1851) 302; Bot. Voy. Herald
(1854) 182, t. 32; STEEN. Thesis (1927) 1008; Bull.
Jard. Bot. Btzg III, 10 (1928) 272; Back. & BAKH.
f. Fl. Java 2 (1965) 542.

Tree to 7m, 20cm @. Leaflets oblong, acumi-
nate, 4-8 by 1'/,-3'!/,cm. Flowers cauliflorous,
nocturnal, white, slightly fragrant. Calyx spatha-
ceous, 3cm; corolla 5-6cm. Berry pale yellow,
pendent, candle-like, smooth, 30-100 by 1%/,-
23/2 cm.

Distr. Panama, cultivated in many tropical
countries, and in many parts of Malesia in parks
and gardens. With its waxy-fleshy, candle-like
fruits a showy plant.

Vern. Candle tree, E, kaarsenboom, D.

The flowers are bat-pollinated, as in Crescentia.
In Panama fruits are utilized as cattle food (SEE-
MANN).

Phryganocydia corymbosa (VENT.) Bur. ex K.SCH.
in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 224, f. 89H;
GENTRY, Ann. Mo. Bot. Gard. 60 (1973) 905;
HerKLots, Fl. Trop. Climb. (1976) 69, col. pl. 6. —
Spathodea corymbosa VENT. Choix (1807) t. 40.

Sometimes pseudostipules. Leaflets (4-)12-15 by
(2-)7-9 cm, triplinerved. Calyx spathaceous. Co-
rolla lavender to blue purple, with a white mouth.

Distr. Panama to Brazil, cultivated at Seria
(Brunei).

A Brunei specimen (VAN NiEL 3826) identified
by A. A. ATCHLEY (1973).

Pithecoctenium cynanchoides DC. Prod. 9 (1845)
193; Fasris, Rev. Mus. La Plata 9, Bot. n. 49
(1965) 353, f. 19.

Leaves reniform-triangular acuminate, ciliate at
the margin, 21/4 by 2-4cm. Corolla 3-6 cm,
white with yellow markings in the mouth.

Distr. Southern Brazil to Argentina, in E. Java
once cultivated (Mt Tengger: BUYSMAN).

Podranea ricasoliana (TANF.) SPRAGUE, FI. Cap. 4,
2 (1904) 450; Sykes, New Zeal. D.S.I.R. Inf. ser. 54
(1966) 39, f. 16; Gentry, Ann. Mo. Bot. Gard. 60
(1973) 916, f. 29; HerkLots, Fl. Trop. Climb.
(1976) 70, f. 92. — Tecoma ricasoliana TANF. Bull.
Soc. Tos. Ort. (1887) 17, t. 1-2. — Pandorea ricaso-
liana BAILL. Hist. Pl. 10 (1891) 40.

Leaflets 7-9, crenate, 2!/,-4 by 1-2 cm. Corolla
pale lavender or pinkish, with magenta patches and
lines in the mouth and tube, 6-8 cm long, in
terminal thyrses.

Distr. South Africa, cultivated elsewhere in the
tropics (e.g. in the Philippines, New Caledonia) and
subtropics (Mediterranean, New Zealand).

Pyrostegia venusta (KER) Miers, Proc. R. Hort.
Soc. Lond. 3 (1863) 188; STEEN. Bull. Jard. Bot.
Btzg III, 10 (1928) 189; BRUGGEMAN, Ind. Tuinb.
(1939) 64, f. 21; BACK. & BAKH. f. Fl. Java 2 (1965)
536; Sykes, New Zeal. D.S.I.R. Inf. ser. 54 (1966)
22, f. 6; HerKLots, Fl. Trop. Climb. (1976) 71,
f. 94. — Bignonia venusta Ker, Bot. Reg. (1818)
t. 249; Bot. Mag. (1819) t. 2050. — Tecoma
venusta LEM. Hort. Univ. (1834) 1, icon. — P. ignea
(VELL.) PRESL, Bot. Bemerk. (1845) 93.

Leaflets ovate-lanceolate, 46 by 3-4cm.
Thyrses dense, rachis 10cm. Calyx 6-7 mm.
Corolla tube c. 7 cm.

Distr. Brazil, commonly cultivated throughout
the tropics and also in subtropics (North I. of New
Zealand), in the temperate zone in glass-houses
since the early 19th century. Widely cultivated in
Malesia.

A richly flowering ornamental, good for walls
and trellis, excellent between 500-1700 m altitude,
not flowering in Java below 250 m and never setting
fruit. The oranje stephanoot, D, is easily propagated
by tjankoks (marcotting) and cuttings.

Saritaea magnifica (STEEN.) DUGAND, Caldasia 3
(1945) 263, fig.; GENTRY, Ann. Mo. Bot. Gard. 60
(1973) 920, f. 31; SANTISUK, Kew Bull. 28 (1973)

1977]

184; Thai For. Bull. Bot. 8 (1974) 46; HERKLoTs,
Fl. Trop. Climb. (1976) 71, f. 95. — Arrabidaea
magnifica STEEN. Rec. Trav. Bot. Néerl. 24 (1927)
830, excl. syn. Bignonia magnifica BULL; Bull. Jard.
Bot. Btzg III, 10 (1928) 191; GUTTENBERG, Ann.
Jard. Bot. Btzg 44 (1934) 195; CHATTERJEE, Bull.
Bot. Soc. Beng. 2 (1948) 78; BACK. & BAKH. f. FI.
Java 2 (1965) 536. — Arrabidaea sp.: DAUBANTON,
Teysmannia 29 (1918) 51-53, phot.

Pseudostipules present. Tendrils simple. Leaflets
obovate, bluntly short-acuminate, 5-11 by 3-6 cm.
Calyx cupular-campanulate, truncate, 6-12 by
3-7 mm. Corolla purple to magenta, 8-9 cm, the
mouth white with magenta markings.

Distr. Colombia and Ecuador, widely culti-
vated through the tropics and common in SE.
Asia and Malesia, never setting fruit. Obviously
first introduced in Singapore; the Bogor Botanic
Gardens received it from Banka in 1911.

A robust climber, good for walls and trellis,
profusely flowering all the year round, especially
in the wet season (Oct.—May), easily propagated
by cuttings, found up to c. 1000 m.

Spathodea campanulata P. Beauv. FI. Oware
Benin 1 (1805) 47, t. 27; Hook. Bot. Mag. 85
(1859) t. 5091; SprRAGuE, Fl. Trop. Afr. 4, 2 (1906)
529: Merr. Fl. Manila (1912) 429; En. Philip. 3
(1923) 447; STEEN. Rec. Trav. Bot. Neéerl. 24
(1927) 945; Bull. Jard. Bot. Btzg III, 10 (1928) 232;
Ho.ttum, M.A.H.A. Mag. 3 (1933) 186, fig.;
IRVINE, Woody Pl. Ghana (1961) 739, t. 7; HEINE,
Fl. Trop. W. Afr. ed. 2, 2 (1963) 386; Back. &
Baku. f. Fl. Java 2 (1965) 540.

Large tree, 7-25 m, 10-50 cm @. Pseudostipules
leafy. Leaves decussate; leaflets in (4~-)5—6(-9)
pairs, oblong, entire, glabrous or puberulous
beneath, 5-14 by 2'/,-5'/,cm. Flowers erect, in
terminal thyrses; calyx navicular, ribbed, beaked,
thinly velutinous, 4-7 cm; corolla scarlet to orange,
wide, 8-14 cm. Capsules erect, lanceolate-oblong,
15-20 by 2!/,-3 cm; valves keeled; seeds 2!/, by
2 cm, hyaline winged.

Distr. Tropical Africa; widely cultivated in the
tropics in parks and as an avenue tree; throughout
Malesia, up to c. 1000 m.

Vern. African tulip, Tulip tree, E, spuitjesboom,
D, panchot, Malaya, djati bélanda, Kangean;
Sabah: anchit anchit, maundi, Banggi.

The flowers are frequented by birds on which
Beumée (Trop. Natuur 14, 1925, 28-30, f. 1) did
observations at Bogor. Flowers remain open for
at least two days and each flower may be visited by
more than one bird, obviously in search of honey.
Possibly the birds play a role in pollination;
corollas are frequently damaged.

AYENSU observed bats visiting the flowers at
night (Ann. Mo. Bot. Gard. 61, 1974, 713).

The calyx, which is closed in bud, contains water
and children play the ‘waterspout’ with it, hence
the Dutch name. Children also use the boat-shaped
valves for making small sailing boats.

Fruiting and flowering takes place throughout
the year. At Bogor I found young seedlings in
hedges and gardens, with a long taproot and dentate
leaves, but there are no records of spontaneous
naturalization. Timber is weak and worthless.

Ripiey (Fl. Mal. Pen. 2, 1923, 547) mentioned
it erroneously from Singapore under the name

BIGNONIACEAE (van Steenis)

185

S. nilotica SEEM.; although this is a closely related
African tree, it is a distinct taxon characterized by a
longer-tomentose, hardly ribbed calyx, leaflets
tomentose underneath, a longer lobed disk and a
long-hairy ovary.

Tabebuia chrysantha (JACQ). NICHOLS. Dict. Gard.
4 (1887) 1; Gentry, Ann. Mo. Bot. Gard. 60
(1973) 941. — Bignonia chrysantha Jace. Hort.
Schoenbr. 2 (1797) 45, t. 211.

Deciduous tree to 25m, 50cm @. Leaflets
elliptic-oblong, abruptly acuminate, 5-25 by 4-11
cm. Calyx shallowly lobed, 5-13 mm. Corolla
glabrous, tubular-infundibuliform, 5-8 cm.

Distr. Mexico to northern Venezuela, in SE.
Asia cultivated, rare in Malesia (Philippines: U.S.
Cemetery, Fort McKinley, Rizal Prov.). No fruit
seen.

A yellow-flowered species has been reported by
Back. & BAKH. f. (Fl. Java 2, 1965, 539) to be
cultivated in Java under the name T. capitata
(Bur. & K.ScH.) SANDW., but in absence of
material I cannot check the identity.

Tabebuia pallida (LINDL.) Miers, Proc. R. Hort.
Soc. 3 (1863) 199; GENTRY, Ann. Mo. Bot. Gard.
60 (1973) 950. — Bignonia pallida LINDL. Bot. Reg.
(1826) 12, t. 695.

Shrub or small tree. Leaves 1—5-foliolate, leaflets
elliptic to elliptic oblong or obovate, obtuse.
Inflorescence few-flowered, often reduced to 1 or 2
flowers. Corolla pinkish lavender to almost white,
the throat opening yellow.

Distr. A common variable West Indian species,
closely related to T. rosea, cultivated in various
parts of the tropics, in Indonesia distributed from
Botanic Gardens, Bogor.

Tabebuia rosea (BERTOL.) DC. Prod. 9 (1845) 215;
SANDwW. Kew Bull. (1953) 454; Gentry, Ann. Mo.
Bot. Gard. 60 (1973) 951, with synonymy. —
Tecoma rosea BERTOL. Fl. Guat. (1840) 25.

Deciduous tree, up to 30 m, | m @. Leaflets and
petiolules often unequal, lepidote, 5-30 by 2-12 cm.
Calyx cupular, bilabiate, densely lepidote, */,—2 cm.
Corolla outside glabrous, 6-10 cm.

Distr. S. Mexico to Venezuela, cultivated in the
tropics in parks and along roadsides; a magnificent
ornamental when abundantly flowering with
blossoms in bunches on the bare twigs; not rare in
SE. Asia, in Malesia only known to me from the
vicinity of Manila, Luzon. No fruit seen.

Tecoma capensis (THUNB.) LINDL. Bot. Reg. 13
(1828) t. 1117. — Bignonia capensis THUNB. Prod.
(1800) 105. — Tecomaria capensis (THUNB.) SPACH,
Hist. Nat. Vég. 9 (1840) 137; SpraAGue, Fl. Cap
4, 2 (1904) 448; STEEN. Thesis (1927) 832; Bull
Jard. Bot. Btzg III, 10 (1928) 193; BRUGGEMAN,
Ind. Tuinb. (1948) 39, 268 f. 273; BAcK. & BAKU.
f. Fl. Java 2 (1965) 538; Brummit, Bull. Jard.
Bot. Nat. Belg. 44 (1974) 421, f. 1 (map); Herk-
Los, Fl. Trop. Climb. (1976) 74, f. 101.

An erect and scrambling shrub, up to c. 3m,
never in my experience a true climber. Leaflets 2-4
pairs, crenate, with domatia, 1-3 by 1-2 cm.
Corolla tubular, red, rarely pale yellow, 4-5 cm.

Distr. South Africa, cultivated and naturalized
in many subtropical and tropical countries, e.g.

186

FLORA MALESIANA

[ser. I, vol. 8?

the Mediterranean; also in South America.
Commonly cultivated in Java and elsewhere in
Malesia, up to c. 1000 m. Flowering all the year
round. No capsules seen; not run wild. Easily
propagated by suckers or cuttings. A nice ornamen-
tal for gardens and often used for hedges.

BrummItT /.c. assumes that there is only one
Tecoma (Tecomaria) in Africa and reduced 7 other
names, distinguishing the tropical taxa as a separate
race, ssp. nyassae (OLIV.) BRUMMITT.

In South African parks and gardens is also
cultivated an even sulphur-yellow variety which was
originally found in the wild, cf. A. JAcoT-GUIL-
LARMOD, Veld en Flora 4 (1974) 36; it hybridizes
eles the red-flowered variety and sets abundant
ruit.

Tecoma capensis, the Cape Honey suckle, is bird-
pollinated; cf. M. S. Evans, Nature 18 (1878) 543;
Scott-E.tiort, Ann. Bot. 4 (1890) 270.

Tecoma stans (L.) H.B.K. Nov. Gen. Sp. 3 (1819)
144; Corner, Wayside Trees (1940) 170, f. 44,
pl. 159; BAckK. & BAKH. f. Fl. Java 2 (1965) 539. —
Stenolobium stans (L.) SEEM. Ann. Mag. Nat. Hist.
10 (1862) 30. — See for elaborate treatment p. 135.

Yellow Bells is a small, erect, ornamental shrub
with showy yellow flowers.

Distr. Southern U.S.A. to southern Argentina,
commonly cultivated through the tropics, in cer-
tain places naturalized. Easily propagated by seed
or by suckers or cuttings. It flowers and fruits
profusely in Malesia the year round, from the
lowland up to c. 1000 m. Suitable for gardens and
parks.

There is a form with hairy leaves (var. velutina
DC. = T. mollis H.B.K.) and one with much
incised leaves (var. incisa G. Don), the latter being
naturalized in Malesia.

CRYPTERONIACEAE (R. J. van Beusekom-Osinga, Leyden)

Evergreen trees. Twigs terete to quadrangular, the younger ones mostly with four
narrow ribs or wings, with thickened nodes, petiole-bases mostly connected by a
faint line. Leaves with minute or rudimentary stipules, opposite, simple, entire,
penninerved, shortly petioled, with arched or almost straight nerves mostly anasto-
mosing in a marginal vein. Inflorescence terminal or axillary, sometimes below the
leaves, paniculate, copiously branched to extremely depauperate, branched up to
the third order, with decussate side axes which are sometimes arranged (sub)-
verticillately or subumbellately by contraction, either ending in profuse to very
poor racemules, or in cymoid florescences. Bracts mostly small to minute, some-
times with rudimentary stipules. Flowers (very) small, shortly pedicelled, bisexual,
sometimes by reduction unisexual and then trees dioecious, actinomorphous, peri-
to epigynous, (4-)5(-6)-isomerous, sometimes with twice the number of stamens;
receptacle widely campanulate. Sepals valvate, triangular to deltoid, mostly per-
sistent. Petals more or less rudimentary, sometimes absent, alternisepalous, inserted
on the margin of the receptacle, inflexed and enveloping the stamens, valvate,
rarely imbricate, sometimes cohering, soon caducous. Stamens if isomerous epi-
petalous, (alternisepalous), inserted on the margin of the receptacle, inflexed in bud,
persistent or caducous; filaments sometimes very short; connective wide, with or
without a tendency to conduplication, or completely conduplicate, sometimes with
a dorsal tubercle or a large outgrowth; anthers adnate, marginal or submarginal,
linear to semiorbicular, lengthwise dehiscent, introrse to latrorse. Ovary superior or
inferior, 2-4(—5)-carpellate, 1-6-locular, septs not, or rarely partly, connate; style
terminal, rather long to short, + terete, mostly persistent; stigma capitate or
punctate. Ovules situated in horizontal or vertical position, 1, 2, 3, or many per
locule, anatropous; placentation parietal, septal, or basal. Fruit superior or '/,- or
3/,-inferior, a chartaceous or woody capsule, subglobose to ellipsoid, small to big,
loculicidally dehiscent with 2-6 valves, on the top often with the persistent style and
stigma. Seeds few or many, flat, usually small, depressed-ellipsoid, situated basally,
apically, centrally, or laterally in its membranous wing in which the raphe is
running freely; endosperm none; embryo straight.

Distribution. Pantropical, 5 genera and 11 spp., 3 genera in Indo-Malesia (of which one
endemic in Borneo), one monotypic genus in S. Africa (Rhynchocalyx) and one in Peru and
Bolivia (Alzatea).

Ecology. Lowland and hill tropical forest, up to 1300 (—1700) m, mostly in rain-forest, but
Crypteronia paniculata not shunning a more or less seasonal climate.

Dactylocladus stenostachys is a characteristic peat swamp forest tree.

Young branches of Crypteronia macrophylla are frequently inhabited by ants.

Young foliage seems often bright coloured: in Axinandra innovations are mauve, while in
Crypteronia paniculata young leaves and twigs are deep blue or violet turning pinkish brown
then green (CorNER), in other species they are purplish.

Morphology. In all genera the nodes of the twigs are thickened and a characteristic trans-
versal ridge or line, sometimes faint (absent in Rhynchocalyx), connects the leaf-bases. On the
internodes four lengthwise raised lines or narrow wings occur, especially distinct in the upper
part of young twigs; they wear off later.

The leaves offer no significant characters, but interesting is the occurrence of ephemerous
rudimentary stipules which can only be observed on innovations, These are common among

(187)

188 FLORA MALESIANA [ser. I, vol. 8?

Myrtalean families, but obviously absent in Melastomataceae, where these structures were not
found in a sampling of fifteen genera.

The petals in Crypteroniaceae are reduced to a varying degree or are even totally absent (in
Crypteronia). Moreover, they are always soon caducous, except in A/zatea where they are almost
invisible and mucilaginous. In all genera they are conduplicate and enveloping the inflexed
stamens as a hood. In Dactylocladus and Rhynchocalyx they are minute and unguiculate; in
Axinandra they are proportionally bigger, and have a broad instead of an unguiculate base. The
petals of Axinandra are complicated and show a highly interesting specialization; they are
coherent to connate, together having the shape of an umbrella or a mushroom. Their wide,
tapering basal parts together form the awning of the umbrella, the narrow, coherent, median
parts form the stem of the umbrella, and often there are wider, frayed, reflexed, apical parts of
the petals together forming the handle of the umbrella (or the ‘root’ of the mushroom). This
whole structure envelops the stamens very closely, and drops when the flower opens and the
inflexed stamens stretch. It is, furthermore, interesting that these petals of Axinandra, depending
on the species, can be valvate-connate, valvate- (or somewhat imbricate-)conduplicate, or imbri-
cate-contorted.

Several petal characters of Crypteroniaceae are found again scattered in other Myrtalean
families. Reduction, absence, as well as caducousness of petals occurs sporadically in almost
all of these families. Unguiculate petals are more or less characteristic for Lythraceae, but are
also found in Sonneratiaceae (Duabanga), and in Rhizophoraceae (e.g. Carallia). Connate petals
with a broad base, sometimes fused to a ‘cap’ occur in Myrtaceae. Coherence of petals is also
present in Rhizophoraceae (Ceriops). Valvate and imbricate petals, both found in one genus,
Axinandra, are usually family characters in Myrtales. Contorted petals are, apart from Axinandra,
only found in Melastomataceae.

The enveloping of the stamens by the petals in all petal-bearing Crypteroniaceous genera is
almost unique in Myrtales, being only found in a few Rhizophoraceae (Rhizophora and Bruguiera).
However, in the latter family the petals do not cover the stamens as a hood, as is the case in
Crypteroniaceae. This is one of the characters upon which the identity of Crypteroniaceae as a
family is based.

The stamens in Crypteroniaceae are arranged in one isomerous epipetalous (alternisepalous)
whorl, except in Axinandra, where one diplostemonous whorl is present. In general number and
position of the stamens in Myrtales can be derived from a situation with two isomerous whorls,
either by reduction or by polymerisation (‘dédoublement’) and multiplication. Arrangement of
stamens in two isomerous whorls, the diplostemonous androecium, is mostly considered to
represent the basic structure of the androecium in Myrtales. MELCHIOR (in Engl. Syllabus 2, 1964,
345) distinguished for the androecium in Myrtales two progressive trends both starting from the
diplostemonous androecium, viz multiplication into many stamens in many whorls and reduction
towards the haplostemonous state and even eventually to 3, 2, or 1 stamen(s). .

In all Crypteroniaceous genera the stamens are inflexed in bud. This is a widespread character
in Myrtales, being rather typical for this order. In some families (Myrtaceae, Rhizophoraceae) it
is not present in all genera, and Lythraceae, Onagraceae, and Haloragidaceae are the only families
in which it is totally absent. The total absence of inflexed stamens in the Lythraceae is another
fact which militates against inclusion of Crypteroniaceous genera in that family.

The gynoecia of the Crypteroniaceous genera are distinguished by cells which are divided by
interrupted septs, though these may touch each other in the centre of the gynoecium. This
(hemi)synplicate condition is very rare and assumed to be primitive within the Myrtales, in which
it is only found in Crypteroniaceae and in a few genera of Myrtaceae.

The capsules of Crypteronia and Axinandra show interesting specialized structures with a func-
tional significance with regard to opening and closing of the capsule. The mechanisms for this are
based upon hygroscopical properties of fibres in vascular bundles.

The morphology of the seed in the Crypteroniaceae is peculiar. The seed-coat forms a flat,
membranous wing, through which the raphe is running from the insertion to the top where it
usually takes a more or less sharp turn, and runs back towards the seed proper, which either
takes a central, apical, lateral, or basal position in the wing. This is another assumedly primitive
character within Myrtales, again only found in Crypteroniaceae and in a few genera of Myrtaceae.

1977] CRYPTERONIACEAE (van Beusekom-Osinga) 189

This character is also rare in other orders. It was first discovered in the Trochodendraceae and is,
therefore, indicated by me as the Trochodendraceous seed type.

Summarizing, we find that almost the whole variety of floral characters in the Crypteroniaceae
is also found scattered in other Myrtalean families. In this respect the Crypteroniaceae are rather
heterogeneous, though not more than for instance the Myrtaceae and the Melastomataceae. On
the other hand, the family is unique in the Myrtales by having petals enveloping the stamens as a
hood. Moreover, the presence of one whorl of epipetalous (alternisepalous) stamens, characteris-
tic for four out of the five Crypteroniaceous genera, is very rare in other Myrtalean families,
being restricted to one or two genera of the Myrtaceae and of the Lythraceae, and to the mono-
typic Oliniaceae. Finally, the conduplication of the connective or the tendency to it in all Crypter-
oniaceae except Dactylocladus, is another important family character, in other Myrtales only
found in a few Melastomataceae. Apart from the above-mentioned characters Crypteroniaceae
are also characterized by the septation of the gynoecium and by the Trochodendraceous seed-
structure, both being only found in other Myrtales in a few Myrtaceous genera. They are, how-
ever, from a practical viewpoint, less useful for easy diagnosis. — C. F. VAN BEUSEKOM.

Taxonomy. Crypteroniaceae belong undoubtedly to Myrtales. Though the family concept in
this order is fairly satisfactory, it can be observed from the above-made remarks that there are
not many exclusive characters, most of them breaking down occasionally in one family, or
occurring also sporadically in another family. Each family in Myrtales seems to be characterized
by a unique character combination in addition to one or two exclusive characters.

For Crypteroniaceae this combination and characters are: swollen nodes with transversal line,
internodes with lengthwise raised lines or wings, petals in bud hood-like enveloping the stamens,
soon caducous (in Crypteronia absent), stamens inflexed in bud and in one epipetalous whorl
(except in Axinandra in two whorls), absence of a perianth tube or of any space between the
insertion of petals and stamens, and furthermore the presence of a (hemi)synplicate gynoecium
and seeds of the Trochodendraceous or related type, both assumedly primitive characters, and
almost exclusive within Myrtales.

Palynological evidence does not fully sustain the recognition of Crypteroniaceae as a distinct
family; it could be accepted, but the evidence may allow other possibilities (MULLER, Blumea 22,
1975, 275).

Anatomical evidence is not much in favour of the family concept as proposed; the genera could
in this respect be divided up among Melastomataceae and Lythraceae, but it remains to be seen
in how far anatomical characters clearly sustain other current family concepts in Myrtales.

Mpyrtales are certainly a very ancient complex and during their evolution advanced characters
have evolved, reduction series occurred in more lines, and primitive characters may have inciden-
tally persisted in various branchings of ancestral tree in taxa which are not necessarily viewed as
closely related. This would also explain the ‘reticulate character distribution’, a condition found
in several families of Myrtales.

None of the Myrtalean families is really homogeneous, but from this can and should not be
concluded that these families are unnatural. For tracing ancestry and evolution naturalness is
more important than homogeneity which, properly, increases always with decreasing taxonomic
rank.

The main subdivision of two subfamilies is supported by wood-anatomical characters (VAN
Vutet, J. Micr. 104, 1975, 65), but other anatomical characters (VAN VLIET & BAAS, Blumea 22,
1975, 175) and palynological data (MULLER, /.c.) do neither support it, nor militate against it. At
tribal level morphological, anatomical and palynological data appear not to agree. The sub-
division adopted here is based on morphology. — C. F. VAN BEUSEKOM.

Anatomy. VAN VuleT, J. Microscopy 104 (1975) 65-82 (wood anatomy and relationships);
VAN Viet & BAAS, Blumea 22 (1975) 175-195 (leaf, nodal and twig anatomy). These two papers
contain a full bibliography to the older literature.

The anatomy of Crypteroniaceae sensu lato is heterogeneous. Distinctive characters are:
cuticle granular (Axinandra, Crypteronia) or smooth (Dactylocladus); stomata paracytic (A xinan-
dra, Crypteronia) or anomocytic (Dactylocladus); hypodermis present (Axinandra, Crypteronia)
or absent (Dactylocladus); petiole with arc-shaped vascular bundle (A xinandra) or with + closed
system (Crypteronia, Dactylocladus); phloem with styloid crystals (Axinandra, Crypteronia) or

190 FLORA MALESIANA [ser. I, vol. 8?

crystal sand (Dactylocladus); cork arising in pericycle (Axinandra, Crypteronia) or subepidermal
(Dactylocladus); node complex, with complete girdling trace (Axinandra, Crypteronia
paniculata) or with common gaps (other species of Crypterania, Dactylocladus); cortical
bundles present (Axinandra p.p.) or absent (other taxa); vesturing of vessel pits in wood confined
to the pit chamber (Axinandra, Dactylocladus) or also on pit apertures (Crypteronia); vessel-ray
pits alternate (Axinandra, Crypteronia) or reticulate to scalariform and larger (Dactylocladus);
parenchyma aliform with narrow wings (Axinandra, Dactylocladus) or chiefly diffuse in aggregates
(Crypteronia); rays heterogeneous Kribs type I (Axinandra, Crypteronia) or Kribs type III
(Dactylocladus); intercellular canal-like spaces present in rays (Crypteronia, Dactylocladus) or
absent (Axinandra).

The entire evidence from vegetative anatomy supports affinities between Crypteronia and
Axinandra — both genera sharing salient features with a number of Melastomataceae. Dactylo-
cladus resembles several Melastomataceae in its anatomy more closely than it does Axinandra or
Crypteronia. The inclusion of Alzatea from S. America and Rhynchocalyx from S. Africa in the
Crypteroniaceae adds to the anatomical heterogeneity of the family. On anatomical grounds,
Rhynchocalyx fits better in Lythraceae, and Alzatea could also be accommodated in that family
with its trilacunar nodes as only aberrant character. The existence of a considerable overlap of
the anatomical range in Melastomataceae with that of Lythraceae, Sonneratiaceae and Oliniaceae,
forbids, however, formal taxonomic decisions on anatomical grounds only. — P. BAAs.

Palynology. Pollen grains are small, ranging in size from 11 um in Crypteronia paniculata to
20 um in Rhynchocalyx lawsonioides, and thin-walled with a smooth or finely verrucate outer
surface. In Alzatea they are tricolporate, in Axinandra, Dactylocladus and Rhynchocalyx hetero-
colpate, while Crypteronia is characterized by bilaterally flattened bisyncolporate grains (MULLER,
1975).

The subdivision of the family according to the pollen types is not correlated with those based
on morphological or anatomical characters.

The relatively unspecialized A/zatea type occurs in many dicotyledonous families. The hetero-
colpate type is found in Combretaceae, Lythraceae, Melastomataceae, Oliniaceae, and Penaea-
ceae. Pollen grains similar to the Crypteronia type only occur in Cunoniaceae, but differ in
sculpture and an affinity with this family, as suggested by ERDTMAN (1952), appears remote. —
References: ERDTMAN, Pollen morphology and plant taxonomy. Angiosperms. Stockholm (1952);
MULLER, Note on the pollenmorphology of Crypteroniaceae s./. Blumea 22 (1975) 275. — J.
MULLER.

Uses. Crypteronia paniculata, which may attain a height of 30m, is said to have durable,
reddish heartwood and is sometimes used in West Java for house-building; also in S. Sumatra
reports are favourable (HEYNE, Nutt. Pl. 1927, 1158), but occurrence is too scattered to have
come into general use.

Dactylocladus stenostachys is one of the most important export timber trees of Sarawak and
Sabah; see under that species.

KEY TO THE GENERA

1. Flowers sustained by 1 bract. Stamens as many as sepals; connective not or only slightly conduplicate.
Capsule small, chartaceous. Seed situated (latero-)centrally in its wing. Tribe CRYPTERONIEAE.

2. Petals absent. Ovary superior with many ovules per cell. Capsule with many seeds per cell. Nerves

distinct, ascending and often anastomosing into a looped marginal nerve . . . . . 1. Crypteronia

2. Petals present, soon caducous. Ovary at least half-inferior, the lower part immersed in the receptacle,
with 3 ovules per cell. Capsule with 1-3 seeds per cell. Leaves coriaceous with vague venation

2. Dactylocladus

1. Flowers sustained by 3 bracts, the two lateral ones often minute. Stamens twice as many as petals.

Connective completely conduplicate. Ovary inferior, with 1 or 2 ovules per cell. Capsule large, woody,

half-inferior. Seed situated basally in its wing. Nerves ascending and anastomosing into a looped
maraimnal nerve. Tribe AXINANDREAE’ .) (i) 2 (2). SU ba ae 5 OU i

1977] CRYPTERONIACEAE (van Beusekom-Osinga) 191

1.CRYPTERONIA

BL. Bijdr. (1826) 1151; HAssk. Cat. Hort. Bog. (1844) 232 (‘Crypterhonia’); BL.
Mus. Bot. Lugd. Bat. 2 (1856) 123, t. 42; B. & H. Gen. PI. 1 (1867) 782; DC. Prod.
16, 2 (1868) 677; CLARKE in Hook. f. Fl. Br. Ind. 2 (1879) 573; KOEHNE, Verh. Bot.
Ver. Brandenburg 22 (1881) 69; O. K. Rev. Gen. Pl. 1 (1890) 250 (‘Cryptoneria’);
NIEDENZU, Bot. Jahrb. 15 (1892) 161; in E. & P. Nat. Pfl. Fam. 3, 7 (1892) 21, t. 8;
HALL. f. Abh. Naturw. Ver. Hamb. 18 (1903) 90; Med. Rijksherb. 1 (1911) 31;
ibid. 35 (1918) 17; Hutcu. Gen. FI. Pl. 2 (1967) 33; BEus.-OsINGA & BEus. Blumea
22 (1975) 258. — Henslowia WALL. PI. As. Rar. 3 (1831) 13, t. 221, non BL. 1850. —
Quilamum BLANCO, FI. Filip. 1 (1837) 851; ed. 2, 1 (1845) 136; ed. 3, 1 (1877) 245;
cf. MERR. Sp. Blanc. (1918) 282. — Fig. 1, 2, 6.

Leaves elliptic or ovate to (ovate-)lanceolate, glabrous or slightly pubescent;
midrib flat or slightly impressed above, prominent beneath; nerves ascending and
often anastomosing in a looped marginal nerve, flat above, + prominent beneath,
intramarginal nerve mostly present. Panicles terminal or axillary, sometimes on
leafless older nodes or ramiflorous, erect to usually pendulous, poorly to rather
copiously branched; axes terete to more or less angular, puberulous; racemules
with very numerous flowers. Flower-bracts persistent. Flowers bisexual or by
reduction unisexual and then trees dioecious, 4~-5(-6)-isomerous, pedicelled.
Receptacle in- and outside puberulous, inside sometimes minutely tomentose,
hardly or not accrescent. Sepals deltoid to triangular, persistent. Petals absent.
Stamens persistent, in 2 flowers staminodial and mostly permanently inflexed;
filaments filiform, somewhat flattened, connective about orbicular, with or without
a tendency to conduplication, dark when dry, anthers apically or laterally on the
connective, semiorbicular to broad-linear, latrorse or + introrse. Ovary superior
or almost so, the lower part adhering to the receptacle, (sub)globose to pyramidal,
2-4-carpellate, 2—4-celled, with free or only basally connate septs, badly developed
in 3 flowers; style filiform to subulate, somewhat longer to shorter than the ovary,
more or less puberulous, persistent; stigma punctate to capitate. Ovules many,
either in horizontal position on the septs or in + vertical position basally between
the septs. Capsule superior or almost so, (sub)globose or more or less (ob)ovoid,
puberulous, upper part dehiscent with 2-4 valves, inside split as far as the basal
connation of the septs; valves at the top kept together by the non-dehiscent part of
style and stigma. Seeds many, very small, in horizontal or vertical position; seed
ovoid-ellipsoid, situated latero-centrally in its narrow, membranous wing, which
has a shorter or longer apical and basal extension, raphe running closely along the
embryo (microscopical!).

Distr. 4 spp., of which one ranges through tropical SE. Asia (Assam, Bengal, Lower Burma, Thailand,
and Indo-China) to Malesia, the other three endemic in Malesia. Fig. 3.

Ecol. Lowland and montane rain-forests below c. 1300 m; C. paniculata also rather frequent in areas

with a more or less seasonal climate. Fl. fr. in almost all taxa Jan.—Dec.
Uses. C. paniculata seems to yield a fairly good timber, but is never found in quantity.

KEY TO THE SPECIES

1. Ovary (and capsule) 2-celled with ~ ovules cq. seeds on the septs. Dioecious tree: flowers by reduction
unisexual, rarely bisexual. Leaves papyraceous. Sect. Crypteronia ...... . 1. C. paniculata

1. oe (and capsule) 3- or 4-celled with - ovules (seeds) inserted basally. Flowers bisexual. Leaves
usually coriaceous. Sect. Basispermia.

192 FLORA MALESIANA [ser. I, vol. 8?

Fig. 1. Crypteronia macrophylla Beus.-OsINGA. a. Inflorescence, X 1/2, b. flower, x 10, c. stamen, x 15,
d. fruit, <x 10 (AsHTON §S 19372).

193

(van Beusekom-Osinga)

1977]

Fig. 2. Crypteronia macrophylla Beus.-OsinGa. Young twig with leaves, x '/; (AsHTON S 19372).

FLORA MALESIANA

[ser. I, vol. 8?

Fig. 3. Range of the genus Crypteronia BL. ; sect. Crypteronia unbroken line, sect. Basispermia broken line.

2. Inflorescences terminal at main or side twigs, 10-25(-40) cm. Ovary (capsule) 3-locular'. Sepals

usually with longitudimalicrestattheunsiders a) 5) sa. 0 eel se oO - one e 2. C. cumingii

2. Inflorescences axillary below the leaves on older nodes, or ramiflorous, 15-90 cm. Ovary (capsule)
3- or 4-celled. Sepals with or without longitudinal crest at the inside.

3. Inflorescences below the leaves on older nodes, 15-50 cm, only with primary, very rarely with a few

secondary side-axes near the base. Ovary (capsule) 3- celled. Sepals without crest . 3. C. griffithii

3. Inflorescences ramiflorous, 45-90 cm?, with primary, secondary and tertiary side-axes. Ovary (cap-

sule) 4-celled. Sepals with a longitudinal crest at the inside

1. Crypteronia paniculata BL. Bijdr. (1826) 1151;
Mus. Bot. Lugd. Bat. 2 (1856) 123, t. 42; DC. Prod.
16, 2 (1868) 679; Kurz, J. As. Soc. Beng. 46, ii
(1877) 86, incl. var. glabra (WALL.) KURZ et var.
pubescens (WALL.) Kurz; Fl. Burma 1 (1877)
519; CLARKE, FI. Br. Ind. 2 (1879) 574; NIEDENZU,
Bot. Jahrb. 15 (1892) 177; K. & V. Bijdr. 1 (1894)
203, incl. var. leptostachya (PLANCH.) K. & V.;

KING, J. As. Soc. Beng. 67, ii (1898) 5; RIDL. oe
Bull. Str. & F. M.S. ns. 1 (1902) 180 (‘Crypto-
rania’); MerR. Philip. J. Sc. 1 (1906) Suppl. 102;
BRANDIS, Ind. Trees (1906) 341; Koorpb. Exk. FI.
Java 2 (1912) 664; Atlas 2 (1914) 318; MerR. Sp.
Blanc. (1918) 282; GAGNEP. & GUILLAUMIN, FI.
Gén. I.-C. 2 (1920) 696, t. 70; RrpL. Fl. Mal. Pen. 1
(1922) 821; Merr. En. Philip. 3 (1923) 140; Crats,
Fl. Siam. En. 1 (1931) 729; Ocuse & BAKH. Ind.
Groent. (1931) 182, f. 111; BurxK. Dict. 1 (1935)
693; CORNER, Ways. Trees (1940) 197, pl. 48, f. 51;
KRAEMER, Trees W. Pacif. Region (1951) 330;
LecompTE, FI. Camb. Laos & Vietn. 4 (1965) 57;
BeEus.-OsINGA & Beus. Blumea 22 (1975) 259. —
Henslowia glabra WALL. Pl. As. Rar. 3 (1831) 14;

4. C. macrophylla

Cat. (1831-32) n. 4903; PLANcH. Hook. Lond. J.
Bot. 4 (1845) 478; Mia. Fl. Ind. Bat. 1, 1 (1856)
716. — C. glabra (WALL.) PLANCH. ex ENDL. Gen.
Pl. Suppl. 4, 2 (1847) 39; BL. Mus. Bot. Lugd. Bat. 2
(1856) 123; DC. Prod. 16, 2 (1868) 678; CLARKE,
Fl. Br. Ind. 2 (1879) 574: F.-VILL. Nov. App
(1880) 91; VIDAL, Sinopsis (1883) 27 eata D2. ake e:
Phan. Cuming. (1885) 48; Rev. Pl. Vasc. Filip.
(1886) 138. — Henslowia pubescens WALL. PI. As.
Rar. 3 (1831) 14, t. 221; Cat. (1831-32) n. 4904;
PLANCH. Hook. Lond. J. Bot. 4 (1845) 477, t.
14 B, f. 1-4; Grrr. Not. 4 (1854) 404; Ic. Pl. As. 4
(1854) t. 562 f. 3, t. 564 f. 2; Mig. Fl. Ind. Bat. 1, 1
(1856) 716. — Quilamum luteum BLANCO, FI. Filip.
1 (1837) 851; ed. 2, 1 (1845) 136; ed. 3, 1 (1877)
245; NIEDENZU, Bot. Jahrb. 15 (1892) 177. —
Henslowia affinis PLANCH. Hook. Lond. J. Bot. 4
(1845) 477; Mia. Fl. Ind. Bat. 1, 1 (1856) 716. —
Henslowia leptostachys PLANCH. Hook. Lond. J.
Bot. 4 (1845) 478; Mig. Fl. Ind. Bat. i, 1 (1856)
716. — Henslowia hookeri WALL. Cat. (1847) n.
8566, nomen. — C. affinis (PLANCH.) PLANCH. ex
EnpDL. Gen. Pl. Suppl. 4, 2 (1847) 39. — C.

(1) Rarely a few 4-locular ovaries may be found among the predominant 3-locular ones in one in-

florescence.

(2) One should be aware that of the big inflorescences of C. macrophylla usually only fragments have

been collected.

1977]

CRYPTERONIACEAE (van Beusekom-Osinga)

195

pubescens (WALL.) PLANCH. ex. ENDL. /.c.; BL.
Mus. Bot. Lugd. Bat. 2 (1856) 123; DC. Prod. 16,
2 (1868) 678, incl. var. affinis (PLANCH.) DC.;
CuarKE, FI. Br. Ind. 2 (1879) 574, incl. var. hookeri
(WALL. ex DC.) CLARKE; F.-VILL. Nov. App.
(1880) 91; NreDENzU, Bot. Jahrb. 15 (1892) 176,
incl. var. typica. — C. leptostachys (PLANCH.)
PLANCH. ex ENDL. Gen. Pl. Suppl. 4, 2 (1847) 39;
BL. Mus. Bot. Lugd. Bat. 2 (1856) 123; DC. Prod.
16, 2 (1868) 679; BaILL. Hist. Pl. 6 (1877) 436,
f. 414, 415 (‘leptostachya’); VIDAL, Phan. Cuming.
(1885) 53; Rev. Pl. Vasc. Filip. (1886) 139;
NIEDENZU, Bot. Jahrb. 15 (1892) 175; Merr. En.
Philip. 3 (1923) 140. — Henslowia paniculata (BL.)
Mia. Fl. Ind. Bat. 1, 1 (1856) 716. — C. lutea
(BLANCO) BL. Mus. Bot. Lugd. Bat. 2 (1856) 123;
DC. Prod. 16, 2 (1868) 679. — C. hookeri WALL. ex
DC. I.c. — C. wallichii DC. l.c.; HANCE, J. Bot. 14
(1876) 259.

Tree up to c. 30 m. Twigs glabrous or puberulous.
Leaves elliptic or oblong or ovate-oblong, (5—)10-
15(—25) by (3—-)5—10(-12) cm, rounded to cuneate
at the base, acuminate to cuspidate at the top,
tip usually obtuse, sometimes acute, glabrous or
puberulous, chartaceous to herbaceous, usually
finely and distinctly reticulate beneath; nerves
8-10(-12) pairs; petiole 5-10mm, 1-2mm @,
glabrous or puberulous. Panicle axillary or termi-
nal, also on leafless older nodes, (5—)10—30 cm,
branched up to the second order, without definite
peduncle, more or less pendulous, not woody;
main axis more or less angular; primary axes 2-5
pairs, the lower ones usually with 1-3 pairs of 5-25
cm long racemules. Bracts of axes triangular or
narrowly triangular, 1-6 mm, usually caducous.
Bracts of the flowers narrowly triangular to linear,
1/,-1 mm, persistent. Pedicels 1-3 mm, c.'/,mm 2.
Receptacle c. ‘|, mm high, c. 2mm wide, puberu-
lous, inside sometimes minutely tomentose. Sepals
deltoid to triangular, (*/,-)'/,-1'/,mm. Stamens

Fig. 4. Range of Crypteronia paniculata BL. var.
paniculata (unbroken line) and var. affinis
(PLANCH.) Beus.-Osinca (broken line).

5 (4), reduced and mostly permanently inflexed in
2 flowers; filaments 2!/,-3'!/, mm, glabrous; con-
nective c. 1/3(—/,) mm, not conduplicate; anthers
linear, 4/,—"/. by c. 4/4mm, latrorse. Ovary
reduced in ¢ flowers, subglobose, 2(—3)-locular,
1-2 mm, puberulous; style 1-2 mm, c. #/,mm @,
puberulous; stigma capitate, c. '/,mm. Ovules
inserted on the surface of the septs. Capsule
(sub)globose or more or less obovoid, 2-3 mm,
usually recurved; valves 2, rarely 3. Seeds */,-1 by
c. 1/, mm; acute apical part of the wing half as long
as the seed proper, basal part mostly short.

Distr. Continental SE. Asia (Assam: Khasya
Hills, Sylhet, Lushai Hills; Bengal; Burma:
Arakan Div. to Rangoon; S. Indo-China) and
West Malesia: eastwards to the Philippines,
Borneo, and Flores. Fig. 4.

Note. Two replacing varieties are distinguished
but it should be mentioned that in Penang and in
the Rangoon area, where the ranges coincide, more
or less intermediate specimens are found which have
almost or practically glabrous leaves and twigs.

KEY TO THE VARIETIES

1. Leaves and branches glabrous. Receptacle
inside more or less puberulous. Sepals c. 1 mm
long. Capsules (sub)globose var. paniculata

1. Leaves at least beneath, and young twigs
puberulous. Receptacle inside tomentose.
Sepals */,—'/,(-*/,) mm long. Capsules (sub)-
globose to obovoid ....... var. affinis

var. paniculata. — C. paniculata, incl. var. glabra et

var. leptostachya — C. glabra — Quilamum luteum

— C. leptostachys — C. lutea — C. wallichii.
Leaves (5-)10-15(-25) by (3-)5-10(-12) cm,

glabrous; petiole glabrous. Buds glabrous. In-

florescence up to c. 30 cm; racemules up to 25 cm.

Receptacle inside puberulous. Sepals c. 1 mm long.

Staminodes */,-1 mm long. Capsule (sub)globose.
Distr. Continental SE. Asia: E. Bengal (Chitta-

gong), Assam (Khasia Hills, Lushai Hills),

Andaman Is., Burma (Arakan, Rangoon, Pegu,

Tenasserim); in Malesia: Malay Peninsula (Kedah,

Perak, Langkawi I., Pahang, Selangor, incl.

Penang I.), Sumatra (Sibolangit, Pajakumbuh,

Painan, Palembang), Java, Lesser Sunda Is. (Bali,

Sumbawa, Flores), Borneo (Sabah, W. Kutai),

Philippines. Fig. 4.

Ecol. Primary and secondary forests on hills
and mountains up to 1700 m. Reported from river-
banks, ridges, ravines and forest borders. Scattered,
locally fairly common.

Field notes. Bole up to 15m, often crooked.
Buttresses sometimes present. Bark smooth or
rough, scaling off. Young twigs and leaves deep
blue or violet (CorRNER), when dried blackish.
Flowers white, pale green, or greenish yellow.

Uses. The timber is reported as of good quality,
used amongst others for house-building purposes
and cart-wheels. In the Philippines the bark is
sometimes used against skin-eruptions. In Java the
young shoots are eaten with rice as lalab (HAss-
KARL).

Vern. Sumatra: kayu kapas, médang ajam, M;
Malaya: békoi, békwoi, berkol, bua babi, ménkuah,
rupal, tukoi, M; Java: kibanén, kitjéngklak, S,
kayu babi, kigandik, ponggokan (Jakarta, M),

196

FLORA MALESIANA

[ser. I, vol. 82

kibakko, kidjarak (Priangan), bang-kongan (Ban-
ten), blis or blisan, kayu tjéléng (Banjumas),
mungur (Madiun), wungu lubu (Kediri), tjéleng(an,)
wrakas or kwakas (Pekalongan), glingsing (Pasu-
ruan), sépat (Besuki), all J; Flores: madja; Sum-
bawa: sarowe; Borneo: kinkidon mantok, Dusun;
Philippines: balinog, bitog, bitok, bongari, bongaui,
bungaing, kilamo, malabayanas, salasan, tiaui, tua,
Tag., agidai, agudai, barabok, barakbak, bungaing,
Ilk., banujo, tolan-manok, P.Bis., baroga, baruga,
kodai, kudai, ladao, \g., kamanok, Bis., malabiong,
Sbl.

var. affinis (PLANCH.) BEuS.-OsINGA, comb. nov. —
Henslowia affinis PLANCH. Hook. Lond. J. Bot. 4
(1845) 477. — C. affinis — C. pubescens, incl. var.
affinis (PLANCH.) DC. Prod. 16, 2 (1868) 678. —
C. hookeri.

Leaves (2-)5—10(-16) by (11/,-)3-6(-8) cm,
sometimes sparsely puberulous above, more or less
puberulous beneath especially on the nerves;
petiole puberulous. Buds puberulous. Inflorescence
up to c. 20 cm; racemules up to 20 cm. Receptacle
inside tomentose. Sepals 1/,—1/,(-*/4) mm long.
Staminodes 0.1-0.3mm long. Capsule (sub)-
globose to obovoid.

Distr. Continental SE. Asia: Burma (Rangoon,
Pegu, Tenasserim Div.), Thailand, Laos, Cam-
bodia, S. Vietnam; in Malesia: Malaya (Kedah;
Penang I.). Fig. 4.

Ecol. Several times reported from riverbanks
and ridges, 90-300 m.

2. Crypteronia cumingii (PLANCH.) PLANCH. ex
ENDL. Gen. Pl. Suppl. 4, 2 (1847) 39; BL. Mus.
Bot. Lugd. Bat. 2 (1856) 123; DC. Prod. 16, 2
(1868) 678; F.-VILL. Nov. App. (1880) 91; VIDAL,
Phan. Cuming. (1885) 20; Rev. Pl. Vasc. Filip.
(1886) 138; NIEDENzU, Bot. Jahrb. 15 (1892) 179;
Me_rr. Philip. J. Sc. 1 (1906) Suppl. 102; En.
Philip. 3 (1923) 139; MERR. & Perry, J. Arn. Arb.
12 (1941) 270. — Henslowia cumingii PLANCH.
Hook. Lond. J. Bot. 4 (1845) 478, t. 14, f. C 1-4. —
C. javanica BAILL. Hist. Pl. 6 (1877) 435, f. 412,
413, nomen. — C. laxa ELMER, nomen in sched., cf.
Me_rr. En. Philip. 3 (1923) 140.

Tree up to 40 m. Twigs glabrous. Leaves elliptic
to ovate-oblong, (5—)8—25 by (3-)4-14 cm, usually
emarginate sometimes rounded or acute at the base,
acute, sometimes acuminate at the top, tip mostly
obtuse, glabrous, coriaceous, distinctly and slightly
prominently reticulate beneath; nerves 6—12(—15)
pairs; petiole 2-10(-15) mm, 1-4 mm @, glabrous.
Inflorescence terminal, at the end of main or side
twigs, 10-25(-35) cm, branched up to the first,
second or third order, whether or not peduncled,
erect, with flaccid to stiff and sublignose axes ; main
axis more or less flattened; primary axes 5-25,
paired or irregular, sometimes in whorls of 4;
racemules up to 15cm. Bracts of axes triangular,
2-10 mm, sometimes present as small or reduced
leaves, persistent or caducous. Bracts of flowers
narrowly triangular, 1/,-1!/, mm, persistent. Pedi-
cels 11/,-2 mm, c. !/, mm @. Receptacle }/,-11/, mm
high, 11/,-2'/, mm wide, densely whitish papillose
inside. Sepals 5, triangular, 1-1*/,(-1/,) mm,
inside mostly with a longitudinal densely puberu-
lous crest. Stamens 5; filaments 1—2!/, mm, glab-
rous; connective c. 1/, mm, slightly conduplicated;

anthers linear or semiorbicular, c. 1/,(-'/,) by
0.1-0.2mm, + latrorse. Ovary subglobose, 3-
celled, 1—-11/, mm, whitish papillose and/or puberu-
lous; style (1—)1*/,-2'/, mm, 0.2-0.4 mm @, more
or less shortly puberulous to papillose; stigma
punctate to slightly capitate. Ovules inserted on the
somewhat conically elevated edges between the
bases of the septs. Capsule (sub)globose, more or
less episeptally impressed, c. 2mm; valves 3.
Seeds c. 3/4 by c. !/4mm, apical and basal part of
wing from '/, to 1 times as long as the seed proper.

Distr. Malesia: Borneo (Sarawak), Philippines
(Luzon), Celebes, Moluccas (Morotai, Halma-
heira), New Guinea (incl. Misool and Louisiades:
Rossel I.). Fig. 5.

Fig. 5. Range of Crypteronia cumingii (PLANCH.)
PLANCH. ex ENDL.

Ecol. Primary and secondary forests, on ridges,
slopes, and along riverbanks, usually at low alti-
tude, but also in the hills to 1200 m, once even
found at 1800 m (Central Celebes, Malili area:
B. Takale Kadju).

Vern. Borneo: ubah, Iban, Sarawak; Philip-
pines: andalai, Tag., ugdu, tigduon, Bik.; Central
Celebes, Malili area: kumba-a, langori tauru,
longari, tomo wanna.

Notes. As appears from his drawings, C.
javanica BAILL. must be referred here: bisexual
flower with basal ovules; very probably they were
made after a Philippine specimen.

In a depauperate collection from New Guinea
(NGF 2958) the panicles are axillary.

In specimens from the Moluccas frequently
6-merous flowers occur among the normally 5-
merous ones.

In specimens from New Guinea there are usually
some 4-celled ovaries and 4-celled fruits present
among the predominantly 3-celled ones.

3. Crypteronia griffithii CLARKE in Hook. f. FI.
Br. Ind. 2 (1879) 574; Kina, J. As. Soc. Beng. 67, ii
(1898) 5; Rip. Agr. Bull. Str. & F. M.S. ms. 1
(1902) 180; Fl. Mal. Pen. 1 (1922) 821; Watson,
Mal. For. Rec. 5 (1928) 176; Burk. Dict. 1 (1935)
693; CORNER, Ways. Trees (1940) 198. — Hens-
lowia sp., GRIFF. Not. 4 (1845) 404; Ic. Pl. As. 4
(1845) t. 564, f. 1. — Fig. 6.

CRYPTERONIACEAE (van Beusekom-Osinga)

Fig. 6. Crypteronia griffithii CLAnke, Singapore watercatchment area, Dec. 1969 (Photogr. VAN BEUSEKOM).

198

Tree up to 40m, 60cm @. Twigs glabrous.
Leaves elliptic-oblong to ovate-oblong, (5-)10-25
(-35) by 5—12(-15) cm, rounded or emarginate at
the base, acute, sometimes acuminate at the top,
tip usually obtuse, sometimes acute, glabrous,
coriaceous, distinctly and slightly prominently
reticulate beneath; nerves (8—)10—15(—18) pairs;
petiole (2-)5-lOmm, 2-5mm_ 4@,_ glabrous.
Inflorescence axillary, one or a few together below
the leaves on older nodes, 15-50 cm, only with
primary side axes, very rarely with a few secondary
side axes near the base, without distinct peduncle,
sublignose, often ferrugineously pubescent, with
pendulous axes; main axis terete or somewhat
flattened, finely ribbed; primary axes 8-12(-15),
not always paired, finely ribbed; racemules up to
50 cm. Bracts of axes triangular, c. 1 mm, mostly
caducous. Bracts of flowers subulate, c. 1 mm,
persistent. Pedicels '/,-1(-11/,) mm, c. ?/,mm @.
Receptacle c. 1mm high, 2-3 mm wide, densely
whitish papillose inside. Sepals triangular, 11/,-2
mm. Stamens 5; filaments 3-4mm, glabrous;
connective '/,—'/,mm, slightly conduplicated;
anthers linear, !/,-'/, by c. 0.1 mm, + latrorse.
Ovary (sub)globose to pyramidal, 3-celled, c. 1 mm,
more or less fine whitish papillose whether or not
with longer hairs in between; style (11/,-)3-4(—5)
mm, c. +/,mm @, sparsely puberulous; stigma
slightly capitate. Ovules inserted on the somewhat
conically elevated edges between the bases of the
septa. Capsule (sub)globose, c. 2mm; valves 3.
Seeds 0.3-0.4 by c. 0.2 mm, apical and basal part
of wing about 1'/, times as long as the seed proper.

Distr. Burma (Moulmein, one coll.); in Malesia:
Central Sumatra (one coll.), Malay Peninsula
(incl. Penang I.), Borneo (W. Sarawak, Sandakan,
Gaya I., E. Kutai, Nunukan I.). Fig. 7.

Ecol. In primary lowland forests, often on sandy
soils, up to 500 m.

Field notes. Bole usually straight. Bark surface

Fig. 7. Range of Crypteronia griffithii CLARKE.

FLORA MALESIANA

[ser. I, vol. 8?

smooth or rough and scaly. Twigs strongly swollen
at the nodes. Young leaves purplish. Flowers dark
blue to purple or magenta, sometimes noted yellow
or yellowish reddish.

U ses. Rmp_ey (1902) reported the wood to be
durable and used for house-building.

Vern. Sumatra: panarahan; Malay Peninsula:
békwoi, kélat tampoi, nyirék bukit, sémpo, sérumpu,
simpoh, sumpu(t ), télingga badak ; Borneo: engkolot,
rambai rambai, Gaya I., ubah sémut, Iban.

4. Crypteronia macrophylla Beus.-OsINGA, Blumea
22 (1975) 261. — Fig. 1, 2.

Tree up to 20 (?) m. Twigs glabrous. Leaves
ovate-oblong to lanceolate, (20—)25—45 by 7-15 cm,
usually emarginate, sometimes rounded at the base,
acute to + acuminate at the top, tip obtuse,
glabrous, coriaceous, distinctly and + prominently
reticulate beneath; nerves 15-25 pairs; petiole
5-10(-15) mm, 2-5 mm @, glabrous. Inflorescence
ramiflorous, 45-90 cm, always branched up to the
third order, without distinct peduncle, pendulous,
woody, with spreading axes; main axis terete;
primary axes many, not always paired; racemules
up to 30cm. Bracts of axes triangular, c. 1 mm,
caducous. Bracts of the flowers subulate, c. 1 mm,
persistent. Pedicels 1-3 mm, c. !/,mm @. Recep-
tacle + flat, 1-1!/,mm wide, puberulous, not
densely papillose inside. Sepals deltoid, c. 1 mm,
inside with a longitudinal densely puberulous crest.
Stamens 4; filaments 11/.-3 mm, glabrous, at the
very base puberulous; connective c. 1/, mm, more
or less conduplicated; anthers + linear, 1/, by
0.1-0.2 mm, + introrse. Ovary pyramidal, 4-celled,
1-2 mm, puberulous, more or less episeptally
impressed; style (1—)11/,-2'/,(-3) mm, c. 1/4 mm @,
puberulous; stigma truncate, hardly wider than
the style. Ovules inserted on the somewhat coni-
cally elevated edges between the bases of the septs.
Capsule (sub)globose to pyramidal, 2-2'/, by
2-21/, mm; valves 4. Seeds c.+/, by c. !/4 mm, apical
and basal part of wing as long as or slightly longer
than the seed proper.

Distr. Malesia: Borneo: Sarawak (Kuching,
Sibu), Kutai (several localities), Sambas region.
Fig. 8.

Fig. 8. Range of Crypteronia macrophylla BeEus.-
OSINGA.

Ecol. In primary forest up to 1200 m. Locally
frequent. Young branches are often inhabited by
ants.

CRYPTERONIACEAE (van Beusekom-Osinga) 199

1977]

Field notes. Tree with semi-pendent branches. Note. The large, complex, woody inflorescence
Bark surface smooth. Young leaves rich purplish. which is always produced below the leaves on older
Flowers greenish with purple-brown stamens. nodes is characteristic.

Fruits dark green.

"
o
Pavan, <2

ey
=
‘a:
a
gq
CG

Fig. 9. Dactylocladus stenostachys OLAV. a. Habit, x 2/s, b. flower, x 10,c. seed, x 16 (a, b Fucus 21186,
cS 9261).

2. DACTYLOCLADUS

Outv. in Hook. Ic. Pl. (1895) t. 2351; Gite in E. & P. Nat. Pfl. Fam. Nachtr. |
(1897) 267; HALL. f. Med. Rijksherb. 35 (1918) 18; BAKH. f. Rec. Trav. Bot.

200 FLORA MALESIANA [ser. I, vol. 8?

Néerl. 40 (1943) preprint 329; Beus.-OsINGA & Beus. Blumea 22 (1975) 261. —
Fig. 9.

asec elliptic or obovate to (obovate-)oblong, glabrous; midrib thickish, + flat
above, prominent beneath; nerves + straight, anastomosing in an indistinct mar-
ginal nerve. Inflorescence terminal or axillary to the highest leaves, erect, poorly
branched; axes flattened, puberulous; racemules with at least a few tens of flowers.
Flower-bracts caducous. Flowers bisexual, 5-—(4-)isomerous, almost sessile.
Receptacle in- and outside puberulous, accrescent. Sepals triangular, persistent.
Petals unguiculate, with suborbicular, irregularly lobed lamina, in bud covering
the stamens as a hood, soon caducous. Stamens persistent; filaments terete, some-
what flattened; connective about orbicular, not conduplicate; anthers inserted
transversally, somewhat below the upper margin of the connective, oblong to
broad-linear, introrse. Ovary half-inferior, the lower part adnate to the recep-
tacle, the top part semiglobose, puberulous, (3-), 4- or 5-carpellate, (3-), 4- or
5-locular, septs not connate; style subulate, somewhat longer than the ovary,
puberulous, persistent; stigma capitate. Ovules 3 per locule, inserted in vertical
position, basally between the septs. Capsule almost inferior, broad-ellipsoid, small,
pericarp chartaceous, puberulous, inside dehiscent down to the bottom with (3) 4
or 5 valves of which only the upper !/, protrudes from that part of the pericarp that
is surrounded by and fused with the enlarged receptacle, at the top often kept to-
gether by the non-splitting stigma. Seeds 3 per locule (1 or 2 sometimes not
developed), small, in vertical position; seed narrow-ellipsoid, flat, situated centrally
in its more or less rectangular, narrow, membranous wing almost 2 times as long
as the body of the seed; raphe running close to the embryo.

Distr. Malesia: Borneo and W. New Guinea (sterile coll.). Fig. 10.
Ecol. Lowland peat swamp forest.

1. Dactylocladus stenostachys OLIv. in Hook. Ic.
Pl. IV, 4 (1895) t. 2351; HALL. f. Med. Rijksherb.
35 (1918) 18; Merr. En. Born. (1921) 452; Die_s &
HACKENBERG, Bot. Jahrb. 60 (1926) 312; Baku. f.
Rec. Trav. Bot. Néerl. 40 (1943) preprint 329;
BROWNE, For. Trees Sarawak & Brunei (1955) 261,
t. 33; ANDERSON, Gard. Bull. Sing. 20 (1963) 178,
pl. 1,2, 6, 7; MEIER, Field Guide Trees W. Mal.
(1974) 205, f. 51, pl. 14; Beus.-OsINGA & BEus.
Blumea 22 (1975) 262. — Fig. 9.

Tree up to 40 m, dbh up to 1'/, m, at the base
producing pneumatophores. Twigs often several
together per leaf-axil, the younger ones often with
ribbed angles. Leaves 4-8(—16) by 2!/,4(-6) cm,
with revolute margin, cuneate at the base, some-
times emarginate, usually rounded up to acuminate
at the top with acute tip, coriaceous; nerves 11-15
pairs, usually rather obscure, flat to prominulent
above and beneath; petiole 3-5 mm, 2-3 mm @.
Inflorescence when axillary 1-3 together, erect, up
to 14cm, consisting of 3 racemules; peduncle up
to 6cm, (sub)glabrous; axes finely ribbed; race-
mules c. 8 cm, from 1 cm above the base + densely
set with flowers. Bracts of axes minute, soon cadu-
cous. Bracts of the flowers linear or narrowly
triangular, c. 1mm. Pedicels up to '!/, mm.
Receptacle c. 2 mm high, c. 2'/, mm wide. Sepals
c. 1 mm. Petals c. 1 mm, puberulous outside and

on the margin. Filaments c. 1 mm, 0.2 mm wide,
puberulous; connective */,—'/, mm; anthers 0.2 by
0.1mm. Style 1!/,.-2mm, c. 0.2mm @; stigma
1/,1/, mm. Capsule 3-4 by 2—3(-31/,) mm. Seed c.
1.4 by 0.2-0.3 mm, including wing 21/,-3 by
3/4-1"/, mm. ;

Distr. Malesia: widely distributed in Borneo.
It was by error incidentally reported from Malaya
(Fl. Mal. Bull. p. 1696, p. 2375). Fig. 10.

Ecol. Durant (For. Rep. Brunei, 1933, 6,
photogr.) reported this species (under the name
Crypteronia) to occur in Brunei as an associate of

Fig. 10. Range of the genus Dactylocladus OLIv.;
D. stenostachys Outv. dots, D. sp. triangle.

1977]

CRYPTERONIACEAE (van Beusekom-Osinga)

201

Dryobalanops and Combretocarpus in considerable
quantity over large areas of freshwater swamp,
often with 12 mature trees (over 30 cm @) per ha.
Diets & HACKENBERG (/.c.) mentioned its occur-
rence in the Sampit swamp forest area in SW.
Borneo together with Combretocarpus, Campno-
sperma, etc. BROWNE (i/.c.) stated that it occurs in
practically all types of peat swamp forest in Sara-
wak. He found its frequency somewhat lower than
DurANnrtT did; he found sometimes 8, but averagely
3 mature trees per ha, but he said that locally,
between Balingian and Bintulu, it was the dominant
tree of the swamps of Sarawak.

BROWNE recorded that the vernacular names
jongkong and tabak allude to characteristic quali-
ties : jongkong referring to the rather stout yellowish
pneumatophores at the stem-base, while tabak
would refer to the characteristic minute perfora-
tions of the wood from radial vessels.

ANDERSON (/.c.) confirmed BROWNE’s observa-
tion that it is one of the most characteristic swamp
forest trees, the only species represented in all
communities throughout Sarawak and Brunei.
He produced photographs of the Gonystylus-
Dactylocladus-Neoscortechinia (l.c. pl. 1 & 2) and
the Combretocarpus-Dactylocladus associations
(l.c. pl. 6 & 7).

Its wide range in the peat swamps is also con-
firmed palynologically in the peat according to
ANDERSON & MULLER (Rev. Palaeobot. & Palyn.
19, 1975, 314-316) where it figures in phases 1-6.
In a Miocene deposit in Brunei, near Berakas,
MULLER reported also its occurrence (/.c. f. 5,
diagram) with the associates as today. It must be
added that its pollen can be confused with that of
Axinandra and some Melastomataceae of which
the latter also occur in peat swamps, albeit in small
numbers.

It is remarkable that whereas this type of peat
forest ranged unbrokenly at least from the
Miocene to the Present, Dactylocladus is not yet
found in Sumatra and Malaya, provinces with
which Borneo was joined, during the Pleistocene
Glacial period, by a huge lowland riverine area
(now the South China Sea); this would have made
dispersal and exchange very probable, as it was for
its associate Combretocarpus and some species of

Gonystylus. It may yet be discovered in the peat
swamps of Malaya and Sumatra, but must then be
very rare.

Uses. According to BROWNE (/.c.) it is the fourth
important export timber tree of Sarawak. Extrac-
tion is facilitated by the fact that the logs float in
water. MEIER (/.c.) termed it a general utility
timber in Sabah where it is, besides Gonystylus,
the most important export timber from the Klias
Peninsula.

Vern. Jongkong, tabak (the most common
names), éntibu, garu buaja, jinjang, (médang)
béladi, m. miang, mélinkat kérangas, mérébong,
térénjangan.

Dactylocladus sp. — Cf. MEER, Field Guide Trees
W. Mal. (1974) 205, in note.

Distr. Malesia: West New Guinea: in peat
forest along Rouffaer R., the only large tree in this
forest type, c. 175 m alt., DocTERS VAN LEEUWEN
9973 (BO, L, etc.), sterile, distributed as Memecylon
sp. Fig. 10.

Notes. Through MerJer (/.c.) attention was
drawn to this collection which was pre-identified
through the uncanny form knowledge of Mr. NEpI
at Bogor. Mr. G. VAN VLIET, Leyden, has examined
the leaves anatomically and found no difference
with the Bornean species. Dr. VAN STEENIS, who
unearthed the Leyden duplicate, has found that it
shares a small but significant vegetative character
with the Bornean species, viz the occurrence of a
shallow, rimmed cavity-like depression at the
extreme base of the petiole, similar to that in e.g.
Garcinia; the two small cups together envelop the
terminal bud. This is not found in Memecylon.

Though in fact he found the sterile macromor-
phology exactly matching, we like to postpone
judgement on specific status until flowers and fruit
are available.

About the considerable geographical gap be-
tween Borneo and West New Guinea it can be said
that true peat forest is not known from this gap
at the present time. This is no explanation, how-
ever, as the same gap occurs in the genus Koom-
passia (Leg.-Caes.) which is not a peat-forest tree
genus; in that genus the New Guinean species is
distinct from the two of the Sunda shelf.

3. AXINANDRA

Tuw. in Hook. J. Bot. 6 (1854) 66, t. 1 C; En. Pl. Zeyl. (1859) 122; B. & H. Gen.
Pl. 1 (1867) 784; Bepp. FI. Sylv. 2 (1869) t. 207; Bait. Adansonia 12 (1876) 84;
CLARKE in Hook. f. Fl. Br. Ind. 2 (1879) 581; CoGNiAux in DC. Mon. Phan. 7
(1891) 1113; KRASsER in E. & P. Nat. Pfl. Fam. 3, 7 (1893) 142, 196; BAKu. f. Rec.
Trav. Bot. Néerl. 40 (1943) preprint 332; Mewer, Ceyl. J. Sc. (Biol. Sc.) 10 (1972)
72; Beus.-OsINGA & Beus. Blumea 22 (1975) 262. — Naxiandra (BAILL.) KRASSER
in E. & P. Nat. Pfl. Fam. 3, 7 (1893) 197, f. 182 A. — Fig. 11.

Leaves elliptic to oblong, sometimes ovate, glabrous; midrib impressed above,
prominent beneath; nerves ascending and anastomosing in a looped marginal
nerve, intramarginal nerve present. /nflorescence terminal or axillary and then at the
end of the twigs, erect, poorly branched; axes more or less angular, puberulous;
racemules with up to some tens of flowers. Flower-bracts 3 per flower, the outer

202 FLORA MALESIANA [ser. I, vol. 8?

S
S$

"TSDIIN
VY

()

Boa
A)

Ale

Fig. 11. Axinandra coriacea BAILL. a. Habit, x 1/,, b. venation undersurface of leaf, nat. size, c. bud,

d. older bud, style protruding, petals separating, e. mature flower, petals dropped, f. old flower, petals and

stamens dropped, all x 7'/,, g. ripe capsule, x 3/4, h. stamen, x 15 (a SAN 57276, c-f,h S 14489, g
MEIER SAN 49845).

ones often minute, persistent during anthesis. Flowers bisexual, 5(—4)-merous,
pedicelled. Receptacle puberulous outside only, glabrescent in fruit, much accres-
cent and lignified in fruit. Sepals 5 (4), deltoid, more or less acuminate, evanescent
in fruit. Petals 5 (4), valvate-connate or valvate- (or somewhat imbricate-)condupli-
cate or imbricate-contorted, more or less connate or cohering and soon dropping
simultaneously in the shape of an umbrella, flimsy, consisting of a wide basal part
tapering into a narrower median part which widens into a whether or not well-
developed, frayed apical part, induplicate and enveloping the stamens which are
situated in pairs between two longitudinal lamellae on the inside of each petal.
Stamens 10 (8), epi- and alternisepalous, (sub)equal, caducous; filaments (rather)
thick and short, terete, to more or less flattened, tapering upwards; connective

1977] CRYPTERONIACEAE (van Beusekom-Osinga) 203

wide, large, conduplicate, mostly provided with a more or less distinct dorsal
tubercle; anthers marginally at the apex of the connective, broad-linear, introrse.
Ovary inferior, immersed in the receptacle, 3-(2—)carpellate, 6-(4-)celled, with
free or towards the base somewhat connate septs, glabrous; style (subulate-)
terete, shorter to distinctly longer than the receptacle, and whether or not pro-
truding from the ripe bud, glabrous, evanescent in fruit; stigma punctate, minute.
Ovules | or 2 per cell, two (one) by two (one) in vertical position inserted basally
on either side of the 3 (2) stouter ones of the 6 (4) septs. Capsule half-inferior, big
and woody, globose to ellipsoid, (sub)glabrous, when young provided with a frag-
ment of the style, inside dehiscent down to the bottom with 2-6 valves of which
only the upper parts protrude from that part of the capsule which is surrounded by
and fused with the enlarged receptacle of which the rim often remains visible as an
irregular more or less conspicuous rib; visible part of the valves triangular, coarse,
solid, deltoid at cross-section, glabrous. Seeds few, in vertical position; seed
depressed-ellipsoid, situated basally and obliquely in its thin, (narrow-)oblong
wing 2-3 times as long as the body of the seed; raphe running from the basal inser-
tion all along the wing margin back to the embryo.

Distr. Ceylon (1 sp.) and Malesia: Malay Peninsula (one record) and Borneo (3 spp.) but nowhere
common. Fig. 12.

A. zeylanica was also mentioned to occur in Borneo by BAKHUIZEN f. (Rec. Trav. Bot. Néerl. 40, 1943,
preprint 332) but HALLIER f. 2683, on which this was based, belongs to A. coriacea. The same author (i.c.)
recorded A. borneensis BAKH. f. (= A. beccariana) from Billiton I.; the sheet TEYSMANN 5.7. was collected,
however, on Mt Blitong in Borneo.

As with Dactylocladus the extreme rarity of the genus on the Sunda-shelf west of Borneo is remarkable
and remains unexplained; sofar known Axinandra is not bound to a rare or very specialized biotope.

Ecol. Lowland and submontane rain-forest, up to 1200 m.

KEY TO THE SPECIES

1. ene pear- or drop-shaped. Filaments 1'/,-2 mm. Style 2-5 mm. Ovules 2 per cell. Sect. Naxian-
a BAILL.
2. Internodes winged upwards. Leaf-base rounded to emarginate. Inflorescences distinctly longer than

3 ell Rite ie ales. Ft! Peat ia east) ted eR A Andel GO RN A oes os ala sal ing i oe 1. A. alata
2. Internodes not winged. Leaf-base shortly attenuate. Inflorescences shorter, or distinctly longer than
5 cm.

3. Leaves chartaceous. Tip of leaf-apex acute. Inflorescence 5—20cm. Petals c. 3mm. Fruit 1'/,-2 by

1-1'/, cm; rim of enlarged receptacle about median 2. A. beccariana

3. Leaves coriaceous. Tip of leaf-apex obtuse. Inflorescence stunted, (1-)2—4(—5) cm. Petals c. 6 mm
(fig. 11d). Fruit 2—3(—3'/,) by 1'/,-2'/, cm; rim of enlarged receptacle infra-median (fig. 11g)

3. A. coriacea

1. Connective quadrate-elliptic. Filaments c. '/, mm. Style shorter than '/, mm. Ovules | per cell. Sect.

eS ae Poe ee Ale rok Velt meg Ayn Wye pew

ov We ee!) Oo id Oe ee Oe ee es ee ee

Axinandra. Species of Ceylon

1. Axinandra alata BAILL. Adansonia 12 (1876) 86;
Bull. Soc. Linn. Paris (1877) 128; CoGNiaAux in
DC. Mon. Phan. 7 (1891) 1114; Merr. En. Born.
(1921) 452; Beus.-OsincA & Beus. Blumea 22
(1975) 263.

Tree. Twigs glabrous; internodes terete at the
base, orn acutely quadrangular upwards with
4 = ually widening wings towards the nodes,
(sub)glabrous; wings 1-3 mm (or more?) wide at
the top, ending in an acute processus often curved
upwards, wearing off when older. Leaves 5—10 by
3-5 cm, rounded at the base, cuspidate at the top
and with acute tip, chartaceous to subcoriaceous,
distinctly reticulate; nerves c. 12 pairs, flat above,
prominent beneath; petiole 3-5 mm, 1-1'/, mm @.
Inflorescence 8-13 cm, consisting of main axis with
1 or 2 pairs of 4-12 cm long primary axes each

A. zeylanica THw.

bearing 10-35 flowers. Bracts of axes deltoid to
triangular, c. 1 mm, acute at the top, ig ce
puberulous, mostly caducous. Floral bracts small,
the middle one only slightly exceeding the lateral
ones, narrowly triangular, c. 1mm. Pedicels
1-2 mm, c. '/,mm @. Receptacle c. 2mm high,
c. 3mm wide, ribbed, densely puberulous. Sepals
c. */4mm. Petals c. 3'/,-4 mm, 1-1'/, mm wide at
the base, valvate to imbricate at the base, valvate-
conduplicate for the rest, more or less connate
especially in the median part, almost completely
enveloping the stamens. Filaments c. 2 mm, 1/,-
3/,mm wide at the base, glabrous. Connective
pear- or drop-shaped, c. 1 mm, with dorsal swelling.
Anther-cells c. 1 mm, c. '/,4 mm wide. Style 3-5 mm,
'/,mm @, protruding from mature bud. Ovules
2 per cell. Fruit not seen.

204

Distr. Malesia: Borneo (Sarawak), only

known from the type.

FLORA MALESIANA

[ser. I, vol. 82

A. maingayi). The differences between this speci-
men and the material from Borneo as mentioned
by CLARKE /.c. are in my opinion of minor impor-
tance; consequently, I have included A. maingayi
in the synonymy of A. beccariana.

Se Sas 2-35" |

Fig. 12. Range of the genus Axinandra Tuw. In
Ceylon 1 sp., in Borneo 3 spp. of which one once
found in Malaya.

2. Axinandra beccariana BAILL. Adansonia 12
(1876) 85; Bull. Soc. Linn. Paris (1877) 127;
COGNIAUX in DC. Mon. Phan. 7 (1891) 1114;
Mere. En. Born. (1921) 452; BrEus.-OsINGA &
Beus. Blumea 22 (1975) 263. — A. maingayi
CLARKE, FI. Br. Ind. 2 (1879) 581; CoGNIAuUx in
DC. Mon. Phan. 7 (1891) 115; Rm. Fl. Mal. Pen.
1 (1922) 826. — A. borneensis BAKH. f. Rec. Trav.
Bot. Néerl. 40 (1943) preprint 332.

Tree. Twigs glabrous or very minutely puberu-
lous. Leaves 5-10 by 2—5 cm, shortly attenuate at
the base, acuminate to cuspidate at the top, with
acute tip, chartaceous, faintly reticulate; nerves
8-12 pairs, flat above, prominent beneath; petiole
5-8mm, 1-2mm @. Inflorescence 5-20cm,
branched up to the second order, with slender axes
each bearing up to c. 15 flowers; primary axes up to
4 pairs, 4-15cm. Bracts of the axes deltoid to
narrowly triangular, 1-3 mm, acute at the top,
subglabrous, mostly caducous, sometimes partly
present as small or reduced leaves. Floral bracts
small, the middle one triangular to linear-lanceo-
late, 1/,-4 mm, the lateral ones minute. Pedicels
1-2!/, mm, c. !/,mm @. Receptacle c. 2 mm high,
c. 3mm wide, sometimes faintly ribbed, sparsely
minutely puberulous. Sepals c. 1/, mm. Petals c.
3mm, c. 1 mm wide at the base, valvate, almost
completely connate, only enveloping the dorsal and
part of the apical side of the stamens. Filaments c.
1'/,mm, c. '/;mm wide at the base, glabrous.
Connective pear- or drop-shaped, c. 1'/, mm, with
dorsal swelling. Anther-cells c. 1 by c. 1/, mm.
Style c. 2 mm, c. !/, mm @, hardly or not protrud-
ing from mature bud. Ovules 2 per cell. Capsule
11/,-2 by 1-1!/,cm, faintly 10-ribbed, rim of
enlarged receptacle about median; valves c. 3/, cm.
Seeds not seen.

Distr. Malesia: Borneo and Malay Peninsula
(Malacca, one old record). Fig. 13.

Ecol. Lowland forests.

Note. From the Malay Peninsula only one
collection is known (MAINGAY 654-2, type of

Fig. 13. Localities of Axinandra beccariana BAILL.
(triangles) and A. coriacea BAILL. (dots).

3. Axinandra coriacea BAILL. Adansonia 12 (1876)
85; Bull. Soc. Linn. Paris (1877) 127; Hist. Pl. 7
(1880) 28, f. 43; CoGNIAUx in DC. Mon. Phan. 7
(1891) 1114; Merr. En. Born. (1921) 452; Beus.-
OsINGA & BEus. Blumea 22 (1975) 264. — Naxian-
dra coriacea (BAILL.) KRASSER in E. & P. Nat. Pfl.
Fam. 3, 7 (1893) 197. — Fig. 11.

Tree up to 35m, 50cm 2g. Twigs glabrous.
Leaves 5-10(-12) by (1'/,-)2-5(-6) cm, shortly
attenuate, sometimes acute at the base, acuminate
to cuspidate at the top, with obtuse tip, coriaceous,
conspicuously reticulate; nerves 8-12 pairs, flat
above, prominent beneath; petiole 5-8 mm, c.
1-2mm @. Inflorescence (1-)2-4(-5) cm, con-
sisting of a few, sometimes one 1-5 cm long rather
coarse, sometimes stunted axes, each bearing 0-15
flowers. Bracts of the axes deltoid to triangular,
c. 1mm, obtuse at the top, subglabrous, mostly
caducous. Floral bracts small, the middle one del-
toid to triangular, c. 1 mm, the lateral ones minute.
Pedicels 1—2!/, mm, c. 1 mm @. Receptacle c. 3 mm
high, 3-4(-5) mm _ wide, densely puberulous.
Sepals c. 1 mm. Petals c. 6 mm, c. 11/, mm wide at
the base, valvate at the base, conduplicate-valvate
for the rest, more or less connate in the median
part, mostly almost completely enveloping the
stamens. Filaments 11/,-2 mm long, 3/,-1 mm wide
at the base, glabrous. Connective pear- or drop-
shaped, c. 11/, mm, with dorsal swelling. Anther-
cells c. 1!/,mm, c. 1/,mm wide. Style 2-5 mm,
c. 1/,mm @, protruding from mature bud. Ovules
2 per locule. Capsule 2-3'/, by 11/,-21/, cm; rim of
enlarged receptacle infra-median ; valves 1/,—1(—11/,)
cm. Seeds 0.7-0.8(-1) by 0.3-0.4(-0.5) cm; wing
c. 1/, cm wide.

Distr. Malesia: Borneo. Fairly rare. Fig. 13.

Ecol. Primary (dipterocarp) forest at low and
medium altitude, up to 1200m, also recorded
from ultrabasic red-brown soil.

Field notes. Buttresses when present up to 1!/,-
2m vertically. Bark surface flaky. Stamens pale
yellow. Flowers greenish; corolla white.

Vern. Ubah, Iban, Sarawak, obah, Sabah.

SYMPLOCACEAE (H. P. Nooteboom, Leyden)!

The family consists of one genus only, Symplocos, which occurred already in the
Eocene over the entire northern hemisphere in the mixed mesophytic forest and in
all probability also in the Indo-Australian tropics.

As proved by abundant fossil endocarps, the Eocene species had already a fruit
structure very similar to that of now living species and the genus existed at that early
time obviously already in optima forma, a reason to assume that it must be of high
antiquity. This is also corroborated by the fact that the tropical subgenus Symplo-
cos has a very disjunct trans-Pacific range; explanation by chance transoceanic
long-distance dispersal must be refuted because it is in contradiction with all
presently known facts.

Although Symplocos has shown a fairly abundant speciation, considering its
present size and 25 fossil species described, it has surprisingly not led to other
generic development and remained in splendid isolation.

Its systematic affinities induced mostly to classify it with Ebenales. In my mono-
graph of the Old World species (1975) I have brought all evidence together and
have concluded that this position is unlikely: pollen structure differs from that in
other families of Ebenales, so do the stomata, the placentation and the structure of
the ovules. This leads to the view that Symplocos is more allied to Cornaceae and
Theaceae, sharing also with both families a primitive wood anatomy. Still the
affinity is not that close, as for example Theaceae have a truly axile placentation.
The chromosome number fits better with Cornaceae sens. lat.

SYMPLOCOS

Jacq. En. Fl. Carib. (1760) 5, 24; Select. Stirp. Am. Hist. (1763) 166, t. 175, f. 68;
Linnf, Gen. Pl. ed. 6 (1764) 272; Miers, J. Linn. Soc. Bot. 17 (1879) 285; BRAND,
Pfl. R. Heft 6 (1901) 13, 9 fig.; Noor. Leid. Bot. Ser. 1 (1975) 33, 7 fig., 21 pl., with
full synonymy. — Fig. 1-20.

For synonyms see under the subgenera.

Shrubs to (rarely) large, (in Mal.) evergreen trees; bark in various spp. bitter;
growth continuous or interrupted (in flushes), in the latter case the buds protected
by often leathery bud-scales; glabrous or hairy (by simple hairs). Leaves simple,
alternate or spirally arranged, rarely pseudoverticillate, estipulate, penninerved,
petioled, rarely almost sessile; when dry often discolouring (often in yellow tinges)in
subg. Hopea. Flowers in spikes, racemes, or panicles, mostly from the upper leaf-
axils, sometimes condensed to clusters, sometimes terminal or from the axils
of fallen leaves, rarely solitary; supported by a bract and 2 bracteoles, rarely
several bracts and bracteoles by abortion of flowers; flowers actinomorphic,
bisexual, rarely by reduction unisexual and plant polygamous, not rarely
fragrant, distinctly so in subg. Symplocos. Calyx with a very short tube above
the inferior ovary, the limb 3-5-lobed, imbricate, persistent, sometimes split into
two parts and seemingly 2-lobed. Corolla sympetalous, but divided nearly to the
base in subg. Hopea; lobes (3—)5(—10 in the New World), quincuncially imbricate,
whitish, bluish or purplish. Stamens 4 to mostly oo, connate in a long monadel-

(1) With co-operation of the General Editor.
(205)

206 FLORA MALESIANA [ser. I, vol. 8?

phous tube, at its base adnate to the corolla and very unequal, but in subg. Hopea
only connate at the very base, monadelphous or pentadelphous and then the
bundles alternipetalous; anthers globose, 2-celled, lengthwise dehiscent, introrse.
Ovary inferior (to + semi-inferior), 2—S-celled, with a complete septation; style 1,
stigma punctiform or peltate. Ovules 2-4 in each cell, pendulous, anatropous-
epitropous or amphitropous, unitegmic, tenuicellular. Drupe monopyrenous,
crowned by the persistent calyx lobes, of various shape: cylindrical to globose,
ampulliform or spindle-shaped; mesocarp usually thin, sometimes thick and then
often quite hard; stone smooth or mostly sculptured in various degree or length-
wise ridged. Seeds straight or curved, 1 in each developed cell, with copious
endosperm; embryo straight or curved, with very short linear cotyledons.

Distribution. About 250 spp., in the eastern parts of the Old World, from Ceylon and Bom-
bay in the Deccan to Fiji in West Polynesia and from Manchuria at 46° N as far as New South
Wales and Lord Howe I. at 32° S; in the New World from the State of Washington in the U.S.A.
to S. Brasil; throughout Malesia. Fig. 1.

>h° Bs

a

ze

Pliocene

Symplocos subg. Hopea AY) - :

a

oO) MIGCEne teense Symplocos subg. Symplocos
e Oligocene
+

Eocene

es 1.

————

Fig. 1. Range of the genus Symplocos, recent and fossil. The fossil localities in Europe, Japan, and E.
North America are all belonging to species of subg. Hopea.

There is no species common to the Old and New World, but the E. Asian S. lucida is closest
allied to the N. American S. tinctoria.

Taxonomy. BRAND (1901) has made an intricate subdivision of the genus, partly based on
former generic names. I believe we cannot go further than a subdivision into two subgenera, in
which macromorphology is supported by chemotaxonomy and palynology, viz subg. Symplocos
and subg. Hopea.

Subdivisions could be based on one important single character: straight versus curved embryo,
spiral versus distichous phyllotaxis, continuous versus flushwise growth from scaly buds, but it
appears that such subdivisions do not coincide. This leads to the view that there is a block of
species with reticulate affinities. This view also emerges from the palynological results.

1977] SYMPLOCACEAE (Nooteboom) 207

Both subgenera occur in the New and the Old World; subg. Symplocos, which is almost strictly
tropical, possesses only 2 spp. in Indo-Malesia, but probably many more in America.

In this revision 58 spp. are distinguished in Malesia; there are more new species, but I have
refrained from describing them as the material is incomplete; I have enumerated them in my
revision /.c. 296.

Fossils. Before the Glacial Epoch Symplocos occurred also in Europe in the mixed mesophytic
subtropical to warm-temperate forest, onwards of the Eocene, obviously as a common con-
stituent of the Tertiary mixed mesophytic forest, as shown from fossil stones. Cf. KIRCHHEIMER,
Palaeontographica 90B (1949) 1-52, t. 1-2. These stones are very similar to endocarps of recent
species; obviously no major changes did occur in the genus during this era. The three fossil
Pliocene species in Japan are almost certainly the same as those that are living there today. One
fossil species is known from the Eocene in the eastern U.S.A. Fig. 1.

Ecology. All species are evergreen, except a single deciduous one, S. paniculata (THUNB.)
Mia. from Kashmir to Manchuria and Japan.

They grow under tropical to temperate conditions in mixed evergreen rain-forest, not under
arid conditions.

Their stature is mostly small and they make part of the undergrowth and lower storeys, in
exceptional cases attaining a maximum height of c. 30m and 60cm @.

In Malesia they are found from sea-level up to the alpine zone at c. 4000 m (Mt Kinabalu; New
Guinea), where they are represented by mostly microphyllous (fig. 12) dwarf shrubs in the dense
elfin and mossy forest on slopes, summits and ridges where they may be common; but they are
almost nowhere recorded as a dominant.

A few species, e.g. S. polyandra, are restricted to the lowland, but most species have a fair
altitudinal range, and are most commonly collected in the hill and mountain forest. A few are
restricted to high altitude, e.g. S. buxifolia, S. deflexa, S. johniana, S. zizyphoides, and several
varieties of S. cochinchinensis.

A fair number seem to be rare and have been seldom collected, others are common and widely
distributed in the archipelago, notably S. cochinchinensis, S. celastrifolia, S. fasciculata, S.
laeteviridis, S. ophirensis, and S. odoratissima.

Especially these species, several of which are variable, grow on a variety of soils, including
young-volcanic; they are scarce on limestone and generally prefer more acid, humous soils, e.g.
S. celastrifolia is common in coastal forests, especially in the transition between mangroves and
freshwater swamps, but it occurs also on kerangas, along river banks, and even in peat swamp
forest.

S. cochinchinensis var. sessifolia is very resistant against poisonous crater gases and acid soil
conditions and can act as a pioneer in crater fields in Java, sometimes dwarfing down to very
small size, although still producing flower and fruit; in the surrounding closed elfin forest it is a
common small tree, growing together with Vaccinium, Myrica, Myrsine, Leptospermum, etc.

Density of species. In fig. 2 the density of species has been indicated for each province and island
(group). The richest areas are those of continental SE. Asia and West Malesia, while the number
of species tapers out towards East Malesia and the SW. Pacific. The greatest number of endemic
species is found in West Malesia, notably (as usually) in Borneo and the Philippines. However, in
East Malesia New Guinea has a fair number of endemic species. The high number of endemics in
New Caledonia is a bit exaggerating the situation as all are certainly derivatives of S. cochin-
chinensis. The same holds for the endemics of New Guinea (with the exception of S. cylindracea)
and for Australia (with the exception of S. cyanocarpa C. T. Wuire).

Flower biology. \n all Symplocos spp. the flowers of an inflorescence open almost simultaneously
and on one tree almost all inflorescences are open at the same time, so that the whole crown is
for a short time gay with the blossoms (fig. 3). Of S. cochinchinensis var. sessifolia flowers are
deliciously scented, as hawthorn, but field records mention other species as scentless or faintly
scented. This varies obviously with the species.

Pollination. Docrers VAN LEEUWEN (Verh. Kon. Ak. Wet. A’dam sect. 2, 31, 1933, 218)
reported of S. cochinchinensis var. sessifolia, on the summit of Mt Pangrango, West Java, at
c. 3000 m, that flowers expand in the morning but open only halfway, the corolla remaining bent
over the sexual organs; at 8 h. anthers are open and often touch the stigma on which the sticky

208 FLORA MALESIANA [ser. I, vol. 82

Fig. 2. Density of species in Old World Symplocos; above the hyphen the endemic species for each island
(group) or country, below the hyphen the non-endemic species.

Fig. 3. Symplocos laeteviridis STAPF var. laeteviridis in full flower, showing also alternate phyllotaxis.—
Sabah (Nooresoom 1017). Photogr. NooTEBOOM, Febr. 1969.

1977] SYMPLOCACEAE (Nooteboom) 209

pollen readily falls; on the 2nd flowering day the corolla is widely open, anthers are empty, and
the stigma is always pollinated. This means self-pollination. DocCTERS VAN LEEUWEN found,
however, also the flowers frequented by various insects, among them bees and bumble-bees. They
are not so much attracted by the little nectar, but are in search of pollen.

Some species may have locally a strict flowering time; e.g. S. cochinchinensis var. sessifolia
flowers, according to DOCTERS VAN LEEUWEN (i.c., fig. 52), from October to January, in the rainy
season, on the summit of Mt Pangrango, West Java.

Hybridization. Though there are in a few instances indications (by high sterile pollen °%) that
hybridization may occur, no clear cases are recorded. It is, however, clear that cross-fertilization
must occur in the polygamous species in East Malesia.

Galls. DOCTERS VAN LEEUWEN (Zoocecidia, 1926, 460) found in S. cochinchinensis var. sessi-
folia small leaf galls, caused by psyllids by which the two halves of the leaf curve upwards till
margins touch and a narrow cavity is formed. He recorded similar galls also from other forms of
this species. In S. fasciculata he found a stem gall caused by a gall-midge and in S. brandisii a
flower gall caused by a gall-midge.

Dispersal. RipLey (Disp. 1930) assumed that bats may be fond of the hard-fleshed drupes
(l.c. 347). He mentioned that in North America tyrant birds (Sayornis phoebe) eat amongst others
fruit of S. tinctoria (1.c. 483) and that in South America a curassow, a sort of turkey, would feed
on the fruit of S. cernua. DOCTERS VAN LEEUWEN (Verh. Kon. Ak. Wet. A’dam sect. 2, 31, 1933,
220) believed Symplocos to be dispersed by birds but did not find endocarps in the stomach of
fruit-eating birds. VAN STEENIS found fruit of S. henschelii abundant on the ground below trees at
Tjibodas, although this species has a fairly thick, hard-fleshed exocarp, in contrast to most
species in which the exocarp is thin. Also in fossils sometimes immense quantities of stones are
found together, about which KIRCHHEIMER reported (Palaeontographica 90B, 1949, 1-52): ina
total mass of c. 3500 m? he estimated the number of endocarps at some 2?/, billions. He assumed
that these were deposited within one century in a site of forest dominated by Symplocos. However,
he added that the layers in which the endocarps were deposited gave no evidence of rivers which
could have transported and accumulated the seeds and he concluded that they have dropped to
the soil in situ. For these reasons abundant dispersal by birds or bats is in Symplocos not very
likely.

Dispersal by water takes place in species in which some fruit cells are barren and remain empty,
e.g. S. celastrifolia.

Morphology. The phyllotaxis is variable but constant for the species; it is either spiral or
alternate (distichous) in which latter case the twigs are often zigzag (fig. 3).

In most species leaves are more or less equally dispersed along the twigs, but in other species
there is a tendency that the leaves are becoming crowded towards the end of the year’s growth,
e.g. in S. macrocarpa, as noted by TRIMEN (Handb. FI. Ceyl. 3, 1895, 103). In Malesian spp. this
occurs also in S. herzogii and S. gigantifolia where the large leaves occur crowded at the end of the
year’s growth.

There is a single species in which all the leaves are in real pseudo-whorls, viz S. verticillifolia
from the Philippine Is. (fig. 20).

The leader-shoots in Symplocos, e.g. S. fasciculata, have spiral phyllotaxis; such shoots may,
however, also carry flowers.

Rejuvenation is in certain species by continuous growth of the twig apex, as is e.g. charac-
teristic in S. fasciculata. In other species, however, there are clear buds with conspicuous bud-
scales, indicating that the growth mode is flushwise and discontinuous, as e.g. in S. costata and
S. lucida (fig. 15). This might be a good character of subdividing subg. Hopea. It can, however,
only be used if one has accurate knowledge of the rejuvenation process of each species. This is
sometimes difficult to ascertain from herbarium material as the bud-scales do not always leave
traces of distinct scars, field data hardly ever mention the character, and material is seldom
collected in the stage of flush. If the growth mode were well examined in all species I believe it
would represent a good key character.

Flushwise, discontinuous growth, with scaly buds could be assumed to be an adaptation to
seasonally cold climates. It is a life form intermediate halfway evergreen and deciduous. It is rare
in the Malesian tropics where it is known e.g. from Acer, some genera of Lauraceae, Fagaceae,

210 FLORA MALESIANA [ser. I, vol. 8?

which also in the tropics are found in the cool, tropical-montane climate, which is however hardly
seasonal. It still could be viewed as an indication of former immigration of taxa of higher latitude.
Once acquired this growth mode must then have been conserved, as it occurs also in S. barring-
toniifolia which is restricted to the tropical lowland.

The inflorescence is either a panicle or a raceme or spike. Morphologically it is cymose, the
flower always being sustained by two bracteoles which may at times carry abortive buds in their
axil (fig. 11b). In some cases the inflorescence is condensed to a fascicle or cluster of flowers
(fig. 20a) or even be reduced to a single flower (fig. 19a). In a few species flowers occur on old
wood, as e.g. in S. polyandra, S. wikstroemifolia (p.p.), S. rubiginosa, and S. tricoccata.

The flowers are bisexual but functionally unisexual flowers are found in several taxa, especially
in New Guinea. Such taxa are either dioecious or polygamous. In male flowers the style is small
and without a stigma, in female flowers the number of stamens is reduced (even to less than 10)
and anthers are sterile. In subg. Symplocos the stamens are monadelphous with a long tube
(fig. 6a, d); in subg. Hopea they are only connate at the base for at most 2 mm (fig. llc), and
intergrading from strictly monadelphous to strictly pentadelphous, the phalanges being alterni-
petalous.

In my revision it has been explained that, in contrast with former opinion, the ovary is initially
1-celled, with the ovules attached close to the centre on the induplicate part of the carpels, each
of the 2-5 compartments having usually 4 ovules; in fruit these appear as cells. In each developed
cell there is usually one seed. The latter and the embryo it contains may be curved or straight.
See fig. 4b, c, g, h, j, k.

Fig. 4. Symplocos ophirensis CLARKE ssp. perakensis (K. & G.) Noor. var. perakensis. a. LS of fruit, out of
centre, b. seed, c. curved embryo, with 2 short apical cotyledons, all x 4. — S. ophirensis CLARKE ssp.
cumingiana (BRAND) Noor. var. cumingiana. d. LS of fruit, seed cavity empty, e. stone, x 4. — S. macro-
phylla WALL. ex DC. ssp. cordifolia (THw.) Noor. var. apicalis (THw.) Noort. f. Ribbed stone, with fold,
g. seed, the curved embryo enveloped by the albumen, x 2. — S. paniculata (THUNB.) Mia. h. LS of seed
showing curved embryo, h’. ditto in CS, showing how such seed may appear deceptively as 2 seeds, x 11/5.
— S. glauca (THUNB.) Koipz. i. Fruit, j. seed in LS showing straight embryo, x 3. — S. paniculata
(THunB.) Mig. k. Germinating seedling with LS of endocarp and seed, showing mode of exist of embryo,
x 11/3, 1. seedling, x 2/,. — S. celastrifolia GRIFF. ex CLARKE. m. U-shaped seed, x 6 (a-c BuRKILL 1013,
d-e NooTeBooM 2229, f-g ASHTON 2480, k—/ after LuBspock). — alb albumen, ec stony endocarp, mc
mesocarp, os outer surface of fruit, sc seed cavity.

1977] SYMPLOCACEAE (Nooteboom) 211

The fruit is a drupe, with a fleshy, corky or woody mesocarp and a very hard stone (endocarp).
The endocarp may be smooth (fig. 10c, 19d) or show outside ridges or irregularities (fig. 4e, f, 9c,
10e, 14d); the same holds for the inside of the endocarp. In the centre of the copious endosperm
the embryo is embedded. It is slender and may be straight or curved. In the tropical subg.
Symplocos it is always straight. In subg. Hopea it is straight in all American spp. and in 80% of
the living species in the Old World and also in all fossil species in Europe. From this it is con-
cluded that a straight embryo seems to be the primitive state in the genus. Only the three Pliocene
fossil species of Japan, which can be matched with living species, have curved seeds and conse-
quently curved embryos.

Curved seeds occur in degree, they may be hook-shaped or U-shaped or even be twice curved
(S-shaped in S. brachybotrys). See fig. 4. This may give some difficulty in studying sections of the
stones to count the number of seeds in a fruit (e.g. fig. 4h-h’).

Although of the living species only 20% have curved seeds the vast majority of the individual
living plants have curved seeds; so it seems that this probably recent trend in the evolution of the
genus was successful although the reason for its origin and advantage of its function remains
obscure.

Seedlings. Few observations are made. LussBock (Contr. Knowl. Seedlings, 1892, 206-208,
fig. 509) noted for S. paniculata (sect. Hopea): the endocarp does not burst during germination;
the radicle emerges by a small hole at the apical narrow end; the hypocotyle elongates, becoming
curved, finally straightening, carrying up the endocarp containing the embryo. As the cotyledons
elongate, they push out at the small hole in the endocarp (so to say throw the latter off), and
finally get free and spread out to the light; they enlarge but remain narrow. The first two leaves
are opposite, hairy on both sides and serrulate which may persist in leaves of saplings (fig.
4l).

Spot-characters. In the herbarium a Symplocos of subg. Hopea can mostly easily be spotted by
spiral, exstipulate, eglandular, serrate or crenate leaves discolouring pale greenish or yellowish or
greenish-brown, a feature connected with a high Al-content of the tissues. At a very young stage,
the just expanding leaves have proportionally conspicuous gland-like teeth on the margin. A
significant character is that in the herbarium the midrib is always sulcate above, with the exception
of 4 spp. in which it is prominent: S. anomala, S. lancifolia, S. lucida, and S. wikstroemifolia.

The cup-like 3 bracts (of which 2 bracteoles) below the flower (fig. 11b) is also characteristic
as is the inferior ovary and fruit.

Innovations and newly expanded leaves are in many species a beautiful violet, afterwards
changing into violet-brown while the drupes are often blue to black-violet, features found in
many aluminium-accumulating plants (Eurya, Helicia, etc.).

Anatomy. For general surveys also covering the older literature, see SOLEREDER, Syst. Anat.
Dicot. Stuttgart (1899) 587-589 (under Styracaceae) and ibid. (1908) 208-210; METCALFE &
CHALK, Anat. Dicot. Oxford (1950) 890-893. Selected references: JANssONIUS, Mikr. 4 (1925)
471-498 (wood anatomy); DEN BerGeR, Determinatietabel Malesié, Veenman, Wageningen
(1949) (wood identification); JANssonius, Blumea 6 (1950) 422-423 & 424 (wood anatomical
affinities); Descu, Mal. For. Rec. 15 (1954) 591-593 (wood); ZAHUR, Mem. Cornell Univ.
Agric. Exp. Stn. 358 (1959) 35 (bark anatomy); Huser, Mitt. Bot. Staatssamml. Miinchen 5
(1963) 1-48; Baas, Blumea 21 (1973) 201-216 (ecological wood anatomy); Nootesoom, Leid.
Bot. Ser. 1 (1975) 20-22 (leaf and wood anatomy).

The wood is characterized by the following primitive set of characters: Vessels solitary and
with many-barred scalariform perforations. Fibre-tracheids with conspicuously bordered pits
on both radial and tangential walls. Parenchyma diffuse or diffuse-in-aggregates. Rays hetero-
geneous, usually of two distinct sizes. The bark is also of a primitive type with compound sieve
plates. Mechanical bark tissue is poorly developed and composed of groups of sclereids (ZAHUR,
Lc.). The leaf anatomy exhibits few constant characters such as paracytic stomata, clustered
crystals and dorsiventral mesophyll. Presence or absence of a hypodermis, of idioblastic leaf
sclereids, of a complex vasculation pattern in the midrib, and of an indumentum varies. The
diagnostic and systematic value of these characters remains to be assessed.

The anatomical evidence is inconclusive with respect to a positive indication of the closest
affinities of Symplocaceae. The traditional treatment of the family as a member of the Ebenales

212 FLORA MALESIANA [ser. I, vol. 8?

close to Styracaceae must, however, be refuted. The anatomy is more compatible with suggestions
of a Cornalean or a Thealean alliance as advocated by NOOTEBOOM I.c.

Palynology. The palynology of the Old World spp. was examined by R. VAN DER MEIJDEN
(Pollen et Spores 12, 1970, 513-551, 1971, suppl. in my Monograph, 1975, 9-15). The essential
results are the following: the two main pollen types coincide with the distinction of the two sub-
genera. In subg. Symplocos there are two minor types, one belonging to the Old World spp., the
other to those of America.

In subg. Hopea there are 9 subtypes, but none is apparently peculiar to American spp. The
distribution of these subtypes is rather complicated and leads to the view of reticulate relation-
ship, which agrees with the impression gained from macromorphology. Another feature is that
within the variable species several subtypes are represented, and furthermore that a number of
subtypes are found in species which are taxonomically not closely related. There is no agreement
between the shape of the embryo, straight or curved, and pollen subtypes. Echinate pollen is
found in the Philippine S. whitfordii and in the East Malesian and Pacific varieties of S. cochin-
chinensis ssp. leptophylla; also the 9 endemic spp. of Symplocos in New Caledonia which are all
related to this subspecies have echinate pollen.

In several taxa a certain amount of pollen is sterile and I have ascribed this to hybridization.

Phytochemistry. Many species of Symplocos, especially from subg. Hopea, contain alu-
minium compounds, a feature which manifestates itself in the yellow colour of dried leaves.
Especially when the plants are dried after having been conserved in alcohol vapour according to
the Schweinfurth method, the yellow colour becomes very intense. The yellow colour is the result
of a reaction of aluminium compounds with flavonols in the drying leaf. The amounts recorded in
literature vary between 0.05 and 4.2% of dry weight of the leaves; barks may contain similar
amounts of aluminium (CHENERY, Kew Bull. 1948, 173-183; Analyst, 1948, 501; NooTEBOoM,
Leid. Bot. Ser. 1, 1975, 19). RADLKOFER (Ber. Deut. Bot. Ges. 22, 1904, 216-224) already men-
tioned that the ash of Symplocos leaves contains c. 50% aluminium oxide. He also described the
so-called ‘‘Tonerdek6rper”’ in the leaves of Symplocos. These are masses of colourless material
filling often large parts of the cells, predominantly in the palissade parenchyma. According to
RADLKOFER these masses consist mainly of aluminium compounds. KRATZMANN (Sitz. Ber. Ak.
Wiss. Wien, 1913, 311-336) found that these aluminium bodies also contain much other material,
for instance silicates, and that the aluminium is also accumulated in other parts of the leaf. NEGER
(Flora N.F. 16, 1923, 326-330) observed that the development of plants of Symplocos lucida
(THuNB.) S. & Z. depends on the amount of aluminium compound in the solution they are culti-
vated on. Plants grew best on a solution containing 1 promille aluminium. Besides aluminium
many other compounds are found (HEGNAUER, Chemotaxonomy der Pflanzen 6, 1973). The
more important are: 1) Phenolic compounds (see also BATE SmiTH, J. Linn. Soc. Bot. 58, 1952,
95-173). Gallic and ellagic acid seem to be rather common. Leucoanthocyanins occur in varying
amount. BATE SMITH /.c. also found quercetin, and caffeic acid. In the bark of S. /ucida (THUNB.)
S. & Z. the lignan glycoside symplocosin has been found, and traces of methylsalicilate were
demonstrated in the bark of several species. True tannins were not yet found in Symplocos.
2) Alkaloids. Only for two species structurally known alkaloids were described. More research is
needed. 3) Saponins. In several species saponin-like compounds were found, as well in the bark
as in the leaves. — R. HEGNAUER.

Chromosomes. In my monograph I have given an account of chromosome numbers, which
are unfortunately too few. However, the majority is n = 11, with some deviations; rarely 2n =
24, and one count of the North American S. tinctoria of 1n = 14, all in subg. Hopea. The one
count known of subg. Symplocos in Malesia yielded 2n = c. 90 (2n = 88 would fit an octoploid).
It would be too rash to conclude that polyploidy would be normal in that subgenus.

On the other hand it may tentatively be concluded that species in subg. Hopea are diploid, with
x = 11. This does not fit the numbers found in other Ebenales families, nor in Theaceae, but it
does agree with Cornaceae sens. lat.

Uses. As timber Symplocos has no great value, according to HEYNE (Nutt. Pl. 1927, 1262).
Leaves and bark of Symplocos contain a fair amount of alum, both in Asian and American spp.
(cf. Ber. Deut. Bot. Ges. 22, 1904, 126). This was commonly used, mostly from decoctions of the
bark, in dyeing processes (red and brown), e.g. in the batik industry in Java. RuMpPuius already

1977] SYMPLOCACEAE (Nooteboom) 213

mentioned this use from the Moluccas. Several species were used for this purpose, e.g. S.
cochinchinensis, S. fasciculata, S. odoratissima (HEYNE, /.c.). The same compound is probably
also the constituent active in medicinal uses against so-called sprue (‘thrush’) as ‘obat seriawan’.

Notes. Identification of material of Symplocos is for several reasons far from easy. Because of
simultaneous flowering flowers and fruits are practically never found together and both stages
are properly needed. Only few species possess well definable vegetative characters. Moreover, a
few widespread species have proved to be rather variable, to a fair degree by racial differentiation.
These are the reasons that besides a general key in which all characters are used, I have found it
useful to add a number of partial keys for islands or island groups in a double series, either for
fruiting or for flowering material.

In fig. 5 a scheme is given elucidating the way in which for this genus descriptive terminology is
used in the keys and descriptions.

Fig. 5. Schemes elucidating descriptive terminology used in the text. — A: a length of leaf, 6 width of
leaf, c divided by b is leaf index, d length of acumen. — B: way of expressing base angle «. — C: deflorated
flower; br bract, bra bracteole, c length of calyx, c/ length of calyx lobes, ov height of ovary.

KEY TO THE SUBGENERA

1. Petals connate at least halfway up. Leaves usually not becoming yellow when drying, not discolouring,
spirally arranged, entire. Flowers very fragrant. Seeds and embryo straight. Spp. 1-2

1. subg. Symplocos

1. Petals connate only at the very base. Leaves usually becoming more or less yellow or greenish yellow

when dried. Leaves spirally arranged or distichous, exceptionally in pseudowhorls. Flowers not or

mostly only faintly fragrant. Seeds and embryo straight or curved. Spp.3-58 . . . . 2. subg. Hopea

1. Subgenus Symplocos

Cf. Noor. Leid. Bot. Ser. 1 (1975) 36. — Cordyloblaste Mor. Bot. Zeit. 6 (1848)
606; RipL. Fl. Mal. Pen. 2 (1923) 307; ALston, Handb. FI. Ceyl. 6 (Suppl.) (1931)
186. — Symplocos sect. Cordyloblaste B. & H. Gen. Pl. 2 (1876) 669; BRAND, Pfl.
R. Heft 6 (1901) 88; Steen. Bull. Bot. Gard. Btzg III, 17 (1948) 429. — Symplocos
subg. Cordyloblaste GAMBLE, J. As. Soc. Beng. 74, ii (1906) 248. — Fig. 6.
Leaves usually not becoming yellow when dry. Corolla tubular, erect, often to
above the middle adherent to the staminal tube and then suddenly expanded;

214 FLORA MALESIANA [ser. I, vol. 8?

margins of the petals free, thus sometimes obscuring the coalescence. Stamens
monadelphous; free part of filaments ribbon-shaped, in several whorls, in the outer
whorl often very short, always suddenly attenuate below the anther. Fruits 2-5-
celled, usually none of the cells aborted. Seeds straight, cylindrical.

Distr. Tropics of Indo-Malesia and South America, largely within 30° N and S, more than 100 spp.

described from the New World, in Malesia 2 spp. Fig. 1.
Ecol. Rain-forest, from the lowland up to c. 3300 m (Mt Kinabalu).

KEY TO THE SPECIES

1. Calyx c. 6(-10) mm long. Corolla 21/,-5 cm long. Fruits 3-Scmlong. ..... .
1. Calyx 3-5 mm long. Corolla */,-13/, cm long. Fruits 1-1'/, cm long

1. Symplocos henschelii (Mor.) BrH. ex CLARKE,
FI. Br. Ind. 3 (1882) 588, quoad nomen et basionym,
excl. stirp.; BRAND, Pfl. R. Heft 6 (1901) 89; Bull.
Herb. Boiss. II, 6 (1906) 750; Koorp. Atlas 2 (1914)
t. 390; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948)
440, f. 2a—l; Nova Guinea n.s. 10 (1959) 210; Back.
& BAKH. f. Fl. Java 2 (1965) 204; STEEN. Mt. FI.
Java (1972) pl. 52-3; Noor. Leid. Bot. Ser. 1
(1975) 37, pl. 1g. — Cordyloblaste henscheli Mor.
Bot. Zeit. 6 (1848) 606. — Eugeniodes henscheli
O. K. Rev. Gen. Pl. 2 (1891) 975. — S. nageli
K. & V. Bijdr. 7 (1900) 159. — S. scortechinii KING
& GAMBLE, J. As. Soc. Beng. 74, ii (1906) 250. —
Cordyloblaste scortechinii Rip. Fl. Mal. Pen. 2
(1923) 309. — S. dolichantha MeErR. Sar. Mus. J. 3
(1928) 545. — S. stenosepala STEEN. Bull. Bot.
Gard. Btzg III, 17 (1948) 444, f. 2 m—n. — Fig. 6a—c.

For further synonyms see under the variety.

Shrub, or mostly a tree, to 30 m; innovations
glabrous to grey or rusty velvety. Leaves glabrous,
sometimes the midrib above and underside hairy,
7-17(-22) by 3-7!/,cm; petiole */,-11/,(—-2) cm.
Racemes up to 10 cm, incl. bracts and flowers grey
or rusty tomentose, short-peduncled, 1-12-
flowered. Bracts narrow-triangular; pedicels 0-6
mm, with 2(-3) tiny bracteoles. Calyx lobes
rounded to triangular, mostly erect, 1—4'/, by
2-3 mm, persistent. Corolla sericeous (in Mal.),
club-shaped in bud, 2!/,-5 cm, connate for 3/,—%/,,
tube 3-4 mm @, lobes spathulate. Staminal tube
1/,cm shorter than corolla, adnate to the corolla
tube except towards apex, free part 1/,-1'/, cm;
anthers 20-110, filaments unequal. Ovary 3-4-
celled; ovules 2-4 per cell, usually only 1 develop-
ing. Fruit obovoid to spindle-shaped, 3-5 by 2-3
cm; mesocarp thick, hard-fleshy to + woody.

Distr. Continental SE. Asia (Burma, Thailand,
Indo-China) and West Malesia (Sumatra, Malay
Peninsula, W. Java, Borneo), a distinct subspecies
in Thailand.

Note. Additional material has shown that
S. stenosepala STEEN. cannot be upheld and, more-
over, that S. maingayi CLARKE deserves only
varietal rank.

KEY TO THE VARIETIES

1. Leaves and twig ends usually glabrous. Free
part of staminal tube 7-15 mm.

a. var. henschelii

1. Twig ends and leaves underneath hairy. Free

part of staminal tube 5-7 mm b. var. maingayi

1. S. henschelii
2. S. pendula

a. var. henschelii. — Fig. 6a—b.

Shrub or tree, up to 25 m, 45 cm @. Twigs glab-
rous, the youngest ones sometimes more or less
grey or rufescent appressedly pubescent to velvety
or tomentose. Leaves glabrous, or the midrib
beneath sparsely short fine-hairy, rarely with same
indument as var. maingayi. Free part of staminal
tube 7-15 mm; anthers (40—)55—-75(—110), in the
upper 5-10mm, ascendent and nearly sessile
above to descendent on a slender filament below,
the lowest ones hanging from a 2-5 mm long fila-
ment. Fruit with + fleshy mesocarp.

Distr. As the species.

Ecol. Below 1100 min mixed dipterocarp forest,
also once in swamp forest, and on podsol (Kalabit),
at higher altitude in oak-chestnut mountain forest,
also on ridges and in mossy forest, 600-2000 m (in
continental SE. Asia at 130-800 m). F/. Jan.—Dec.,
fr. Febr.—Sept.

Vern. Sumatra: kayu djaram-djaram bosi,
Batak; Borneo: té baradang, Sarawak, Kalabit,
yum, Kenyah lang., /amau-lamau, Brunei.

b. var. maingayi (CLARKE) Noor. Leid. Bot. Ser. 1
(1975) 39. — S. maingayi BTH. ex CLARKE, FI. Br.
Ind. 3 (1882) 588; BRAND, Pfl. R. Heft 6 (1901) 90;
K. & G. J. As. Soc. Beng. 72, ii (1906) 249; STEEN.
Bull. Bot. Gard. Btzg III, 17 (1948) 445. — Euge-
niodes maingayi O. K. Rev. Gen. Pl. 2 (1891) 975.
— Cordyloblaste maingayi Rip. Fl. Mal. Pen. 2
(1923) 309. — Fig. 6c.

Tree up to 21 m, 40cm @. Twigs densely rusty
tomentose or velvety, glabrescent. Leaves sparsely
fine-hairy beneath, especially on midrib and nerves,
to greyish tomentose or velvety. Free part of
staminal tube 5-7 mm; anthers 20-60, in the upper
5 mm, ona very short (!/,—1/, mm) thin free part of
the filaments. Fruit with -- woody mesocarp.

Distr. Malesia: Malay Peninsula and Borneo
(Sarawak, Brunei).

Ecol. Evergreen primary and depleted lowland
forest, 15-150 m; in Borneo often on low sandy
ridges, raised beaches, and large sandy podsols
(kerangas). F/. April-May, fr. Jan.

2. Symplocos pendula WiGuHT, Ic. 4 (1848) 10,
t. 1237; Ill. Ind. Bot. 2 (1850) t. 151-b, 7-12;
CLARKE, FI. Br. Ind. 3 (1882) 587; BRAND, Pfl. R.
Heft 6 (1901) 88; STEEN. Bull. Bot. Gard. Btzg III,
17 (1948) 437; Noor. Leid. Bot. Ser. 1 (1975) 40,
pl. lh. — S. scortechinii (non K. & G.) RIDL. J.
Linn. Soc. Bot. 38 (1908) 315. — S. pulcherrima

1977]

SYMPLOCACEAE (Nooteboom)

Fig. 6. Symplocos henschelii (Mor.) BTH. ex CLARKE var. henschelii. a. Flower, b. fruit, exocarp halved. —

S. henschelii var. maingayi (CLARKE) Noor. c. Fruit. — S. pendula WiGut var. pendula. d. Flower. —

S. pendula var. hirtistylis (CLARKE) Noor. e. Fruit. All nat. size (@ WILSON 2547, 6 after STEEN. 1972, pl.
52-3b, c KOSTERMANS 9328, d father ANGLADE s.n., e MEER 3618).

Ript. J. Fed. Mal. St. Mus. 6 (1915) 160. —
Cordyloblaste pulcherrima Rip. Fl. Mal. Pen. 2
(1923) 308. — Fig. 6d-e.

For further synonyms see under the variety.

var. pendula. — Fig. 6d.

Small shrub '/,—-3 m or tree up to 27 mand 50 cm
@. Twigs glabrous to rusty tomentose. Leaves
glabrous or nearly so, elliptic to obovate or orbicu-
lar, entire to crenate, apex rounded to acuminate,
(1-)2#/,-12'/, by (1-)1'/.-6cm; nerves 4-8(-11)
pairs; petiole (1-)5-15 mm. Racemes very short,
sometimes flowers solitary. Bracts to 1 mm. Brac-
teoles 2-4, narrow-triangular. Pedicels 0-5 mm,
longer in solitary flowers. Calyx lobes very short
and rounded, ciliate. Corolla tubular-trumpet-
shaped, (5-)10-17 mm, fleshy, silver-white to
creamy, fragrant, the petals connate halfway up,
spathulate, rounded at apex, glabrous to tomentose.
Staminal tube adnate to corolla except for upper
3-5 mm, hairy to glabrous inside; anthers 30-50
(-80). Ovary semi-inferior, glabrous, the apex
semi-globose, c. 1'/, mm high, densely grey-hairy;
style c. 1 cm, more or less hairy at the base to

brous at the apex. Fruits spindle-shaped, 10-15

y 3-6cm, green pinkish red, the enlarged calyx
lobes surrounding the hairy, conical, persistent
style-base.

Distr. Continental SE. Asia (Ceylon, Deccan,
Hainan), in Malesia: Malay Peninsula.

Ecol. Mountain forests and open heath and
scrub, often on ridges, 600-1750 m. Fl. Febr.-
March, /r. Sept.

var. hirtistylis (CLARKE) Noor. Leid. Bot. Ser. |

(1975) 42, f. 2a, with full synonymy. — S. henschelii
(non BTH.) CLARKE, FI. Br. Ind. 3 (1882) 588, pro
stirp., incl. var. hirtistylis CLARKE. — S. confusa
BRAND, Pfl. R. Heft 6 (1901) 88; Bull. Herb. Boiss.
II, 6 (1906) 750; K. & G. J. As. Soc. Beng. 74, ii
(1906) 248; BRAND, Philip. J. Sc. 3 (190%) Bot. 3;
Me_rr. En. Philip. 3 (1923) 297; Steen. J. Arn.
Arb. 28 (1947) 423; Bull. Bot. Gard. Btzg III, 17
(1948) 432. — S. albifrons BRAND, Pfi. R. Heft 6
(1901) 88; Bull. Herb. Boiss. II, 6 (1906) 750; Nova
Guinea 14 (1924) 189. — S. capitellata BRAND,
Pfi. R. Heft 6 (1901) 88; Bull. Herb. Boiss. II, 6
(1906) 750; Nova Guinea 14 (1924) 188. — S. fox-
worthyi BRAND, Philip. J. Sc. 3 (1908) Bot. 3;
Merr. En. Philip. 3 (1923) 299. — Styrax obovatus
Rip. J. Str. Br. R. As. Soc. m. 61 (1912) 8. —
S. obovata Rio. J. Fed. Mal. St. Mus. 6 (1915) 51.
— §. crenulata Riwv. l.c. — S. novoguineensis
Gisss, Arfak (1917) 176. — Cordyloblaste obovata
Rip. Fl. Mal. Pen. 2 (1923) 308. — Cordyloblaste
crenulata Riwi. lc. 309. — S. atrata BRAND,
Nova Guinea 14 (1924) 188. — S. topica BRAND,
l.c. 189. — Fig. 6e.

Ovary hairy.
Distr. Continental SE. Asia (N. Burma, Indo-
China, China, Japan, Formosa), throughout

Malesia, except Java and Lesser Sunda Is.

Ecol. Primary and secondary montane and sub-
alpine forest, mossy forest, often common on
ridges, or in open fern thickets (Tamrau), on sand
or clay, 1500-3300 m, but in kerangas forest in
Sarawak at 800 m. Fi. March—Aug. (Sept.—Febr.),
Sr. Febr.—April, July-Sept.

At higher altitude often a dwarf shrub with small
leaves, but sometimes also a dwarf shrub with large
leaves in high forest.

2. Subgenus Hopea

CLARKE, FI. Br. Ind. 3 (1882) 572; BRAND, Pfl. R. Heft 6 (1901) 25; Noor. Leid.
Bot. Ser. 1 (1975) 43, with full synonymy. — Hopea Linné, Mant. (1767) 105, nom.

216 FLORA MALESIANA [ser. I, vol. 8?

a
000 0486000
3 3 4 8 9 10 10 1a 12

a \

‘ 65° 0000086

14 16-1a 16-1b 16-1C 16-1d 16-2e 16-3f 16-4g 16—4g

OHo00008

16-49 16-49 16-49 16-4h 116-41 16-41 16-4m 16-4mM 16-4nN 16-40

605608 0 4 6 8 o

16-4p 16-4Q 16-4rF 16-4F 16—4S 16-4t 16-4U 16-4U 16-4W

0 9
00.0()¢

20C
24 27 28 30 31 32 33a 33eE 34 37
40 41a 41b

42-1a 42-3f 42-3f 42-3g 42-2Cc 43

0000500

Fig. 7. Fruits in outline, in the dried state. Of each fruit the voucher specimen is cited by the number of the
taxon. If for showing variability more fruits of the same taxon are drawn, read from left to right corres-
ponding with the voucher numbers. All drawings natural size. — 3 CHEw WEE LEK 938 — 3 CF 104879 —
4 SAN 56690 — 5 DE Wipe 13773 — 6 A. Ernst 736 — 7 KING’s Coll. 6179 — 8 MEUER 7581 — 9

45

eos

1977] SYMPLOCACEAE (Nooteboom) 217

rejic. — Dicalix Lour. Fl. Coch. 1 (1790) 663; BL. Bijdr. (1826) 1116 (‘Dicalyx’). —
Sariava REINW. Syll. Ratisb. 2 (1825) 12. — Carlea Pr. Epim. Bot. (1851) 216. —
Baranda Luianos, Mem. Ac. Cienz. Madrid 3, 2 (1857) 502. — Eugeniodes O. K.
Rev. Gen. Pl. 2 (1891) 409, 975, nom. illeg. — Fig. 7-20.

Leaves usually becoming more or less yellow when drying. Petals glabrous, or
hairy in only few species, connate only at the very base, mostly expanded. Stamens
monadelphous to pentadelphous, only connate at the very base (for at most 2 mm);
filaments cylindrical, slender to rather stiff, often gradually attenuate towards the
anther. Fruits 2—3(—5?)-celled, often 1-celled by abortion. Seeds either straight or
curved, and then with curved embryo.

Distr. About 150 spp., as for the genus. Fig. 1.

Note. As explained in the note under the genus, a general overall key is given to all species, as much as
possible based on vegetative characters and on flowering material.

To facilitate identification additional local keys are given for the main Malesian islands or island groups,
one each for flowering and for fruiting material.

In addition in fig. 7 fruits are drawn of all species as far as available in the dried state. They have been
ate thor according to the number of the taxa. The following terminology has been adopted for fruit-
shapes:

globose 34, 41b ampulliform 23, 42c, 43
ellipsoid 4, 19, 33e, 46 spindle-shaped 38: fig. 19d.
ovoid 48 cylindrical 20c, 21

obovoid la, 1b: fig. 6c

It should be observed that the shape of the stone may differ from the shape of the fruit and that for
instance ovoid fruits may possess an ampulliform stone.

There is no strict relation between the shape of the seed and the shape of the fruit or stone, but ampulli-
form fruits have always a curved seed and curved embryo and spindle-shaped and cylindrical fruits have
always a straight seed and embryo.

Besides the overall-shape of the drupe, the shape of the stone can be important: sometimes it bears
lower or higher ridges, which ornamentation provides good characters.

KEY TO THE SPECIES

1. Leaves (pseudo-)verticillate.
Zaeerpes Mae OF Jeaves glabrous, Twigs hirsute. . .. . % \ ©. °WeU ee 55. S. verticillifolia
eee mice Of leaves'hairy. Twigs tomentose ... . . » » » «wemplue eeuuele 30. S. herzogii
1. Leaves not verticillate.
3. Midrib prominent on the upper surface.
4. Twigs glabrous.
5. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath
37. S. wikstroemifolia
in menves evenly distributed, glabrous... . «).6) sss 50) eye ee 35. S. lucida
4. Twigs hairy.

HALuieR f. 2197 — 10 CLemens 32525 — 10 CLEMENS 32478 — lla bb 23324 — 12 SAN 46543 — 14
KoOSsTERMANS 9158 — 15a KOSTERMANS & ANTA 527 — 16—la Forbes 861 — 16—1b MEWER 1690 — 16-Ic
CLEMENS 17224 — 16-1d BS 4476 — 16-2e LARSEN c.s. 887 — 16-3f NGF 33643 — 16-4g ANU 2027 —
16-4g A. C. Smitu 1054 — 16-4g BW 4970 — 16—4g GILLespie 3918 — 16—-4g NGF 28481 — 16—4h VINK
17308 — 16—4i Brass 28343 — 16-41 Brass 29919 — 16-4m LEDERMANN 8946 — 16-4m T. G. HARTLEY
13135 — 16-4n PULLEN 479 — 16-40 NGF 49168 — 16-4p PULLEN 7783 — 16-4q KosTERMANS &
WIRAWAN 878 — 16—4r NICOLAS 19 — 16—4r VAN BALGOoy 862 — 16-4s KALKMAN 5128 — 16-4t VINK
16079 — 16-4u KosterMAns 2375 — 16~—4u Forses P. P. 652 — 16-4w @rass 28191 — 16-4w CLEMENS
1661 — 16-4x NGF 23728 — 17 CLemens 33706 — 19 Koorpers 15596 — 20c VAN BEUSEKOM c.s. 837—
20d ENpert 2580 — 20f CF 97832 — 21 bb 22503 — 22 Jacoss 5766 — 23 SAN A2240 — 24 MERRILL
6148 — 27 BS 45592 — 28 KEP/FRI 8236 — 30 T. G. Hart.Ley 12509 — 31 Noorrenoom & ABAN 1500
— 32 HILDEBRAND 55 — 33a SAN 65017 — 33e SAN 44386 — 34 PNH 18483 — 35 BURGER s.n. — 36
Cel. 11-374 — 37 F. C. How 73506 — 40 BS 26447 — 4la Ja 7723 — 41b SAN 57045 — 42-la BurRN
Murpocnu 340 — 42-3f Dinc Hou 274 — 42-3f Nooresoom 2229 — 42-3g BS 83753 — 42-2c CF 98890
— 43 Kasewsx1 1208 — 44 § 17287 — 45 Carr 12782 — 46 Rip_ey 16102 — 47a ROBINSON & KLoss 199
— 47b Mever 7665 — 48 Ismaet 9 — 50 Beccari P. S. 106 — 51 Koeiz 29538 — 52 CLemMeNns 32559 —53
S 26305 — 55 PNH 14397 — 56 BS 45675 or 45775 — 57 Jacons 7484 — 58 Nooresoom 1491,

218 FLORA MALESIANA [ser. I, vol. 8?

6. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath
37. S. wikstroemifolia
6. Leaves evenly distributed, glabrous or sparsely fine hairy beneath.

7. Underside of leaves glabrous. Corolla:4—6 nm: j x5... - ee ea 4. S. anomala
7, Underside o1 leaves hairy: ‘Corolla 2*/-4 mm iF 2 ek kt ls . 34. S. lancifolia

3. Midrib sulcate above.
Pe Gea len flee la ieee gwen eee Oo ee ee re oe 41. S. odoratissima

8. Corolla glabrous.
9. Twigs hairy.
10. Underside of leaves glabrous. (When petiole and leaf margin beset with closely spaced vesicular
glands: 3. S. adenophylla).
fiaideaves:disuchonus Viluitioee fie. C2. 2S SL WeDo A ZEB 33. S. laeteviridis
11. Leaves spirally arranged.
12. Calyx and ovary glabrous.

13. Petole:0-5:tmim:, Fae. ser oie a. ca) ON a 16-4. S. cochinchinensis ssp. leptophylla
13. Petiole more than 5 mm.
14. Leaves shorter thanScm .......... 16-4. S. cochinchinensis ssp. leptophylla
14. Leaves longer than 5 cm.
1 DEY. Ss ck © de rentaien eee be 16-4. S. cochinchinensis ssp. leptophylla

15. Disk glabrous.

16. Twigs (appressedly) pubescent, puberulous or pilose. Seeds not straight.

16. S. cochinchinensis
16. Twigs tomentose or tomentellous.
17. Petiole 12-17 mm. Acumen 2-7 mm long. Nerves 8-12 pairs. Fruits more than 10 mm
long ede cme 8A cantina fhoue) 2 os cece > gh ee 5. S. atjehensis
17. Petiole 5-12 mm. Acumen longer than 7 mm. Nerves 10-16 pairs. Fruits to c. 10mm long.
28. S. glomerata
12. Calyx and/or ovary hairy.
18. Leaves crowded towards the end of the twigs, the latter tapering off towards the apex.

S. polyandra
18. Leaves evenly distributed, twigs not obviously tapering off.
19. Ovary glabrous.
20-“Inflorescence only A=flowered”) = 65 FP SONS ee: 38. S. multibracteata
20. Inflorescence more-flowered.
PPebiskinaryer is ot ete GO, oR 16-4. S. cochinchinensis ssp. leptophylla

21. Disk glabrous.
22. Seed and embryo uncinately curved towards the base
16-4. S. cochinchinensis ssp. leptophylla

22. Seed and embryo twice curved .. . . 16-1. S. cochinchinensis ssp. cochinchinensis
19. Ovary hairy.
DACA E SLA DTOUS gy ets 0d eS) sg) ihe ae 16-4. S. cochinchinensis ssp. leptophylla
23. Calyx hairy.

24. Bracts caducous.
25. Inflorescence an often branched raceme to 4 cm. Calyx 1-2 mm long 47. S. robinsonii

25. Inflorescence a 1—3-flowered short spike. Calyxc.3mm ... . 10. S. brachybotrys
24. Bracts persistent.
262 Penole 055 maven. 2 OR) ara. 16-4. S. cochinchinensis ssp. leptophylla
26. Petiole more than 5 mm.
213: BECKS SUBICNE SS 5c FRO a Bok ee ee ee 24. S. filipes
D272 Seeds riot 'straignt: | ors ee a. ee 16-4. S. cochinchinensis ssp. leptophylla

10. Underside of leaves hairy.
28. Leaves distichous.
29. Nerves up to 6 pairs.

30; Argie:of leaf base more than 00? 0) 29% £00 2s Ae, Bee © 31. S. johniana
30. Angle of leaf base less than 90°.
31. Dek Gatiious 25-612. 2,06 wench) ghd TER Wa ate 52. S. trichomarginalis
31 Diskhatty——- £28 aged fe! SS eee IS no Ee 33. S. laeteviridis

29. Nerves (5-)6 pairs or more.
32. Leaves longer than 5 cm (mean length).
33. Flowers c. 3 in an up to 3 cm long lax raceme. Fruits 10-14 mm long. Stamens c. 90 or more.
17. S. colombonensis
33. Inflorescence usually different. Fruits to c. 12 mm long. Stamens c. 70 or less.

34. Inflorescence a fascicle. Bracts to c. 1 mm long, persistent, bracteoles persistent. Ovary
c. 1 mm high, calyx c. 1 mm long, lobes not becoming longer by tearing. Corolla c. 2-4"/,
min. Style base hairy. Fruits: ampulliform” 2A +. 0 See 23. S. fasciculata

34. Inflorescence not a fascicle. Bracts longer than 1 mm, caducous, bracteoles caducous.
Ovary more than 1 mm high, calyx longer than 1 mm, lobes becoming longer by tearing.
Corolla more than 4 mm long. Style base glabrous. Fruits ovoid to ellipsoid

33. S. laeteviridis

1977] SYMPLOCACEAE (Nooteboom) 219

32. Leaves shorter than 5 cm.
35. Inflorescence only 1-flowered.
36. Reticulation not prominent. Ovary c. 1 mm high, calyx longer than 2 mm, lobes c. 3 mm.

Bracts several. Corolla c.4 mm. Fruits 8-9 mm long . . 252. S. trichomarginalis
36. Reticulation present beneath. Ovary 1—11/, mm high, calyx c. 2mm long, lobes 1—1!/, mm
long. Bract 1. Corolla 4-6 mm long. Fruits 10-12mmlong .... 58. S. zizyphoides
35. Inflorescence more-flowered.

Beeracts.and bracteoles caducous . .. . . «asgte Bodeeeee avo: 33. S. laeteviridis

37. Bracts and bracteoles persistent.
ae narys hrmsisc:..10. by 5 mm ..< \3 eae. Boe Pee ee eee se 22. S. deflexa
38. Disk glabrous. Fruits 10-12 by 5-6mm. .........+... 58. S. zizyphoides

28. Leaves spirally arranged.
39. Upper side of leaves hairy.

feerenele of leaf base more than 90°. . . 26. 3/3 sche ieee 13. S. calycodactylos
40. Angle of leaf base less than 90°.
41. Leaf margin (and petiole) beset with closely spaced glands. ..... 3. S. adenophylla
41. Leaf margin (and petiole) often glandular but glands not closely spaced.
42. Ovary hairy. Fr. cylindrical, 13-18 by 3-5 mm. Embryo straight. . . . 20. S. crassipes

42. Ovary glabrous. Fr. ampulliform, 6 by 4 mm. Embryo twice curved 43. S. paucistaminea
39. Upper side of leaves glabrous.
43. Calyx and ovary glabrous.
44. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath
37. S. wikstroemifolia
44. Leaves evenly distributed, glabrous or longer hairs beneath.
45. Seeds straight

46. Leaf index 2-3. Fruits less than20mmlong............ 5. S. atjehensis
46. Leaf index more than 3. Fruits longer than 20 mm.
Ari ereesiicss than 1@ipans -. . . .-ey oe De eee 15. S. cerasifolia
a9." Nexves more‘than 10 pairs ~~... . . Se 15b. S. cerasifolia var. grandifolia
meemerais tie Stone ee 16-4. S. cochinchinensis ssp. leptophylla
43. Calyx and/or ovary hairy.
48. Leaves crowded towards the end of the twigs ......... 37. S. wikstroemifolia

48. Leaves evenly distributed.
49. Ovary glabrous.

50. Disk hairy.
51. Inflorescence only 1-flowered. Seeds straight. ........ 38. S. multibracteata
51. Inflorescence more-flowered. Seeds not straight.
we, Pee O-) WIT ss ck aur pk ee 34. S. lancifolia
52. Petiole morethanSmm......... 16—4. S. cochinchinensis ssp. leptophylla
50. Disk glabrous.
BE emy toc, tenon =... ws 16-1. S. cochinchinensis ssp. cochinchinensis

53. Ovary more than | mm high.
54. Twigs (appressedly) pubescent, puberulous or pilose.
16-4. S. cochinchinensis ssp. leptophylla
54. Twigs not appressedly pubescent or puberulous.
55. Leaf index 2-3. Bracts persistent, shorter than 3 mm, bracteoles persistent. Calyx
lobes not becoming longer by tearing. Corolla shorter than c. 4 mm. Stamens less than
30. Fruits to c. 10 mm long. Mesocarp fleshy (shrivelled when dry). Seeds not straight.
16-4. S. cochinchinensis ssp. leptophylla
55. Leaf index more than 3. Bracts caducous, longer than 3 mm, bracteoles caducous.
Calyx lobes becoming longer by tearing. Corolla c. 5mm long. Stamens more than
30. Fruits more than 20 mm long. Mesocarp woody or corky. Seeds straight
15. S. cerasifolia
49. Ovary hairy

56. Calyx ibrous 6. te.0\6. of tense Tele 16-4. S. cochinchinensis ssp. leptophylla
56. Calyx hairy.
57. Bracts caducous.
58. Leaves longer than 15 cm.
59. Fruits more than 10 mm long, 2-5-celled. Mesocarp woody or corky. Stone with high
lengthwise not interrupted ridges. Seeds straight . 15b. S. cerasifolia var. grandifolia
59. Fruits to c. 10 mm long, 1-celled. Mesocarp thin, friable in dry state. Stone with a
transverse constriction at one side. Seeds not straight .... . 48. S. rubiginosa
58. Leaves shorter than c. 15 cm.
60. Calyx lobes longer than 1'/, mm. Style base hairy.

61.) Leaves aaprter than 5 ott «a: 05) aoe ees 10. S. brachybotrys
61. Leaves longer than 5 cm.
Gz. Tloreacenes @ Teeth os Seve: eR we 2 50. S. sumatrana

62. Inflorescence a 1-3-flowered short spike or a spike to 4.cm.

eS

220 FLORA MALESIANA [ser. I, vol. 8?

63. Petiole 3-4 mm. Ovary c. 1 mm high, calyx c. 3 mm, lobes longer than 2!/, mm.
Ovary (appressedly) pubescent. Disk inconspicuous. Fruits c. 5mm _ broad,
(-celled?Scedsinotistraisht] Were 2) aes OS CO: 10. S. brachybotrys

63. Petiole 5-7 mm. Ovary c. 1!/, mm high, calyx 2 mm long, lobes 1'/,-21/, mm long.
Ovary sericeous. Disk clearly present. Fruits more than 5mm broad, 3-celled.

Seeds isttaiehias 51-341. 4 BO Se eee era . 1. AA 50. S. sumatrana
60. Calyx lobes */,-1*/, mm long. Style base ele
64. Nerves more than 10 pairs ..... 1... 6-2. 5 5. 47. S. robinsonii
64. Nerves less that 10 pairs.
65. Ovary c. 1 mm high, lobes triangular. . 42-1b. S. ophirensis var. densireticulata
65. Ovary 1-2 mm high, lobes not triangular. ......... 47. S. robinsonii

57. Bracts persistent.

66. Inflorescence only 1-flowered.
Gi-wAngle ofleafibase.more than\90°° >... 33. ts SO See ee. ee 31. S. johniana
67. Angle of leaf base less than 90°.
68. Leaf index 4-7. Ovary c. 1'/,mm high. Stamens less than 30. Disk hairy, incon-

spictousastone:smoothiad tstabsale nehOoie toe ee 49. S. salicioides
68. Leaf index 1.3-4. Ovary to c. 1 mm high. Stamens more than 30. Disk glabrous,
clearly present. Stone with ridges or grooves ........ 20. S. crassipes

66. Inflorescence more-flowered.
69. Seeds straight.
70. Bracts to c. 1 mm long.
71. Ovary to c. 1 mm high.
72. Reticulation fine. Calyx longer than 1 mm. Inflorescence a much reduced often

clustered spike. Fruits not ampulliform, 13-18 mmlong . . . 20. S. crassipes
72. Reticulation coarse. Calyx c. 1 mm long. Inflorescence a fascicle. Fruits ampulli-
form; 5—7'mmlongee sash wean aed «fnew: scien aaa 23. S. fasciculata
71. Ovary more than 1 mm high.
73. Leaf index more than 3. Calyx lobes longer than!/,mm. . . 3. S. adenophylla
13- seat index. 2-3 Calyx lobes toc: 4/,mm jong, -.5)2.. . 3 cee 24. S. filipes
70. Bracts longer than 1 mm.

(4-eAnelejof leaf base more than90>, 405 teeter sa Ga ee 20. S. crassipes

74. Angle of leaf base less than 90°.
75. Underside of leaves especially hairy onthe margin. .... . 20. S. crassipes

75. Underside of leaves not especially hairy on the margin.
76. Calyx lobes longer than 1!/, mm. Disk hairy. Stamens more wie Pat Style base

hairy: Eruits’more than (Ommilons ~ . 3). . . . . se . S. crassipes
76. Calyx lobes 1-1!/, mm long. Disk glabrous. Stamens less than 0. Style base
glabrous se ruts.cs1 0mm Jong). |. .2 Sy tay ee ee 56. S. vidalii
69. Seeds not straight.
Ue ISkeClADTOUS. /~ an ope h od Gees 2 os bw ae 16-4. S. cochinchinensis ssp. leptophylla
77. Disk hairy.
Peer secdtandlembryOsU-snapeG a a seg spe, es 34. S. lancifolia

78. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla
9. Twigs glabrous.
79. Underside of leaves hairy.
80. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath
37. S. wikstroemifolia
80. Leaves evenly distributed.
81. Calyx and ovary glabrous.
2) Diskhiaye ss. £0 fo BOTS, BALES NO Wom, 16-4. S. cochinchinensis ssp. leptophylla
82. Disk glabrous.
83. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla

83. Seed and embryo not uncinately curved towards the base ...... 5. S. atjehensis
81. Calyx and/or ovary hairy.

84." Weaves. distichotis < Pain, «ag oT ahs YS Pia. a. 5 ee 33. S. laeteviridis
84. Leaves spirally arranged.

$5. Leaves shorter than Ss cat “4 4a oe ee. dun 2 eel sie ee 10. S. brachybotrys

85. Leaves longer than 5 cm.
86. Calyx glabrous.
87. Calyx lobes becoming longer by tearing. Seeds straight ....... 20. S. crassipes
87. Calyx lobes not becoming longer by tearing. Seeds not straight.
16—4. S. cochinchinensis ssp. leptophylla
86. Calyx hairy.

88. Petiolé 3=4. mm. 2 82.22 Se ee ee ee 10. S. brachybotrys
88. Petiole more than 5 mm.
89. Ovary glabrous 00 < wo ote hese 16—4. S. cochinchinensis ssp. leptophylla

1977] SYMPLOCACEAE (Nooteboom) 221

89. Ovary hairy.
nt Bracts ane bracteoles caducous«.) .yc of 4. Wetted paces 48. S. rubiginosa
90. Bracts and bracteoles persistent.
Sieeeacwsite.c. tm long. Seeds straight .. . .. . . seecwsles ok wets 24. S. filipes

91. Bracts longer than 1 mm. Seeds not straight.
16-4. S. cochinchinensis ssp. leptophylla
79. Underside of leaves glabrous.
RS CISTICHOUS MU Sr are kite SE) op a 5 sy! op hayh sadaeen oy eee 33. S. laeteviridis
92. Leaves spirally arranged.
93. Calyx and/or ovary hairy.
94. Ovary glabrous.
95. Bracts caducous.

ERA RO CE LRA CM... 6). 0s: asa nies Coed Se ee 12. S. buxifolia
SP OAVES (OL GHN Ls PSF a! s/s.) sx. La ces 6 SR eee 21. S. cylindracea
PP CANCSNONCEr HAN MO CMs) ys. «woes sue © eee 7. S. barringtoniifolia
95. Bracts persistent.
me senorescence Only I-flowered . . . . . . « + seisudlp ue eee 12. S. buxifolia
97. Inflorescence more-flowered.
Op DISET gS So a oor 16-4. S. cochinchinensis ssp. leptophylla
98. Disk glabrous.
99. Calyx lobes toc.*/,mmlong....... 16-4. S. cochinchinensis ssp. leptophylla
99. Calyx lobes longer than /, mm.
foe Ovary toiemymmobighe soo ooo. ace Cl cl Re, ee 16. S. cochinchinensis
100. Ovary more than 1 mm high.
Pe IIE RITEADDS ile oe eos: «deus wt ole eee 42. S. ophirensis

101. Seeds not straight.
102. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla

102. Seed-and embryo different .. . «6.» volte 42. S. ophirensis
94. Ovary hairy.
103. Leaves shorter than 5cm. Petiole3-4mm ........... 10. S. brachybotrys

103. Leaves longer than 5 cm.
104. Calyx glabrous.
105. Disk glabrous.
106. Calyx lobes not becoming longer by tearing. Seeds not straight.
16-4. S. cochinchinensis ssp. leptophylla
106. Calyx lobes becoming longer by tearing. Seeds straight .... . 20. S. crassipes
105. Disk hairy.
107. Seeds not straight. Bracts and bracteoles persistent.
16-4. S. cochinchinensis ssp. leptophylla
107. Seeds straight. Bracts and bracteoles caducous.
Ess SOOTONAL SOMME sys) Kee a eh» set wt Re 21. S. cylindracea
A ee oe er 32. S. junghuhnii
104. Calyx hairy.
109. Bracts caducous.
110. Calyx longer than 1 mm. Style base hairy.
111. Petiole 3-4 mm. Inflorescence a (basally branched) spike. Ovary c. 1 mm high. Disk
inconspicuous. Fruits c. 10 mm long, I-celled. Seeds not straight .10. S. brachybotrys
111. Petiole more than 5 mm. Inflorescence a panicle. Ovary 1-1'/, mm high. Disk clearly

present. Fruits 15 mm long, 3-celled. Seeds straight .... . 21. S. cylindracea

110. Calyx to c. 1 mm long. Style base glabrous .......... 42. S. ophirensis
109. Bracts persistent.

RAMEN P SITING Guy te 8 8 eo» 4) ew 42. S. ophirensis

112. Petiole more than 5 mm.
113. Leaves crowded towards the end of the twigs, the latter tapering off, at least 5 mm @
IER. 5g x01 che nilesun > ip eee eee 44. S. polyandra
113. These characters not combined.
114. Nerves 13-20 pairs. Intramarginal vein absent. Leaves 21-62 cm 26. S. gigantifolia
114. Nerves 4-13 pairs. Leaves 4-23 cm.
115. Disk glabrous.

116. Bractslongerthanimm ...... 16-4. S. cochinchinensis ssp. leptophylla
116. Bracts to c. 1 mm long.
117. Calyx lobes to c. '/, mm long. Stone with low ridges .... . 24. S. filipes

117. Calyx lobes longer than '/, mm. Stone with high lengthwise interrupted ridges.
42. S. ophirensis
115. Disk hairy.

118. Nerves 4-7 pairs. Fruits ampulliform with long beak, c. 7 by 5mm
42. S. ophirensis
118. These characters not combined. . . . 16-4. S. cochinchinensis ssp. leptophylla

——- © =

222 FLORA MALESIANA [ser. I, vol. 8?

93. Calyx and ovary glabrous.
119" Inflorescence fetmindl. nn... te Pe eee oe ees 46. S. pyriflora
119. Inflorescence axillary.
120. Bracts caducous.
121. Nerves more than 10 pairs.
122. Inflorescence a (basally branched) spike, forming a cone in bud. Fruits more than 20 mm

[Steere ere ye a ae Get oi, ee nines LS) 5) OPS ee es
122. Inflorescence not a spike. Fruits less than 20 mm long.
123) Beactssana bracteoles plabrous 2. 92... ss use ... . 46. S. pyrifiora
123. Bracts and bracteoles hairy.
124. Leaf margin entire. Disk inconspicuous ........... 36. S. maliliensis

124. Leaf margin not entire. Disk clearly present.
125. Bracts shorter than 3 mm. Stamens less than 100. Corolla c. 4mm
11. S. brandisii
125. Bracts longer than 3 mm. Stamens more than 100. Corolla c. 5 mm long.
11b. S. brandisii var. pseudoclethra
121. Nerves less than 10 pairs.
126. Disk hairy.
I27= Stamens 15-40 betiole | —sstittie ses aot. so es ee 34. S. lancifolia
127. Stamens more than 40. Petiole more than 5 mm.
128. Ovary to c. 1 mm high.
129. Inflorescence a (basally branched) lax spike. Bracts to c. 1 mm long
25. S. gambliana
129. Inflorescence not a spike. Bracts longer than 1 mm.
130. Inflorescence a (basally branched) raceme. Stamens 40-c. 60. Calyx lobes becoming

longer by tearing. Style base glabrous .......... 14. S. celastrifolia
130. Inflorescence a panicle of racemes. Stamens more than 100. Calyx lobes not becom-
me longer by tearing. style base hairy... - =. + «. - =. «eee 39. S. nivea

128. Ovary more than 1 mm high.
SI Petiolers Arie re te ke ser eae as toe ’.. . . 9. S. borneensis

131. Petiole more than 5 mm.
132. Terminal buds glabrous.
133. Inflorescence a (basally branched) raceme. Calyx 11/, mm long. Style base glabrous.

EUUISEG ol OMIM ONPG cue usec yn) «, cdaaice camseees ee 8. S. batakensis
133. Inflorescence a fascicle or very short spike. Calyx c. 2 mm long. Style base hairy.
Bris NGO E, ee fee, ees, ce, ek, peo sie 53. S. tricoccata

132. Terminal buds hairy. Inflorescence a panicle. Calyx longer than 2 mm, lobes longer
than 1!/, mm. Style base hairy. Fruits morethanl10 mmlong . 21. S. cylindracea
126. Disk glabrous.
134. Fruits 2—5-celled.
135. Inflorescence a fascicle or a very short spike. Ovary more than 1 mm high.
53. S. tricoccata
135. Inflorescence a (basally branched) raceme. Ovary to c. 1 mm high.
136. Inflorescence axis hairy. Corolla more than 4mm long. Calyx lobes becoming
longer by tearing. Stone smooth. Seeds not straight .... . 14. S. celastrifolia
136. Inflorescence axis glabrous. Corolla shorter than c. 4 mm. Calyx lobes not becoming
longer by tearing. Stone with ridges or grooves. Seeds straight 27. S. glabriramifera
134. Fruits 1-celled.

137. Reticulation fine. Ovary 2-3 mm high ............ 2. S. buxifolia
137. Reticulation coarse. Ovary 1-2 mm high.
138. Inflorescence much branched ..... 16-3. S. cochinchinensis ssp. thwaitesii
USE... Itloresrence Semple ais Fe. fe A ae fen oo Se Ce 51. S. sumuntia

120. Bracts persistent.
139. Leaves shorter than 5 cm.
140. Inflorescence only 1-flowered. Bracts several.
141. Bracts shorter than 3 mm. Corolla shorter than c. 4 mm. Ovary 1-2 mm high. Stamens
less than 30. Stone smooth. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla
141. Bracts longer than 3 mm. Corolla more than 4 mm long. Ovary more than 2 mm high.
Stamens more than 50. Stone with ridges or grooves. Seed and embryo not uncinately
curved: towardsithe: base:. s.0.°5. *. eae, eee censor tee ties 12. S. buxifoli
140. Inflorescence more-flowered. Bract 1.
142. Petiole 0-5 mm.
143; Corolla’ S=fimm eo ee eee ee ee 57. S. whitfordii
143. Corolla shorter.
144. Leaf index less than 2. Acumen shorter than 5 mm. Bracts longer than 1 mm.

144. Leaf index more than 2. Acumen longer thanSmm...... 34. S. lancifolia
142. Petiole more than 5 mm.

1977} SYMPLOCACEAE (Nooteboom) 223

145. Inflorescence a basally branched raceme. Corolla 5-7 mmlong . . 57. S. whitfordii
145. Inflorescence a (basally branched) spike. Corolla shorter than c. 4 mm.
16-4. S. cochinchinensis ssp. leptophylla
139. Leaves longer than 5 cm.

are PIO Mer SATII ea fc. wy Sud. we us 5. ag Sk Sh 00 choap hace oe ee ee 34. S. lancifolia
146. Petiole more than 5 mm.
147. Inflorescence not a spike.
148. Inflorescence not a fascicle.
149. Reticulation fine. Ovaryc. 1 mmhigh. ........ 18. S. composiracemosa
149. Reticulation coarse. Ovary more than I mm high
16-3. S. cochinchinensis ssp. thwaitesii
148. Inflorescence a fascicle.
150. Disk glabrous.
151. Petiole 12-17 mm. Ovary more than | mm high. Nerves 8-12 pairs. Fruits 10-12 mm
MBTIP elie a eile eo es 8 ew od eee 5. S. atjehensis
151. Petiole 5-12 mm. Ovary c. 1 mm high. Nerves 10-16 pairs. Fruits 7-10 mm long.

28. S. glomerata
150. Disk hairy.

152. Leaves obovate, longer than 10 cm. Acumen longer than 5 mm. Inflorescence axis
hairy. Calyx regularly 5-lobed. Fruits ovoid or obovoid, 1I-celled. Seeds 1, not
REERINNE pers Sire Be. cin cis wok). (Sear& 16-4. S. cochinchinensis ssp. leptophylla

152. Leaves elliptic or circular, shorter than c. 10 cm. Acumen shorter than 5 mm. In-
florescence axis glabrous. Calyx 2-4-lobed or symmetrically cleft. Fruits cylindric or
ellipsoid, 2—S-celled. Seeds more than 1, straight. . ... . 40. S. obovatifolia

147. Inflorescence a (basally branched) spike.
153. Twigs (exceptionally) thick.
154. Terminal buds hairy. Disk hairy . . . . 16-4. S. cochinchinensis ssp. leptophylla
154. Terminal buds glabrous. Disk glabrous.
155. Inflorescence axis hairy. Bracts 2-3 mm, hairy. Bracteoles hairy. Calyx 11/,-2 mm

long. Prots:c210'mm longs. . .c.1 2s So eee 6. S. barisanica
155. Inflorescence axis glabrous. Bracts 5-7 mm, glabrous. Bracteoles glabrous. Calyx
longer than;2'mm. Fruits c:-13.mm long: .. . os nis kee 45. S. pulvinata

153. Twigs not (exceptionally) thick.
156. Calyx 2-4-lobed or symmetrically cleft.

Baie REOOIEAD SAAN sk ke wt oe 54. S. trisepala
DP PCHOIG: =-e-Ots Gok Gd ea ee kw ce 40. S. obovatifolia

156. Calyx regularly 5-lobed.
158. Base angle to 20—30°. Leaf index 3'/,—5. Nerves 11-13 pairs. . 29. S. goodeniacea
158. Base angle more than 30°. Leaf index less than 3'/,. Nerves at most 11 pairs.
159. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla
159. Seed and embryo twice curved. ..... 16-2. S. cochinchinensis ssp. laurina

KEYS TO FLOWERING MATERIAL ARRANGED BY ISLANDS AND ISLAND GROUPS

Sumatra

1. Midrib prominent on the upper surface.
hes SitenckGiniviens 7.15) . + « 2 + 4 0.6, #, © ¥, Wee ee 4. S. anomala

5. Twigs hairy. ’ ;
6. Leaves distichous. Calyx 2—4-lobed or symmetrically cleft, calyx lobes becoming longer by tearing.
33. S. laeteviri
6. Leaves spirally arranged. :
7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Petiole more
than 20 mm. Corolla more than 7 mm long. Apex of leaves rounded or acute. 44. S. poly.
7. Leaves evenly distributed.
- Colvx end ovmry. giabrous .. . . . «+> ‘es ion ote ee eee ee 5. S. atjehensis
8. Calyx and/or ovary hairy. :

9. Leaves longer than 10 cm. Nerves more than 10 pairs. Inflorescence a (basally branched) spike.
Bracts persistent. Ovary glabrous, to c. | mm high, calyx lobes longer than 1'/, mm. Disk
glabrous, clearly present. Fruits ampulliform, —— P

. S. cochinchinensis ssp. cochinchinensis

224 FLORA MALESIANA [ser. I, vol. 8?

9. Leaves shorter than c. 10cm. Nerves less than 10 pairs. Inflorescence a raceme. Bracts
caducous. Ovary hairy, 1-2 mm Heh ates lobes iiilinia mm es Disk hairy, inconspicuous.

Fruits ellipsoid, 3-celled . . : . . . . 47. §. robinsonii
5. Twigs glabrous.
POMINET Ves 4-5, DAMS 0 yes x Mey segieme, so we 8) es eee oes ae vie as oA OL

10. Nerves more than 5 pairs.
11. Calyx and/or ovary hairy.

12: dkeavesiGistichous 12 «cele .Shariccles ack BE Ae. Deol SSS eiserceruiis
12. Leaves spirally arranged.

13. Leaves crowded towards the end of the sean -off me: ..... . 44. §. polyandra
134 Plantidiiierentes.... Bad SE a alk thal - oo oe « « AZ SS ODMIeNSIS

11. Calyx and ovary glabrous.
14. Inflorescence not a spike.

15 inflorescence. a‘fascicle. Stamens'c./50!' sou view. ..eo. si. 2S Se eiensis
15. Inflorescence a raceme.
16s Stamensi40=60; 211-40 eecw gels eam 1 oe ter) wie) 4) 31-2 e
16. Stamens c. 100 . ee Peres) fe Lec Sie
14. Inflorescence a (basally branched) spike.
ijemuivwies thick. betminaljbudsilarce, sa.) ie) Seales . 6. S. barisanica
17. Twigs not thick. Terminal buds small ....... 16-2. S. cochinchinensis ssp. laurina

4. Underside of leaves hairy.
18. Twigs glabrous.
19. Leaves distichous. Petiole 1-4mm. Corolla3-5 mm .......... . 33. S. laeteviridis
19. Leaves spirally arranged. Petiole more than 5 mm.
20. Inflorescence a fascicle. Bracts persistent, c. 2mm. Calyx glabrous. Calyx lobes not becoming

longer DY: tearing ty.c Sieecticg ‘aya ks bcc oa ee Cee EP ie? Be 5. S. atjehensis
20. Inflorescence a spike forming a cone in bud. Bracts caducous, 3-5 mm. Calyx hairy. Calyx lobes
becoming longer by tearing. ..... » Uae) ceereielg.cnu lanl? Aa Sarbioinss

18. Twigs hairy.
21. Leaves distichous.
22. Calyx usually hairy. Inflorescence a fascicle. Bracts persistent. Ovary c. 1 mm long. Calyx
c. 1 mm long. Calyx lobes not becoming longer by tearing. Style base hairy . 23. S. fasciculata
22. Calyx often glabrous. Inflorescence a raceme or panicle of racemes. Bracts caducous. Ovary
1-1'/, mm high. Calyx 2-3 mm. Calyx lobes becoming longer by tearing. Style base glabrous
33. S. laeteviridis
21. Leaves spirally arranged.
23. Calyx and ovary glabrous.

D4niLeaf index. 2.to.32 Nerves!8=12 pairs, ...¢) ies) a 0 Si Soak ae. APE Se Seaehensis
24. Leaf index more than 3. Nerves 6-9 Dae

25. Nerves less than 10 pairs. . . . artnet se : . S. cerasifolia
255 INetvesumoreithan: 10) pairs? ., .. « sch aves en _ 15b. S. poe var. grandifolia

23. Calyx and/or ovary hairy.
26. Upper side of leaves hairy (pulverulent). Leaf margin and petiole beset with many closely
SpacedavesiculanmclandSin-t.a ee ceiiia cucees Moen sees ete e . . 3. S. adenophylla
26. Upper side of leaves glabrous. Leaf margin and petiole different.
27. Ovary glabrous.
28. Nerves less than 10 pairs. Bracts caducous. Ovary 1—1!/, mm high. Calyx ee mm. Calyx
lpbes/becominelonger by teaming . ete ee oe ee . S. cerasifolia
28. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm long. Guise t to 2 mm long.
Calyx lobes not becoming longer by tearing.
16-1. S. cochinchinensis ssp. cochinchinensis
27. Ovary hairy.
29. Leaves longer than 15 cm.
BO MMICHVESHIOSISsDAITS us Sais 54.0 oat one eee eek aes} 15b. S. cerasifolia var. grandifolia
SOMINGHVESEL2—IVAPallSs Fe Naaincuk: Piecet Seeerals «os ss) 6 Fo ee ee 48. S. rubiginosa
29. Leaves shorter than c. 15 cm.
31. Bracts to c. 1 mm long.
32. Leaf margin (and petiole) beset with closely spaced glands. . . . . 3. S. adenophylla
32. Leaf margin (and petiole) often glandular but glands not closely spaced.
33. Nerves less than 10 pairs. Reticulation coarse. Inflorescence a fascicle. Bracts persistent.

Ovaryie.f muni high. Stylebase hairy’) 3 sao. o9) ep) et DS 23. S. fasciculata
33. Nerves more than 10 pairs. Reticulation fine. Inflorescence a (basally branched) raceme.
Bracts caducous. Ovary more than 1 mm high. Style base glabrous . 47. S. robinsonii

31. Bracts longer than 1 mm.
34. Angle oljleanbase.c. 90° 1 sa oer es es 3 OS Pe 50. S. sumatrana

4 Angleton leat base less'than'607).Gai)) 6. Ra ae eee 47. S. robinsonii

1977] SYMPLOCACEAE (Nooteboom) 225

Malay Peninsula

1. Midrib prominent on the upper surface.
2. Twigs hairy.
3. Leaves evenly distributed, underside glabrous ........2.2.2.2.2.4.2. 4. S. anomala
3. Leaves crowded towards the end of the twigs, minutely sparsely appressedly hairy beneath.
37. S. wikstroemifolia
2. Twigs glabrous.

4. Leaves crowded towards the end of the twigs ............. 37. S. wikstroemifolia

EaAUCuen IY Cistripnted (bows) «ke se oe eee SD ee ee 35. S. lucida
1. Midrib impressed on the upper surface.
Ty. ec SOS ee Se. 8 eS eae 41. S. odoratissima
5. Corolla glabrous.

6. Twigs hairy.

7. Leaves distichous.
eemeorsice Of leaves SlabrouS . . . . «ics ay ce © « ommcgeinelne Wee 33. S. laeteviridis

8. Underside of leaves hairy.
9. Bracts persistent.
10. Inflorescence a true fascicle. Ovary c. 1 mm high. Calyx 1 mm long . . . 23. S. fasciculata
10. Inflorescence a short, often clustered spike. Ovary 1-2 mm high. Calyx 1'/,-4 mm long
20. S. crassipes
SMP EAUICOUS ba > si ERAT. Wins) Se <5) oh os dee Se ee ee 33. S. laeteviridis
7. Leaves spirally arranged.
11. Upper side of leaves hairy.
12. Angle of leaf base more than 90°. Bracts and bracteoles caducous. Hairs on twigs more than
PenarMREYTTI HF tg stats ede ie bree eee 13. S. calycodactylos
12. Angle of leaf base less than 90°.

13. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 mm long. Calyx to
c. 1 mm long, calyx lobes */,-1 mm long. Disk glabrous. Style base not conical. Fruits to c.
meemiong PP Mae a. 3S re 3. S. adenophylla

13. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than
1 mm. Calyx 11/,4 mm, calyx lobes longer than 1'/, mm. Disk hairy. Style base conical.

Bee ts- 1S mint long Oe 20. S. crassipes
11. Upper side of leaves glabrous.
14. Leaves crowded towards the end of the twigs... ......... 37. S. wikstroemifolia

14. Leaves evenly distributed.
15. Underside of leaves glabrous.
16. Calyx glabrous. Inflorescence a fascicle. Disk pulvinate or cylindric. Fruits cylindrical. Seeds

nee 2. UP Webbe wood «les eater ae 28. S. glomerata
16. Calyx hairy. Inflorescence a (basally branched) spike. Disk annular. Fruits ampulliform.
Becas notetraicnt-; Foo. Ss . eeG@adls 16-1. S. cochinchinensis ssp. cochinchinensis

15. Underside of leaves hairy (pulverulent, nearly glabrous, in S. adenophylla).
17. Calyx and ovary glabrous.

ae pecmoreacence’a spiké.. . . . . . mie ee ee A 15. S. cerasifolia
Deameeenrenee a igeticle. 2. nis « caite eee eee 28. S. glomerata

17. Calyx and/or ovary hairy.
19. Ovary glabrous.

20. Nerves less than 10 pairs. Bracts and bracteoles caducous. Ovary 1-1'/, mm high, calyx
longer than 2 mm, calyx lobes 1-1'/, mm long, becoming longer by tearing. Fruits 22-40
ge ee aera ee 15. S. cerasifolia

20. Nerves more than 10 pairs. Bracts and bracteoles persistent. Ovary to c. 1 mm high, calyx
c. 2mm long, calyx lobes longer than 1'/, mm, not becoming longer by tearing. Fruits
sya some, A-colled «5. + 2 hm 2 16-1. S. cochinchinensis ssp. cochinchinensis

19. Ovary hairy.

21. Ovary more than 1 mm high.

22. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 oc long. Calyx

teccbmmlong 3 es! oh. eels Ce SET ee Pee 3. S. adenophylla
22. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than
1 mm.

23. Nerves 6-11 pairs. Reticulation faintly prominent. Bracts and bracteoles persistent.
Calyx lobes not becoming longer by tearing. Fruits 13-18 mm long . 20. S. crassipes
23. Nerves 12-17 pairs. Reticulation much prominent. Bracts and bracteoles caducous.

Calyx lobes becoming longer by tearing. Fruits toc. 10 mmlong . . 48. S. rubiginosa
21. Ovary to c. 1 mm high.
24.\ heaves longer than:i3.0m. «sis a2 «nan elt 6 nod? re ah 20. S. crassipes
24. Leaves shorter than c. 15 cm.
25. (Reticulation fine.) Bracts and bracteoles caducous ....... 42. S. ophirensis

25. (Reticulation coarse.) Bracts persistent.

226 FLORA MALESIANA [ser. I, vol. 8?

26. Inflorescence an often clustered short spike. Bracts 1-4 mm. Calyx 2!/.-3 mm. Stamens

more than 60. Fruits not ampulliform, 13-18 mm long ... .. 20. S. crassipes
26. Inflorescence a fascicle. Bracts to c. 1 mm long. pes c. 1 mm long. Stamens 12-35.
Fruits ampulliform, 5-7 mmlong ............. . 23. S. fasciculata

6. Twigs glabrous.
27. Nerves more than 10 pairs.
28. Underside of leaves hairy.
S90 inflorescence aspike. .. 6 2645 4 ede ey a
99. Inflorescence afascicle .. 2.0.0 62 © 6 1 oe es hw ow «ogee eA RE ONIMCIE TSE
28. Underside of leaves glabrous.
30. Petiole more than 20 mm. Inflorescence a spike or a cone. Fruits 3-celled
7. S. barringtoniifolia
30. Petiole less than 20 mm.

31 Inflorescence terminal =") =). es ie. we es es OUT AGS aor
31. Inflorescence axillary.
32. Calyx andjor ovary hairy. Corolla. 2-5mm . . . ws ww. CS 42. S. ophirensis

32. Calyx and ovary glabrous.

33. Intramarginal vein far from margin. Inflorescence a fascicle. Bracts persistent, hairy, shorter
than 3 mm. Ovary c. 1 mm high, calyx 1-2 mm long, calyx lobes not becoming longer by
tearing. Corolla 4-5 mm. Stamens less than 50. Fruits 7-10 mm long. . 28. S. glomerata

33. Intramarginal vein close to margin. Inflorescence a raceme or panicle of racemes. Bracts
caducous, glabrous, longer than 3 mm. Ovary 1'/,-2 mm high, calyx 3-5 mm, calyx lobes
becoming longer by Eee Corolla 8-10 mm long. Stamens c. 100 or more. Fruits c. 15 mm
lONS Vie ae eo a ae ete Sen ee Se a

27. Nerves less than 10 pairs.
34. Underside of leaves hairy.

35. Leaves crowded towards the end of the twigs .. ..... . . . . 37. S. wikstroemifolia
35. Leaves evenly distributed.
36. Bracts persistent. Disk glabrous. Fruits 13-18 mm long Jw) 70) ZO Saerassines
36. Bracts caducous. Disk hairy. Fruits 7-12mmlong ....... . . . 33. S. laeteviridis

34. Underside of leaves glabrous.
37. Calyx and/or ovary hairy.
38. Disk hairy.

39. Leaves distichous . . fy eee PE 8 Gren) GG
39. Leaves spirally arranged Sh oto gg eee ab ae. oa: th 42. S. ophirensis
38. Disk glabrous.

40. Inflorescence a short, often clustered spike. Bracts persistent. . . . . . 20. S. crassipes
AQ: Inflorescence’a raceme? Bractsicaducous . . . . /S0.0) ee Cee 51. S. sumuntia

37. Calyx and ovary glabrous.
41. Inflorescence a (basally branched) spike. Bracts persistent
16-2. S. cochinchinensis ssp. laurina

41. Inflorescence not a spike. Bracts caducous.

42. Inflorescence a panicle of racemes. Stamens more than 100 ........ 39. S. nivea
42. Inflorescence a (basally branched) raceme. Stamens 25-60.
43. Calyx lobes becoming longer by tearing. ............ 414. S. celastrifolia
43. Calyx lobes not becoming longer by tearing .... . &! & =o eS Sy snes

Java & The Lesser Sunda Islands

PrietoachnaIty oe Sono te es Boe pea de A A eee 41. S. odoratissima
1. Corolla glabrous.
2) Midrib proniinent on theiupper suriace?;\"> 202". pe, ey) es RD) 35. S. lucida

2. Midrib impressed in the upper surface.
3. Twigs hairy.
4° Leaves distichous) 5.7) Deco meee Wey Gi) co gs SR OE 23. S. fasciculata
4. Leaves spirally arranged.
5. Underside of leaves glabrous. (If leaf ene and Hg he beset with closely spaced vesicular glands:

35 Sy adenopnylia:) ss.) ee 16-1. S. cochinchinensis ssp. cochinchinensis
5. Underside of leaves hairy.
6.“ Upper side of leaves hairy (pulverulent).5" s We. GOkA. 2st. i228 See 3. S. adenophylla

6. Upper side of leaves glabrous.
7. Ovary glabrous, calyx longer than 1 mm. Bracts longer than 1 mm
16-1. S. cochirichinensis ssp. cochinchinensis
7. Ovary hairy.
8. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Ovary
12mm high . 2. se ts Spee oe Se nee aoe eee 3. S. adenophylla
8. Leaf ndee 2-3. Leaf pena (and petiole) often glandular but glands not closely spaced.
Ovary’c. 1! murvhigh . . . . . . UR eee eee eee 23. S. fasciculata

1977] SYMPLOCACEAE (Nooteboom) 227

3. Twigs glabrous.
9. Calyx and/or ovary hairy.
10. Ovary hairy. Inflorescence a raceme. Calyx glabrous .......... 32. S. junghuhnii
10. Ovary glabrous. Inflorescence a spike. Calyx hairy.
16-1. S. cochinchinensis ssp. cochinchinensis
9. Calyx and ovary glabrous.
11. Inflorescence axis glabrous.

Pree IesSitnans2ONM aie i SG See.) TR 16-2. S. cochinchinensis ssp. laurina

i@ereuoic more than 20mm... .........-. 16-4. S. cochinchinensis ssp. leptophylla
11. Inflorescence axis hairy.

mecemesoess than iO.pairs 6. is. 6 ed SI ee 16. S. cochinchinensis

13. Nerves more than 10 pairs.
14. Angle of leaf base 25—40°. Inflorescence a (basally branched) spike, forming a cone in bud.
19. S. costata

14. Angle of leaf base more than 60°. Inflorescencea raceme ........ 11. S. brandisii
Borneo
SS a a co a a rr mri eS Re 41. S. odoratissima
1. Corolla glabrous.
Sessa fiat ‘or prominent on the upper surface .......:..2.082..-. 4. S. anomala

2. Midrib impressed in the upper surface.
3. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands:
3. S. adenophylla).

4. Twigs hairy.
ReEIPEPECISUICHOUS, . ceo sss wes ww ws vee 33. S. laeteviridis
5. Leaves spirally arranged.
femeeaves 4-6.em. Petiole 3-4mm. . . . . .. «5 a os el ee 10. S. brachybotrys

6. Leaves longer than 6 cm. Petiole more than 5 mm.

7. Leaves evenly distributed. Twigs not thick, cylindrical. Leaf margin not entire. Petiole less than
20 mm. Ovary glabrous, to c. 1 mm high, calyx 1-2 mm long. Corolla 3-5 mm. Fruits ampulli-
form, 1-celled. Seed 1, not straight. Apex of leaves acuminate.

16-1. S. cochinchinensis ssp. cochinchinensis

7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Leaf margin
entire. Petiole more than 20 mm. Ovary hairy, c. 2mm high, calyx 2-3 mm long. Corolla
8-10 mm long. Fruits ellipsoid, 3-celled. Seeds more than 1, straight. Apex of leaves rounded or
MS ke fe ee 44. S. polyandra

4. Twigs glabrous.
8. Calyx and/or ovary hairy.
ees cisticnous,..Calyx SIabrOUS. «. «= «. ++. «lels) ll) an 33. S. laeteviridis
9. Leaves spirally arranged.
10. Inflorescence a 1—3-flowered spike. Bracts caducous. Stamens c. 100. Petiole 3—-4 mm. Leaves

ae CON o 50 (tI PRIN 58 O.5 5. 4) i) et SR 10. S. brachybotrys
10. These characters not combined.
eaves 15—SO.mmilong . 0... 2 os os 1 Wt eae 12. S. buxifolia

11. Leaves longer than 5 cm.
12. Petiole more than 20 mm. Twigs thick. Inflorescence a spike. Calyx 3-3!/, mm
7. S. barringtoniifolia
12. Petiole less than 20 mm. Inflorescence a raceme or a spike. Calyx to c. 1 mm long
42. S. ophirensis
8. Calyx and ovary glabrous.
a peeves more wen TO paite.. . ww tt nw + ke 29. S. goodeniacea
13. Nerves less than 10 pairs.
14. Bracts persistent.
Seer waeawee suorier than 5 Cm... «i «4s « «©, peel eee 12. S. buxifolia
SRR CM IMIG TROT OC ws) gk ie oe 16-2. S. cochinchinensis ssp. laurina
14. Bracts caducous.
16. Ovary to c. 1 mm high. Apex of leaf mostly abruptly acuminate.
17. Leaf margin entire. Inflorescence axis glabrous. Petiole 5-10 mm . . . 25. S. gambliana
17. Leaf margin not entire. Inflorescence axis hairy. Petiole 3-I15mm_ . 14. S. celastrifolia
16. Ovary more than | mm high. Leaf apex usually not or faintly acuminate.
18. Leaf margin entire. Calyx */,-1 mm long. Disk hairy. Petiole 3-4mm . 9. S. borneensis
18. Leaf margin not entire. Calyx longer than 1 mm. Disk glabrous or the style base hairy.
19. Leaves shorter than 5 cm and acumen shorter than 5 mm. Reticulation fine. Inflorescence a
few-flowered raceme. Bracts longer than 3 mm. Ovary 2-3 mm high, calyx 2—5 mm long.
Style base glabrous. Fruits 1-celled 9) o/5)4 Gla ake 12. S. buxifolia
19. Leaves longer than 5 cm and acumen longer than 5 mm. Reticulation coarse. Inflorescence
a fascicle or very short spike. Bracts c. 1'/, mm. Ovary c. 2 mm high, calyx c. 2 mm long.
Style base hairy. Fruits 3-celled. Stone smooth. Seeds more than! . . 53. S. tricoccata

228 FLORA MALESIANA [ser. I, vol. 8?

3. Underside of leaves hairy.
20. Leaves distichous.
21...\Calyx symmetrically teared whenvolder ...). Grace 6 4s ee he . . 33. §. laeteviridis
21. Calyx regular.
22. Stamens 25-35. Inflorescence a true fascicle. Petiole 2-8 mm. Leaves 5-18 cm. Ovary c. 1 mm
High ave 6.0 MN IONS te isles) ws «0 we cf ook 2) 3 se 23. S. fasciculata
22. Stamens more than 40. pipers solitary or ina raceme or panicle. Leaves 2-9 cm. Calyx 1-3 mm.
23. Nerves 7-11 pairs. Petiole 3-4 mm. Leaves 4-9 cm. Stamens more than 90. Calyx 13/,-3 mm
17. S. colombonensis
23. Nerves 3-8 pairs. Stamens 40-100, but when more than 90 Bonde 1-2 mm.

24. Petiole 2-4 mm. Leaves 2- 31/5 cm. Calyxcc.Sitiim) .o. eee . . 52. S. trichomarginalis
24. Petiole 1-2 mm. Leaves 21/,—7 cm. aes 1-2 mm.
25 4Galyaccuommilonewicaiset. vies) 4 sarktovotie |. OR -oner. ae. 58. S. zizyphoides
25. Calyx 1-1"/, mm long.
26 Ovary, limm nich) weer & oe. Sanit, “te wat) eae. ead Leask 31. S. johniana
DOmOV ALY ales —2 meh Nun. ei A ea sensin Silos 8s (ce es hele cate ee 22. S. deflexa

20. Leaves spirally arranged.
27. Twigs glabrous.

PR MIBCAVESTA—OIGIT) wa ciultt we gies) clog curts. Pets nisi! aseecs: oumblenite Tacit re 10. S. brachybotrys
28. Leaves longer than 6 cm.
29e Teaves: /—16cm., Nerves 6-9) palTS... «. -, cusses Mtr Mucmacnel. bereaaleieain. 15. S. cerasifolia
DO eaves 1 5—45/ CIN NEVES I 2-7, Dalts 0.) 5, oasems Deen een) Otten 48. S. rubiginosa
27. Twigs hairy.
30. Upper side of leaves hairy (pulverulent). .........2.2.2.... 3. S. adenophylla
30. Upper side of leaves glabrous.
S@Alwe AUG OVALY .SIADTOUS: 1.4. 4 soa « & 5 a 2 » GS = = CRE 15. S. cerasifolia

31. Calyx and/or ovary hairy.
32. Ovary glabrous.
33. Nerves less than 10 pairs. Bracts caducous. Ovary 1—1*/, mm high, calyx longer than 2 mm.
Rrutsiellipsoida22—40immilones3-celled) 7) 4.5 suis eerie ariel) Saleuee 15. S. cerasifolia
33. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm high, calyx 1-2 mm long.
Fruits ampulliform, 5—7 mm long, 1-celled.
16-1. S. cochinchinensis ssp. cochinchinensis
32. Ovary hairy.
Sdseeavesslongenthanilisycom: ©. side: “erly ..mta dS. cout) Suse: ome 48. S. rubiginosa
34. Leaves shorter than c. 15 cm.
35. Inflorescence only 1-flowered.

36, Angleton leaf base more than OO kkk se eo a 31. S. johniana
36. Angle of leaf base less than 90°.
Sie bracts. le 'Calyxsl—Z2 mmMVONg, cto <i oui: eee OLS eee 20. S. crassipes
Bie, bractsisevetals Calyx e:,3)IM >, « c: ahs... %. « 1.1 cee 10. S. brachybotrys
35. Inflorescence more-flowered.
38. Bracts caducous. Petiole 3-4 mm. Nerves 6-9. Stamens c. 100 . . 10. S. brachybotrys

38. Bracts persistent. Petiole 2-12 mm. Nerves 3-12. Stamens 12-more than 100.
39. Calyx to c. 1 mm long. Stamens 12-50.
40. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands.

Ovanyal=2 mmhighiot Aosta as Boies wae. -ees UE eel Eee oe 3. S. adenophylla

40. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced.

Ovary scalimmihish: ps wenene. A eres tek ose) Bt oe 23. S. fasciculata

39) Galyx longer than 1mm. Stamens 25-100: 7 >. 5). we ne 20. S. crassipes
Philippines

ieelbeavesverticiilate... . . « «hy 4 . uw aha a in" od ey) GR Se oe 55. S. verticillifolia

1. Leaves not verticillate.
2. Midrib prominent in the upper surface.

5; ULWwigs glabrous) Petiolesmorethanpormim) eiemes see ee Sens este 35. S. lucida

Sihwies hairy. Petiole:l!—Simm ainiss: sels ate tye Geta nt ee 34. S. lancifolia
2. Midrib impressed on the upper surface.

4. Corolla hairy (in the Philippines sometimes nearly glabrous!) ...... . 41. S. odoratissima

4. Corolla glabrous.
5. Twigs hairy.

Ovwleeaves distichous ls» ..1.....08 ain 4 hele eee ee Do Eee 23. S. fasciculata
6. Leaves spirally arranged.
74 Galyx divided into: three 23/,,-mm long lobes) asus o-aesiee ee 54. S. trisepala

7. Calyx not so.
8. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards the apex. Fruits
3-celled. Apex of leaves rounded or acute... 5. «1.5.8 os 44. S. polyandra

1977] SYMPLOCACEAE (Nooteboom) 229

8. Leaves evenly distributed.
9. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular
glands: 3. S. adenophylla).
10. Ovary hairy, c. 1'/, mm high. Inflorescence a lax raceme. Bracts to c. 1 mm long. Calyx lobes
omni lone: stamens ¢.25 . . . . ks fs so 5 ne” Ree 24. S. filipes
10. Ovary glabrous (hidden between bracts!). Inflorescence a spike.
16-1. S. cochinchinensis ssp. cochinchinensis
9. Underside of leaves hairy.
11. Upper side of leaves hairy (pulverulent) ..........2... 3. S. adenophylla
11. Upper side of leaves glabrous.
12. Ovary glabrous.

13. Leaves longer than 10cm. Petiole more than 5 mm. Nerves more than 10 pairs. Calyx
lobes longer than 11/, mm. Stamens more than 30. Disk glabrous. Style base glabrous.
enusranipuitorn 2s! rk) i 16-1. S. cochinchinensis ssp. cochinchinensis

13. Leaves shorter than c. 10 cm. Petiole 1-5 mm. Nerves 4-11 pairs. Calyx lobes 1/,-11/, mm
long. Stamens 15-40. Disk hairy. Style base hairy. Fruits not ampulliform

34. S. lancifolia
12. Ovary hairy.
14. Leaf margin (and petiole) beset with closely spaced glands .. . . 3. S. adenophylla
14. Leaf margin (and petiole) often glandular but glands not closely spaced.
15. Style base glabrous.
16. Bracts to c. 1 mm long. Calyx lobes c. '!/, mm long, not triangular . . 24. S. filipes
16. Bracts 2-3 mm. Calyx lobes 1-1!/, mm, triangular ......... 56. S. vidalii
15. Style base hairy.
17. Intramarginal vein present. Inflorescence a fascicle. Fruits ampulliform
23. S. fasciculata
17. Intramarginal vein absent. Inflorescence a (basally branched) spike. Fruits ellipsoid to
BEcHiatiM: walutsitis matic (fic sc Jee, (a Se ee ee 34. S. lancifolia
5. Twigs glabrous.
18. Calyx and ovary glabrous.
19. Inflorescence a (basally branched) spike.
en eran Sring? )sis) oS. lec! oro, ye, a 34. S. lancifolia
20. Acumen shorter than 5 mm.
21. Angle of leaf base less than 60°. Bracts 2-3 mm. Calyx lobes c. 2 mm long. Style base hairy.
40. S. obovatifolia
21. Angls of leaf base c. 90°. Bracts 3-5 mm. Calyx lobes longer than 2'/, mm. Disk glabrous.
54. S. trisepala
19. Inflorescence not a spike.
22. Bracts and bracteoles persistent.
23. Leaves 2-5?/, cm. Inflorescence a (basally branched) raceme. Bracts longer than 3 mm. Calyx
regularly 5-lobed, calyx lobes semi-ovate. Fruits ovoid, 5-7 mm long, 1-celled.
57. S. whitfordii
23. Leaves 7!/,-11 cm. Inflorescence a fascicle. Bracts 2-3 mm long. Calyx 3-lobed, the lobes
semi-elliptic. Fruits ellipsoid, 11 mm long, 3-celled ........ 40. S. obovatifolia
22. Bracts and bracteoles caducous.
24. Inflorescence axis glabrous. Corolla 3-4 mm. Calyx lobes not becoming longer by tearing.
27. S. glabriramifera
24. Inflorescence axis hairy. Corolla4-6mm...........+.-. 14. S. celastrifolia
18. Calyx and/or ovary hairy.
25. Inflorescence a spike. Ovary glabrous.
Seevaty*/—immbhigh .......+..-s 16-1. S. cochinchinensis ssp. cochinchinensis
Teawvere 2 iste mem kk we tw te ee eg 8 ee ee cee 42. S. ophirensis
25. Inflorescence a (sometimes compound) raceme. Ovary hairy.
27. Inflorescence a very lax raceme of 4-10 cm. Pedicels slender, 2-15 mm. Axis of raceme sparsely
pulverulent-puberulous. Stamensc.25 ..... + ee eee eee es 24. S. filipes
27. These characters not combined. ...... 5.5626 + sseeerevee 42. S. ophirensis

Celebes & The Moluccas

CE EE eer Me Pe a 41. S. odoratissima
1. Corolla glabrous.
2. Midrib prominent on the upper surface ©. 6 6 ee ee 35. S. lucida
2. Midrib impressed in the upper surface.
3. Twigs hairy.

4. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands:
3. S. adenophylla).
5. Leaves distichous. Bracts caducous .......+ + ss ee etree 33. S. laeteviridis
5. Leaves spirally arranged.

230 FLORA MALESIANA [ser. I, vol. 8?

6. Calyx and ovary slabrous. ... <2 )<o Se eee eee 16-4. S. cochinchinensis ssp. leptophylla
6. Calyx and/or ovary hairy.
7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex.

44. S. polyandra
7. Leaves evenly distributed.
2. Owary ligity:.. acu « « soe’ See oe 16-4. S. cochinchinensis ssp. leptophylla
8. Ovary glabrous.
9. Ovarytoc.1mmhigh ...... . . . 16-1. S. cochinchinensis ssp. cochinchinensis
9. Ovary morethani mmhigh ......... 16—4. S. cochinchinensis ssp. leptophylla

4. Underside of leaves hairy.
10. Leaves distichous.
11. Inflorescence a fascicle. Bracts persistent. Ovary c. 1 mm high, calyx c. 1 mm long, calyx lobes
not becoming longer by tearing. Style base hairy. Fruits ampulliform . . 23. S. fasciculata
11. Inflorescence not a fascicle. Bracts caducous. Ovary 1-11/, mm high, calyx 2-3 mm, calyx lobes
becoming longer by tearing. Style base glabrous. Fruits not ampulliform . . 33. s. laeteviridis
10. Leaves spirally arranged.

42, Colyeand ovary glabrous... ws 6s wie oe 16-4. S. cochinchinensis ssp. leptophylla
12. Calyx and/or ovary hairy.
43.) Umer side of leaves batty aia, cies) Sih seed fever buey cage 2 3. S. adenophylla
13. Upper side of leaves glabrous.
14." Cire Se ee eae ace ak Be Sk 16-4. S. cochinchinensis ssp. leptophylla
14. Ovary hairy.
15 Calyx clhabrows.. 2 - seipucans vie Oh ak 16-4. S. cochinchinensis ssp. leptophylla

15. Calyx hairy.
16. Bracts to c. 1 mm long.
17. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced —_— Ovary

1-2 mm high. Stone not ampulliform .............. 3. S. adenophylla
17. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced.
Ovary c.2mm high. Stone ampulliform .........2... 23. S. fasciculata

16. Bracts longer than 1 mm.
18. Bracts caducous. Ovary c.1 mmhigh . . . . 42-1b. S. ophirensis var. densireticulata

18. Bracts persistent. Ovary more than 1 mm high 16-4. S. cochinchinensis ssp. leptophylla
3. Twigs glabrous.
19. Underside of leaves hairy.

20. Leaves distichous. Petiole 0-5 mm. Bracts and bracteoles caducous . . . . 33. S. laeteviridis
2h) Veavesispiralyarranced: 3) 202%) yale. ees G2. 16-4. S. cochinchinensis ssp. leptophylla

19. Underside of leaves glabrous.
21. Calyx and ovary glabrous.
22. Ovary to c. 1 mm hi
23. Inflorescence a raceme. Bracts caducous ............. 14. S. celastrifolia
23. Inflorescence a (basally branched) spike. Bracts persistent
16-2. S. cochinchinensis ssp. laurina
22. Ovary more than 1 mm high.

PA Uo es re ea” bal hn AG cat eth 16-4. S. cochinchinensis ssp. leptophylla
24. Twigs not thick.

25. Inflerescence:a raceme. Bracts'caducous, .. . 2... & woke ae ace 36. S. maliliensis

25. Inflorescence a (basally branched) spike. Bracts persistent.
26 Pasha 6 ee Pe ns oes 16-4. S. cochinchinensis ssp. leptophylla
26 BIS ORIG ee te re ee Oe. a hae a. wm oo 16-2. S. cochinchinensis ssp. laurina

21. Calyx and/or ovary hairy.

Ris iPaaves custiengear sen els. c's Sl hs Ba ws RES es 33. S. laeteviridis

27. Leaves spirally arranged.
28. Ovary glabrous.

29 Ovaryia ce: lamnnthigh ooteis cissiiea) sera 2 16-1. S. cochinchinensis ssp. cochinchinensis
29. Ovary more thanlmm.high......... 16-4. S. cochinchinensis ssp. leptophylla
28. Ovary hairy.
0) Gales Ciaran ene ee a ak ang 16-4. S. cochinchinensis ssp. leptophylla
30. Calyx hairy.
33. Brects to ¢..) mer $0 ee ae eC ee ret td 5 re Be 42. S. ophirensis
31. Bracts longer than Pim sk te 16-4. S. cochinchinensis ssp. leptophylla

New Guinea
(incl. New Ireland & New Britain)

hs sReaves (pscudo-)verticillate.: 6s 26a) he a ea eee a ee ee 30. S. herzogii
1. Leaves not verticillate.
2. Inflorescence only 1-fiowered.
2 Coren ane Ovary iabrom N.S 16-4. S. cochinchinensis ssp. leptophylla

1977] SYMPLOCACEAE (Nooteboom) 231

3. Calyx and/or ovary hairy.

I ate tees Sapa bo fat ce wy mu) ya ini lia ao ot 49. S. salicioides

TS Se 2 rer 38. S. multibracteata
2. Inflorescence more-fiowered.

5. Calyx and ovary glabrous.

6. Twigs hairy.
Sans items WSS as (hc! eae Sue) 2 16-4. S. cochinchinensis ssp. leptophylla
7. Petiole more than 5 mm.
8. Underside of leaves glabrous .......... 16-4. S. cochinchinensis ssp. leptophylla
8. Underside of leaves hairy.
9. Upper side of leaves hairy. Ovary c.*/, mmlong. ......... 43. S. paucistaminea
9. Upper side of leaves glabrous. Ovary more than | mm high.
10. Bracts caducous. Calyx lobes becoming longer by tearing. . ..... 15. S. cerasifolia

10. Bracts persistent. Calyx lobes not becoming longer by tearing.
16-4. S. cochinchinensis ssp. leptophylla
6. Twigs glabrous.
Smeeenderside of leaves hairy .......2..4.+5:. 164. S. cochinchinensis ssp. leptophylla
11. Underside of leaves glabrous.
12. Bracts caducous.

fen bracts longerthan3mm ............ 16-4. S. cochinchinensis ssp. leptophylla
13. Bracts shorter than c. 3 mm.
14. Ovary c. 1 mm long. Inflorescence a (basally branched) raceme . . . . 14. S. celastrifolia
14. Ovary more than 1 mm high. Inflorescence different.
15. Disk hairy. Ovary 1-1!/, mm. Inflorescence a panicle ....... 21. S. cylindracea

15. Disk glabrous. Inflorescence an often branched spike.
16-3. S. cochinchinensis ssp. thwaitesii
12. Bracts persistent.
16. Inflorescence not a spike.
MEEIETINOTESCENCE ANIASCIGIG © 6 fk es 16—4. S. cochinchinensis ssp. leptophylla
Es 18. S. composiracemosa
16. Inflorescence a (basally branched) spike.
18. Twigs not thick.
19. Leaves 6-12 cm. Petiole 5-25 mm. Inflorescence an (often branched) spike. Flowers bisexual,
gvary 1—1"/, mm, calyx */,—"/, mm 2 6 se ke 16-3. S. cochinchinensis ssp. thwaitesii
19. Plants different. Flowers usually functional unisexual.
16-4. S. cochinchinensis ssp. leptophylla
18. Twigs thick.
20. Terminal buds hairy, small. Acumen longer than 5 mm. Bracts hairy, shorter than 3 mm.

Disk hairy. Leaves elliptic or circular. . ... 16-4. S. cochinchinensis ssp. leptophylla
20. Terminal buds glabrous, large. Acumen shorter than 5 mm. Bracts glabrous, 5-7 mm. Disk
Mmeorous..Leaves ObOvate . . . . . . + » « & « a oe ee 45. S. pulvinata

5. Calyx and/or ovary hairy.
21. Ovary glabrous.

22. Ovary hidden by bracts and bracteoles ... . 16-1. S. cochinchinensis ssp. cochinchinensis
22. Ovary not hidden by bracts and bracteoles.
23. Inflorescence a spike, forming a short coneinbud .......... 15. S. cerasifolia
23. Inflorescence sometimes a spike, but never forming a cone in bud.
16-4. S. cochinchinensis ssp. leptophylla
21. Ovary hairy.
2A. Twigs at least 8 mm thick. Leaves 21-62cm . .... 51.21 se eee 26. S. gigantifolia

24. Twigs thinner. Leaves at most 33 cm, but usually much smaller.
25. Calyx 2- to 4-lobed or symmetrically cleft. Calyx lobes becoming longer by tearing.
16-4. S. cochinchinensis ssp. leptophylla
25. Calyx regularly 5-lobed.

A” a 16-4. S. cochinchinensis ssp. leptophylla
26. Disk hairy.
Zia eects and bractecles caducous: i.) eos a. Sle vee 21. S. cylindracea
NOT hehe vive Sodan 6 16-4. S. cochinchinensis ssp. leptophylla

KEYS TO FRUITING MATERIAL ARRANGED BY ISLANDS AND ISLAND GROUPS

Sumatra
1. Midrib prominent on the upper surface.
2. Twigs hairy. Terminal buds hairy, small. Seeds straight... .......4.4. 4. S. anomala
2. Twigs glabrous. Terminal buds glabrous. Seeds curved . © 6. 1 ee ee 35. S. lucida

1. Midrib impressed on the upper surface.
3. Underside of leaves glabrous.

232 FLORA MALESIANA [ser. I, vol. 8?

4. Twigs hairy.
5. Leaves distichous . . ee ee a es ee ee eee me Ss foe ees
5. Leaves spirally arranged.
6. Leaves crowded towards the end of the twigs. Twigs thick, per towards apex. Petiole more
than 20 mm. Apex of leaves rounded or acute. .. . ; j . . . . 44. S. polyandra
6. Leaves evenly distributed.
7. Fruits ampulliform. Inflorescence a (basally branched) spike. Seeds not straight.
16-1. S. cochinchinensis ssp. cochinchinensis
7. Fruits ellipsoid.
8. Fruits 10-12 mm long. Inflorescence a fascicle. Bracts persistent . . . . 5. S. atjehensis
8. Fruits 7-10 mm long. Inflorescence a (basally branched) raceme. Bracts caducous
47. S. robinsonii
4. Twigs glabrous.

Oh NervesilessithancS) palrSanune). at ache mre. we) A owns a oe eS ahinensis
9. Nerves more than 5 pairs.
10. Leaves distichous ..... ee er a ee eee ne ee. i. See iis

10. Leaves spirally arranged.
11. Inflorescence a (basally branched) spike.
12. Fruits ovoid to orbicular, 9-10 mm long. Twigs thick. Terminal buds large 6. S. barisanica
12. Fruits ampulliform to globose, 5-7 mm long... . 16-2. S. cochinchinensis ssp. laurina
11. Inflorescence not a spike (rarely a cone in bud).
13. Bracts persistent.
14. Petiole 12-17 mm. Fruits ellipsoid. Inflorescence a fascicle ...... . 5. S. atjehensis
14. Petiole 3-9 mm. Fruits ampulliform. Inflorescence a raceme or panicle . 42. S. ophirensis
13. Bracts caducous.

15. Fruits 1-celled. Inflorescence a rusty tomentellous panicle ..... 41. S. odoratissima

15. Fruits 2—3-celled. Inflorescence a raceme.
16. Leaf margin entire. Fruits c. 10 mm. Seeds straight 8. S. batakensis
16. Leaf margin not entire. Fruits 4-10 mm, the sterile cells larger than the fertile ones, towards
the base*fillediwith air. 6 ws a so ns i, GB, augthn tt as a ae 14. S. celastrifolia

3. Underside of leaves hairy.
17. Twigs glabrous.

18. Leaves distichous. Petiole 1-4mm ...... 14 ike. AL 33. S. laeteviridis
18. Leaves spirally arranged. Petiole more than 10 mm.
19. Inflorescence a rusty tomentellous panicle .......... . . . . 41. S. odoratissima

19. Inflorescence a spike or fascicle.
20. Leaves 8-21 cm. Reticulation faintly prominent. Inflorescence a fascicle. Bracts persistent.
Fruits 10-12 mm long. Seeds straight. Stone different from the following . . 5. S. atjehensis
20. Leaves 15-45 cm. Reticulation much prominent. Inflorescence a (basally branched) spike (a
cone in bud). Bracts caducous. Fruits 8-10 mm long. Seeds not straight. Stone with a transverse
CONStHICHONVAt ONE SIGE ==, aos ve ch as cs sy ve tw pay 48. S. rubiginosa
17. Twigs hairy.
21. Leaves distichous.
22. Fruits ampulliform. Inflorescence a fascicle. Bracts persistent. Stone ampulliform.
23. S. fasciculata
22. Fruits not ampulliform. Inflorescence not a fascicle. Bracts caducous. Stone not ampulliform.
33. S. laeteviridis
21. Leaves spirally arranged.
23. Upper sideofleavesihairy 5. 4... w. = = Sy NC ate he . . 3. S. adenophylla
23. Upper side of leaves glabrous.
24. Bracts persistent.
25. Fruits0=12;mmilongtand S=6immibroad ors au.) ce Lean Sere 5. S. atjehensis
25. Fruits to c. 10 mm long.
26. Stone ellipsoid. Leaf margin (and petiole) beset with closely spaced glands.
3. S. adenophylla
26. Stone ampulliform. Fruits 5-7 mm.
27. Nerves more than 10 pairs. Inflorescence a (basally branched) spike.
16-1. S. cochinchinensis ssp. cochinchinensis
27. Nerves less than 10 pairs. Inflorescence a fascicle. ......... 23. S. fasciculata
24. Bracts caducous.
28. Inflorescence not a spike.

29) infiorescence a Panicle SccGs MOL Stale itt amet ears se ron a 41. S. odoratissima
29. Inflorescence a (basally pranches) raceme.
30; -Anoleiof leafibase-cs 90°: so. es erated tole emcee reset tv ecm ae me 50 S. sumatrana
30% Angle: of leafibase 20=60° .. .. 5)... a) ne Se 47. S. robinsonii

28. Inflorescence a (basally branched) ‘spike or a cone.
31. Fruits to c. 10 mm long; mesocarp fleshy (shrivelled when dry) or thin, coriaceous.
32. Leaves longer than 15 cm. Fruits 1-celled. Angle of leaf base 20-40°. Stone with low ridges
and a depression or transverse groove near the base. Seeds not straight. . 48. S. rubiginosa

1977] SYMPLOCACEAE (Nooteboom) 233

32. Leaves 6-14 cm. Fruits 3-celled. Angle of leaf base c. 90°. Stone with low not interrupted

ridges or grooves or brain-like grooved. Seeds straight. ....... 50. S. sumatrana

31. Fruits 2240 mm long; mesocarp woody or corky.
ES he 15b. S. cerasifolia var. grandifolia
PRINCE OEP DANES ck es kk cn ek ees Cn ea 15. S. cerasifolia

Malay Peninsula
1. Midrib prominent on the upper surface.

2. Twigs hairy.
3. Leaves evenly distributed. Fruits ellipsoid ................. 4. S. anomala
3. Leaves crowded towards the end of the twigs. Fruits ovoid ....... 37. S. wikstroemifolia
2. Twigs glabrous.
4. Leaves crowded towards the end of the twigs. Fruits ovoid ....... 37. S. wikstroemifolia
a7 beaves evenly distributed. Fruits ellipsoid ......4...092 00.2004 35. S. lucida
1. Midrib impressed on the upper surface.
5. Twigs hairy.

6. Leaves distichous.
7. Inflorescence a raceme or panicle. Fruits ovoid to ellipsoid, 7-12 mm. Bracts caducous.
33. S. laeteviridis
7. Inflorescence a fascicle. Fruits ampulliform. Bracts persistent ....... 23. S. fasciculata
6. Leaves spirally arranged.
8. Upper side of leaves hairy.
9. Angle of leaf base more than 90°. Bracts caducous. Hairs on twigs more than 2 mm long.

13. S. calycodactylos
9. Angle of leaf base less than 90°.
10. Fruits 8-10 mm long. Leaf margin (and petiole) beset with closely spaced glands.
3. S. adenophylla
10. Fruits more than 10 mm long. Leaf margin (and petiole) often glandular but glands not closely
oT ee a aes ee ye hm 20. S. crassipes

11. Leaves crowded towards the end of the twigs. Inflorescence a spike. Fruits ovoid, 10-12 mm.

37. S. wikstroemifolia
11. Leaves evenly distributed.
12. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular
ds: 3. S. adenophylla.)
13. Fruits cylindrical. Inflorescence a fascicle ............4.. 28. S. glomerata
13. Fruits ampulliform or ovoid. Inflorescence a spike or a panicle.
14. Fruits ampulliform. Inflorescence a (basically branched) spike. Seeds not straight.
16-1. S. cochinchinensis ssp. cochinchinensis
14. Fruits + ovoid. Inflorescence a panicle. Seeds curved ....... 41. S. odoratissima
12. Underside of leaves hairy (or pulverulent, nearly glabrous, in S. adenophylla).
15. Bracts caducous.

16. Stone smooth. Fruit 5-8 mm, ellipsoid .. . . 42-1b. S. ophirensis var. densireticulata
16. Stone with ridges or grooves.
17. Fruits 3-celled with 8 high ridges, 2240mm ........... 15. S. cerasifolia
17. Fruits 1-celled.
enor a Dames (SR. Ge. 4S Severe 41. S. odoratissima
18. Inflorescence a (basally branched) spike (a cone in bud). ...... 48. S. rubiginosa
15. Bracts persistent.
aueeee more tua 13-1Smmiong ......+s+s+v»rversuae 20. S. crassipes

19. Fruits to c. 10 mm long.
20. Stone ellipsoid. Leaf margin (and petiole) beset with closely spaced glands 3. S. adenophylla
20. Stone ampulliform.

. 21. Nerves more than 10 pairs. Inflorescence a (basally branched) spike. Seeds not straight.
16-1. S. cochi is ssp. nensis
21. Nerves 6-8 pairs. Inflorescence a fascicle. ..........444-. 23. S. fasciculata
5. Twigs glabrous.
22. Underside of leaves hairy.
23. Leaves crowded iienate the end of the twigs. Nerves 8-10 pairs. Stone smooth. Inflorescence a
e+ © we 6 @ wa © x Be oats ie dunes ee ee 37. S. wikstroemifolia
23. Leaves evenly distributed or nerves more than 10 pairs. Stone with ridges or grooves.
24. Leaves distichous. Inflorescence a raceme or panicle .......... 33. S. laeteviridis
24. Leaves spirally arranged. f .
25. Nerves less than 10 pairs. Bracts persistent. Fruits 13-18 mm long. Stone with low not inter-
rupted ridges or grooves or brain-like grooved. Seeds straight .... . 20. S. crassipes

25. Nerves 12-17 pairs. Bracts caducous. Fruits 8-10 mm long. Stone with low ridges and a depres-
sion or transverse groove near the base. Seeds not straight ....... 48. S. vebidioenn

234 FLORA MALESIANA [ser. I, vol. 8?

22. Underside of leaves glabrous.

Je, Pires . >. ddeunie hale Leeeein aaleaae 1 Aororg ao: 33. S. laeteviridis
26. Leaves spirally arranged.
STE antiGresccncemcrminiais 4 FOS. eo. OS ea en eee se SP 46. S. pyriflora

27. Inflorescence axillary.
28. Bracts caducous.
29. Inflorescence a spike. Fruits 25-40mm. .......... . . . 7. §. barringtoniifolia
29. Inflorescence not a spike. Fruits shorter than 25 mm.
30. Inflorescence a panicle.

Si wetiole 10 mines 2 SOR SRS 2 FL. eee eh) 20.0 39. S. nivea

46. S. pyriflora

Seewetriaie 10-0 mint 4 ais. sch: Mets oe SS SIG EL: branialiage 41. S. odoratissima

30. Inflorescence a (basally branched) raceme (rarely a cone in bud).

EHTS EY CONEOIE his. A Af Ghote SE et ee! a a eh a 14. S. celastrifolia
32. Fruits 1-celled.

33. Nerves 5-8 pairs. Terminal buds hairy ..........-24464-. 42. S. ophirensis

51. S. sumuntia

33. Nerves 9-14 pairs. Terminal buds glabrous. ........... 46. S. pyriflora

28. Bracts persistent.
34. Fruits ovoid to cylindrical.
35. Acumen shorter than 5 mm. Apex of leaves rounded or acute. . . . . 42. S. ophirensis
35. Acumen longer than 5 mm.
36. Nerves 10-16 pairs. Reticulation coarse. Inflorescence a fascicle. Fruits 7-10 mm long.
28. S. glomerata
36. Nerves 3-11 pairs. Reticulation fine. Inflorescence a (basally branched) spike. Fruits 13-18

ACCME EE Es Sih ayee. eer S re re PM eh ai ae ks 20. S. crassipes

34. Fruits ampulliform.
37. Petiole 2-9 mm. Inflorescence a raceme or panicle. ......... 42. S. ophirensis
37. Petiole 10-15 mm. Inflorescence aspike ...... 16-2. S. cochinchinensis ssp. laurina

Java & The Lesser Sunda Islands

ij. Midnb promment on the upper sariice 2.5. es ss. 2 La EA 35. S. lucida
1. Midrib impressed in the upper surface.
2. Twigs hairy.
3: Deavesidistichous 9 caces Cosine sins < WE Feet 23. S. fasciculata
3. Leaves spirally arranged.
4. Underside of leaves glabrous.

5 Riraltsis eo HTiniien ae, 12. et abyidiete «seers, lame avin wa. uh peeeet 41. S. odoratissima

Seb eitis sith Meee eee eS 16-1. S. cochinchinensis ssp. cochinchinensis
4. Underside of leaves hairy.

6. Upper side of leaves hairy (pulverulent) .............4.. .. 3. S. adenophylla

6. Upper side of leaves glabrous.
7. Seeds not straight. Embryo at least U-shaped curved.
8. Fruits ampulliform. Inflorescence a (basally branched) spike. Bracts persistent. Seeds twice
erinvedye so. OS) oS. Got ee ph ees oc) apg 16-1. S. cochinchinensis ssp.
8. Fruits not ampulliform. Inflorescence a panicle. Bracts caducous. Seeds U-shaped

41. S. odoratissima
7. Seeds straight. Embryo at most slightly curved.
9. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone not

CATHAY UTES 0 ies Poeea et G, cane el Seceme- a cle Vs. es EU kao
9. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not a spaced. Stone
PO HNOTtine ee Pk SL ae ee tet RR UES BES . S. fasciculata
2. Twigs glabrous.
RO a Rie ITN ERSERIALSRSR 5 DBS OF 1 USS © PGI og. sae ea 16. S. cochinchinensis

10. Fruits spindle-shaped or otherwise not ampulliform.
11. Inflorescence a (basally branched) spike or a cone.
12. Angle of leaf base less than 60°. Fruits 20-40 mmlong .......... 19. S. costata
12. Angle of leaf base more than 60°. Fruits to c. 10 mm long.
16-4. S. cochinchinensis ssp. leptophylla
11. Inflorescence not a spike (rarely a cone in bud).

13 sbetmimnalbads/largs:(7—10 sii). ese a eres! oe ey Dae 32. S. junghuhnii
13. Terminal buds small.
14. Inflorescence a panicle. Embryo curved ...........20468-. 41. S. odoratissima

14. Inflorescence a (basally branched) raceme. Embryo straight ...... 11. S. brandisii

a ia

1977] SYMPLOCACEAE (Nooteboom) 235

Borneo
1. Midrib flat or prominent on the upper surface .........:.2.2.2.4.. 4. S. anomala
1. Midrib impressed in the upper surface.
2. Twigs glabrous.
3. Underside of leaves hairy.
Deeeneeenous. Seeds Straight . . . .. :.... 2 «sb aebeeme eases 33. S. laeteviridis
4. Leaves spirally arranged
5. Leaves 4-6 cm. Nerves 6-9 pairs. Petiole34mm ........... 10. S. brachybotrys
5. Leaves 7-45 cm. Nerves 5-17 pairs. Petiole 10-50 mm.
6. Leaves 15-45 cm. Nerves 12-17 pairs. Inflorescence a spike ....... 48. S. rubiginosa
6. Leaves 7-40 cm. Nerves 5-16 pairs. Inflorescence a panicle ...... 41. S. odoratissima
3. Underside of leaves glabrous.
SP nePIMMIG (SN ees Ss kk eo el oe OS ea 33. S. laeteviridis
7. Leaves spirally arranged.
8. Bracts persistent.
nate EATAT SCT ow fc)... le Sar oe eee 12. S. buxifolia
9. Leaves longer than 5 cm.
PnCaeence NOL A SPIKG: (5 we ss es ne se 1 SS ee 42. S. ophirensis
10. Inflorescence a (basally branched) spike or a cone.
11. Petiole 3-10 mm. Leaves 6-22 mm. Nerves 6-13 pairs ........ 42. S. ophirensis

11. Petiole 10-25 mm.
12. Petiole 10-15 mm. Leaves 4/,-21 cm. Nerves 6-9 pairs.
1

6-2. S. cochinchinensis ssp. laurina
12. Petiole 15-25 mm. Leaves 17-30 cm. Nerves 11-13 pairs. . .... 29. S. goodeniacea

8. Bracts caducous.
13. Inflorescence a (basally branched) spike or a cone in bud.
SY Soe SETI est Se Se ss ee a, GS Se ee 10. S. brachybotrys
14. Petiole more than 5 mm. (When dubious under “15’’, look under 53. S. tricoccata.)
15. Leaves longer than 15cm. Nerves more than 10 pairs. Twigs thick. Terminal buds hairy.

Petiole more than 20 mm. Fruits254Omm........... 7. S. barringtoniifolia
15. Leaves shorter than c. 10cm. Nerves less than 10 pairs. Twigs not thick. Terminal buds
en ee ee 25. S. gambliana

13. Inflorescence not a spike (rarely a cone in bud).
16. Inflorescence a short spike, fascicle or panicle.
17. Fruits 1-celled. Inflorescence a panicle. Stone with ridges or grooves. —_= 1, not straight.
. S. odoratissima
17. Fruits 3-celled. Inflorescence a fascicle. Stone smooth. Seeds more than L ‘straight.

53. S. tricoccata
16. Inflorescence a (basally branched) raceme.
18. Leaves 1'/,-5 cm. Nerves 6-9 pairs. Fruits ellipsoid, 10-15 mm long . ._ 12. S. buxifolia
18. Leaves longer than 5 cm.
19. Apex of leaf rounded to faintly acuminate.
20. Leaves 5-8 cm. Nerves 6-9 pairs. Petiole 34 mm. Fruitunknown . 9. S. borneensis
20. Leaves 6-18 cm. Nerves 6—13 pairs. Petiole 3-10 mm. Fruit ovoid to ellipsoid, 5-12 mm.
42. S. ophirensis

19. Apex of leaf rather abruptly acuminate.

21. Fruit globose, 3-celled, the sterile cells filled withair. ...... 14. S. celastrifolia
21. Fruit ovoid to ellipsoid, | ey 42. S. ophirensis
2. Twigs hairy.
22. Leaves distichous.
23. Underside of leaves glabrous... ...-. 2 - - eee eee ees 33. S. laeteviridis

23. Underside of leaves vairy.
24. Leaf base 90-130°. Flowers solitary. Petiole 1-3 mm. Leaves ovate 2'/,-7 cm. Nerves 3-6 pairs.
Fruit narrowly flask-shaped,c.13 mmlong .......- 2+ + +e ees 31. S. johniana
24. These characters not combined. Fruits ellipsoid to ovoid.
25. Petiole 3-4 mm. Leaves 4-12 cm. Nerves 7-11 pairs. Base of leaves 40-90°. Inflorescence more
flowered. Bracts caducous.
26. Leaves 4-9 cm, acumen 9-16 mm, base 40-90° ........2.4-. 17. S. colombonensis
26. Leaves 9-12 cm, acumen 7-12 mm, base c.90° ... 1... 2 wees 33. S. laeteviridis
25. Petiole '/,-3 mm, if up to 4mm, inflorescence 1-flowered. Nerves 4-9 pairs.
27. Acumen 1-3 mm. Inflorescence 1-flowered. Leaves 2-3'/, cm. Bracts persistent.

52. S. trichomarginalis
27. These characters not combined.
28. Angle of leaf base 50-90°. Flowers solitary or in a few-flowered raceme. Bracts persistent.
Petiole 1-2 mm. Leaves 2'/,-5'/, cm. Nerves 5-9 pairs. Acumen c. 5 mm.

28, These characters not combined ......-..+0 60008085 33. S. laeteviridis
22. Leaves spirally arranged.

236 FLORA MALESIANA [ser. I, vol. 8?

29. Leaves crowded towards the end of the twigs. Twigs thick, with large leaf-scars.
44. S. polyandra
29. Leaves evenly distributed.
30. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular
glands: 3. S. adenophylla.)
31. Inflorescence a spike.

Bete Vedves 4-6 Cll PETOle 3 —4 INI aes eis. bos Ps on ERR Ae ee 10. S. brachybotrys

32. Leaves 6-25 cm. Petiole more than 5mm. Nerves more than 10 pairs. Fruits ampulliform. Stone

AEIUOL Es as Bake is o> «ss vs. oe Oe we 16-1. S. cochinchinensis ssp. cochinchinensis

31 SInniGrescence:a panicle’. 4. «AOE BOS, 2bes ti 2 seee, . . 41. S. odoratissima
30. Underside of leaves hairy.

Sora pper side of leaves hairy (pulverulent)mw 2. fae) oS a 3. S. adenophylla

33. Upper side of leaves glabrous.
34. Seeds not straight.

ST ECMO lees — Fate oy sess Pele a 5 ee es Sens as Go aa 3 10. S. brachybotrys
35. Petiole more than 5 mm.
36. Fruits ampulliform. Bracts persistent . . . 16-1. S. cochinchinensis ssp. cochinchinensis
36. Fruits not ampulliform.
Sp THHOIESCENICE a nPANICIG <2 ¢- -c.0. os. 8c, -< 4s. fo) vers, + en Se ee 41. S. odoratissima
37. Inflorescence a (basally branched) spike (a cone in bud). ...... 48. S. rubiginosa

34. Seeds straight.
38. Fruits to c. 10 mm long.
39. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone

PIIPSCIAM AN GON se eis ney nbcd a eke oh as See 3. S. adenophylla
39. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced.
Stone AMmpullitonities. t,o cs, ence welany fs) eat eee cote ct 23. S. fasciculata

38. Fruits more than 10 mm long.
40. Angle of leaf base more than 60°.
41. Angle of leaf base less than 90°. Twigs and underside of leaves (appressedly) pubescent,
puberulous or pilose. Fruits not ampulliform, 2—3-celled. Stone with ridges or grooves.

20. S. crassipes
41. Angle of leaf base more than 90°. Twigs and underside of leaves not appressedly pubescent
or puberulous. Fruits ampulliform, I-celled. Stone smooth ...... 31. S. johniana
40. Angle of leaf base less than 60°.
42>. Petiole 1—10 mim: Bruits 3-18 mmilong Ss eeesewsas fy. 8. eee 20. S. crassipes
42. Petiole 15-25 mm. Fruits 2240 mmlong ............ 15. S. cerasifolia
Philippines
cee eer Siew eae 242) Dicenlion shit wseq rere gop f- 55. S. verticillifolia
1. Leaves not verticillate.
2. Midrib prominent on the upper surface.
Seabee elabrous. Pchole shore than Simi << 6s! den Oe a a Ee 35. S. lucida
ge Er et le A ee oe eee ee er 34. S. lancifolia
2. Midrib impressed in the upper surface.
4. Twigs hairy.

5. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands:
3. S. adenophylla.)
6. Fruits spindle-shaped or otherwise not ampulliform.
7. Leaves evenly distributed, 41/,-71/, cm. Acumen 12-20 mm. Twigs not thick, cylindrical. Nerves
5—6 pairs. Petiole 7-8 mm. Fruits 1-celled. Seed 1. Apex of leaves acuminate . . 24. S. filipes
7. Leaves crowded towards the end of the twigs, longer than 10 cm. Acumen shorter than 5 mm.
Twigs thick, tapering towards apex. Nerves more than 10 pairs. Petiole more than 20 mm. Fruits

3-celled. Seeds more than 1. Apex of leaves rounded or acute ...... 44. S. polyandra

G (brens ampuiiforn >. 2} cia Sb pee ie 16-1. S. cochinchinensis ssp. cochinchinensis
5. Underside of leaves hairy.

ee mnen So 5 one SS) eS Oe See Eee 23. S. fasciculata
8. Leaves spirally arranged.

9. Leaf margin and petiole beset with closely spaced glands ........ 3. S. adenophylla

9. Leaf margin and petiole not so.
10. Fruits ampulliform.
11. Nerves more than 10 pairs. Inflorescence a (basally branched) spike.
16-1. S. cochinchinensis ssp. cochinchinensis

11. Nerves less than 10 pairs. Inflorescence a fascicle .......... 23. S. fasciculata
10. Fruits spindle-shaped or otherwise not ampulliform.
IES eR ee ol. ne pod ee Re ee ee oP eae 34. S. lancifolia

12. Petiole more than 5 mm.
13. Bracts and bracteoles caducous. Seeds not straight. Petiole 10-50 mm. 41. S. odoratissima

ee —

1977] SYMPLOCACEAE (Nooteboom) 237

13. Bracts and/or bracteoles persistent. Seeds straight. Petiole 7-8 mm .. . . 24. S. filipes
Bete 5 cig ber 8 ope 120s A et eee a Bae eee: 56. S. vidalii

4. Twigs glabrous.
DCR PRM Gk coke wo '. ns ee Os eee. 24. S. filipes

14. Underside of leaves glabrous.
15. Inflorescence a (basally branched) spike or a cone.
16. Petiole 1-3 mm. Fruit ellipsoid to globose, 3-5 mm. Inflorescence a spike . 34. S. lancifolia
16. Petiole longer than 3 mm.
17. Petiole 15-25 mm. Inflorescence a spike to 11/,mmlong. ........ 54. S. trisepala
17. Petiole shorter than 15 mm or inflorescence longer than 11/, cm.
18. Inflorescence a raceme or panicle of racemes. Embryo curved ..... 42. S. ophirensis
18. Inflorescence a spike. Embryo straight or curved.
19. Inflorescence a short spike to 1'/,cm. Embryo straight ...... 40. S. obovatifolia
19. Inflorescence a spike, longer than 1'/, cm. Embryo curved.
16-1. S. cochinchinensis ssp. cochinchinensis
15. Inflorescence not a spike (rarely a cone in bud).
20. Fruits 2—5-celled.
Peete With TOpes OF Grooves... « %24.%.'. GTi. +. <. xNemsals serene 27. S. glabriramifera

42. S. ophirensis
21. Stone smooth.

22. Acumen shorter than 5 mm. Inflorescence a fascicle to 1'/,cm. Bracts persistent. Fruits

more than 10 mm long. Seeds straight. .........2...4.. 40. S. obovatifolia

PREEEeTICSY LONPET COATS TOM hs os, 3 she ae Ose I ee 14. S. celastrifolia
20. Petiole 10-15 mm. Leaves 7-20 cm. Inflorescence a panicle. Fruit 8-25 mm, ovoid

41. S. odoratissima
an meaves 2-5*/,cm. Fruits ovoid; 5—-7'mm. .. .. .% <6 S)<esiset Shoe 57. S. whitfordii
23. These characters not combined.

24. Embryo straight. Leaves 4'/,-7!/, cm. Petiole7-8mm .......... 24. S. filipes
24. Embryo curved. Leaves 5-22 cm. Petiole 1-1Omm.......... 42. S. ophirensis

20. These characters not combined.

Celebes & The Moluccas

ipaeesame prominent on the upper surface .........6s5s8682s6 5s s¥ 58% 35. S. lucida
1. Midrib impressed in the upper surface.
2. Twigs hairy.
3. Leaves distichous.
rem Ot leaves CIADTOUS) | 4k «ie so ole wile) x Uae 33. S. laeteviridis
4. Underside of leaves hairy.
5. Fruits ampulliform. Inflorescence a fascicle. Bracts persistent ....... 23. S. fasciculata
5. Fruits not ampulliform. Inflorescence not a fascicle. Bracts caducous. . . . 33. S. laeteviridis
3. Leaves spirally arranged.
6. Leaves crowded towards the end of the twigs ............24-. 44. S. polyandra

6. Leaves evenly distributed.
7. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands:
3. S. adenophylla.)
8. Seed andembryotwicecurved ....... 16-1. S. cochinchinensis ssp. cochinchinensis
8. Seed and embryo uncinately curved towards the base.

16-4. S. cochinchinensis ssp. leptophylla
7. Underside of leaves hairy.

9. Upper side of leaves hairy (pulverulent) .........+4++64+4e46-s 3. S. adenophylla
9. Upper side of leaves glabrous.
10. Bracts caducous.
11. Stone smooth. Inflorescence a (basally branched) raceme.
42-1b. S. ophirensis var. densireticulata

11. Stone with ridges or grooves. Inflorescence a panicle of 5S-30cm_ . . 41. S. odoratissima
10. Bracts persistent.
OEE ES | a ee 16-4. S. cochinchinensis ssp. leptophylla

12. Seeds straight.
13. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone
GOES Geis. o Us he. eel ee) ei eee 3. S. adenophylla
13. Leaf i index 2-3. Leaf margin (and petiole) often glandular but glands not closely s a
EE ee ae 23. S.

2. Twigs glabrous.
14. Underside of leaves hairy.
15. Leaves distichous. Petiole 1-5 mm. Bracts caducous. — eo . . . . 33. S. laeteviridis
15. Leaves spirally arranged. ......-+5655856458 -4. S. cochinchinensis ssp. leptophylla

14. Underside of leaves glabrous.

238 FLORA MALESIANA [ser. I, vol. 8?

16. Fruits 2—3-celled.
17. Leaves longer than 15 cm. Fruits more than 10 mm long. Leaf margin entire. Nerves 9-14 pairs.
Stone with ridges or grooves. Seeds straight. ...........+ee-. 36. S. maliliensis
17. Leaves shorter than c. 15 cm. Fruits to c. 10 mm long. Seed and embryo U-shaped.
14. S. celastrifolia
16. Fruits 1-celled.

See eaves disticuouss eae a se ke, Os, AO fH, ©: RGA . . . 33. S. laeteviridis
18. Leaves spirally arranged.
19. Inflorescence a panicle of 5-30cm........ bee a aes 41. S. odoratissima

19. Inflorescence not a panicle or shorter than 5 cm.
20. Fruit stone with high, interrupted ridges which often protrude from the base.

42. S. ophirensis
BOMStone diferent ates seek IO AOS «5 el See & RG 16. S. cochinchinensis

New Guinea
(incl. New Ireland & New Britain)

1eeavesi(pseudo-)verticillate-" «.. Havel, 6. creer. ae ¢.coel) tats ee 30. S. herzogii
1. Leaves not verticillate.
2. Twigs glabrous.

3, Undersideiofileaves hairy). Sane & Oral. 2. 16-4. S. cochinchinensis ssp. leptophylla
3. Underside of leaves glabrous.
4. Nervesils 20 pais: weaves 21-620m ..... >. .sabltes oe Same 26. S. gigantifolia

4. Nerves less than 15 pairs. Leaves usually much smaller.
5. Twigs thick.
6. Acumen shorter than 5 mm. Fruits c. 13 mm long. Nerves 8-12 pairs . . . 45. S. pulvinata
6. Acumen longer than 5 mm. Fruits to c. 10 mm long.
16-4. S. cochinchinensis ssp. leptophylla
5. Twigs not thick.
7. Bracts persistent.
8. Inflorescence a panicle to 5 cm. Petiole 13-15 mm. Nerves 5-9 pairs . 18. S. composiracemosa

Svblantvdificrente) faci, 7 en 4. Gl eet. cite. as COR Se lt 16. S. cochinchinensis
7. Bracts caducous.
OMRTUISTC A MTON Rites, Guns voices Aeisti outs savas ek sUaeies te ae 21. S. cylindracea
9. Fruits to c. 10 mm long.
10-4b rts) 3-celled'(often'l or 2 aborted))<) sans . 5. SS CO 14. S. celastrifolia
Sa Sort (BES [150 EMR A am i oe 16-3. S. cochinchinensis ssp. thwaitesii

2. Twigs hairy.
11. Underside of leaves glabrous.
12. Inflorescence only 1-flowered.
seelceavesishorter than’. cm jie, & tills... . anal & 16-4. S. cochinchinensis ssp. leptophylla
13. Leaves longer than 5 cm.

14. Leaf index less than 2. Acumen shorter than 5 mm. Angle of leaf base less than 90°. Nerves less
than 5 pairs. Reticulation coarse. Fruits to c. 10 mm long. Seeds not straight. Apex of leaves
TOUNded Onacutomintwe: (er Vert U2) aah Awe 16-4. S. cochinchinensis ssp. leptophylla

14. Leaf index more than 2. Acumen longer than 5 mm. Angle of leaf base more than 90°. Nerves
more than 5 pairs. Reticulation fine. Fruits 17-22 mm long. Seeds straight. Apex of leaves

agminate) » <2: rebel Cane Uy aie orks 22.08) ies, Sea 38. S. multibracteata
12. Inflorescence more-flowered.
15; Petiolei0-Sinim 4otee ss, ie. tor Ps eee. 16-4. S. cochinchinensis ssp. leptophylla
15. Petiole more than 5 mm.
16. Seed and embryo (twice) curved ...... 16-1. S. cochinchinensis ssp. cochinchinensis

16. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla
11. Underside of leaves hairy.
i. Upper side wi leavesbalfy) 05 os 6 GH ew ae BO we SE 43. S. paucistaminea
17. Upper side of leaves glabrous.
18. Inflorescence only 1-flowered.
19. Leaf index more than 3. Angle of leaf base less than 90°. Reticulation coarse. 49. S. salicioides

19. Leaf index 2-3. Angle of leaf base more than 90°. Reticulation fine . . 38. S. multibracteata
18. Inflorescence more-flowered.
20). “Bractsicaducous;:Séeds straights. ers < Des eee ee ee ee 15. S. cerasifolia
20. Bracts persistent. Seeds not straight.
21. Seed and embryo (twice) curved ...... 16-1. S. cochinchinensis ssp. cochinchinensis

21. Seed and embryo uncinately curved towards the base.
16-4. S. cochinchinensis ssp. leptophylla

——————— ti“ il

1977]

3. Symplocos adenophylla WALL. (Cat. 1831,
n. 4427A, nomen) ex G. Don, Gen. Syst. 4 (1837) 3;
DC. Prod. 8 (1844) 257; Mia. Fl. Ind. Bat. 1, 2
(1859) 466; CLarKE, Fl. Br. Ind. 3 (1882) 575;
BRAND, Pfi. R. Heft 6 (1901) 48, incl. var. virgata
WALL. (Cat. 1831, m. 4427B, nomen) ex BRAND;
K. &G. J. As. Soc. Beng. 74, ii (1906) 240; BRAND,
Bull. Herb. Boiss. II, 6 (1906) 747, incl. var. atrata
BRAND, /.c. 748; MERR. Philip. J. Sc. 2 (1907) Bot.
298; BRAND, Philip. J. Sc. 3 (1908) Bot. 7, incl. var.
merrittii BRAND; RIDL. Fl. Mal. Pen. 2 (1923) 303,
t. 101, incl. var. montana RIDL.; Noor. Leid. Bot.
Ser. 1 (1975) 121. — S. bancana Mia. FI. Ind.
Bat. Suppl. 1 (1861) 476. — S. iteophylla Mia. l.c.,
incl. var. rostrata Mia. et var. elliptica M1Q.; MERR.
En. Born. (1921) 486. — Eugeniodes adenophyllum
O. K. Rev. Gen. Pl. 2 (1891) 410. — S. beccarii
BRAND, Pfi. R. Heft 6 (1901) 49. — S. constricta
BRAND, /.c. 41; MERR. En. Born. (1921) 486. —
S. fulvosa KING & GAMBLE, J. As. Soc. Beng. 74, ii
(1906) 233; Rip. Fl. Mal. Pen. 2 (1923) 300. —
S. palawanensis BRAND, Philip. J. Sc. 3 (1908) Bot.
10; Merr. En. Philip. 3 (1923) 301. —S. pruniflora
Rip. J. Fed. Mal. St. Mus. 4 (1909) 46; Fl. Mal.
Pen. 2 (1923) 304. — S. brandii Ever, Leaf.
Philip. Bot. 4 (1912) 1477. — S. pahangensis BRAND
in Fedde, Rep. 14 (1916) 326. — Fig. 7.

hrub or tree to 20m, 50cm @. Young twigs
pulverulent-puberulous or rarely tomentellous,
glabrescent, often dark-brown to blackish. Inno-
vations light redbrown. Leaves chartaceous to
coriaceous, often dark brown when dry, pulverulent
beneath or on both faces, soon glabrescent, elliptic,
acuminate, with cuneate base and recurved to
revolute margin with many pellucid glands, 4!/,-
16 by 1*/,-4°/, cm; nerves 4-12 pairs, meeting in a
looped intramarginal vein; petiole 6-12 mm.
Flowers in a spike, raceme or panicle to 6 cm; in-
dument of axis as twigs. Bracts and bracteoles with
same indument persistent in fruit, '/,-1 mm. Pedicel
mostly only under older flowers, to 3 mm. Calyx
nearly entirely divided into '/,-1 mm long lobes.
Corolla 2-5 mm. Stamens (20—)25-S0. Disk gla-
brous or rarely hairy. Ovary with same indument as
that of twigs, 1-2 mm high; style glabrous or with
some hairs towards the base, 2-4 mm. Fruit ellip-
soid to cylindrical, sometimes with c. 6 ridges when
dry, blue or black-purple, soon glabrescent,
crowned by the incurved calyx lobes, with only one
developed cell, 8-10(-11) by 3-5(-6) mm. Seed 1,
with straight embryo.

Distr. Continental Asia (China incl. Hainan,
Indo-China, Thailand), throughout Malesia, except
Java (but found in Bawean I.), the Lesser Sunda Is.
and New Guinea. A variety in Indo-China.

Ecol. Usually in montane rain-forest, in moun-
tain heaths, on ridge-crests and ridges, and mossy
forest, also in Baeckea-Leptospermum heath forest,
often on granite, but also on ultra-basic (Trusmadi),
from sea-level to m, but at low altitude
oe gl on podsolized sand (Banka; Bako N. P.)
and in heath forest on humid podsol. Fi. Sept.
(Febr.—Oct.), fr. May (Jan.—Dec.).

As is the case with more species, dwarfed speci-
mens or hardly 1 m high may already come into
flower.

Uses. The timber can be used for light construc-
tions (Descu, Mal. For. Rec. 15”, 1954, 593).

Vern. & Uses. Méndong, ménugan, Malaya,

SYMPLOCACEAE (Nooteboom)

239

kaju lattan, k. porugis, Sumatra, Batak, kayu kain,
W. Borneo, G. Klamm; the latter name alluding
to the use for tanning cloth in dyeing.

The Besisi (Mal. Pen.) believe that the leaves of
certain plants, e.g. S. adenophylla, if carried in the
quiver with their darts, act as charms bringing them
success in hunting (BuRK. Dict. 1935).

4. Symplocos anomala BRAND, Bot. Jahrb. 29
(1900) 529; Pfi. R. Heft 6 (1901) 67; Noor. Leid.
Bot. Ser. 1 (1975) 126, pl. la—-f, with full synonymy.
— §. concolor BRAND, Pfl. R. Heft 6 (1901) 65;
K. & G. J. As. Soc. Beng. 74, ii (1906) 242; Rip.
Fl. Mal. Pen. 2 (1923) 304. — Fig. 7.

Shrub or tree to 21 m, 40cm @. Young twigs
tomentellous to tomentose or appressedly pubes-
cent, glabrescent. Leaves glabrous, brownish or
olive to yellowish green glossy above, elliptic,
acuminate with cuneate-attenuate base and more
or less revolute finely glandular dentate to nearly
entire margin, 2'/,-12 by 1'/,-3 cm; midrib promi-
nent above or flat, rarely flat and sunken; nerves
5-11 pairs, meeting in a looped intramarginal vein;
petiole 2-7 mm. Raceme to 2 cm long, axis tomen-
tose to appressedly pubescent. Bracts 1-2 mm,
bracteoles !/,-1'/, mm, both persistent, with same
indument as axis. Pedicels 2-5 mm. Calyx lobes
rounded, ciliolate +/,-2mm. Corolla 4-6 mm.
Stamens 50 to more than 100. Disk tomentose or
shortly soft hairy. Ovary tomentose to (finely)
appressedly pubescent, c. '/,-1'/, mm high; style
glabrous or hairy towards the base, 4-7 mm. Fruit
3-celled, ellipsoid, violet, almost black, c. 10 by
6mm in Malaya, 10-13 by 6-8 mm in Borneo.
Seed 1 in each cell, straight with straight embryo.

Distr. Continental Asia (Burma, Thailand,
Indo-China, China incl. Hainan, Japan, Ryu
Kyu Is., Formosa) and Malesia: Malaya (incl.
Penang), N. Sumatra (incl. Banka), and Borneo.

Ecol. Mixed evergreen montane forest, also on
ridges and along streamsides, 700-2200m (in
continental Asia to 3000 m), but also found on
podsolized sands at very low altitude, 20-50 m, in
Banka. Fi. June—Oct., fr. Jan.—Dec.

Vern. Rénak, Banka.

5. Symplocos atjehensis Noor. Leid. Bot. Ser. 1
(1975) 128. — Fig. 7, 8.

Treelet to c. 8m, 10 cm @. Twigs glabrous or
tomentose. Leaves glabrous or sparsely appressedly
hairy, especially on midrib and nerves, elliptic,
acuminate, with acute to rounded base and dentate
margin, 8-21 by 3'/,-6cm; nerves 8-12 pairs,
meeting in a looped intramarginal vein; petiole
12-17 mm. Flowers in fascicles with persistent red-
brown tomentose to pubescent c. 2 mm long bracts
and bracteoles. Calyx 2mm, the (ciliate) lobes
1-1'/, mm. Corolla c. 5mm. Stamens c. 50. Disk
glabrous. Ovary glabrous without, c. 1 mm high;
style glabrous, 4-6 mm. Fruit ellipsoid, 10-12 by
5-6 mm, 3-celled, but only one cell developing;
stone shallowly lengthwise ribbed. Seed 1, straight
with straight embryo.

Distr. Malesia: N. Sumatra (Gajo Lands).

Ecol. Mixed evergreen mountain forest, 1700—
2850 m. Fil. Aug.—Sept., /r. July.

6. Symplocos barisanica Noor. Leid, Bot. Ser. 1
(1975) 130, — Fig. 7.

240 FLORA MALESIANA [ser. I, vol. 82

i

A

ah
NN

er «ws
ry

; a
GS
Lin

Sebi
;
Hi

Fig. 8. Symplocos atjehensis Noort. a. Habit, nat. size, b. flower, c. ditto after removal of petals and stamens,
both x 6, d. fruit, x 3 (a—c VAN STEENIS 6529, d DE WILDE 13773).

241

SYMPLOCACEAE (Nooteboom)

1977]

oT Sa,
ae, ‘£ TRS 4
SLOT Re
QQ USA
ae fe E OSA es *
Cy a Qs re - \

KER
Li A
<s : | \ *

Fig. 9. Symplocos batakensis Noor. a. Habit, nat. size. — S. barringtoniifolia BRAND. b. Fruit, c. ditto in

CS, both x 1'/, (a Ropinson & Kxoss 125, b-c KEP/FRI 10736).

242

Small tree, 6-10 m, 25cm @. Twigs glabrous.
Leaves glabrous, elliptic, with acute to nearly
rounded base, denticulate margin and acuminate to
rounded apex, 12-20 by 5-12cm; nerves 8-12
pairs, whether meeting in an intramarginal vein or
not; petiole 10-30 mm. Spike branched, to 5 cm
with minutely appressedly hairy axis. Bracts and
bracteoles persistent, + ovate, with same indument,
2-3 mm long. Calyx divided into semi-orbicular,
glabrous but ciliolate 1'/,-2 mm long lobes. Corolla
c. 6mm. Stamens c. 50. Disk glabrous. Ovary c.
1 mm high, glabrous; style glabrous, 4-5 mm, or
reduced. Fruit ovoid to globose, c. 10 by 9 mm with
globose to ampulliform stone of c. 8 by 7 mm (the
neck c. 2 mm long and the belly irregularly length-
wise grooved, c. 6mm high). Seed 1, U-shaped
with U-shaped embryo.

Distr. Malesia: Central
Kerintji and Merapi).

Ecol. Montane rain-forest, on Mt Kerintji in
Gleichenia woodland, 2000-2600 m. F/. June-July.

W. Sumatra (Mts

7. Symplocos barringtoniifolia BRAND, Ann. Cons.
Jard. Bot. Genéve 4 (1904) 283; Noor. Leid. Bot.
Ser. 1 (1975) 131, pl. 4. — Doxomma rigidum
Miers, Trans. Linn. Soc. II, Bot. 1 (1875) 104. —
Barringtonia rigida CLARKE, Fl. Br. Ind. 2 (1879)
510. — S. rigida CLARKE, FI. Br. Ind. 3 (1882) 581,
non G. Don, 1837; BRAND, Pfi. R. Heft 6 (1901)
52; K. & G. J. As. Soc. Beng. 74, ii (1906) 246;
Riwi. Fl. Mal. Pen. 2 (1923) 306. — Eugeniodes
ey O. K. Rev. Gen. Pl. 2 (1891) 976. — Fig. 7,
9b-c.

Tree to 25m, 40cm @. Twigs glabrous, often
marked with prominent orbicular scars of fallen
leaves; growth discontinuous, terminal buds pro-
tected by leathery scales, leaving conspicuous scars.
Leaves glabrous, elliptic to obovate with cuneate
base and acuminate apex, 15-35 by 6-11 cm;
nerves 10-14(-16) pairs; petiole 2-5cm. Spike
resembling a cone in bud because of the large
bracts, becoming 51/,(-8)cm; axis tomentose.
Bracts and bracteoles tomentellous to appressedly
pubescent, both soon caducous, broadly ovate,
6-10 by 6 mm and narrowly ovate, 2!/.-5 mm long
respectively. Calyx tomentellous, 3-31/, mm long,
the 5 lobes originally c. 1 mm long but the calyx
becoming 2-3-lobed by tearing. Corolla 4-6 mm.
Stamens c. 60 to more than 100. Disk glabrous.
Ovary glabrous, 1—-11/, mm high; style c. 5 mm,
with soft hairy conical base. Fruit ovoid or ellipsoid,
royal blue, 2'/,-4 by 11/,-2 cm, with chartaceous
mesocarp; stone stellate in cross-section with 8
very high ridges; cells 3, mag only 1 fertile. Seed
straight with straight emb

Distr. Continental Auia (Indo-China), in
Malesia: Malay Peninsula and Borneo (only once:
W. Kutei).

Ecol. Lowland rain-forest, river valleys in low
undulating country, on hillsides on clay, on dry
hillocks in Dryobalanops forest, but also on sand-
stone or granite, mostly below 300 m, but also in
Malaya more rarely in montane forest up to
1500 m. Fi. July—Aug., fr. Febr.—-May (July).

VERN. Médang, Malaya.

8. Symplocos batakensis Noor. Leid. Bot. Ser. 1
(1975) 132.— Fig. 7, 9a, 10a-d.
Twigs glabrous. Leaves often coriaceous, glab-

FLORA MALESIANA

[ser. I, vol. 82

rous, elliptic (to obovate) with acute base, entire
margin and acuminate apex, 6-10 by 2-41), cm;
nerves 7-10 pairs, meeting in an intramarginal
vein; petiole 5-8 mm. Raceme to 8 cm, axis glab-
rous or sparsely minutely pilose. Bracts and brac-
teoles with same indument, ovate, caducous, 11/,
and 1mm respectively. Pedicel to 2(—5) mm.
Calyx glabrous, 1'/, mm long, the semi-orbicular
lobes 1—1!/,mm long. Corolla c. 6mm. Stamens
c. 100. Disk shortly pilose. Ovary glabrous, 1!/, mm
high; style glabrous, c. 5 mm, sometimes reduced.
Fruit nearly globose, c. 10 by 8 mm, or ellipsoid-
ampulliform, c. 10 by 5 mm, 3-celled, often only 1
cell fertile. Seed often only 1, straight with straight
embryo.

Distr. Malesia: Central W. Sumatra (Tapanuli
and Westcoast Res.).

Ecol. Montane rain-forest on low ridges, 1200-
1700 m. Fr. Jan., Aug.

Vern. Loala lola, sihondung, Tapanuli.

Fig. 10. Symplocos batakensis Noor. a. Flowers and
buds, b. fruit, c. endocarp, d. ditto in CS, all x 2.
— S. cerasifolia WALL. ex DC. e. Fruit in CS, nat.
size (a-d ROBINSON & KLoss 125, e SAN 45168).

9. Symplocos borneensis BRAND, Pfl. R. Heft 6
(1901) 56; Merr. En. Born. (1921) 486; Noor.
Leid. Bot. Ser. 1 (1975) 134. — Fig. 7.

Twigs glabrous. Leaves narrowly elliptic, glab-
rous, with acute base, entire margin and rounded
to faintly acuminate apex (the acumen with broad
rounded tip), 4°/,-8 by 13/,-2"/, cm; nerves 6-9
pairs, faintly prominent beneath, meeting in an
intramarginal vein; reticulation hardly prominent ;

1977]

SYMPLOCACEAE (Nooteboom)

243

petiole 3-4 mm. Raceme lax, to 5 cm, axis minutely
sparsely hairy to glabrous. Bracts and bracteoles
caducous, glabrous, ciliolate, c. 1 and c. '/;mm
long respectively. Pedicel to 2 mm. Calyx glabrous,
divided into 3/,-1 mm long ciliolate lobes. Corolla
c. Smm. Stamens 60-80. Disk shortly pilose.
Ovary glabrous, 1-1'/, mm _ high; style glabrous
except the conical shortly pilose base, c. 5 mm long.
Fruit unknown.

Distr. Malesia: Borneo (Sarawak and W. Bor-
neo: Kenepai), 2 collections.

Ecol. Lowland rain-forest.

Note. A sterile collection from Central Celebes
(Malili) possibly belongs to this species.

10. Symplocos brachybotrys Merr. J. Str. Br. R.
As. Soc. n. 76 (1917) 110; En. Born. (1921) 486;
Herne, Pfi. Samml. Clemens (1953) 87; Noor. Leid.
Bot. Ser. 1 (1975) 134. — Fig. 7.

Twigs (sparsely) appressedly pubescent in inno-
vations, soon glabrescent. Leaves sparsely appres-
sedly fine hairy when young, soon glabrescent,
ovate to elliptic, with acute to rounded base,
denticulate margin and acute to acuminate apex,
4-6 by 2-3'/,cm; nerves 6-9 pairs, meeting in an
intramarginal vein; petiole 3-4 mm. Spike short,
1-3-flowered, axis at most 7mm, appressedly
pubescent, or flowers solitary, sessile from the leaf
axils and then several appressedly pubescent 2-4
mm long bracts. Bracts and bracteoles caducous,
in the spikes not seen. Ca/yx appressedly pubes-
cent, divided into the c. 3 mm long lobes. Corolla
6mm. Stamens c. 100. Disk glabrous, inconspi-
cuous. Ovary appressedly pubescent, 1 mm high;
style glabrous, c. 4mm. Fruit ovoid to ellipsoid,
intense indigo-blue, c. 10 by 5 mm, stone shallowly
lengthwise grooved. Seed 1, ovoid, slightly curved
with S-shaped embryo.

Distr. Malesia: Borneo (Sarawak and Sabah:
Mt Kinabalu).

Ecol. Mixed,
1500-1800 m.

11. Symplocos brandisii K. & V. Bijdr. 7 (1900)
157; BRAND, Pfi. R. Heft 6 (1901) 90; Koorp.
Atlas 2 (1914) t. 381; Back. & BAKH. f. Fl. Java 2
(1965) 206; Noor. Leid. Bot. Ser. 1 (1975) 135,
pl. 5. — S. koordersiana BRAND, Bull. Herb. Boiss.
II, 6 (1906) 748. — S. pseudoclethra HALL. f. Med.
Rijksherb. 14 (1912) 41.
Tree to 30 m, 40cm @. Twigs glabrous. Leaves
brous, mostly narrowly elliptic, with attenuate
, (coarsely) crenate margin and hardly acumi-
nate apex, (5'/,-)7-13(—22) by 2'/,-5(-6'/,) cm;
nerves (7—)10-16(-18) pairs, meeting in a looped
intramarginal vein; petiole 6-15 mm. Raceme to
10 cm, but often shorter, axis (sparsely) pubescent.
Bracts very soon caducous, appressedly pubescent,
obovate or elliptic, |-4 mm; bracteoles falling after
the bracts, sometimes rather long persistent, less
hairy, (broadly) ovate to narrowly elliptic, 1-2'/,
mm. Pedicels pubescent, at most 6 mm but often
shorter. Calyx glabrous, or some appressed hairs
on the base of the tube, 1'/,-2'/, mm long, the
lobes c. */,mm_ shorter, sometimes ciliolate.
Corolla c. 4-5) mm. Stamens 60 to more than 100.
Disk 5-glandular, glabrous. Ovary glabrous,
1-1'/, mm high; style glabrous or with few hairs,
but the conical base soft-hairy, 4-5 mm. Fruit

evergreen mountain forest,

ovoid to ellipsoid, slightly narrowed towards the
apex, 10-16 by 5—7 mm;; stone 1-celled, smooth or
faintly ribbed. Seeds 1 (or 2), filling the whole
stone, ovoid, with straight embryo.

Distr. Malesia: Java and Lesser Sunda Is.
(Lombok).

Ecol. From sea-level to 1800 m.

KEY TO THE VARIETIES

1. Leaves 5'/,-13 cm. Bracts obovate, 1-3 mm.
Stamens c. 60 . a. var. brandisii
1. Leaves 11-22 cm. Bracts elliptic, 3-4 mm.
Stamens more than 100 . b. var. pseudoclethra

a. var. brandisii. — S. brandisii K. & V. — S. koor-
dersiana BRAND. Cf. Noor. Leid. Bot. Ser. 1 (1975)
136, pl. Sa—h. — Fig. 7.

Leaves 5'/,-13cm long. Nerves 7-16 pairs.
Bracts obovate, 1-3mm; bracteoles ovate to
narrowly elliptic, 1-2!/,mm. Corolla c. 4mm.
Stamens c. 60.

Distr. Malesia: West Java (Udjung Kulon
Peninsula, Peutjang I. and Depok), East Java
(Besuki: Pantjur Idjén), and Lesser Sunda Is.
(Lombok: Mt Rindjani).

Ecol. Lowland primary and secondary forest, in
P. Peutjang on raised coral, in Java below 200 m,
in Lombok in montane forest at ‘800-1800 m’. Fi.
March-June, Nov., fr. July.

b. var. pseudoclethra (HALL. f.) Noor. Leid. Bot.
Ser. 1 (1975) 136, pl. Si-j. — S. pseudoclethra HALL.

Leaves 11-22 cm. Nerves 11-18 pairs. Bracts
elliptic, boat-shaped, 3-4 mm; bracteoles broadly
ovate, c. 2mm. Corolla 1'/; mm. Stamens more
than 100.

Distr. Malesia: Lesser Sunda Is. (Lombok: Mt
Rindjani). Only known from the type.

Ecol. Montane high forest, 800-950 m. F/. April.

12. Symplocos buxifolia Stapr, Trans. Linn. Soc.
Bot. 4 (1894) 206; BRAND, Pfl. R. Heft 6 (1901) 64;
Mere. En. Born. (1921) 487; Noor. Leid. Bot.
Ser. 1 (1975) 136, pl. 6a—d. — Fig. 7.

Shrub or treelet, 2-10 m; crown dense, globular,
fastigiate. Twigs glabrous, dark, + zigzag. Leaves
glabrous, closely placed, elliptic to nearly orbicular
with more or less attenuate base, finely glandular
dentate or crenate margin and rounded to acute or
slightly acuminate apex, 15-50 by 7-25 mm; nerves
4-6 pairs, meeting in an intramarginal vein; petiole
3-7 mm. Inflorescence an axillary few-flowered
raceme or often a 1I-flowered shoot with several
miniature sparsely pubescent to glabrous bract-
like leaves of 3 by 1 to 10 by 5 mm; axis glabrous or
minutely appressedly hairy. Bracts and bracteoles
caducous; pedicel between them to 2 mm. Calyx
glabrous or finely appressedly hairy, 2-5 mm long,
the lobes ciliate, 1-3 mm. Corolla 5-8 mm. Stamens
70 to more than 100. Disk glabrous. Ovary glab-
rous or rarely finely appressedly hairy, 2-3 mm
high; style amon 3-7 mm. Fruit ellipsoid to
ovoid, 10-15 by 6-8 mm; stone with low length-
wise ridges. Seed 1, straight with straight embryo,

Distr. Malesia: N. Borneo (Sabah; Mt Kina-
balu).

244

Ecol. Mixed, evergreen, subalpine low forest
and scrub, common, 2400-4000 m. F/. March—July,
Oct., Dec., fr. Febr.—Aug.

Note. This species can hardly be distinguished
from the mountain forms of S. cochinchinensis ssp.
leptophylla in New Guinea, especially those with
small orbicular leaves.

13. Symplocos calycodactylos BRAND, Pfl. R.
Heft 6 (1901) 63; Noor. Leid. Bot. Ser. 1 (1975)
137i iGe:

Shrub, 3m. Twigs densely spreadingly long-
hairy, hairs to 3 mm. Leaves long-hairy on both
surfaces, ovate to elliptic with rounded to sub-
cordate base, dentate, long-ciliate margin and
acuminate apex, 6-14 by 2!/,-5cm; nerves 7-8
pairs, meeting in an intramarginal vein; petiole
3-5 mm. Inflorescence a fascicle (or flowers soli-
tary?) or raceme to 10cm; axis long-hairy.
Bracts and bracteoles soon caducous, to 7 mm long,
narrowly elliptic clothed with long hairs. Pedicels
from ?5 mm in fascicles to 13 mm in racemes.
Calyx entirely divided into the narrow-elliptic to
linear, 4-6 mm long pubescent lobes. Corolla c. 6
mm. Stamens c. 100. Disk pilose. Ovary obscured
by the 3 mm long hairs, 11/, mm high; style glab-
rous, c. 8 mm. Fruit + cylindrical, densely long-
hairy, crowned by the persistent calyx (only young
fruits seen).

Oe
IRN

FLORA MALESIANA

[ser. I, vol. 8?

Distr. Malesia: Malay Peninsula (Perak and
Kedah), 2 collections.

Ecol. Evergreen hill forest, 900-1000 m. FY.
Febr.

Note. Closely allied to the Indian-Ceylonese
S. pulchra WiGuT with which there are hardly any
vegetative differences ; in flower easily distinguished
by the extremely long calyx lobes.

14. Symplocos celastrifolia Grirr. ex CLARKE, FI.
Br. Ind. 3 (1882) 575; BRAND, Pfl. R. Heft 6 (1901)
48; K. & G. J. As. Soc. Beng. 74, ii (1906) 239;
RID. Fl. Mal. Pen. 2 (1923) 302; MerR. Un. Cal.
Publ. Bot. 15 (1929) 248 ; FLETCHER, FI. Siam. En. 2
(1938) 385; Noor. Leid. Bot. Ser. 1 (1975) 138. —
Eugeniodes celastrifolius O. K. Rev. Gen. Pl. 2
(1891) 975. — S. nigricans BRAND, Pfl. R. Heft 6
(1901) 49. — S. candicans BRAND, |.c. — S. hutchin-
sonii BRAND, Philip. J. Sc. 4 (1909) Bot. 109; MErRrR.
En. Philip. 3 (1923) 299. — S. peninsularis BRAND,
Philip. J. Sc. 4 (1909) Bot. 110. — Fig. 4m, 7, 11.

Shrub or small tree, rarely up to 30 m high and
60cm @. Twigs glabrous. Leaves glabrous, or
rarely sparsely fine-hairy on midrib and nerves
beneath, often the upper surface dark coloured to
nearly black when dry and the undersurface olive
brown, + elliptic, with cuneate-attenuate base,
crenate margin and mostly abruptly acuminate
apex, 51/,-15 by 2'/,-6cm; nerves 6-9 pairs,

Fig. 11. Symplocos celastrifolia GrirF. ex CLARKE. a. Habit, x 2/3, b. bud, with bract and bracteoles,
c. corolla and stamens, both x 3, d. anther, e. stigma, both x 9, f. LS of flower, 5, g. CS of fruit, h.
LS of fruit, both « 9 (a MAIN 1258, b-h KOsTERMANS 1144, all from Morotai I.).

1977]

usually meeting in the intramarginal reticulation;
petiole 3-15 mm. Raceme often basally branched,
axis fine-hairy to appressedly pubescent, 3-12 cm.
Bracts and bracteoles soon caducous, 2—3(-4 in
Morotai) and c. 1'/, (or 2-2'/, in Morotai) mm
long respectively. Pedicels with same indument as
axis, 1-S mm. Calyx glabrous, 1'/,-2!/,(-3 in
Morotai) mm; lobes ciliate, when young I—1'/,mm,
becoming longer by tearing apart. Corolla 4-6 mm.
Stamens 40-c. 60. Disk glabrous, with some hairs or
pilose, especially after anthesis. Ovary glabrous, c.
1 mm high; style glabrous,4—-5 mm. Fruit orbicular,
pink, green, yellow or dark blue (sec. coll.), 4-10(—
20) by 3—-8(-15) mm; stone smooth, cells 3, but
usually only 1 fertile, the sterile cells larger than the
fertile ones, towards the base filled with air. Seed
and embryo U-shaped.

Distr. Peninsular Thailand and throughout
Malesia, except in Java, the Lesser Sunda Is., the
northern islands of the Philippines, the northern
half of Celebes, and most of the Moluccas. The
number of collections in Sumatra and East Malesia
(E. of Makassar Straits) is small compared with
those in Malaya and especially Borneo.

Ecol. Usually in coastal, primary and secondary
lowland forests especially in the transition zone
between mangrove (Nypa) and freshwater swamps,
mostly in deep marshy, sandy soils, but in a variety
of other habitats: sandy beaches, sandbanks near
the sea, kerangas, Casuarina peat swamp, in lalang
fields on white sandy soils, open heath forest
behind the mangrove, in Shorea laevifolia forest
(Nunukan), on a dry bamboo ridge at 300 m, also
on red or yellow sandy loams, exceptionally as
high as 750 m, and even 1900 m. Fi. March-May
(June-Jan.), fr. June-Aug. (Sept.—Jan.). Flowers
are noted to be fragrant. The fruits are obviously
buoyant, the sterile cells being filled with air.

Vern. Sumatra: kéndung, Palemb., krunjing,
Banka; Borneo-Sarawak: purup, Lundu; Sabah:
kayu tanyong, kulimbabok, tandjong jawa, tanjong-
tanjong, M, mangkasugoi, Mub., songal, Tengara,
inderatan, Bajau, balas, Banggi, enadak, inderopis,
lamai-lamai, mata kinai,_ tukil-tukil, Dusun;
Kalimantan: adad, Nunukan, bintangur pantai, E.
Kutei, mangkinang tikus, Kahajan, tawi, Sampit.

Notes. In Morotai I. a differing population is
found, with tomentose axis of raceme and bracts
and calyx lobes longer than in other specimens, and
growing at 800-1000 m. Fig. 11.

Also in West New Guinea (Vogelkop Peninsula)
deviating specimens are found with large, thicker-
walled fruits at c. 1900 m.

15. Symplocos cerasifolia Watt. (Cat. 1831,
n. 4434, nomen) ex DC. Prod. 8 (1844) 257; Mia.
Fl. Ind. Bat. 1, 2 (1859) 466, excl. stirp. Zoll.;
CLARKE, FI. Br. Ind. 3 (1882) 580; BRANp, Pfl. R.
Heft 6 (1901) 52; K. & G. J. As. Soc. Beng. 74, ii
(1906) 245; Ripv. Fl. Mal. Pen. 2 (1923) 306; Noor.
Leid. Bot. Ser. 1 (1975) 140, pl. 7c-f. — Bobua
cerasifolia Miers, J. Linn. Soc. Bot. 17 (1879) 304,
— Eugeniodes cerasifolium O. K. Rey. Gen. Pl. 2
(1891) 975. — Fig. 7, 10e.

a. var. cerasifolia. — Fig. 7, 10e. P
Tree to 25 m, 35cm @. Twigs often spreadingly

thin-pilose in innovations ; growth discontinuous;

terminal buds with many leathery scales, the latter

SYMPLOCACEAE (Nooteboom)

245

leaving conspicuous scars. Leaves long spreadingly
to more or less appressedly pilose beneath, espe-
cially on midrib and nerves, sometimes entirely
glabrous, with cuneate base, sharply dentate mar-
gin and acuminate apex, 7—16(—22) by 2-S(-7) cm;
nerves 6—9 pairs, meeting in a distinct looped intra-
marginal vein; petiole slender, 15-25 mm. Spike
resembling a short cone in bud as in S. barringtonii-
folia, becoming at most 3 cm long; axis + appres-
sedly long pilose to densely pubescent. Bracts
broadly ovoid to orbicular, boat-shaped, appres-
sedly (silky-)pubescent on the back, at least in the
middle, c. 5 by 5 mm; bracteoles with same indu-
ment, narrowly elliptic, c. 3 mm long, both soon
caducous. Calyx glabrous or slightly pubescent,
2'/,-4 mm, the lobes initially 1—11/, mm, becoming
often as long as the calyx by tearing apart. Corolla
c. 5mm. Stamens 30 to more than 100. Disk
glabrous. Ovary glabrous, 1-1!/, mm high; style
glabrous, but the conical base sometimes hairy.
Fruit ellipsoid, shiny blue, 22-40 by 8-18 mm;
stone with 8 high ridges, 3-celled with a central
canal, often only one cell developed. Seed cylindri-
cal, with straight embryo.

Distr. Extreme south of Peninsular Thailand;
in Malesia: Sumatra (also Banka), Malay Penin-
sula, Borneo, and West New Guinea (once, near
Merauke), showing a most unusual disjunction in
range.

Ecol. Lowland rain-forest, hillsides on granite,
on granitic sand, low ridges with sandy soil, also
sandy loam with lime, mostly below 200 m, rarely
ascending to 1000 m. Fi. June, fr. April—Oct.

Vern. Sumatra: séséham, Pakanbaru, ménta-
pung, méntépung, Banka.

b. var. grandifolia Noor. Leid. Bot. Ser. 1 (1975)
141

Leaves c. 30 by 8 cm. Nerves 10-14 pairs.
Distr. Malesia: NE. Sumatra (Asahan), 2 collec-
tions. Flowers unknown.

16. Symplocos cochinchinensis (LouR.) S. Moore,
J. Bot. 52 (1914) 148; GurLLaAumin, Bull. Soc. Bot.
Fr. 71 (1924) 277; Fl. Gén. L.-C. 3 (1933) 998;
Merr. Comm. Lour. (1935) 304; HAND.-MAzz.
Beih. Bot. Centralbl. 62 B (1943) 32; H. L. Lt, J.
Wash. Ac. Sc. 43 (1953) 107; Noor. Leid. Bot.
Ser. 1 (1975) 141, with full synonymy. — Dicalix
cochinchinensis Lour. Fl. Coch. 1 (1790) 663,
excl, syn. Arbor rediviva RuMPH. — Fig. 12, 13.

For the many synonyms see under the varieties.

Small shrub to large tree. Leaves very variable in
all characters. /nflorescence usually a spike, rarely a
raceme, but in ssp. leptophylla sometimes reduced
to a fascicle in the axils of the leaves or beneath
them, in ssp. thwaitesii sometimes a panicle of
racemes. Fruits ampulliform to globose, in ssp.
leptophylla and ssp. thwaitesii from globose to
ellipsoid, ovoid or ampulliform, in ssp. cochinchin-
ensis var. imbricata ovoid to ellipsoid. Seed and
embryo curved.

Distr. Continental Asia (India, Burma, Thai-
land, Indo-China, China, Japan, Ryu Kyu Is.,
Hainan, Formosa), throughout Malesia to Austra-
lia (Queensland, New South Wales, Lord Howe I.),
the Solomons, New Hebrides, and Fiji.

Notes. The oldest name for this species is
Myrtus laurinus Retz, 1786. However, its epithet

246

FLORA MALESIANA

[ser. I, vol. 8?

can not be used because of the heterotypic synonym
S. laurina WALL. ex G. Don, 1837.

This is the most widely distributed and also most
variable species of the genus. The two main forms
of the western part of its distribution, ‘cochinchinen-
sis’ and ‘laurina’, have usually been treated as
different species, the main difference being hairy
versus glabrous calyx lobes; in addition the bract
and bracteoles in cochinchinensis form a cup
appressed to and concealing the ovary while the
calyx lobes often enlarge in fruit forming a conical
beak. In /aurina the cup formed by the bract and
bracteoles is more platter-shaped, while the calyx
lobes form a small crown on top of the fruit, but
they can also be closed.

These two forms can be kept rather well apart in
large parts of the range, but in other parts they keep
less well separate and this results in a great varia-
bility, in part intergrading, which I have ascribed
to hybridization, while it is possible that from these
hybrid swarms new small local taxa may have
evolved through environmental conditions, e.g.
var. sessifolia and var. imbricata.

Towards the eastern end of the range, in New
Guinea, Australia, and the Pacific Islands forms
occur which often have no resemblance any more
to the two main western forms, but in the inter-
mediate area they are linked with them in a con-
tinuous variation, and thus break down any defin-
able distinction between them.

In these eastern forms, which I assume are
‘derived’ during the former eastward extension of
the range, some new tendencies have developed, in
that seed and embryo are only curved at the base
and are uncinate and that there is a tendency to-
wards unisexuality of the flowers. Several New
Guinean forms are further characterized by a
condensed fascicle-like inflorescence, while the
disk often becomes hairy.

Within the species 5 of the 9 pollen subtypes
known from subg. Hopea are found. The pollen
type is only constant for ssp. laurina and for ssp.
cochinchinensis and its varieties philippinensis and
sessifolia.

Instead of giving a lengthy discussion on the
variability I have found it more convenient and
clear to subdivide the species in formally named
subspecies and varieties, although I am aware that
it will not always be possible to name odd deviating
or intermediary specimens.

KEY TO THE SUBSPECIES

1. Seeds and embryo twice curved. Inflorescence a
basally branched spike, rarely a raceme. Flowers
bisexual. Disk always glabrous. Fruit ampulli-
form (ovoid to ellipsoid in ssp. cochinchinensis
var. imbricata).

2. Calyx lobes hairy (except on Mt Diéng in
Central Java), often enlarged in fruit, forming
a conical beak . . 1. ssp. cochinchinensis
2. Calyx lobes glabrous, often ciliate, not en-
larged in fruit 2. ssp. laurina

1. Seeds and embryo once curved. Inflorescence a
basally branched spike or raceme, or flowers
solitary or in a fascicle. Flowers bisexual or
functionally unisexual (or plant polygamous).
Disk glabrous or hairy. Fruit ellipsoid to ovoid
or ampulliform.

3. Seeds and embryo once curved. (Disk glabrous
or rarely pilose.) Calyx lobes glabrous, often
ciliate. Flowers bisexual . 3. ssp. thwaitesii

3. Seeds and embryo uncinately curved towards
the base. (Disk glabrous to densely pilose.)
Calyx lobes glabrous to densely hairy. Flowers
functionally unisexual or polygamous (in male
flowers the stigma is absent) 4. ssp. leptophylla

1. ssp. cochinchinensis.
For synonyms see under the varieties.

KEY TO THE VARIETIES

1. Leaves usually pubescent or tomentose beneath;
nerves 10-14 pairs, much prominent beneath,
strictly parallel to each other, nearly reaching the
margin; petiole (2—)5—17(—35) mm.

a. var. cochinchinensis

1. Leaves glabrous; nerves 4-11 pairs, usually not
strictly parallel to each other, anastomosing or
meeting in an intramarginal vein at some dis-
tance of the margin.

2. Leaves 3-12 by 1'/,-6 cm, index 11/,-3; nerves
4-8 pairs; petiole 0-3 mm. Fruit at most 7 mm
long . . b. var. sessifolia

2. Leaves 6-18 by 14/,- 6'/, cm, index 11/,—4/,;
nerves 5-11 pairs; petiole 3-25 mm. Fruit at
most 7 mm long c. var. philippinensis

2. Leaves 4-9 by 2!/,-51/, cm, index 1-2; nerves
5-7 pairs; wpetiae 4-7 mm. Fruit 10-12 mm
long . . d. var. imbricata

a. var. cochinchinensis. — Dicalix cochinchinensis
Lour. Fl. Coch. 1 (1790) 663, excl. syn. Arbor
rediviva RuMPH. — Dicalyx aluminosus BL. Bijdr.
(1826) 1117, p.p. — Dicalyx javanicus BL. l.c.
1117. — S. ferruginea Roxs. (Hort. Beng. 1814, 40;
WALL. Cat. 1831, 1. 4412, nomen) Fl. Ind. ed. Carey
2 (1832) 542; Mia. Fl. Ind. Bat. 1, 2 (1859) 466;
CLARKE, FI. Br. Ind. 3 (1882) 574; K. & V. Bijdr. 7
(1900) 141; BRAND, Pfi. R. Heft 6 (1901) 40; K. &
G. J. As. Soc. Beng. 74, ii (1906) 238; Koorp.
Atlas 2 (1914) t. 384; Rip. Fl. Mal. Pen. 2 (1923)
302. — S. mollis Wai. (Cat. 1831, n. 4433,
nomen) ex G. Don, Gen. Syst. 4 (1837) 3. — S.
spicata Roxs. var. platystachya G. Don, l.c. 2. —
S. polystachya WALL. (Cat. 1831, n. 4428, nomen)
ex DC. Prod. 8 (1844) 254; Mor. Syst. Verz. (1854)
43; ZoLL. Syst. Verz. 2 (1854) 136; Mia. FI. Ind.
Bat. 1, 2 (1859) 465. — S. verhuellii JuNGH. &
DE Vr. Pl. Ind. Or. 3 (1845) 12; Mia. Fl. Ind. Bat.
1, 2 (1859) 467. — S. horsfieldiana Mia. Sum. (1861)
475. — S. lachnobotrys Mia. l.c., incl. var. glabrior
Mia. — S. javanica Kurz, J. As. Soc. Beng. 40, ii
(1871) 64; ibid. 46, ii (1877) 239, excl. syn. S. rubi-
ginosa; MERR. Int. Rumph. (1917) 420; HEyYNeg,
Nutt. Pl. (1927) 1263; Burx. Dict. (1935) 2114;
Back. & BAKH. f. Fl. Java 2 (1965) 205. — Lodhra
javanica Miers, J. Linn. Soc. Bot. 17 (1879) 302. —
Lodhra ferruginea Miers, l.c. 299. — Lodhra
polystachya Miers, l.c. 300. — Lodhra verhuellii
Miers, /.c. 302. — S. ferruginea ROxB. var.
polystachya CLARKE, FI. Br. Ind. 3 (1882) 575. —
Eugeniodes ferrugineum O. K. Rev. Gen. Pl. 2
(1891) 975. — Eugeniodes lachnobotryum O. K. l.c.
— S. delectans BRAND, Bot. Jahrb. 54 (1916) 219.
— S. ferruginea Roxs. var. delectans KANEH. &
Hatus. Bot. Mag. Tokyo 56 (1942) 487. — Fig. 7.

1977]

Shrub or small tree, 9-22 m by 30 cm @, rarely a
large tree to 45 m by 80 cm @. Twigs rusty tomen-
tose or velvety, glabrescent, rarely pubescent,
appressedly pilose, or glabrous. Leaves rusty or
brownishly pubescent or tomentose beneath,
especially on midrib and nerves, rarely glabrous,
(ovate to) elliptic (to obovate) with cuneate, rarely
rounded or (in New Guinea) cordate base, glandu-
lar dentate or crenulate margin and more or less
acuminate apex, (6-)12-25 by (2'/,-)3-10cm;
nerves (8—)10—-14(-16) pairs, very prominent
beneath, parallel to each other, mostly quite
straight, curved upwards towards the margin and
nearly reaching it, whether forming an intramargi-
nal vein or not; petiole (2-)5-17 mm (rarely to
35 mm in New Guinea). Spike usually branched,
3-15 cm, in topodeme morobeensis up to 3 cm, axis
densely rusty tomentose or pubescent, in New
Guinea sometimes sericeous. Bracts and bracteoles
persistent, with same indument, the former at least
2 mm long and broad, but usually longer, excep-
tionally up to 10mm long, with the 2 smaller
bracteoles forming a calycle hiding the ovary.
Flowers faintly scented to fragrant. Calyx appres-
sedly pubescent (in topodeme morobeensis indument
only towards the apex), divided into (1-)2(-3) mm
long lobes. Corolla white (according to some collec-
tors with a yellow spot on each lobe), from 2
(sometimes in New Guinea) to 3-5 mm long.
Stamens 30-70 (in New Guinea from 10 at high
altitudes to more than 70 at low altitudes). Disk
glabrous. Ovary glabrous, */,-1 mm high; style
glabrous, 3-5 mm. Fruit ampulliform or globose,
5-7 by 45mm, more or less ribbed when dry,
often narrowed into a cylindrical neck, crowned by
the usually closed, enlarged, calyx lobes which form
a conical beak on top. Seed 1, twice curved with
similar curved embryo.

Distr. Continental SE. Asia (India, Burma,
Thailand, Indo-China, China, Hainan, Formosa,
Ryu Kyu Is., Japan) and throughout Malesia
except the Lesser Sunda Is., Celebes, and the
Moluccas, scarce in the Philippines.

Ecol. A variety of habitats over a considerable
altitudinal range, from the lowland up to c.
2500 m, in New Guinea even to 3000 m, in the
understorey of rain-forest, primary and secondary,
in the hills often associated with Eugenia and
Fagaceae, extending to a few exceptional condi-
tions, e.g. in Banka and Billiton on granite sands.
Fl. (Jan.-May) June-Sept. (Oct.-Dec.), fr. Oct.-
July. Ripe fruit dark blue. In Malaya crown shape
often called deep, domed, narrow and dense.

Vern. Sumatra: digéra, kédung, kémbang lonah,
Djambi, kayu njari badok, Lampong, kaju salon-
dung, k. si hondung, Padanglawas, kékaput,
Pasemah, /loba-loba, Batak, harbo, Tapa-
nuli, ménkéndung, Banka, sékéndum, sépandong,
Palembang; Java: djirak, S, ki huiit, Bantam; kayu
ara, Kota Belud, habo, Sg. Baru, kayu (h)abu,
Bandjar, Martapura; Philippines: fabu, Ifiago;
New Guinea: kumén, Wigote, Wapi lang., kutomi,
Wandammen lang., mirik, Sepik, Waskuk lang.

Notes. Var. cochinchinensis possesses rather
constant characters in large parts of its area,
especially in continental Asia. In Java glabrous
leaves become rather common, towards East Java
the number of nerves decreases, and the leaves
begin to resemble those of ssp. laurina. Here we

SYMPLOCACEAE (Nooteboom)

247

find the gradual transition to var. philippinensis.
The latter variety replaces var. cochinchinensis in
the Lesser Sunda Is., Celebes, the Moluccas, and
most of the Philippine islands.

A conspicuous population from the Morobe
District, New Guinea, is named topodeme moro-
beensis (petioles 15—35 mm, inflorescence up to 3 cm,
indument of calyx only towards the apex or on the
Margin).

b. var. sessifolia (BL.) Noor. Leid. Bot. Ser. 1 (1975)
153. — Dicalyx sessifolius BL. Bijdr. (1826) 1118.
— Dicalyx salaccensis Bu. l.c. — S. laurina (non
WALL.) Mor. Syst. Verz. (1845) 42. — S. sub-
sessilis CHoIsy (ex ZOLL. Syst. Verz. 2, 1854, 136,
nomen) ex Mia. FI. Ind. Bat. 1, 2 (1859) 467. — S.
sessi(li)folia GURKE in E. & P. Nat. Pfi. Fam. 4, 1
(1890) 170; BRAND, Pfl. R. Heft 6 (1901) 35;
Koorp. Atlas 2 (1914) t. 388; Back. & BAKH. f.
Fl. Java 2 (1965) 205. — Eugeniodes sessilifolius
O. K. Rev. Gen. Pl. 2 (1891) 409. — Eugeniodes
salaccense O. K. l.c. — Eugeniodes diengense O. K.
l.c. — S. spicata Roxs. f. subsessilis K. & V. Bijdr.
7 (1900) 146. — S. cochinchinense ssp. sessifolia
Noort. ex STEEN. Mt. Fl. Java (1972) pl. 52-4. —
Fig. 7, 12, 13c-e.

Shrub 1-5m to small tree, 10m, 10cm @.
Twigs glabrous or nearly so. Innovations purple.
Leaves glabrous, coriaceous, with cuneate-
attenuate base and faintly acuminate apex, 3-12 by
1'/,-6cm; nerves 4-8 pairs, meeting in a faint
intramarginal vein; petiole O-3(-5) mii. Spike
often branched, up to 6 cm, often crowdeu towards
the end of the twigs, axis densely appressedly
pubescent; flowers purplish. Bracts, bracteoles and
flowers as in var. cochinchinensis, but on Mt Diéng
the calyx only ciliate, or only pubescent towards
the margin. Calyx lobes on the fruit not enlarged
and closed.

Distr. Malesia: West & Central Java (Mts Salak
eastward to Sumbing).

Ecol. A constituent of the summit forest of the
volcanic peaks, often associated with Myrsine,
Leptospermum, Eurya, Schima, Photinia, and
Myrica, on stony ridges and summits, able to
invade exposed sterile rocky places in the vicinity
of craters as a dwarf pioneer shrub, 1700-3050 m.
Fl. mainly Oct.-Jan. (Febr.—March), fr. July-Aug.

For the ecology and flower biology see the
general paragraphs under the genus. Fruit blue-
black when ripe. Flush purple or blue-volet.

veh Flush is sometimes eaten as /alab (veget-
able).

Vern. Djirak, putat, S, djirik mélowo, sasah, J.

c. var. philippinensis (BRAND) Noor. Leid. Bot.
Ser. 1 (1975) 154. — Dicalyx aluminosus BL. Bijdr.
(1826) 1117, p.p. — S. spicata (non Roxs.) F.-VILL.
Nov. App. 4 (1880) 127. — S. syringoides BRAND,
Pfi. R. Heft 6 (1901) 41; S. Moore, J. Bot. 52 (1914)
148; Merr. Int. Rumph. (1917) 421. — S. fer-
ruginea Rox. var. philippinensis BRAND, Philip. J.
Sc. 3 (1908) Bot. 6. — S. ahernii BRAnv, L.c.;
Merr. En Philip. 3 (1923) 297. — S. ramosii MerRr.
Philip. J. Se. e (1917) Bot. 293; En. Philip. 3
(1923) 302. — S. ferruginea Roxs. var. syringoides
HALL. f. Beih. Bot, Centralbl. 39 B (1923) 92, —
ten (non Kurz) Merr, En. Philip. 3 (1923)

248

FLORA MALESIANA

[ser. I, vol. 8?

Fig. 12. Symplocos cochinchinensis (Lour.) S. Moore ssp. cochinchinensis var. sessifolia (BL.) Noort. in
fruit (and 1 flower), on summit of Mt Pangrango, West Java, at 3000 m (NooTEeBooM 906). Photogr.
Nooresoom, Febr. 1969.

Tree to 25 m high, 50cm @. Twigs glabrous or
appressedly pubescent. Leaves glabrous, or the
midrib sparsely appressedly fine-hairy, + elliptic
with cuneate base and acuminate apex, (4-)6-18 by
11/,-61/, cm, but 5-13 by 2!/,-51/, cm in the Philip-
pines; nerves 5—10(-15) pairs; petiole 3-15, in the
Philippines 10-25 mm. Spike with tomentellous to
pubescent axis. Bracts 1-2 mm, to 3 mm in the
Lesser Sunda Is., with the bracteoles with same
indument as the spike, further as in ssp. /aurina.
Calyx finely appressedly pubescent, divided into
c. 1mm long lobes. Corolla 3-6 mm. Stamens
35-70. Disk glabrous. Ovary glabrous, '/,-1 mm
high; style glabrous, 2-5 mm. Fruit as in ssp.
laurina.

Distr. Malesia: Central & East Java, Lesser
Sunda Is. (Bali, Sumbawa, Flores), Philippines
(common, throughout), Celebes, Moluccas (Tidore,
Ternate, Buru, Ambon, Ceram).

Ecol. In Java in mountain rain-forest, also in
tjemara forest, 700-2600 m, in the Lesser Sunda Is.
500-2400 m, in the Philippines from low altitude
up to 2000 m, also recorded from primary Diptero-
carp forest, in the Moluccas from low altitude to
1400 m. Fi. (Jan.-June) July—Dec., fr. Jan.—Aug.
Flowers said to be scented; fruit turning through
red to blue.

Vern. Java: kayu djurang, tjirug, J; Philippines:
abuabu, chaniusiu, gudik, \g., banatong-babde, Tag.,
tarangisi, Bag., ngarau-ngarau, Neg.; Moluccas:
bunga ajang, Ambon.

Note. In East Java this variety has probably
originated by hybridization between ssp. /aurina

var. laurina and ssp. cochinchinensis var. cochin-
chinensis.

d. var. imbricata (BRAND) Noor. Leid. Bot. Ser. 1
(1975) 155. — S. imbricata BRAND, Philip. J. Sc. 4
(1909) Bot. 109; ibid. 7 (1912) Bot. 31; MERR. En.
Philip. 3 (1923) 299. — Fig. 7.

Shrub or tree, 8-10 m. Twigs glabrous. Leaves
glabrous, more or less coriaceous, usually broadly
ovate, with cordate or slightly acuminate base,
glandular dentate margin, and acuminate apex,
4-9 by 21/,-5!/, cm; nerves 5-7 pairs. Spikes axil-
lary or pseudoterminal. Bract and bracteoles per-
sistent. Calyx more or less appressedly pubescent,
divided into 2 mm long lobes. Corolla c. 6 mm.
Stamens c. 60. Disk glabrous. Ovary glabrous,
c. 1mm high. Fruits black, ovoid to ellipsoid,
10-12 by c. 8 cm with smooth stone.

Distr. Malesia: Philippines (Luzon).

Ecol. In subalpine one-storey, mossy forest,
2000-2600 m. F/. Dec.—April, fr. Aug.—Sept., Jan.—
March. Innovations glossy redbrown, ripe fruit
dark blue.

Note. This mountain form is probably directly
derived from var. philippinensis, from which it
differs in the shorter leaves and larger fruit.

2. ssp. laurina (RETZ.) Noort. Leid. Bot. Ser. 1
(1876) 156. — Myrtus laurinus Retz. Obs. Bot. 4
(1786) 26.

Note. This subspecies ranges from Ceylon
eastwards to Celebes, China and Japan. Besides
the type variety there is only one local stenophyl-
lous variety in Indo-China and S. China.

1977]

SYMPLOCACEAE (Nooteboom)

249

Fig. 13. Leaf size and shape in Symplocos cochin-
chinensis (Lour.) S. Moore at different altitudes,
from a to e at 100, 1600, 71800, 2000, and 3000 m
respectively. a—b. ssp. laurina (RetTz.) Noor. var.

, c-€. ssp. cochinchinensis var. sessifolia (BL.)
Noor., all x +/,; (a LUTseEHARMS 4561, b BLOKHUIS
7-12-21, c Blume 1965, d Ja 4010, e Docrers VAN

LEEUWEN 8425).

e. var. laurina. — Laurus serrata floris spicatis
Burm. Thes. Zeyl. (1737) 139, t. 62. — Myrtus
laurinus Retz. Obs. Bot. 4 (1786) 26, non S. laurina
WALL. ex G. Don, 1837. — Drupatris cochinchin-
ensis Lour. Fl. Coch. 1 (1790) 314. — Decadia
aluminosa Lour. l.c. 315. — Eugenia laurina
WILLD. Sp. Pl. 2 (1799) 967, p.p. — Dicalyx
spicatus BL. Bijdr. (1826) 1118. — Dicalyx acumi-
natus BL. l.c. 1119. — S. spicata Roxs. (Hort.
Beng. 1814, 40; WALL. Cat. 1831, n. 4417, nomen)
Fl. Ind. ed. Carey 2 (1832) 542; Cuolsy in Zoll.
Syst. Verz. 2 (1854) 136; Mig. Fl. Ind. Bat. 1, 2
(1859) 465; CLARKE, Fl. Br. Ind. 3 (1882) 573,
incl. var. malasica l.c. et var. laurina l.c. p.p.; K. &
V. Bijdr. 7 (1900) 144, incl. f. javanica l.c. et f.
acuminata et f. xanthophylla l.c. 145, excl. f. sub-
sessilis; BRAND, Pfl. R. Heft 6 (1901) 39, incl. var.
acuminata l.c. 41; K. & G. J. As. Soc. Beng. 74, ii
(1906) 236; Koorp. Atlas 2 (1914) t. 386, 387;
RIDL. Fl. Mal. Pen. 2 (1923) 301; S. Moore, J.
Bot. 63 (1925) Suppl. 65; HEYNE, Nutt. Pl. (1927)
1263; Burk. Dict. (1935) 2115; BAck. & BAKH.
Sf. Fl. Java 2 (1965) 205. — S. laurina WALL. (Cat.
1831, n. 4416, nomen) ex G. Don, Gen. Syst. 4
(1837) 3; Rep. & Wis. in Sargent, Pl. Wils. 2
(1916) 594; Renp. J. Arn. Arb. 15 (1934) 298;
MErR. Comm. Lour. (1935) 303; CorNER, Ways.
Trees (1940) 623 ; HAND.-MAzz. Beih. Bot. Central-
bl. 62 B (1943) 33; Sreen. Fl. Mal. I, 5 (1957)
clxxxi, f. 4. — §. polycarpa WALL. (Cat. 1831,
n. 4423, nomen) ex G. Don, Gen. Syst. 4 (1837) 3;
Mia. Fl. Ind. Bat. 1, 2 (1859) 465. — S. ribes
JUNGH. & DE VR. Pl. Ind. Or. 3 (1845) 11; Mia. Fl.
Ind. Bat. 1, 2 (1859) 468. — S. acuminata Mia.
l.c. 467. — Lodhra ribes Miers, J. Linn. Soc. Bot.
17 (1879) 302. — Lodhra xanthophylla Miers,
l.c. — S. flavida Mig. (Pl. Hohenacker n. 1053) ex
CLARKE, Fl. Br. Ind. 3 (1882) 573, in syn. —
Eugeniodes ribes O. K. Rev. Gen. Pl. 2 (1891) 976.
— Fig. 7, 13a-b.

Shrub, 3 m, to tree, 6-14 m by 30cm @. Twigs
and leaves glabrous, except sometimes the very
youngest parts. Leaves + elliptic with cuneate base
and acuminate apex, 41/,-21 by (11/,-) 21/,-8 cm;
nerves 6-9 pairs (but in forms transitional to var.
cochinchinensis up to 13 pairs), not strictly
parallel, anastomosing at some distance of the
margin, often meeting in an intramarginal vein;
petiole (S—)10—15(—20) mm (in transitional forms
the leaves are like those of var. cochinchinensis
except for the indument). Spike 1'/,-14 cm, axis
glabrous to more or less appressedly puberulous or
pubescent. Bracts and bracteoles persistent, at
most 2 mm long and broad, but usually only 1 mm,
only enveloping the base of the ovary. A short
pedicel exceptionally present. Calyx glabrous or
nearly so, divided into 1-2 mm long, often ciliate
lobes, not elongating in fruit, whether or not
closed after anthesis. Rest of flower and fruit as in
var. cochinchinensis.

Distr. Continental Asia (India, Ceylon, Burma,
Thailand, Indo-China, China, Hainan, Formosa,
Japan); in Malesia; Sumatra (also Enggano L.),
Malay Peninsula (rare), Java (very common),
Borneo (rare), Celebes (rare), } ’

Ecol. Substage tree in rain-forest, sometimes in
coastal vegetation, near waterfall, in Malaya
found also in sandy, tidal gelam (Melaleuca) forest,
in continental Asia, Sumatra and Celebes from low

250

altitude to c. 2000 or 3000 m, in Java only above
c. 1000m. Fl. Sept.April, fr. Febr.—Sept.
Flowers are said to be slightly foetid to strongly
smelling, opening early in the morning. Fruit turns
black via blue.

Vern. Sumatra: kayu djari manuk, Batak,
dadak putih, diera, Enggano; Java: djirak, d. sasak,
S, djirék, J.

3. ssp. thwaitesii (F.v.M.) Noor. Leid. Bot. Ser. 1
(1975) 159, with full synonymy. — S. thwaitesii
F.v.M. Fragm. 3 (1862) 22.

Distr. This subspecies consists of 4 varieties
which occur in Queensland, New South Wales, and
Lord Howe I. One of these is also found in New
Guinea.

Note. There is one sheet (LAE 54751) which is
not identified to a variety; it might belong to the
Queensland var. montana (C. T. WHITE) Noort.

f. var. stawellii (F.v.M.) Noort. Leid. Bot. Ser. 1
(1975) 161. — S. stawellii F.v.M. Fragm. 5 (1865)
60; BRAND, Pfi. R. Heft 6 (1901) 37, excl. var. —
S. spicata Roxs. var. australis Bru. Fl. Austr. 4
(1869) 292. — Fig. 7.

Tree up to 30 m high, 80 cm @. Twigs glabrous.
Leaves glabrous, elliptic with broadly cuneate base
and not or faintly acuminate apex, 6-16 by 2!/,-10
cm; nerves 8-11 pairs; petiole 5-25 mm. Spike
often branched, rarely exceeding 6cm, the axis
glabrous or appressedly puberulous. Bracts and
bracteoles usually persistent, 1-1!/, and +/,-1 mm
long respectively. Calyx glabrous, divided into
1/,—3/, mm long lobes. Corolla 3-5 mm. Stamens
25-50. Disk glabrous. Ovary glabrous, 1—11/, mm
high; style glabrous. Fruit ellipsoid-ovoid, 5-7 mm.

Distr. Australia: Queensland, New South
Wales, and Lord Howe I.; in Malesia: New Guinea
(Papua).

Ecol. Two habitats are recorded, viz in the
lowlands with influence of a dry season, on edge of
savannah forest, and on the Oriomo R. in associa-
tion with Acacia, and in the middle mountains at
c. 2000-2300 m, in secondary forest, tall mixed
rain-forest, and in Podocarp-dominated forest on
peaty soil. F/. June, Sept., fr. June—Oct. Flowers
are recorded to be fragrant. Fruit develops from
green via blue to purple-black.

Vern. New Guinea: tuliper, Poio, Enga lang.,
kun’gum, Yogoo, Enga lang., truom, Oriomo R.,
Kiunga lang.

4. ssp. leptophylla (BRAND) Noort. Leid. Bot. Ser. 1
(1975) 162. — S. stawellii F.v.M. var. leptophylla
BRAND, Pfl. R. Heft 6 (1901) 37. — S. leptophylla
TURRILL, J. Linn. Soc. Bot. 43 (1915) 30.— S. mam-
beramo BRAND, Nova Guinea 14 (1924) 186.

For further synonyms see under the varieties.

Notes. This is a rather heterogeneous subspecies
ranging from the Lesser Sunda Is. and Moluccas
through New Guinea (incl. Bismarcks) to Mela-
nesia (Solomons, New Hebrides) and W. Poly-
nesia (Fiji), the type having been described from
Fiji. The varieties are rather reticulately allied and
are often connected by intermediate specimens
among which may be some hybrids. Some collec-
tions I could not refer to a variety, in part due to
inadequate material, e.g. the type of S. mamberamo.

In most varieties the flowers are functionally

FLORA MALESIANA

[ser. I, vol. 8?

unisexual or bisexual in the same variety. In the
functionally female flowers the number of stamens
is low, while the style is large, with peltate stigma.
In the functionally male flowers the number of
stamens is high and the style is small, without
stigma.

KEY TO THE VARIETIES

1. Underside of leaves hairy.
2. Twigs glabrous
2. Twigs hairy.
3. Calyx and ovary glabrous.
4. Disk hairy g. var. leptophylla
4. Disk glabrous. Twigs sericeous or tomen-
tose, % S. var. ovata
3. Calyx and/or ovary hairy.
5. Ovary glabrous.
6. Disk aeaas Twigs sericeous or tomen-
LOSE Sue a Sane eae Ss. var. ovata
6. Disk hairy.
7. Twigs (appressedly) pubescent.
g. var. leptophylla
7. Twigs tomentose or pilose.
v. var. versteegii

g. var. leptophylla

5. Ovary hairy.
8. Calyx glabrous.
9. Twigs sericeous or tomentose.
s. var. ovata
9. Twigs (appressedly) pubescent.
g. var. leptophylla
8. Calyx hairy.
10. Disk glabrous.
11. Bracts shorter than 3mm, fruits to
cnlOimm long” > eases s. var. ovata
11. Bracts longer than 3 mm, fruits more
than 10 mm long t. var. revoluta
10. Disk hairy.
12. Twigs (appressedly) pubescent.
g. var. leptophylla
12. Twigs not appressedly pubescent or
puberulous, e.g. tomentose.
13. Petiole more than 20.
j. var. tomentosa
13. Petiole less than 20 mm.
14. Inflorescence a (basally branched)
spike.
15. Bracts shorter than 3 mm
0. var. reginae
15. Bracts longer than 3 mm
t. var. revoluta
14. Inflorescence not a spike.
16. Nerves in 7-11 pairs.
m. var. molobros
16. Nerves in 4-8 pairs.
0. var. reginae
1. Underside of leaves glabrous.
17. Calyx and ovary glabrous.
18. Twigs hairy.
19. Petiole 0 to 5 mm.
20. Leaves shorter than 5 cm.
r. var. orbicularis
20. Leaves longer than 5 cm.
l. var. longilobata
19. Petiole more than 5 mm.
21. Leaves obovate, 10-25cm. Petiole 10-
40 mm. . i. var. insularis
21. Leaves ovate or elliptic, 2+/,-23 cm.
Petiole 5—25 mm.

ie

1977]

SYMPLOCACEAE (Nooteboom)

258

22. Leaves ovate or elliptic, 2'/,-11 cm.
Twigs sparsely appressedly pilose.
u. var. sogeriensis
22. Leaves + elliptic, 5-23 cm. Twigs
appressedly pubescent. g. var. leptophylla
18. Twigs glabrous.
23. Inflorescence a very slender, often branched
spike (or raceme) of 2-10 cm.
24. Twigs (exceptionally) thick.
p. var. schumanniana
24. Twigs not (exceptionally) thick.
25. Intramarginal vein far from margin.
p. var. schumanniana
25. Intramarginal vein close to margin.
w. var. maculata
23. Inflorescence a fascicle or a (reduced) often
branched, stout spike (or raceme).
26. Petiole 0 to 5mm . ©. var. orbicularis
26. Petiole more than 5 mm.
27. Bracts and bracteoles caducous. New
Hebrides.
var. aneityensis (BRAND) Noor.
27. Bracts persistent.
28. Reticulation fine, usually prominent on
both under and upper surface.
u. var. sogeriensis
28. Reticulation fine or coarse, usually only
prominent on the undersurface.
29. Leaves usually less than 5(-8) cm long.
h. var. monticola
29. Leaves usually more than 5 cm long.
30. Inflorescence axis glabrous.
q. var. floresana
30. Inflorescence axis hairy.
31. Leaves obovate. . i. var. insularis
31. Leaves elliptic or circular.
g. var. leptophylla
17. Calyx and/or ovary hairy.
32. Petiole 0 to 5mm
32. Petiole more than 5 mm.
33. Ovary glabrous.
34. Disk glabrous. Twigs glabrous. Reticula-
tion fine, usually prominent on both under
and upper surface, calyx lobes to c. !/, mm
pate ee ace ae u. var. sogeriensis

x. var. parvifolia

h. var. monticola
35. Leaves usually more than 5 cm long.
36. Inflorescence axis glabrous.
k. var. doormanensis
36. Inflorescence axis hairy.

g. var. leptophylla
33. Ovary hairy.
37. Twigs hairy.
38. Calyx glabrous . . _ g. var. leptophylla
38. Calyx hairy.
39. Calyx symmetrically cleft.

n. var. pedicellata
39. Calyx regular.
40. Leaves obovate . . . i. var. insularis
40. Leaves elliptic or circular.
g. var. leptophylla
37. Twigs glabrous.

41. Calyx —- . . . & var. leptophylla
41. Calyx hairy.

42. Inflorescence a very slender, often

branched spike (or raceme) of 2-10 cm.

p. var. schumanniana

42. Inflorescence a fascicle or a (reduced),
often branched, stout spike (or raceme).
43. Calyx 2-4-lobed or symmetrically cleft,
calyx lobes becoming longer by
tearing | (ooo n. var. pedicellata
43. Calyx regularly 5-lobed.

44. Leaves obovate . i. var. insularis

44. Leaves elliptic or circular.
g. var. leptophylla

g. var. leptophylla. — S. stawellii F.v.M. var.
leptophylla BRAND, Pfl. R. Heft 6 (1901) 37. —
S. leptophylla TurriLu, J. Linn. Soc. Bot. 43 (1915)
30, incl. f. compacta TuRRILL, I.c. 31. — S. pal-
marum BRAND, Bot. Jahrb. 54 (1916) 220. — S.
trifurceps BRAND, Nova Guinea 14 (1924) 186. —
S. rémeri BRAND, l.c. — S. aggregata WHITE &
FRANCIS, Proc. R. Soc. Queensl. 38 (1927) 256,
t. 17. — S. luteifolia KANEH. & Hatus. Bot. Mag.
Tokyo 56 (1942) 487. — S. turrilliana A. C. SMITH,
J. Arn. Arb. 33 (1952) 111. — Fig. 7.

Shrub 2-3 m to tree 20-28 m by 20-45 cm @.
Twigs glabrous or pubescent. Leaves glabrous or
pubescent to finely appressedly pilose beneath, +
elliptic, with cuneate to cordate base, entire to
dentate margin and acuminate apex, 5-23 by 2-12
cm; nerves 6-12 pairs, meeting in an intramarginal
vein; petiole 5-25 mm. Inflorescence a fascicle or a
reduced, branched spike, sometimes a spike or
raceme to 5cm, axis appressedly puberulous to
pubescent or sericeous. Bracts and bracteoles
persistent, with same indument, 1-10 and 1-4 mm
long respectively. Pedicels 0-2 mm. Flowers 3, 2, or
3. Calyx 3/,-3 mm, either entirely divided into the
hairy or glabrous lobes or not. Corolla 2-5 mm.
Stamens c. 10 to more than 100, in 2 flowers less
than 20. Disk softly hairy. Ovary glabrous or
pubescent to sericeous, 1—2'/,mm _ high; style
glabrous or with few hairs towards the base, small,
without stigma in functionally ¢ flowers, with
peltate stigma in 9 and §% flowers. Fruit glabrous or
sparsely pubescent, sessile in a fascicle or infruc-
tescence up to 5 cm or even more, ovoid to ellipsoid
or ampulliform, often globose, 6-15 by 4-9 mm.

Distr. W. Polynesia (Fiji), Melanesia (Solomons
and Santa Cruz Is.); in Malesia: Moluccas (Buru,
Ambon, Ceram) and very common in New Guinea
(incl. Jappen, Normanby, and Goodenough
Is.) and the Bismarck Archipelago (New Britain,
New Ireland).

Ecol. Very variable, rare in the lowland, mostly
from 900-3360 m (Mt Otto), in the lauro-faga-
ceous forest, transition of conifer-Castanopsis-
Nothofagus forest to grassland, mossy forest on
ridge tops, in forest relicts of Quercus-Dacrydium
forest (Arfak), once noted as a dominant on upper
ridges, in association with Podocarpus pilgeri in
New Britain, and in Casuarina forest there. FI.
Jan.-Aug., fr. Jan.-Dec. Flowers said to be fra-
grant. Fruit dark blue to purple black when mature,

Vern. New Guinea: aibeh, Minj, arilth, Non-
dugl, kelekende, Mt Ambua, koka, Telefomin,
Nah lang., guguma, konguma, kunguma, Mt Hagen,
Wankl lang., lelicop, Waria, matala, Mt Talawe,
New Britain, navako, New Britain, paiwiediedie,
Tari, Huli lang., peiwadidi, Mt Ne, Habono,
pungali, Wabag, tulifaro, ypap, Enga lang., toma,
Saidor, utu-utu, Cycloop Mt, Ormu lang., wapi,
Sepik, Wagu lang.

252

h. var. monticola Noor. Leid. Bot. Ser. 1 (1975) 166.
— Fig. 7.

Shrub 2m to tree to 16m by 22cm @. Twigs
glabrous. Leaves glabrous, + elliptic, with cuneate
base, entire or denticulate margin and acute or
rounded apex, 2-8 by 1-3 cm; nerves 5-7 pairs,
meeting in an intramarginal vein; petiole 4-10 mm.
Spike to 11/,cm long, axis glabrous or sparsely
appressedly hairy. Bracts and bracteoles persistent,
glabrous, 1-2 and 1-1'/, mm long respectively.
Flowers functionally unisexual or bisexual as in
var. leptophylla. Calyx appressedly pubescent or
puberulous to glabrous, usually divided into 1-1/,
mm long, often purple-tinged lobes. Corolla 1—2'/,
mm. Stamens 15-35. Disk hairy. Ovary glabrous,
1-1!/, mm high; style glabrous. Fruit ovoid to
ellipsoid, 8-10 by 4-6 mm.

Distr. Malesia: East New Guinea.

Ecol. Substage tree in mossy forest and secon-
dary forest with much climbing bamboo, 2700-
3500 m. Fl. April-Sept., fr. July—Aug.

Vern. Ped-ped, Giluwe, Mendi lang.

i. var. insularis Noor. Leid. Bot. Ser. 1 (1975) 167.
— Fig. 7.

Tree up to 15 m by 25 cm @. Twigs glabrous or
appressedly pubescent. Leaves glabrous, mostly
broadly ovate with attenuate base and acuminate
apex, 10-25 by 5'/,-15cm; nerves 6-8 pairs;
petiole 10-40 mm. Flowers not seen. Infructes-
cence a fascicle or spike to 51!/,cm long; fruit
aaa pubescent, ovoid to globose, 8-13 mm
ong.

Distr. Malesia: East New Guinea (Louisiades:
Sudest, Rossel & Misima Is.).

Ecol. Substage of rain-forest, along stream-
bank, also on a summit where dwarfed to 1!/, m
tall shrub; from the lowland to 800 m. Fr. July-
Oct. Ripe fruits black.

j. var. tomentosa Noor. Leid. Bot. Ser. 1 (1975) 167.

Tree to 20m. Twigs and midrib tomentose
beneath. Leaves mostly obovate, pubescent be-
neath, with cuneate base and rather abruptly
acuminate apex, 18-23 by 10-12cm; nerves 8-9
pairs; petiole 2-3 cm. Fascicle in the axils of the
leaves or often beneath them, including the broadly
boat-shaped 5mm long bracts, the 3 mm long
bracteoles and the calyx appressedly pubescent.
Flowers unisexual or bisexual as in var. leptophylla.
Calyx divided into 1-2 mm long lobes. Corolla
4-6 mm. Stamens 25 to more than 100. Disk softly
pilose. Ovary pubescent, 1-2 mm high. Fruit not
seen.

Distr. Malesia: East New Guinea (Fergusson

“Ecol. Montane rain-forest dominated by oaks,
in the substage, 700-900 m. Fi. June. Flowers said
to be very fragrant rose-scented.

k. var. doormanensis (BRAND) Noor. Leid. Bot.
Ser. 1 (1975) 168. — S. doormanensis BRAND, Nova
Guinea 14 (1924) 187. — S. dalmannensis KANEH.
& Hartus. Bot. Mag. Tokyo 56 (1942) 487.

Shrub or small tree, 11/, m. Twigs sparsely pilose
to glabrous. Leaves glabrous, coriaceous, elliptic,
with cuneate base, entire to glandular denticulate
margin and not or faintly acuminate apex, 6-12 by
21/.-6 cm; nerves 5-10 pairs; petiole 7-10 mm.

FLORA MALESIANA

[ser. I, vol. 8?

Fascicles in the axils of the leaves or on wood, in-
cluding the 5 mm long broadly boat-shaped bracts
and the 3-4 mm long bracteoles appressedly (long)
pilose to pubescent; bracts and bracteoles per-
sistent. Flowers unisexual or bisexual as in var.
leptophylla. Calyx appressedly pilose to pubes-
cent, 2-3 mm long, divided into the lobes. Corolla
4-5 mm. Stamens 30-50. Disk softly pilose. Ovary
glabrous, 11/,-2 mm high; style glabrous. Fruit
(immature) ellipsoid.

Distr. Malesia: New Guinea.

Ecol. Montane rain-forest, also in mossy forest,
1800-2700 m. Fi. Jan., fr. Oct.—Nov.

l. var. longilobata Noor. Leid. Bot. Ser. 1 (1975)
169. — Fig. 7.

Shrub or small tree, !/,-8 m by 15cm @. Twigs
sparsely appressedly fine-pilose, glabrescent.
Leaves glabrous, elliptic (to orbicular) with
rounded to more often cuneate base, crenulate
margin and acute (to rounded) apex, 10-23 by
6-14mm; nerves 2-4 pairs; petiole 2-3 mm.
Flowers unisexual or bisexual, solitary or c. 3 ina
condensed spike to 1 cm, axis pubescent. Bracts
and bracteoles persistent, 3-5S—6 together, narrowly
triangular, 3-S mm long. Calyx glabrous, 21/,-
41/, mm long, the lobes (ovate to) triangular, ciliate,
glandular, 2-4mm. Corolla 34mm. Stamens
14-24. Disk shortly pubescent. Ovary glabrous,
1-2 mm high, style glabrous, 2-4 mm. Fruits ovoid
to ellipsoid, c. 10 by 6 mm, stone rather smooth.

Distr. Malesia: East New Guinea (Mt Wilhelm).

Ecol. Alpine shrubberies and forest edges, in
subalpine tussock grassland, along creek in peaty
grassland, a stiff, fastigiate, microphyllous race, in
sterile exposed places often dwarfed, 3200-3400 m.
Fl. June-July, fr. July. Ripe fruit blue-black.

m. var. molobros (BRAND) Noor. Leid. Bot. Ser. 1
(1975) 169. — S. molobros BRAND, Bot. Jahrb. 54
(1916) 217. — Fig. 7.

Small shrub */,-11/,m to slender tree, 4-6 m.
Twigs densely (woolly) pilose. Leaves softly pilose
beneath, (broadly) elliptic, with cuneate to rounded
or even subcordate base, entire to glandular den-
tate margin and apex whether or not acuminate,
6-18 by 31/,-8 cm; nerves 7—11 pairs, meeting in a
looped intramarginal vein; petiole 5-10 mm.
Inflorescence a much reduced, branched, spike or a
fascicle in the axils of the leaves or on wood, up to
2cm long; axis rusty patently sericeous-pilose.
Bracts and bracteoles persistent, rusty long pilose
to appressedly sericeous, 2-4 and 1-3 mm respec-
tively. Calyx appressedly rusty sericeous or long
pubescent, divided into 1-2 mm long lobes. Corolla
2'/,-5 mm. Stamens 20-60. Disk pilose. Ovary
greyish sericeous, 1-2 mm high; style glabrous.
Fruit ovoid to globose, 10-15 mm long, pubescent,
becoming glabrous.

Distr. Malesia: New Guinea.

Ecol. Substage treelet in montane rain-forest, on
sandy clay, on limestone or sandstone ridges, 700—
2200 m. Fi. April—Nov., fr. Sept.

Vern. Chandujant, Wabag, Enga lang.

n. var. pedicellata Noor. Leid. Bot. Ser. 1 (1975)
170. — Fig. 7.

Shrub 2-4!/,m to slender tree 8-16 m. Twigs
glabrous. Leaves glabrous, stiff, + elliptic, with

sa... he eee eee

eben

1977]

cuneate to rounded base and (abruptly) acuminate
apex, 5-11 by 2'/,-6cm; nerves 6-10 pairs;
petiole 5-16 mm. Raceme up to 4 cm; axis sparsely
appressedly puberulous as the persistent 1-2 mm
long bracts and the 1-3 mm long pedicel. Calyx
appressedly puberulous, c. 2mm long, wholly
symmetrically cleft. Corolla 3-4mm. Stamens
c. 40 in 3 flowers, c. 10 in Q flowers. Disk softly
pilose. Ovary appressedly puberulous, 2 mm high;
style c. 3 mm, with conical pubescent base. Fruit
ovoid to ampulliform, 10-15 by 7-9 mm. Seed
strongly ruminate, embryo probably curved.

Distr. Malesia: East New Guinea.

Ecol. Substage of mossy forest and subalpine
forest dominated by Nothofagus-Weinmannia or
conifers (Araucaria, Podocarpus, Papuacedrus),
sometimes abundant on ridges, also on limestone,
2100-2900 m. Fi. (Jan.) April—Oct., fr. June.

Vern. Ypap, Wabag, Enga lang., keh, kepilam,
Enga lang.

0. var. reginae (BRAND) Noot. Leid. Bot. Ser. 1
(1975) 171. — S. reginae BRAND, Bot. Jahrb. 54
(1916) 214. — Fig. 7.

Shrub 1-2 m to small tree to 10m by 10cm @.
Twigs densely short and long pilose, only long-
pilose, or woolly to tomentose; growth discon-
tinuous. Leaves pubescent beneath, especially on
the nerves, elliptic, with acuminate to rounded
base, entire to glandular denticulate margin and
acuminate apex, 11/,-11 by 3/,—6'/,cm; nerves
4-8 pairs; petiole 2-10 mm. Flowers solitary or
few together in the axils of the leaves or below
them, or on the apical part of an up to 3(-7) cm
long spike; axis patently pilose. Bracts and brac-
teoles persistent, appressedly pilose, 2-4 mm and
1-2 mm respectively. Calyx appressedly pilose,
divided into 1—-1'/, mm long lobes. Corolla 2-3 mm.
Stamens 10-25. Disk pilose. Ovary appressedly
pilose, */,-2 mm high; style glabrous or with pilose
base. Fruit ovoid, pubescent, 9-15 by 7-8 mm. Seed
1-2, curved towards the base.

Distr. Malesia: New Guinea.

Ecol. Oak and beech forest, also on ridges, and
in river gorge, 900-2000 m. Fi. June-Aug., /r.
Jan.-Oct. Fruit from cream through purple to
purplish-blue when ripe.

Vern. Dorso, Kassam Pass, Kainantu, mongutl,
Hagen, harkomerinkey, Okapa, mamele, Morobe,
Wagau.

p. var. schumanniana (BRAND) Noor. Leid. Bot.
Ser. 1 (1975) 171. — S. rhynchocarpa K.Scu. ex
BRAND in K.Sch. & Laut. Nachtr. (1905) 347; Bot.
Jahrb. 54 (1916) 223. — S. schumanniana BRAND,
Lc. 347 et 224. — S. schlechteri BRAND, I.c. 348 et
224. — S. rupestris BRAND, Bot. Jahrb. 54 (1916)
220. — S. myrmecophila SCHLTR ex BRAND, L.c.
224. — S. pusilliflora S. Moore, Trans. Linn. Soc.
II, Bot. 9 (1916) 107. — S. cyclops BRAND, Nova
nae 14 (1924) 188. — S. lamii Branp, I.c. —

hrub 2m to tree 10-18m by 12-37cm @.
Twigs sometimes very thick, glabrous, sometimes
innovations app pubescent, often the
branches thickened in some places, hollow, lodging
ants. Leaves + elliptic, glabrous, with cuneate
base, + entire margin and acuminate apex, 9-33 by
3'/,-14 cm; nerves 8-15 pairs, meeting in intramar-

SYMPLOCACEAE (Nooteboom)

259

ginal vein far from the margin; petiole 5-22 mm.
Inflorescence a slender spike (or rarely a raceme) to
6 cm, often branched towards the base, rarely for
its whole length; axis pubescent or puberulous to
glabrous. Bracts and bracteoles mostly persistent,
rarely caducous, pubescent or puberulous, 1—2!/,
mm and ?/,-11/, mm long respectively. Pedicel if
present at most | mm. Calyx glabrous or puberu-
lous, entirely divided into c. !/, mm long lobes, or
11/, mm long and then the lobes c. 1 mm. Corolla
1'/,-S mm. Stamens 10-30 in 2 and 8% flowers,
30-80 in ¢ and § flowers. Disk pilose. Ovary glab-
rous or puberulous, 1—1'/, mm high; style glabrous
or with some hairs towards the base. Fruit ampulli-
form, 5-6 by 3-4 mm, sometimes with rather long
neck; stone ampulliform, rather smooth. Seed 1,
curved, U-shaped with U-shaped embryo.

Distr. Malesia: Moluccas (Morotai),
Guinea, New Ireland, and New Britain.

Ecol. In high lowland rain-forest, sometimes
with climbing bamboo, montane rain-forest on
ridges, also on sandy clay, in Nothofagus dominated
rain-forest on peaty soil, in New Britain also on
limestone, from sea-level to 2100(—2820) m. Fi.
Jan.—Dec., fr. July-Nov. Flowers are said to be
faintly fragrant. Fruits turn from green through red
to bluish when mature.

Vern. Moluccas: reha, Morotai; New Guinea:
pai, Wandammen, tembek, Telefomin.

q. = — Noor. Leid. Bot. Ser. 1 (1975) 172.
— Fig. 7.

Small, glabrous tree, up to 7m by 15cm @.
Leaves (broadly) elliptic with cuneate to rounded
base and not or slightly acuminate apex, 9-16 by
5-10 cm; nerves 7-12 pairs, meeting in an intra-
marginal vein; petiole stout, 2'/,-4'/, cm. Spike
basally branched, to 7 cm, axis glabrous. Bracts
and bracteoles persistent, glabrous or appressedly
pubescent, often ciliate. Calyx glabrous, divided
into c. 1 mm long lobes. Corolla 3-4 mm. Stamens
25-35. Disk glabrous. Ovary glabrous, */,-3/, mm
high; style glabrous. Fruit c. ovoid, 5-6 by 4-5 mm.

Distr. Malesia: Lesser Sunda Is. (Flores).

Ecol. Montane rain-forest, 1000-1500 m. Fi.
May-July, fr. April. Ripe fruit blue.

New

r. var. orbicularis (HEMSL.) Noor. Leid. Bot. Ser.
1 (1975) 173. — S. orbicularis Hems_. Kew Bull.
(1899) 105, — S. englishii Hemst. I.c. — S. klossii
S. Moore, Trans. Linn. Soc. II, Bot. 9 (1916) 108.
— Fig. 7.

Stiff, often compact, microphyllous treelet, with
densely foliaged twigs and patent, brittle, thick
(living + fleshy) leaves; 20-50 cm to 3-10 m by
35 cm @. Twigs glabrous or hairy. Leaves glabrous,
orbicular to elliptic, with cuneate to rounded or
slightly cordate base, dentate to denticulate margin
and rounded or acute apex, '/,—3(—3'/,) by */,-2 cm;
nerves 2-7 pairs; petiole 1-3 mm. Flowers solitary
or in a spike to 4.cm; bracts 1-3 mm, several when
flowers solitary, or 1. Bracteoles mostly persistent,
glabrous or hairy, '/;-3 mm long. eb me
entirely divided into 1—1*/, mm long lobes or a tube
of '/,-1mm_ present. Corolla 2'/,-4(-6) mm.
Stamens from less than 10 in @ flowers to 25 in ¢
and ¥% flowers. Disk glabrous. Ovary glabrous,
(‘/,-)1-2 mm high. Fruit ellipsoid, 7-15 by 4-6 mm.

Distr. Malesia; New Guinea.

254

Ecol. Subalpine grassland shrubberies (often
ericoid), sparse ridge top scrub, in moss-mounds
in ridge thickets, associated with Eurya, Dimorphan-
thera, Drimys, on creviced faces and ridges of
sandstone, also in subalpine moss forest, bank of
a mountain torrent, still recorded as a tree of 10 m
at 3300 m, 2500-3800 m, in Arfak as low as 1900 m.
Fl. June—Aug., fr. June—Sept.

Vern. Dibenkur, Chimbu, pombor, Giluwe,
Mendi lang.
s. var. ovata Noor. Leid. Bot. Ser. 1 (1975) 173. —
Fig. 7.

Shrub 3/,-4m to tree 12-21m by 15cm @.
Twigs appressedly sericeous to pubescent or tomen-
tose, glabrescent, rarely glabrous. Leaves appres-
sedly thin-hairy underneath, ovate to elliptic, with
cuneate to cordate base and acuminate apex, 4-12
by 2-7. cm; nerves 5-10 pairs; petiole 5-20 mm.
Spike basally branched, axis finely pubescent to
tomentose. Bracts and bracteoles persistent, with
same indument as axis or less hairy, 1-3 and 1-2
mm long respectively. Calyx glabrous but ciliate,
or appressedly fine-hairy, divided into */,—1+/,(—2)
mm long lobes. Corolla 2-3(-4)mm. Stamens
8-25. Disk glabrous. Ovary glabrous or sparsely
appressedly fine-hairy, 1—11/,(-2) mm high; style
glabrous. Fruit ellipsoid to ovoid, 5-10 by 3-8 mm;
stone ovoid, rather smooth. Seeds 1-2, ruminate,
fitting into the grooves of the stone.

Distr. Malesia: East New Guinea, very
common. ;
Ecol. Substage tree in tall mossy montane

forest, in association with Phyllocladus, in alpine
shrubberies, sometimes fire-induced, on margin of
bog grasslands, 1900-3700 m. Fi. Jan.—Dec., fr.
July-—Jan. Flowers are said to have a slightly fetid
fragrance.

Vern. Bolbeh, Chimbu, Masul, gongigl, miluad,
Chimbu, holai, Asaro, Kefamo, iamuga, Minj,
Togoba, kumbag, Togoba, kungum, Poio, Enga
lang., Kunguma, Goroka, Togoba, ontkumanip,
Wahgi, Minj, paiwadedie, Mt Ne, Huli lang.,
paiweriedie, Margarima R., Huli lang., pohn,
Hagen, Togoba, uinyambangau, Kubor, Minj,
wanépape, Sirunki, winjabunggawont, Minj, mara,
ypap, Wabag, Enga lang.

t. var. revoluta Noor. Leid. Bot. Ser. 1 (1975) 174.
— Fig. 7.

Shrub 1-3 m to tree 10m. Twigs appressedly
pubescent to villous or tomentose. Leaves
appressedly sericeous to pubescent or tomentose
beneath, especially on midrib and nerves, glabres-
cent, ovate to elliptic, with cuneate to cordate base,
strongly revolute or recurved margin and rounded
to acuminate apex, (2!/,-)4-10 by (1-)2!/,-6 cm;
nerves (4-)7-10 pairs; petiole (2-)10-15 mm. Spike
basally branched, to 3 cm, becoming much longer
in fruit, axis densely pubescent to villous or
tomentose; bracts often broadly boat-shaped,

—4mm. Bracteoles 2 mm, both persistent,
appressedly long pubescent to villous. Calyx with
same indument, (nearly) entirely divided into 1-2
mm long lobes. Corolla 2—4(—5)mm. Stamens
10-60. Disk glabrous, with few hairs, or densely
pilose. Ovary with same indument as calyx, 1-2 mm
high. Style glabrous. Fruit ovoid to ellipsoid, 10-11
by 6-7 mm. Seed more or less curved towards the

FLORA MALESIANA

[ser. I, vol. 8?

base, embryo from nearly straight to U-shaped.

Distr. Malesia: New Guinea.

Ecol. Mossy forest, alpine shrubberies, on
ridges and in valleys, constituent of subalpine
forest of Xanthomyrtus, Papuacedrus, Quintinia,
and Ericaceae, sometimes on peaty soil, 2200-
3600 m. Fi. Febr.-Aug., fr. July-Dec. Ripe fruit
purple blue.

Vern. Bug-bakl, Minj.

u. var. sogeriensis (BRAND) Noor. Leid. Bot. Ser. 1
(1975) 175. — S. sogeriensis BRAND, Pfi. R. Heft 6
(1901):49. — S. angiensis KANEH. & Hatus. Bot.
Mag. Tokyo 56 (1942) 485. — Fig. 7.

Shrub 2-5 m to tree 22m by 25cm @. Twigs
glabrous (or appressedly pilose in innovations).
Leaves glabrous, ovate or elliptic, with cuneate to
rounded base, mostly crenate margin and rounded
to faintly acuminate apex, 2'/,-11 by 1*/,-7 cm;
nerves 5-9 pairs; petiole 5-20 mm. Spike basally
branched to c. 3 cm, axis glabrous or appressedly
pilose. Bracts and bracteoles persistent, glabrous
or appressedly pilose, !/,-11/, and +/,-1 mm long
respectively. Calyx glabrous, or lobes shortly pilose
towards the apex, '/,-11/, mm, lobes !/, mm long.
Corolla 2-3 mm. Stamens less than 10 in 2 flowers,
to 30 in 3 flowers. Disk glabrous (or with few hairs).
Ovary glabrous, */,-11/, mm high; style glabrous.
Fruit (ovoid to) ellipsoid, 5-9 by 3-5 mm; stone
shallowly lengthwise or irregularly grooved.

Distr. Malesia: New Guinea.

Ecol. Montane to subalpine rain-forest and
subalpine scrubberies, in stunted Nothofagus-
Myrtaceae mossy forest, or forest dominated by
Castanopsis or by Podocarpus-Papuacedrus, scat-
tered in subalpine grasslands, on Mt Wilhelmina
even at 3560 m in sheltered places still a con-
stituent of 8-10m high stunted forest; (1950-)
2100-3560 m. Fi. Sept.—April, fr. Jan.—Nov. Fruit
turns bluish black when mature. Underside of
leaves has sometimes globular, pea-sized galls.

vy. var. versteegii (BRAND) Noor. Leid. Bot. Ser. 1
(1975) 176. — S. versteegii BRAND, Nova Guinea
14 (1924) 188.

Shrub or treelet to 5 m. Twigs densely tomentose
or pilose. Leaves elliptic, except the tomentose or
pilose midrib and nerves glabrous, or the whole
surface covered by a cobweb-like or a long-pilose
indument, base cuneate, apex not or slightly acumi-
nate to mucronate-caudate, 10-16 by 4-6'/, cm;
nerves 6-14 pairs; petiole 6-18 mm. Fascicles in
the axils of the upper leaves or on wood. Bracts
and bracteoles persistent, appressedly long pubes-
cent or sericeous, 4-5 and 2-3 mm long respectively.
Calyx with same indument, divided into 2-3 mm
long lobes. Corolla c. 5mm. Stamens c. 50. Disk
pilose. Ovary glabrous, 1-2 mm high; style glab-
rous. Fruit not seen.

Distr. Malesia: New Guinea.

Ecol. Rain-forest, 100 and 1300 m. Fi. Febr.,
June-July.

w. var. maculata (BRAND) Noort. Leid. Bot. Ser. 1
(1975) 176. — S. maculata BRAND in K.Sch. &
Laut. Nachtr. (1905) 348; Bot. Jahrb. 54 (1916)
222. — S. margarita BRAND, Bot. Jahrb. 54 (1916)
215. — S. pisifera BRAND, l.c. 216, incl. var.
miophylla BRAND. — S. ensicuspis BRAND, I.c. 219.

1977]

SYMPLOCACEAE (Nooteboom)

255

— S. arfakensis Gripss, Arfak (1917) 175. —
S. morobeensis SLEUM. in Fedde, Rep. 42 (1937)
265. — Fig. 7.

Shrub 1-2 m to small or moderate tree up to
15 m by 25 cm @. Twigs glabrous. Leaves glabrous,
+ elliptic with cuneate, decurrent base, mostly
entire margin and acuminate apex, 2-13 by 1'/,4
cm; nerves 4-10 pairs; petiole 3-15 mm. Spike very
slender, often branched, 2-10 cm, axis pubescent
or puberulous to glabrous. Bracts caducous or

’ persistent, 1-1'/, mm long, with the c. */, mm long

hi

bracteoles pubescent or puberulous to glabrous.
Calyx glabrous, divided into '/,-1 mm long ciliate
lobes. Corolla 2-4 mm. Stamens from less than 10
and sterile in 2 flowers to 25 in ¢ flowers. Disk
pilose. Ovary glabrous, */,-1'/,mm high; style glab-
rous. Fruit ovoid to ampulliform, 4-6 by 3-4 mm.

Distr. Malesia: New Guinea (incl. Sudest, Misi-
ma & Rossel Is.); common in New Guinea.

Ecol. Both in the lowland rain-forest at 150-
300 m (Louisiades) as well as in montane rain-forest
at 1600-2800 m, where associated with Nothofagus,
Araucaria and Castanopsis, on narrow crests
sometimes said to be abundant, also in secondary
forests. F/. Aug.—Jan. (June), fr. Aug.—Jan. Ripe
fruit blue-black.

Vern. Comogu, Mendi, kunguma, Minj, Togoba,
a Hagen, Togoba lang., ouksanok, Tele-
omin.

x. var. parvifolia Noor. Leid. Bot. Ser. 1 (1975)
177. — Fig. 7.

Shrub 1'/,-4 m to tree up to 10 m, often bushy
and much-branched. Twigs (appressedly) pubes-
cent or puberulous. Leaves glabrous, + elliptic,
with cuneate, attenuate base, denticulate or dentate
margin and acute or acuminate apex, 1'/,-4 by
3/,-1°/, cm; nerves 5-7 pairs ; petiole 2-4 mm. Spike
small, few-flowered, to 1 cm, axis puberulous.
Bracts and bracteoles persistent, puberulous, 1-2
and '/,-1 mm long respectively. Calyx appressedly
puberulous, divided into c. 1 mm long lobes.
Corolla 2-2'/, mm. Stamens c. 10 in 9 flowers to 25
in 3 flowers. Disk densely soft hairy. Ovary glab-
rous or appressedly puberulous, 1—1'/, mm high;
style glabrous or hairy towards the base. Fruit
ovoid to ellipsoid, 7-10 by c. 4mm.

a:

CY

Distr. Malesia: East New Guinea.

Ecol. Understorey treelet in lower montane to
subalpine rain-forest dominated by Nothofagus
and conifers (Podocarpus and Papuacedrus), often
mossy, also on forest edges, 1850-3300 m. Fi.
June-Oct., fr. Aug.

Uses. Flush is sometimes eaten as vegetable.

Vern. Gili, Ebenda, Mendi lang.

17. Symplocos colombonensis Noort. Leid. Bot.
Ser. 1 (1975) 177. — Fig. 7, 14a-c.

Small tree to 10 m. Twigs appressed-pubescent,
dark brown. Leaves alternate, sparsely appressedly
pilose beneath, especially on the margin, ovate,
with cuneate to rounded base often revolute margin
and acuminate apex, 4-9 by 1*/,-3'/; cm; nerves
7-11 pairs; petiole 3-4 mm. Raceme c. 3-flowered,
to 3cm long, axis finely appressedly pubescent.
Bracts and bracteoles soon caducous, pubescent.
Pedicel 1-5 mm. Calyx appressedly brown-pilose,
13/,-3 mm, lobes triangular, 1'/,-2'/, mm. Corolla
glabrous, or thinly red-hairy on the outside in bud,
c. 5mm. Stamens c. 90 or more. Disk glabrous or
with some hairs. Ovary appressedly brown-pilose,
1'/,-2 mm _ high; style glabrous, 4-5 mm. Fruit
(obliquely) ovoid to ellipsoid, 10-14 by 6mm;
stone except the apical 2-3 mm brain-like grooved.
Seed not seen, but embryo probably straight.

Distr. Malesia: Borneo (Mt Kinabalu).

Ecol. Mountain forest, 2100-2800m. Fi.
Febr.—March, June-July, fr. July, Dec.

Note. Resembles S. zizyphoides, but a tree with
less zigzag twigs, larger leaves with longer acumi-
nate apex, and with calyx lobes longer in propor-
tion to the tube.

18. Symplocos composiracemosa Noor. Leid. Bot.
Ser. 1 (1975) 178.

Twigs glabrous. Leaves glabrous, elliptic, with
cuneate, acute base, entire or slightly undulate
margin and acuminate apex, 8-13'/, by 2'/,-7 cm;
nerves 5—9 pairs, meeting in a looped intramarginal
vein; petiole 13-15 mm. Raceme compound, to
5cm; axis sparsely minutely pilose. Bracts and
bracteoles persistent, with same indument, | and
'/, mm long respectively. Pedicels at most 1 mm.
Calyx glabrous, divided into the rounded, semi-

Fig. 14. Symplocos colombonensis Noor. a. Habit, nat. size, b. fruit, c. CS of fruit, both « 3. — S. costata
(BL.) Cuorsy. d. CS of fruit, nat. size. — S. deflexa Starr. ¢. Habit, nat. size, f. deflorated flower, « 4
(a~c CLEMENS 33706, d Koorvers 10996, e-f NooTeBoom 1489).

256

FLORA MALESIANA

[ser. I, vol. 8?

elliptic, recurved, 1/.-1 mm long lobes. Corolla
c. 2mm. Stamens 15-25, rather stiff. Disk glab-
rous. Ovary glabrous, c. 1 mm high; style glabrous,
1 mm. Immature fruit elliptic.

Distr. Malesia: East New Guinea (Morobe
Distr.).

Ecol. Slender substage tree, 1300-1800 m, once
mentioned in understorey of Nothofagus dominated
ridge. Fi. Aug., Nov.

19. Symplocos costata (BL.) CHotsy in Zoll. Syst.
Verz. 2 (1854) 136; Mia. FI. Ind. Bat. 1, 2 (1859)
467; K. & V. Bijdr. 7 (1900) 153; BRAND, Pfl. R.
Heft 6 (1901) 52; Koorp. Atlas 2 (1914) t. 380;
Back. & BAKH. f. Fl. Java 2 (1965) 206; Noor.
Leid. Bot. Ser. 1 (1975) 179, pl. 8a—d, phot. 1-2. —
Dicalyx costatus BL. Bijdr. (1826) 1117. — S. cera-
sifolia (non WALL. ex DC.) Cuotsy in Zoll. Syst.
Verz. 2 (1854) 136; Mia. Fl. Ind. Bat. 1, 2 (1859)
466, pro stirp. Zoll. — S. caryophylloides ZO...
(Syst. Verz. 2, 1854, 136, nomen) Nat. Tijd. N. I. 14
(1857) 161; Mia. Fl. Ind. Bat. 1, 2 (1859) 467. —
Eugeniodes costatum O. K. Rey. Gen. Pl. 2 (1891)
975. — S. arcuata BRAND, Pfi. R. Heft 6 (1901) 58.
— §. sericea BRAND, /.c. 58; Bull. Herb. Boiss. II, 6
(1906) 748. — Fig. 7, 14d, 15, 16.

Tree to 20m, 40 cm @. Twigs glabrous, often
with cushion-shaped conspicuous leaf-scars, ter-
minal buds with many scales, 5-10 mm long.
Leaves glabrous, narrowly ovate to elliptic, with
cuneate, acute base, slightly dentate, nearly entire

margin and acuminate apex, 6-21 by 2-7 cm;
nerves (8—)10-13(-14) pairs; petiole 10-25 mm.
Spike from the axils of the leaves or on wood, in
bud resembling a cone like in S. barringtoniifolia,
becoming at most 4cm long, axis tomentose to
pubescent. Bracts and bracteoles densely sericeous
to pubescent, broadly boat-shaped, 5-8 mm long,
soon caducous, and 2-3 mm long, later caducous
respectively. Calyx glabrous, entirely divided into
(narrowly) ovate to triangular, 2'/,-3 mm long
lobes. Corolla 3-5 mm. Stamens 60 to more than
100. Disk shortly pilose. Ovary glabrous, c. */, mm
high; style glabrous except sometimes the very base,
3-6 mm long. Fruit ellipsoid to cylindrical, often
slightly curved, azure blue, 20-40 by 8-20 mm;
mesocarp thick, corky, stone with c. 8 high ridges,
3-celled with a central canal, often only 1 cell
developed. Seed cylindrical; embryo straight.

Distr. Malesia: West & Central Java (E as far
as G. Telemojo). Fig. 17.

Ecol. High mountain forest, 900-2000 m,
scattered. Fl. Aug.—Nov., fr. Aug.—-March.

Vern. Ki glédog (Tjibodas), ki télor, ki tomkil, S.

20. Symplocos crassipes CLARKE, Fl. Br. Ind. 3
(1882) 580; BRAND, Pfi. R. Heft 6 (1901) 52; K. &
G. J. As. Soc. Beng. 74, ii (1906) 245; RipL. FI.
Mal. Pen. 2 (1923) 305; Noor. Leid. Bot. Ser. 1
(1975) 180, pl. 9-10.

For synonyms see under the varieties.

Shrub or small tree to 18 m. Twigs glabrous, or

. Fig. 15. Symplocos costata (BL.) Cuotsy. Left a tree at Tjibodas Botanic Garden, West Java, 1450 m; right
a twig in bud (NooTEBOoM 885). Photogr. NooTEBooM, Febr. 1969.

1977]

SYMPLOCACEAE (Nooteboom)

257

Fig. 16. Symplocos costata (BL.) CHoisy. Close-up
of flowers in anthesis (NOOTEBOOM 885). Photogr.
Nooresoom, Febr. 1969.

(obliquely) pubescent to appressedly or spreadingly
long-hairy, sometimes with a double indument of a
short tomentum and long spreading hairs. Leaves
(narrowly) elliptic to ovate, beneath sparsely
appressedly pilose, nearly glabrous, to densely
appressedly to spreadingly long-hairy, rarely also
hairy above, with cordate to cuneate base, re-
curved, entire to glandular denticulate margin and
acuminate apex, 6-27 by 11/,-81/, cm; nerves 3-11
pairs; petiole 1-10 mm. Spike short, often clus-
tered, to 1(-2)cm, from the axils of the upper
leaves, rarely flowers solitary, axis subglabrous to
appressedly pubescent, or with long, spreading to
appressed, stiff, brown to rusty hairs 1-4 mm long.
Bracts and bracteoles persistent, (broadly) ovate,
triangular or semi-elliptic, rarely acuminate, hairy.
Calyx hairy or glabrous, whether or not entirely
divided into the lobes. Corolla 2'/,-6 mm. Stamens
c. 30 to c. 100. Disk glabrous. Ovary hairy, 1-2 mm
high; style glabrous, but often with conical, hairy
base. Fruits mostly 1-2 from each inflorescence,
glabrous or sparsely long-hairy, bright blue in
vivo, cylindrical, narrowed towards the apex,
13-18 by 3-5 mm; stone with c. 12 lengthwise
grooves; cells 1-3. Seed usually 1, straight with
Straight embryo.

Distr. Continental Asia (Peninsular Thailand),
in Malesia: Malay Peninsula (incl. Penang) and
Borneo.

KEY TO THE VARIETIES

1. Leaves ovate, to 6cm long and 2°/,cm wide;
nerves 3-6 pairs. Flowers solitary.
e. var. havilandii
1. Leaves (narrowly) ovate to elliptic, 5'/,-27 by
2-8'/, cm; nerves 3-11 pairs.

2. Leaf-base cordate, base angle 90-180°.
3. Leaves 5'/,-14 cm long. Petiole 1-2 mm
b. var. brandiana

3. Leaves 16-18 cm long. Petiole c. 5 mm.

a. var. crassipes
2. Leaf-base not cordate. Base angle 25-90°.

4. Leaves sparsely appressedly pilose beneath,
but the indument inconspicuous and leaves
seemingly glabrous.

5. Twigs glabrous, rarely appressed-pubescent.
Calyx often glabrous or nearly so, rarely
appressed-pubescent. Style-base glabrous,
rarely pilose -... . var. curtisii

5. Twigs appressed-long-hairy, rarely glabrous.
Calyx appressed-pubescent. Style-base pilose.

d. var. ernae

4. Leaves densely appressed-hairy to sparsely
more or less appressed-long-hairy beneath,
indument always evident.

6. Twigs densely patently brown hairy (hairs
often c. 2 mm). Leaves sparsely (appressedly)
long-hairy beneath. Nerves 6-11 pairs.

f. var. penangiana

6. Twigs densely obliquely pubescent. Leaves
densely appressed-pilose beneath. Nerves
4-6 pairs g. var. rufomarginata

a. var. crassipes.

Twigs sparsely appressedly long-hairy. Leaves
sparsely appressed-pilose beneath, the hairs incon-
spicuous, elliptic, with cordate base and acuminate
apex, 16-18 by 6-8 cm; nerves c. 10 pairs; petiole
much swollen, 5 mm. Inflorescence and flowers as
in var. brandiana (sec. CLARKE).

Distr. Malesia: Malay Peninsula (Johore), only
known from the type.

b. var. brandiana (K. & G.) Noor. Leid. Bot. Ser. 1
(1975) 182. — S. brandiana KiNG & GAMBLE, J. As.
Soc. Beng. 74, ii (1906) 245.

Small tree, 3-8 m. Twigs patently dark brown
pubescent to tomentose and long-hairy. Leaves
(appressedly) long-hairy beneath, but midrib and
nerves patently hairy, with cordate base, 5'/,-14 by
13/,-5 cm; nerves 6-10 pairs ; petiole 1-2 mm. Spike
often on a reduced twig with many cataphylls.
Bracts and bracteoles narrowly ovate, appressedly
long-hairy, 3-8 mm. Calyx divided into ovate,
acuminate, appressedly brown hairy, 2'/,-3 mm
long lobes. Stamens 60 or more, Style with hairy
conical base, 4mm. Fruit hairy.

Distr. Malesia; Malay Peninsula.

Ecol. Mixed forests, 100-1500 m.

c. var. curtisii (OLIv.) Noor. Leid. Bot. Ser. 1 (1975)
183, pl. 9b-c. — S. curtisii OLtv. in Hook. Ic. Pl. 18
(1888) t. 1757. — S. monticola Kinc & GAMBLE, J.
As. Soc. Beng. 74, ii (1906) 235; Ript. Fl. Mal. Pen.
2 (1923) 301. — Fig. 7.

Treelet or shrub to 10 m, 35cm @. Twigs glab-
rous or rarely appressed-pubescent. Leaves usually
sparsely appressedly pilose, nearly glabrous be-
neath, with cuneate, slightly attenuate base,
8'/,-18 by 3-8'/,cm; nerves 4-9 pairs; petiole
3-7 mm. Spike contracted, often branched, axis
glabrous to appressedly pubescent. Bracts and
bracteoles ovate to triangular, appressedly pubes-
cent, 1-1'/, and c. 1 mm long respectively. Calyx

*

258

glabrous or nearly so, rarely appressedly pubes-
cent, 11/,-2 mm, the lobes '/,-1'/, mm, becoming
longer by tearing apart when older. Corolla
31/,4mm. Disk glabrous. Ovary appressedly
pubescent, often narrowly funnel-shaped, 1*/,-2
mm high; style glabrous, the base glabrous or
pilose. Fruit glabrous, deep blue.

Distr. Continental Asia (Peninsular Thailand),
in Malesia: Malay Peninsula (Johore, Selangor).

Ecol. Hill rain-forest, 200-1400 m. Fl. Aug.-
Jan., fr. Febr._May, Oct., flowers scented.

Vern. Malaya: kayu jenerku, Selangor: Temuan.

d. var. ernae (BRAND) Noort. Leid. Bot. Ser. 1
(1975) 184, pl. 10b. — S. ernae BRAND, Pfli. R.
Heft 6 (1901) 58; Merr. En. Born. (1921) 486. —
Fig. 7.

Shrub or slender tree to 18 m, 15cm @. Twigs
appressedly (long—)hairy, rarely glabrous. Leaves
sparsely appressedly pilose, nearly glabrous
beneath, with cuneate base, 6-15(-18) by 2'/,-6
(-7) cm; nerves 3-6 pairs; petiole 3-5 mm. Spike
basally branched, contracted, axis appressedly
pubescent. Bracts and bracteoles broadly ovate,
often boat-shaped, appressedly pubescent, c. 1 mm
long. Calyx appressedly pubescent, 11/,-2 mm long,
lobes 1-1!/, mm, often becoming longer in older
stage. Corolla 3-5 mm. Stamens c. 30 to c. 70.
Disk glabrous. Ovary appressedly pubescent,
1-11/, mm high; style glabrous. Fruit glabrous.

Distr. Malesia: Borneo (Sarawak, Brunei,
Sabah; also in W. Kutei: G. Kemul).

Ecol. Lowland mixed Dipterocarp forest, also
in a swamp forest, and in hill rain-forest on sandy
clay, from sea-level to 1500m. Fi. Sept.—Oct.,
Febr.—June, fr. July, Nov.

e. var. havilandii (BRAND) Noor. Leid. Bot. Ser. 1
(1975) 184, pl. 10c. — S. havilandii BRAND, Pfl.
R. Heft 6 (1901) 41; Merr. En. Born. (1921) 486.

Treelet. Twigs pubescent. Leaves ovate, rather
densely appressed-pilose, especially on midrib and
nerves and along the margin, with rounded base,
23/4-6 by 11/,-23/, cm; nerves 3-6 pairs; petiole 2-3
mm. Flowers solitary, sessile from the axils of the
leaves. Bracts and bracteoles appressedly (long-)
pubescent, semi-orbicular 2mm long and ovate
11/,mm _ long respectively. Calyx appressedly
pubescent, 2 mm, the lobes 1!/, mm long. Corolla
2'/,mm. Stamens c. 35. Disk glabrous. Ovary
appressedly pubescent, c. 1 mm high; style with
pilose base. Fruit pale blue.

Distr. Malesia: Borneo (Sarawak).

Ecol. Hill rain-forest, 600-900 m. Fl. fr. July.

f. var. penangiana (K. & G.) Noor. Leid. Bot. Ser. 1
(1975) 185, pl. 9d. — S. penangiana KING &
GAMBLE, J. As. Soc. Beng. 74, ii (1906) 245; RIDL.
Fl. Mal. Pen. 2 (1923) 306. — Fig. 7.

Shrub or treelet to 10 m. Twigs densely patently
dark brown hairy. Leaves narrowly elliptic, sparsely
(appressedly) long-hairy beneath, especially on
midrib and nerves, rarely also long-hairy above,
with rounded to acute base, 6-27 by 2!/,-8 cm,
margin often sharply glandular dentate, appres-
sedly long-hairy beneath; nerves 6-11 pairs;
petiole 2-10 mm. Spike contracted, branched, axis
densely more or less appressedly villous, hairs
1-4 mm. Bracts and bracteoles appressedly dark

FLORA MALESIANA

[ser. I, vol. 8?

brown long-hairy, narrowly elliptic to ovate,
sometimes caudate, 1-7 mm. Calyx densely
appressedly dark brown hairy, entirely divided into
1'/,-4 mm long lobes. Corolla 3-6 mm. Stamens
30 to more than 100. Disk pilose. Ovary with same
indument as calyx, 1-1'/;mm high; style with
pilose base. Fruits hairy, pink.

Distr. Malesia: Malay Peninsula (incl. Penang).

Ecol. Lowland rain-forest, 150-500 m. Fi. May,
Sr. Nov., April.

g. var. rufomarginata Noor. Leid. Bot. Ser. 1 (1975)
185, pl. 10a.

Shrub or treelet to 5 m. Twigs densely pubescent.
Leaves rather densely appressedly hairy beneath,
ovate to elliptic, with cuneate base and margin
densely appressedly rufous-hairy beneath, 51/,-
11'/, by 2-3'/,cm; nerves 4-6 pairs; petiole
2-3 mm. Spike much contracted, axis hairy.
Bracts and bracteoles (broadly) elliptic, c. 3 mm.
Calyx densely, appressedly long sericeously pubes-
cent, entirely divided into 2mm long rounded
lobes. Corolla c. 2'/,mm. Stamens c. 25. Disk
glabrous. Ovary with same indument as calyx, c.
1 mm high; style hairy halfway up.

Distr. Malesia: Borneo (Sarawak, near Kuch-
ing).

21. Symplocos cylindracea Noor. Leid. Bot. Ser. 1
(1975) 187. — Fig. 7.

Tree 10-30m, 35cm @. Twigs glabrous, or
pubescent in innovation. Leaves glabrous, or mid-
rib (and nerves) minutely hairy beneath, + elliptic,
acuminate with acute to rounded, attenuate base
and crenate or crenulate margin, 9-15 by 31/,-
91/,cm; nerves 6-9 pairs, meeting in a looped
intramarginal vein; petiole 7-20 mm. Flowers inan
up to 8cm long panicle with minute or shortly
pilose axis. Bracts and bracteoles caducous, glab-
rous or minutely hairy, ciliate, ovate, c. 3 and c.
2mm long respectively. Pedicel !/,-3 mm, some-
times seemingly much longer when only one flower
is left on a small branch. Calyx 2-31/, mm long,
entirely divided into elliptic to nearly semi-orbi-
cular lobes, sparsely appressedly pilose to glabrous,
ciliate. Corolla 5-6 mm. Stamens more than 100.
Disk 5-glandular, pilose except the glands. Ovary
glabrous or pubescent, 1-1'/,mm _ high; style
(minutely) pilose, 2-4 mm. Fruit cylindrical, 15 by
5-6 mm, mesocarp fleshy, stone with low length-
wise ridges, 3-celled, 1, 2, or all 3 cells developed.
Seed 1 in each fertile cell, straight with straight
embryo.

Distr. Malesia: New Guinea (West and North,
Morobe Distr., Central Div., and New Britain).

Ecol. Plain rain-forest, also in Anisoptera forest
ri oe top, 60-800 m. Fi. Jan.—July, fr. Febr.—

arch.

22. Symplocos deflexa Stapr, Trans. Linn. Soc. 4
(1894) 205; BRAND, Pfl. R. Heft 6 (1901) 64;
Gisps, J. Linn. Soc. Bot. 42 (1914) 109; Merr. En.
Born. (1921) 487; Noor. Leid. Bot. Ser. 1 (1975)
188. — Fig. 7, 14e-f.

Treelet to 6 m high and 8 cm @. Twigs obliquely-
patently brown hairy, + zigzag. Leaves alternate,
glabrous above, rather densely pilose beneath,
especially towards the margin, elliptic, acuminate,
obtuse or acute, with rounded or sharply attenuate

1977]

base and recurved to revolute, sharply glandular
dentate margin, 3-5 by 1'/,-2'/,cm; nerves 5-7
pairs, usually merging into the reticulation; petiole
1-2 mm, densely patently brown hairy. Flowers
fragrant, in an up to 6-flowered, 1-4 cm long lax
raceme which is appressedly to patently brown
pilose in all parts except the corolla. Bracts and
bracteoles persistent, c. 5 by 3 and c. 3 by 1'/, mm
respectively. Pedicel 2-4 mm. Calyx divided into
obtuse and semi-elliptic to acute and triangular
lobes, c. 1'/, mm long. Petals 5—7, glabrous or the
outer ones minutely appressedly hairy, 4-6 mm
long. Stamens 60-90. Disk low, 5-glandular,
sparsely long-pilose. Ovary 1'/,-2 mm high; style
c. 5 mm, gradually thickened towards its base, the
lower half sparsely long-pilose. Fruit ovoid, often
curved, including the persistent calyx c. 10 by 5S mm;
stone c. 8 by 4 mm with shallow grooves and large
apical pore. Seed straight with straight embryo.

Distr. Malesia: Borneo (Sabah, only found on
Mt Kinabalu near Paka cave).

Ecol. Low subalpine forest and mountain

scrub, 2400-3200 m. Fi. Oct.—Febr., fr. March,
Aug.—Oct.
23. Symplocos fasciculata ZoLt. Syst. Verz. 2
(1854) 136; Nat. Tijd. N. I. 14 (1857) 161; Mia.
Fl. Ind. Bat. 1, 2 (1859) 467; Suppl. 1 (1861) 474,
incl. var. minor Mia. l.c. 475; CLARKE, FI. Br. Ind. 3
(1882) 574; K. & V. Bijdr. 7 (1900) 150, incl. var.
blumeana K. & V. l.c. 151; BRAND, Pfl. R. Heft 6
(1901) 34; K. & G. J. As. Soc. Beng. 74, ii (1906)
235; Koorpb. Atlas 2 (1914) t. 383; Ripv. Fl. Mal.
Pen. 2 (1923) 301; Heyne, Nutt. Pl. (1927) 1262;
Mekr. Un. Cal. Publ. Bot. 15 (1929) 248; Burk.
Dict. (1935) 2113; Corner, Ways. Trees (1940) 622,
t. 231; Back. & BAKH. f. Fl. Java 2 (1965) 205;
Noort. Leid. Bot. Ser. 1 (1975) 191, f. 2c, pl. 13. —
Sariava Reinw. Syll. Ratisb. 2 (1825) 12. —
Dicalyx tinctorius Bu. Bijdr. (1826) 1116, non
S. tinctoria L’Hérit. 1791. — Eugeniodes fascicu-
latum O. K. Rev. Gen. Pl. 2 (1891) 409. — S.
phanerophlebia Merr. Philip. J. Sc. 9 (1914) Bot.
382; J. Str. Br. R. As. Soc. n. 76 (1917) 112; En.
Philip. 3 (1923) 301. — Fig. 7.

Shrub, or less often a tree to 22 m high and 50
cm @. Twigs sparsely pilose, puberulous, or appres-
sedly pubescent, glabrescent, often zigzag. Leaves
alternately or (on the leaders) spirally arranged,

brous above, sparsely appressedly fine-hairy

th, rarely patently hirsute, especially on mid-

rib and nerves and towards the margin, (narrowly)
elliptic or sometimes ovate, acuminate to caudate
with acute to rounded base, 5-13(-18) by 2-4'/,
(-6) cm; nerves (4-)6-8(-11) pairs, meeting in a
looped intramarginal vein; petiole 2-8 mm. Flowers
in a fascicle of reduced, often branched, racemes to
2'/,cm long. Bracts and bracteoles persistent,
minute (rarely to 3mm), as the axis pubescent;
often several bracts present, indicating the origin
from a more branched inflorescence. Pedicel
1-5 mm, pubescent. Calyx divided into (4-)5(-6)
broadly ovoid, rounded, appressedly pubescent or
brous lobes, c. 1 mm long but sometimes the
bes different in size, often some of the lobes
petaloid. Corolla glabrous or more often with
minute hairs towards the outer base, rarely some
hairs on the back too, 2-4'/, mm. Stamens 12-35.
Disk glabrous to more or less pilose, low annular.

SYMPLOCACEAE (Nooteboom)

259

Ovary appressedly hairy, c. 1 mm high; style hairy,
especially towards the thickened base, rarely
glabrous, 2-3!/, mm. Fruit broadly or narrowly
ampulliform, often curved, the belly globose or
ovoid, the neck broadly conical, dark violet-blue
or cobalt-blue, 5-7 by 3-5 mm; stone brain-like
grooved without or with c. 10 shallow grooves.
Seed 1, much lobed, with slightly curved embryo.

Distr. Extreme South Peninsular Thailand
(Pattani) and throughout Malesia, except the
Lesser Sunda Is., the Moluccas, and New Guinea.
One of the most common Symplocos species in
Malesia. Fig. 17.

Fig. 17. Ranges of a. Symplocos costata (BL.)
Cuolisy, b. S. cerasifolia WALL. ex DC. var. cerasi-
folia, c. S. fasciculata ZOLL.

Ecol. In primary high and open secondary
forest and thickets, common in disturbed forest,
rather indifferent to soils, besides on latosols,
recorded from sand (Banka), in Borneo from sand-
stone, black soils, seasonally swampy land and
Dipterocarp forest, also riparian, in Udjong Kulon
from raised coral limestone, from sea-level up to
c. 2200 m. Fl. June-Sept. (Nov.-April), fr. Sept.-
March. Several times flowers are noted to be scent-
ed, but once recorded as emitting a pervasive sour
smell (Malaya, WHITMORE).

Vern. Malaya: kérénang, nasi-nasi, ménasi
(obviously referring to the often unripe white
fruit, resembling grains of cooked rice), M, Kepong,
sébiak, Selangor; Sumatra: kayu loba-loba,
Asahan, djarak bulau, Pajakumbu, djirok, Kerintji,
kékatja, lélébah, Bengkalis, pipi udan, Karo,
léebomélukut, M. Ulu, Palembang, gente on,
havu-havu, h. h. délok, h. h. itam, h. h. uding,
kareut kareut uding, lihai-lihai uding, Simalur,
gia, Kepahiang, kayu lebeu, Palembang, djarok,
Banka; Java; djarak, djirék, d. leutik, d. prit,
d. sasag, d. wulu, J, S, ki piit, S; Borneo: Sarawak:
Jirah, Iban, périaboh, Murut; Sabah: labah, léboh,
loboh, Kinabatangan, Kadasan lang., giak,
Kedayan, idabo, Dusun; Brunei: pachal ambok;
Kalimantan: njam-njam, Bulungan, gumiting puteh,
Balikpapan.

260

Notes. The fruit is of a type usually containing a
curved seed with curved embryo; here it is, how-
ever, only slightly curved.

In the herbarium sterile sheets are sometimes
confused with Eurya acuminata which has, in
Malaya, often the same vernacular names; cf.
CORNER (1940).

Normally lateral shoots are collected which have
a characteristic alternate phyllotaxis, but I have
also found leader-shoots which have a spiral phyl-
lotaxis flowering in Borneo.

In habit S. fasciculata is very similar to S.
laeteviridis but its flowers are truly fascicled with
more than 3 bracts under each flower and these
persistent, a regular 5-lobed calyx, an ampulliform
fruit with a ruminate seed and curved embryo.
In S. laeteviridis the inflorescence is a raceme or
panicle with 1 bract and 2 bracteoles under each
flower and these caducous, a calyx which splits
into a 3-lobed and a 2-lobed part, while the fruit is
ellipsoid to ovate, with a non-ruminate seed and a
straight embryo.

24. Symplocos filipes Noort. Leid. Bot. Ser. 1
(1975) 193, pl. 14a—d. — Fig. 7.

Twigs glabrous or sparsely pulverulent-puberu-
lous, the terminal buds small, with pulverulent-
puberulous scales which often bear large vesicular
glands on the margin. Leaves glabrous or sparsely
pulverulent-puberulous beneath, + elliptic, long
acuminate, with acute often attenuate base and
entire or slightly denticulate margin which contains
a row of large vesicular glands, 41/,-7!/, by 2-3 cm;
nerves 5—6 pairs, meeting in a looped intramarginal
vein; petiole 7-8 mm. Flowers in a lax raceme of
4-10 cm, the axis sparsely pulverulent-puberulous.
Bracts and bracteoles persistent, with same
indument, 1/, and 1 mm long respectively. Pedicel
slender, 2-15 mm. Calyx sparsely pulverulent-
puberulous, divided into semi-ellliptic !/, mm long
lobes. Corolla c. 3 mm. Stamens c. 25. Disk annu-
lar, glabrous. Ovary with same indument as calyx,
c. 11/,mm high; style glabrous, c. 3mm. Fruit
ellipsoid, c. 10 by 4 mm, the small calyx incurved;
stone spindle-shaped, with shallow lengthwise
grooves, l-celled. Seed 1, straight with straight
embryo.

Distr. Malesia: Philippines (Mindoro: Mt
Halcon), two collections.

25. Symplocos gambliana BRAND, Bull. Herb.
Boiss. II, 6 (1906) 748; MerrR. En. Born. (1921)
484; Noor. Leid. Bot. Ser. 1 (1975) 195. — S. havi-
landii KiNG & GAMBLE, J. As. Soc. Beng. 74, ii
(1906) 251, non BRAND, 1901.

Twigs glabrous. Leaves glabrous, + elliptic,
abruptly oblique acuminate with acute, attenuate
base and entire, recurved margin, 6—9 by 3—41/, cm;
nerves 6-8 pairs meeting in a looped, faintly
prominent intramarginal vein; petiole 5-10 mm.
Flowers in a lax spike or raceme to 6cm; axis
glabrous. Bracts and bracteoles ?minute, soon
caducous. Pedicel less than 1 mm. Calyx entirely
divided into semi-orbicular, ciliate, 3/,-11/, mm
long lobes. Corolla ciliolate, often with some minute
hairs on the outside, c. 5mm. Stamens c. 50. Disk
5-glandular, with the style base minutely pilose.
Ovary glabrous, c. 1mm high; style glabrous
except the base, 4 mm. Fruit not known.

FLORA MALESIANA

[ser. I, vol. 82

Distr. Malesia; Borneo (Sarawak), only known
from the type.

26. Symplocos gigantifolia Noor. Leid. Bot. Ser. 1
(1975) 195.

Twigs glabrous, very thick. Leaves glabrous,
obovate, shortly acuminate, the base cuneate but
truncate at its lowermost part, margin + entire,
21-62 by 7-19 cm; nerves 13-20 pairs, merging
into the venation; petiole c. 1 cm. Flowers in a
fascicle or very short spike on wood. Bracts and
bracteoles persistent, appressedly pubescent, semi-
elliptic, rounded, 1-2 mm. Calyx minutely ap-
pressedly pubescent, 2mm, the 3 semi-elliptic,
rounded lobes c. 11/; mm long. Corolla 4-5 mm.
Stamens c. 50. Disk 5-glandular, glabrous, but
style base pilose. Ovary with same indument as
calyx, c. 1 mm high; style glabrous, reduced (only
3 flowers seen). Fruit very young. Seeds not seen.

Distr. Malesia: East New Guinea (Central
Division, Southern Highlands and Western Dis-
trict), 3 collections.

Ecol. In high forest, once along a riverbed, 90,
500, and 800 m. Fi. April-May.

Notes. Brass (3894) noted that it is a ‘striking
tree with erect branching habit and flowers between
the whorls.’ In the three collections studied the
‘whorled’ position of the leaves could not be
checked. Possibly the main leaves may be conspi-
cuously crowded at the end of the year’s growth
(flush).

A similar situation is reported to occur in S.
herzogii, which is the closest related species,
differing in having smaller, hairy leaves, hairy
twigs, and larger bracts.

27. Symplocos glabriramifera Noort. Leid. Bot.
Ser. 1 (1975) 196, pl. 15Sa—d. — Fig. 7.

Twigs glabrous. Leaves glabrous, elliptic to
obovate, (faintly) acuminate, with acute, attenuate
base and crenate or crenulate apex, 4-6!/, by 11/,-
2!/,cm; nerves 6-8 pairs, meeting in a looped
intramarginal vein; petiole 5-7 mm. Flowers in a
short lax raceme to 11/, cm, axis glabrous. Bracts
and bracteoles caducous, glabrous, ciliolate, 11/2
and 1 mm long respectively. Pedicel 1-2 mm. Calyx
glabrous, c. 11/, mm long, the lobes 3, semi-elliptic,
rounded, c. 11/,mm long. Corolla probably 3-
merous, 3-4 mm. Stamens 30-50. Disk glabrous,
3—5-glandular. Ovary glabrous, c. 1mm high;
style glabrous. Fruit ellipsoid, truncate at both
ends, 8-12 by 4-6 mm; stone shallowly lengthwise
grooved without, 3-celled. Seed 1 in each cell,
straight with straight embryo.

Distr. Malesia: Philippines (Luzon: Benguet &
Nueva Vizcaya Prov.).

Ecol. Mountain forest, 1900 m. F/. Febr., May.

28. Symplocos glomerata KING ex CLARKE, FI. Br.
Ind. 3 (1882) 577; BRAND, Pfl. R. Heft 6 (1901) 69;
BRANDIS, Ind. Trees (1906) 438; HAND.-MAZz.
Beih. Bot. Centralbl. 62 B (1943) 30; Noor. Leid.
Bot. Ser. 1 (1975) 199, pl. 16a—b, with full syno-
nymy. — Fig. 7.

var. glomerata.

Small tree, 6 m. Twigs glabrous, or tomentellous
and then soon glabrescent. Leaves elliptic, acu-
minate, with glandular dentate margin, 7-20 by

ee aaa

1977]

SYMPLOCACEAE (Nooteboom)

261

2-4!/, cm; nerves 10-16 pairs meeting in a looped
intramarginal vein; petiole 5-12 mm. Flowers ina
fascicle from the axils of the leaves or from wood.
Calyx glabrous, 1-2 mm, the ciliate lobes slightly
shorter. Corolla 4-S mm. Stamens c. 25 to c. 50.
Disk cylindrical, c. 1 mm high. Ovary glabrous,
c. 1 mm high. Fruit 7-10 by c. 3 mm.

Distr. Continental Asia (India, Burma, Indo-
China, China, Hainan, Hong Kong, Formosa); in
Malesia: Malay Peninsula (Trengganu, once found
on G. Lawut Besut).

Ecol. Montane forest, 1500 m. Fr. April.

Note. There is a considerable synonymy in-
volved in this widely spread continental SE. Asian
species which I have subdivided into two subspecies
and several varieties.

29. Symplocos goodeniacea Noor. Leid. Bot. Ser. 1
(1975) 204.

Small tree to 7!/,m. Twigs glabrous. Leaves
narrowly elliptic, shortly acuminate with cuneate
base and recurved entire or denticulate margin,
17-30 by 31/,-7 cm; nerves 11-13 pairs, at least in
the apical part of the leaf meeting in a looped
intramarginal vein close to the margin; petiole
15-25 mm. Flowers in a spike to 4 cm; axis puberu-
lous. Bracts and bracteoles persistent, glabrous but
ciliate, c. 2mm. Calyx glabrous, divided into the
broadly rounded 1'/,-2 mm long lobes. Corolla
6-8 mm. Stamens more than 100. Disk annular,
minutely pilose. Ovary glabrous, 1!/,-2 mm high;
style glabrous. Fruit not known.

Distr. Malesia: Borneo (Sabah), only known
from the type.

Ecol. Lowland rain-forest, 150 m.

30. Symplocos herzogii SLEUM. in Fedde, Rep. 42
oy 264; Noor. Leid. Bot. Ser. 1 (1975) 207. —
see

Small tree or leaning shrub, 4-6 m high. Twigs
thick, densely tomentose. Leaves pseudoverticillate,
but between the whorls the scars of fallen spirally
arranged leaves visible in at least one collection,
rather densely hairy beneath, especially on midrib
and nerves, + elliptic, acute to acuminate with
cuneate base (the very base truncate) and sharply
dentate margin, 13-20 by 5—9'/, cm; nerves 10-17
pairs; petiole with same indument as twigs, very
thick, 7-20 mm. Flowers in a fascicle or spike to
2cm from the axils of the leaves or from wood.
Bracts and bracteoles persistent, densely redbrown
sericeous, c. 5mm and c. 3mm respectively.
Calyx appressedly redbrown hairy, 2—2'/, mm, the
lobes + ovate, acute, 1'/,-2 mm. Corolla 3-4 mm.
Stamens c. 40 in 3 flowers (according to SLEUMER
Le. absent in @ flowers). Disk pilose. Ovary

brous, '/,-1 mm high; style reduced in 3 flowers,
in 2 flowers 3'/, mm (according to SLEUMER /.c.).
Fruit globose to ampulliform, c. 8 by 6 mm, the
stone ribbed. Seed 1, curved with curved embryo.

Distr. Malesia: East New Guinea (Morobe
Distr.).

Ecol. Midmountain rain-forest, 1500-1800 m.
Fl, Dec.—April.

Notes. I have only seen ¢ flowers and fruits.
According to SLEUMER & flowers are few, at the base
of the inflorescence.

This species is allied to S. gigantifolia, see the
notes under that species.

31. Symplocos johniana Stapf, Trans. Linn. Soc.
Bot. 4 (1894) 206; BRAND, Pfl. R. Heft 6 (1901) 65;
M_erR. En. Born. (1921) 487; H. Herne, Pf.
Samml. Clemens Kinabalu (1953) 88; Noor. Leid.
Bot. Ser. 1 (1975) 208, pl. 17f-g. — Fig. 7.

Shrub or small tree, to 3m. Twigs densely
obliquely to patently rusty hirsute. Leaves spirally
arranged or alternate, rather densely patently
hirsute beneath, or only midrib and nerves hairy,
acuminate to caudate with rounded to cordate base
and usually rather coarsely sharp-dentate margin,
ovate, 2'/,-7 by 1'/,-3'/,cm; nerves 3-6 pairs
meeting in a looped intramarginal vein; petiole
1-2 mm. Flowers in 1-flowered raceme, axis !/, mm,
with 1'/,mm long bract and the c. 1 mm long
bracteoles loosely appressedly rusty hirsute.
Pedicel c. 1 mm. Calyx rusty hirsute, divided into
the semi-elliptic rounded 1-1'/, mm long lobes.
Corolla c. 5mm. Stamens 60-90. Disk stellate,
densely hirsute. Ovary rusty hirsute, c. 1 mm high;
style glabrous, c. 6mm. Fruit narrowly flask-
shaped, often sparsely hairy, intense indigo-blue,
c. 13 by 4 mn, the persistent calyx not included.
Seed 1, straight, narrowly elliptic, embryo straight.

Distr. Malesia; Borneo (Sabah: Mt Kinabalu;
W. Kutei: G. Kemul).

Ecol. In forest, in damp shady places, often in
crevices of granite rocks, 1500-3200 m. Fi. Febr.—
Oct., fr. Febr.-May, Sept.—Oct.

Note. The distribution is interesting because this
species, which was assumed to be a Kinabalu
endemic, is also found on an old, worn-down,
rather low summit in W. Kutei, in a mountain
range which is probably older than Mt Kinabalu.
This feature is also found in some other mountain
plants, e.g. Lobelia borneensis, which were found
on Mt Murud, in Sarawak (cf. STEEN. Proc. R. Soc.
Lond. B 161, 1964, 16). VAN STEENIS concluded
that Kinabalu plants possibly in the past had a
wider distribution in Borneo when there were more
higher peaks in the island, and that the few present
stations on the low mountains are relict stations
(cf. also STEEN. Mal. Nat. J. 20, 1967, 39).

32. Symplocos junghuhnii Koorp. Proc. Kon.
Acad. Wet. A’dam 10 (1908) 160; Noor. Leid. Bot.
Ser. 1 (1975) 209. — Fig. 7, 18.

Twigs glabrous. Leaves glabrous, or with some
appressed hairs beneath, acuminate with cuneate to
cordate base and entire to denticulate margin,
obovate to elliptic, 9-13 by 4'/,-5cm; nerves
7-10 pairs; petiole 10-17 mm. Flowers in a raceme
to 6 cm, axis pubescent to tomentose, glabrescent.
Bracts and bracteoles soon caducous, not seen.
Calyx glabrous, divided into + semi-orbicular
cordately based c. 2 mm long lobes. Corolla 8-10
mm. Stamens more than 100. Disk 5-glandular,
with the broadly conical style base soft hairy.
Ovary tomentose, 2-3 mm high; style glabrous,
c. 7mm. Fruit (only young fruits seen) + elliptic,
15 by 8 mm. Embryo probably straight.

Distr. Malesia; West Java (Preanger: Tjigen-
teng).

Ecol. Mixed montane rain-forest, 1750 m.

Note. It is not clear why this species was omitted
from Back, & BAKH, /.’s Flora of Java.

33. Symplocos laeteviridis Starr, Trans. Linn, Soc.
Bot. 4 (1894) 205; Branp, Pfl. R. Heft 6 (1901) 53;

262

FLORA MALESIANA

[ser. I, vol. 8?

Fig. 18. Symplocos junghuhnii Koorp. a. Leaf

underside, nat. size, b. deflorated flower from

above, showing 5-glandular disk, x 4, c. fruit, x 2
(KOORDERS 26420).

Merr. En. Born. (1921) 487; Airy SHAW, Kew
Bull. (1939) 408; H. Herne, Pfl. Samml. Clemens
Kinabalu (1953) 88; Noor. Leid. Bot. Ser. 1 (1975)
208, pl. 18-19. — Fig. 3.

For synonyms see under the varieties.

Shrub or tree to 10(—21) m. Twigs glabrous or
clothed by a much variable indument, often faintly
zigzag. Leaves alternate, glabrous to more or less
pilose beneath, acuminate to caudate with acute to

cordate base and nearly entire finely glandular
dentate or sharply dentate, flat or recurved, margin,
(narrowly) ovate to elliptic, 19/,-12 by 1-4!/, cm;
nerves (3—)4-11 pairs, usually meeting in a looped
intramarginal vein. Flowers in a raceme or panicle
to 4!/, cm, the axis clothed with hairs. Bracts and
bracteoles hairy, soon caducous. Pedicels 0-5 mm.
Calyx glabrous or hairy, 2-3 mm long, symmetri-
cally cleft, the lobes 1-3 mm. Corolla 3-5 mm,
often with minute hairs on the outside. Stamens
25-70. Disk 5-stellate, shortly minutely pilose.
Ovary (appressedly) hairy, 1-1'/, mm high; style
glabrous, as long as the corolla. Fruit white to
bluish-black, (obliquely) ovoid to ellipsoid, 7-12
by (3-)5-—6 mm. Seed 1, cylindrical to ellipsoidal or
ovoid with straight embryo.

Distr. Malesia: N. Sumatra, Malaya, Borneo,
and Celebes.

Note. See for differences with S. fasciculata
under that species.

KEY TO THE VARIETIES

1. Leaf base distinctly cordate.
2. Twigs with an indument of c. 2 mm long hairs.
Leaves 5-12 cmlong . . . e. var. mjébergii
2. Twigs with an indument of !/,-1 mm long hairs.
Leaves 13/,—4!/, cm long. d. var. kinabaluensis
1. Leaf base cuneate to rounded.
3. Twigs velutinous.
4. Leaves c. 4cm long . f. var. pauciflora
4, Leaves 9-12 cm long. . . g. var. velutinosa
3. Twigs glabrous, or pubescent, hairs much
shorter than 2 mm.
5. Twigs glabrous or appressed-pubescent.
Nerves 6-9 pairs. . a. var. laeteviridis
5. Twigs loosely appressed-pubescent. Nerves
3-6 PaltS 0a) eos b. var. alternifolia
3. Twigs obliquely to patently long-pilose, hairs
of the indument c. 2 mm long.
c. var, basirotunda

a. var. laeteviridis. — Cf. Noor. Leid. Bot. Ser. 1
(1975) 211, pl. 18e-f, 19b. — S. forbesii BRAND,
Pfl. R. Heft 6 (1901) 63. — Fig. 3, 7.

Shrub or tree to 10(-21) m. Twigs glabrous or
appressedly pubescent. Leaves often yellowish
green above, brownish beneath in sicco, acuminate
to caudate with cuneate to rounded base (narrowly)
elliptic to ovate, 4-11 by 11/,-4cm; nerves 6-9
pairs, usually meeting in a looped intramarginal
vein; petiole 1-3(-4) mm. Flowers in a predomi-
nantly basally branched, often very short lax
panicle of racemes, rarely a simple raceme, to 3 cm
long; axis pubescent. Bracts and the 0-1 bracteoles
very soon caducous. Pedicel with same indument as
axis, 0-2(—5) mm. Fruit black-blue.

Distr. Malesia: N. Sumatra, Banka, Malay
Peninsula (Perak, once), Borneo (throughout,
many collections from Mt Kinabalu), SW.
Celebes (Bonthain, Todjambu).

Ecol. In hill and montane rain-forest, in a
variable set of conditions, on rich clay in mixed
Dipterocarp forest near a river, on stony hillsides,
on black soil on ridge top, on a basalt ridge under
Dipterocarp forest (Sarawak), and even on ultra-
basic; 500-2000 m. F/. Jan.—Oct., fr. almost Jan.—
Dec.

1977]

SYMPLOCACEAE (Nooteboom)

263

Vern. Sumatra: alleban, Karolands, kayu loba-
loba, k. sae-sae, Asahan; Borneo: Sarawak: luroh,
Kayan.

b. var. alternifolia Noort. Leid. Bot. Ser. 1 (1975)
211, pl. 18a.

Shrub or treelet. Twigs densely loosely appres-
sedly brown-pubescent. Leaves rather densely to
sparsely appressed-pilose beneath, especially on the
margin, acuminate to caudate with cuneate shortly
attenuate base and ciliate, recurved, entire to
finely glandular dentate margin, + elliptic, 4—5'/,
by 1*/,-2'/, cm; nerves (3—)4—6 pairs, meeting in a
looped intramarginal vein but sometimes obscured
by the indument; petiole c. 2 mm. Flowers in a
(sometimes branched) raceme to 3 cm or solitary,
axis red-brown pilose. Pedicel 0-'/, mm (to 4mm
in solitary flowers).

Distr. Malesia: Borneo (Sabah: Mt Kinabalu).

Ecol. Montane rain-forest, 1000-1500 m. Fi.
May.

c. var. basirotunda Noort. Leid. Bot. Ser. 1 (1975)
212, pl. 18b.

Shrub or treelet. Twigs obliquely to patently
long-pilose. Leaves glabrous to sparsely appres-
sedly long-pilose beneath, acuminate to caudate
with rounded to subcordate base and sharply
glandular dentate to nearly entire margin, elliptic,
3-11 by 17/,-3'/, cm; nerves 6-9 pairs, meeting ina
looped intramarginal vein; petiole 1-2 mm.
Flowers in a raceme or panicle to 2cm, axis
pilose. Pedicels 0O—2(—3) mm. Fruit blue.

Distr. Malesia: Borneo (Sarawak: Kalabit Up-
lands).

Ecol. Montane rain-forest, on humus on sand-
stone, and on podsolized sand (kerangas), 1000-
1700 m. Fil. March-April, fr. April, Aug.

d. var. kinabaluensis (HEINE) Noort. Leid. Bot.
Ser. 1 (1975) 212, pl. 19c. — S. kinabaluensis
ce , Mitt. Bot. Staatssamml. Minchen 6 (1953)
Shrub or small tree to 4 m. Twigs shortly
obliquely hairy. Leaves acuminate with cordate
base and finely glandular-dentate margin, ovate to
elliptic, 1°/,-4'/, by 1-23/,cm; nerves 4-6 pairs;
petiole c. '/, mm. Flowers ina + 3-flowered raceme
to 3. cm, axis with same indument as twigs. Bracts
4 mm, leaf-like, soon caducous. Pedicel '/,—
mm.
Distr. Malesia: Borneo (Sabah: Mt Kinabalu).
Ecol. Montane rain-forest, also secondary
forest, and in landslip regrowth, on black or clay
soils, 1400-2300 m. Fi. Febr., May-Sept., /r.
March, Aug., Nov.—Dec.

e. var. erg (Merr.) Noor. Leid. Bot. Ser. |
Al 212, a 18g. — S. mjdbergii Merr. Sar.
us. J. 3 (1928) 546, — Fig. 7.
Small tree. Twigs patently brown or rusty pilose.
Leaves (narrowly) elliptic or ovate, acuminate,

cordate with 2-10 mm long lobes, margin
finely glandular dentate, 5-12 2'/,-4'/, cm;
nerves strongly impressed above, in 6-9 pairs,

meeting in a conspicuous looped intramarginal
vein; petiole c. 1'/, mm. Flowers in a predomi-
nantly basally branched panicle to 4cm, the axis
+ patently brown or rusty pilose. Bracts often

leaflike, and then up to 10 mm. Pedicels 1-5 mm.
Fruit from green to purple, finally bluish.

Distr. Malesia: Borneo (Sabah: Mt Kinabalu:
Sarawak: Mt Murud).

Ecol. Montane rain-forest, also in secondary
forest, along hillsides and streams, in Agathis-
Podocarpus-oak forest, sometimes on_ blackish
soil, 1200-2400 m. F/. Aug.—Nov., fr. Dec.—June.

f. var. pauciflora Noor. Leid. Bot. Ser. 1 (1975)
213, pl. 18c—d.

Shrub. Twigs velutinous. Leaves glabrous except
the appressedly pilose midrib and the recurved
finely dentate margin underneath, or appressedly
fine-pilose beneath, acuminate with rounded base,
elliptic, c. 4 by 2 cm; nerves c. 5-7 pairs, meeting
in a looped intramarginal vein; petiole with same
indument as twigs, c. 2mm. Flowers in a 1-5-
flowered raceme up to 3 cm; axis patently pubes-
cent. Pedicel 0-'/,mm, but much longer when
flowers solitary. Fruit blue.

Distr. Malesia: Borneo (Sabah: Mt Kinabalu;
Sarawak: Mt Murud).

Ecol. Montane rain-forest, often mossy, on
ridges, also in scrub forest, 1700-2570 m. Fi. April,
July, Oct., fr. April.

e- = velutinosa Noor. Leid. Bot. Ser. 1 (1975) 213,
pl. 19a.

Treelet to c. 10m. Twigs velutinous. Leaves
glabrous above, more or less appressedly pilose
beneath, especially on the nerves and the sharply
dentate flat margin, acuminate with rounded base,
(narrowly) elliptic, 9-12 by 3-4cm; nerves 7-11
pairs, meeting in a conspicuous looped intra-
marginal vein; petiole 3-4 mm. Flowers in a panicle
to 3 cm, axis patently pilose. Bracts and the 0-3 mm
long pedicels with same indument.

Distr. Malesia: Borneo (Sabah: Mt Kinabalu;
Sarawak: Kapit area).

Ecol. Primary and old secondary rain-forest,
1000-1500 m. F/. Aug.—Oct.

34. Symplocos lancifolia S. & Z. Fam. Nat. 2
(1846) 133; CLARKE, Fl. Br. Ind. 3 (1882) 577;
BRAND, Pfi. R. Heft 6 (1901) 41; Noor. Leid. Bot.
Ser. 1 (1975) 214, pl. 2la—d, with full synonymy. —
S. montana VIDAL, Rev. PI. Vasc. Filip. (1886) 179,
non BRONGN. & Gris, 1866. — S. luzoniensis
Ro re, J. Bot. 24 (1886) 348; BRAND, Pfl. R. Heft 6
(1901) 61; Philip. J. Sc. 3 (1908) Bot. 9; Rote,
Kew Bull. (1912) 157; BRAND, Philip. J. Sc. 7 (1912)
Bot. 35; Merr. En. Philip. 3 (1923) 300. — S.
depauperata MerRR. Publ. Gov. Lab. Philip. n. 29
(1905) 45; BRAND, Philip. J. Sc. 3 (1908) Bot, 10,
incl. var. sordida BRAND; ibid. 7 (1912) Bot. 36,
incl. var. angustissima BRAND; Merr. En. Philip. 3
(1923) 298. — S. merrilliana BRAND, Philip. J.
Sc. 3 (1908) Bot. 9. — S. betula BRAND, l.c. 8;
Merr. En. Philip. 3 (1923) 297; Noor. Leid. Bot.
Ser. 1 (1975) 133. — S. inconspicua BRAND, ae
J. Sc. 4 (1909) Bot. 110; Merr. En. Philip, 3 (1923)
299. — S. zamboangensis BRAND in Fedde, Rep. 14
(1916) 325; Merr. En. Philip. 3 (1923) 303, —
Fig. 7.

Low shrub 1-2 m or tree to 20 m. Twigs appres-
sedly to patently hairy, soon glabrescent. Leaves
often sparsely appressedly fine-hairy beneath,
acuminate, with cuneate to nearly rounded base

264

FLORA MALESIANA

[ser. I, vol. 8?

and mostly finely glandular dentate or undulate
margin, (narrowly) ovate, 2-10 by 1'/,-4'/, cm;
midrib above prominent to slightly sulcate; nerves
(4-)6-11 pairs, often meeting in a looped intra-
marginal vein; petiole 1-3(—5) mm. Flowers in a
raceme to 3(—7) cm. Bracts and bracteoles persistent
but falling in fruit, 1/,-2 and 4/,-1'/, mm respec-
tively. Pedicel 0-1 mm. Calyx usually sparsely
appressedly fine short-hairy or pubescent, rarely
glabrous, divided into '/,-1'/;mm long lobes.
Corolla 2'/,4mm. Stamens 15-40. Disk 5-
glandular, mostly hairy including the style base.
Ovary with same indument as calyx or glabrous,
1/,-11/, mm high. Fruit ellipsoid to globose, 3-5 by
2-5 mm, the calyx forming a blunt beak on top;
stone smooth. Seed 1, filling the whole fruit, with
U-shaped embryo.

Distr. Continental SE.-E. Asia (N. India,
Indo-China, China, Hainan, Hong Kong, Ryu Kyu
Is., Formosa); in Malesia: Philippines (Luzon,
Mindoro, Panay, Negros, Mindanao).

Ecol. In a variety of habitats, also in dense
mossy forest at higher altitude, 400-2500 m. FY.
ei ag diam fr. May—Dec. Flowers noted as scent-
ess.

35. Symplocos lucida (THUNB.) S. & Z. Fl. Jap. 1
(1835) 55, t. 24, excl. syn. Myrtus laevis; OHW1, FI.
Jap. (1965) 727; Noor. Leid. Bot. Ser. 1 (1975) 217,
with full synonymy. — Laurus lucida THUNB. FI.
Jap. (1784) 174. — Hopea lucida THuns. Ic. FI. Jap.
(1800) t. 4. — S. theaefolia D. Don, Fl. Nepal.
(1825) 145; BRAND, Pfl. R. Heft 6 (1901) 66
(‘theifolia’); HALL. f. Med. Rijksherb. 14 (1912) 40;
Back. & BAKH. f. Fl. Java 2 (1965) 205. —
Dicalyx ciliatus BL. Bijdr. (1826) 1119. — S. ciliata
Mia. FI. Ind. Bat. 1, 2 (1859) 466; K. & V. Bijdr. 7
(1900) 155; BRAND, Pfl. R. Heft 6 (1901) 65. —
S. ridleyi KING & GAMBLE, J. As. Soc. Beng. 74, ii
(1906) 239; RipL. Fl. Mal. Pen. 2 (1923) 302. —
S. loheri BRAND, Philip. J. Sc. 7 (1912) Bot. 32;
Merr. En. Philip. 3 (1923) 300. — S. laeviramulosa
ELmer, Leafli. Philip. Bot. 7 (1914) 2323; MEeErR.
En. Philip. 3 (1923) 300. — Fig. 7.

Shrub or tree to 20 m, 25 cm @. Twigs glabrous,
yellowish green, angular when dry. Leaves coria-
ceous, glabrous (sometimes quite thin), acute or
obtuse with cuneate base and entire or glandular
dentate revolute margin, + elliptic, 5-12 by 2-4!/,
cm; midrib more or less prominent on the upper
surface, often sulcate towards the base; nerves
5-15 pairs, prominent on both upper and under-
surface; petiole 5-15 mm. Flowers in a basally
branched short dense raceme or condensed spike
of 11/,-4 cm; axis puberulous or pubescent. Bracts
and bracteoles persistent under the fruit, glabrous,
or sometimes pubescent or puberulous on midrib
and base, 1-3 mm. Pedicels 0-5 mm. Calyx mostly
glabrous, nearly divided into 5 lobes, 1-3 mm.
Corolla 3-5 mm. Stamens 10-70. Disk densely
hairy. Ovary glabrous, *'/,-2mm_ high; style
glabrous, or hairy, mostly towards the base. Fruit
ellipsoid (to rarely nearly orbicular), 1—3-celled,
5-18 by 4-15 mm, the wider ones with 2 seeds.
Seeds usually U-shaped with U-shaped embryo, in
the 3-celled fruits the seeds abortive or (at most)
V-shaped; the legs of the U are either separated by
a septum or not.

Distr. Continental SE.-E. Asia (N. India, N.

Burma, N. Thailand, Indo-China, China, Hong
Kong, Hainan, Japan, Ryu Kyu Is., Formosa);
throughout Malesia, except Borneo, the Moluccas,
and New Guinea.

Ecol. High and low mountain forest, elfin forest,
and mossy forest at higher altitude, also in tjemara
forest, 1500-3000 m. Fi. (July) Oct.-Nov., fr. July
(April—Oct.). In habit very much resembling the
Theaceous Pyrenaria serrata BL. which grows in
similar forest.

Vern. Sumatra: kayu hotir, Asahan; Java:
djarak lulub, S, djirék, J.

36. Symplocos maliliensis Noor. Leid. Bot. Ser. 1
(1975) 237. — Fig. 7.

Tree, 25-30m, 30-40cm @. Twigs glabrous.
Leaves acuminate, base cuneate, often the very
base rounded, margin entire, recurved, (narrowly)
obovate, 15-22 by 4'/,-8'/, cm; nerves 9-14 pairs,
meeting in a looped intramarginal vein; petiole
8-15 mm. Flowers in a raceme to 8 cm, axis pubes-
cent. Bracts and bracteoles caducous, pubescent,
ovate, 3-4 and 2-3 mm long respectively. Pedicel
to 2mm. Calyx glabrous, oblique, 3-4 mm, the
lobes ovate, 2-3 mm. Corolla c. 6 mm. Stamens c.
100 or more. Disk shortly pilose. Ovary glabrous,
1-2 mm high; style with broadly conical shortly
pilose base, the rest glabrous, c. 5mm. Fruit
ellipsoid, 15-20 by 10-12 mm, stone with c. 6
lengthwise ridges, mostly 2-celled. Seeds not seen.

Distr. Malesia: Central Celebes (Mailili).

Ecol. Primary high rain-forest, at low altitude,
c. 200 m. Fil. June-July, fr. Febr., Sept.

Vern. Lako, kandoa, Tobela lang.

37. Symplocos wikstroemifolia HAyATA, Ic. PI.
Form. 5 (1915) 119, t. 25b; Mort, Sylvia 5 (1934)
249; KANEH. Form. Trees rev. ed. (1936) 602,
t. 560. — S. microtricha HAND.-MAzz. Beih. Bot.
Centralbl. 62 B (1943) 17; Noor. Leid. Bot. Ser. 1
(1975) 239. — Fig. 7.

Shrub 1*/, m, or tree to 20 m. Twigs sometimes
soon thickened, tapering towards the apex. Leaves
often only towards the end of the twigs, minutely
sparsely appressedly fine hairy beneath, acuminate,
with cuneate base and nearly entire margin,
(narrowly) elliptic to obovate, 6-151/, by 13/,—4/,
cm; midrib above prominent or sunken, flat or
slightly sulcate; nerves 8-10 pairs, joined in an
intramarginal looped vein 1-3mm from the
margin; petiole 3-10mm. Flowers in an often
branched spike from the axils of the leaves, the
lower ones from wood. Bracts and bracteoles soon
caducous, appressedly pubescent, 1—11/, and 1 mm
respectively. Flowers 3 or %, probably all flowers
on one plant alike. Calyx divided into c. 1 mm long
semi-orbicular or semi-elliptic lobes, glabrous, or
the outer lobes appressedly fine pubescent. Corolla
2-3 mm. Stamens 15-20 in ¢ flowers, 5, alterni-
petalous, in % flowers (observed once). Disk pul-
vinate, glabrous or (minutely) shortly pilose.
Ovary glabrous or finely appressedly short hairy,
1/,mm high in g, 1-1'/,mm in & flowers; style
glabrous, 2 mm, with thick, knob-like stigma, but
aborted in ¢ flowers. Fruit ovoid, or slightly con-
stricted towards the apex, 10-12 by 6-8 mm. Seed
1, curved, with curved embryo.

Distr. Continental SE. Asia (Indo-China,
China, Hainan, Formosa); in Malesia: Malay

1977]

SYMPLOCACEAE (Nooteboom)

265

Peninsula (Pahang: G. Paking,
Fraser’s Hill, G. Tahan).

Ecol. In hill forest, on mossy or exposed ridges,
1300-1500 m. Fi. Febr.—March, fr. Oct. In elfin
forest noted to assume a fastigiate habit. Flowers
often in part ramiflorous.

G. Benom,

38. Symplocos multibracteata Noor. Leid. Bot. Ser.
1 (1975) 241. — Fig. 19.

Small shrub, 3/,—11/, m, or treelet to 4m. Twigs
densely appressedly to patently (softly) pilose to
nearly glabrous. Leaves acuminate with rounded to
cordate base and glandular denticulate to dentate
margin, elliptic to ovate, 5-14 by 2!/,-5!/,cm;
nerves 7-13 pairs, meeting in a looped intramargi-
nal vein; petiole 2-10 mm. Flowers in a reduced
spike of at most 2 cm, usually only 1 (subterminal)
flower left, sometimes another flower present in
bud, axis glabrous. Bracts many, appressedly
pubescent, 4-8 cm. Calyx divided into the appres-
sedly pubescent 3-5 mm long lobes. Corolla 5-8
mm. Stamens 80 to more than 150. Disk softly
pilose. Ovary glabrous, 2-3 mm high; styie gla-
brous, to 2'/, mm long. Fruit obliquely ovoid to
ellipsoid to spindle-shaped, 17-22 by 8-10 mm.
Seed 1, filling the whole stone, with the embryo
straight or slightly curved.

Distr. Malesia: East New Guinea (W. & E.
Highlands).

Ecol. Montane rain-forest and depleted
Castanopsis-Nothofagus forest, 2000-2300 m. Fi.
July, Sept., fr. Aug., Jan.

Vern. Chandujant, Wabag, Enga lang.

39. Symplocos nivea BRAND, Pfi. R. Heft 6 (1901)
36; K. & G. J. As. Soc. Beng. 74, ii (1906) 234:
RIDL. Fl. Mal. Pen. 2 (1923) 300; Noor. Leid. Bot.
Ser. 1 (1975) 241.

Tree to 18 m. Twigs glabrous. Leaves acuminate
with cuneate, attenuate base and entire to obscurely
undulate-crenate margin, glabrous, (narrowly)
elliptic, 7-11 by 2-4'/,cm; nerves 5-8 pairs,
meeting in an intramarginal vein 2-5 mm from the
margin; petiole 7-10 mm. Flowers in a panicle of
racemes, axis villous. Bracts and bracteoles
glabrous, soon caducous, 2!/,-3 and c. 2!/, mm
long respectively. Pedicel pubescent, to 5 mm long.
Calyx glabrous, 2'/,-3 mm, the lobes 1-2 mm long,
becoming longer by tearing apart. Corolla c. 5 mm.
Stamens more than 100. Disk 5-glandular, with the
broadly conical style base soft hairy. Ovary
glabrous, c. 1 mm high; style glabrous, c. 5 mm.
Fruit not known.

Distr. Malesia: Malay
Johore), 2 collections.

Ecol. Hill rain-forest.

Note. Closely allied to S. pyriflora RIDL.,
differing in the number of nerves and with shorter
corolla. May in future prove to be conspecific.

Peninsula (Penang,

A)
Ry

>

Fig. 19. Symplocos multibracteata Noor. a. Leaf and flower, nat. size, b. deflorated flower, spec 5-lobed

hairy disk, c. fruit, d. endocarp, e. fruit in CS, all « 2 (a-b HOOGLAND 5882, c-e HOOGLAND

887).

266

40. Symplocos obovatifolia MERR. Philip. J. Sc. 12
(1917) Bot. 290; En. Philip. 3 (1923) 300; Noor.
Leid. Bot. Ser. 1 (1975) 242. — Fig. 7.

Twigs glabrous. Leaves glabrous, rounded or
shortly acuminate with cuneate, attenuate base and
entire or glandular denticulate apex, obovate,
7'/,-11 by 3!/,-6 cm; nerves 7-9 pairs, meeting ina
looped intramarginal vein; petiole 7-12 mm.
Flowers in a fascicle or short spike to 1'/, cm, axis
glabrous. Bracts and bracteoles glabrous, per-
sistent, 2-3 mm. Only fruits seen. Calyx 3-lobed,
glabrous, elliptic, rounded, c. 2 mm. Disk glabrous,
style base shortly pilose. Fruit (obliquely) ellipsoid,
c. 11 by 5 mm, the persistent calyx not included;
stone smooth, 3-celled. Seed 1 in each cell, straight
with straight embryo.

Distr. Malesia: Philippines (Luzon, Mt Umin-
gan, Nueva Ecija), 2 collections.

Ecol. Hill rain-forest. Fr. Aug.—Sept.

41. Symplocos odoratissima (BL.) CHoIsy ex ZOLL.
Syst. Verz. 2 (1854) 136; Miq. Fl. Ind. Bat. 1, 2
(1859) 468; GUrkKE in E. & P. Nat. Pfl. Fam. 4, 1
(1891) 170; K. & V. Bijdr. 7 (1900) 148, incl. var.
aluminosa K. & V. l.c. 150; BRAND, Pfi. R. Heft 6
(1901) 35, incl. var. divaricata BRAND; K. & G. J.
As. Soc. Beng. 74, ii (1906) 233; Koorp. Atlas 2
(1914) t. 382; RipL. Fl. Mal. Pen. 2 (1923) 299;
S. Moore, J. Bot. 63 (1925) Suppl. 65, incl. var.
leptocarpa S. Moore; Heyne, Nutt. Pl. (1927)
1263; Burk. Dict. 2 (1935) 112; BAck. & BAKH. f.
Fl. Java 2 (1965) 205; Noor. Leid. Bot. Ser. 1
(1975) 245. — Dicalyx odoratissimus Bu. Bijdr.
(1826) 1116. — Eugeniodes odoratissima O. K. Rev.
Gen. Pl. 2 (1891) 975.

For further synonyms see under the varieties.

Tree (shrub) to 30 m high and 50cm @. Twigs
glabrous or tomentellous to tomentose or pubes-
cent. Leaves glabrous or pubescent beneath,
especially on midrib and nerves, with blunt, usually
acuminate apex, acute to rarely rounded base and
entire or mostly crenulate or dentate margin,
(narrowly) elliptic to obovate, 7—20(-40) by
(21/,-)5—10(-20) cm; nerves 5-13(-16) pairs;
petiole stout, 1-5 cm. Flowers mostly many in a
5-30 cm long panicle which is sometimes only
branched towards the base, the axes rusty tomen-
tellous. Bracts at the base of the 3-7 mm long
pedicel, 3-5 mm, bracteoles directly under each
flower, both tomentellous on both surfaces,
caducous. Calyx tomentellous, the lobes blunt,
1/,-11/, mm. Corolla usually tomentellous, at least
in bud, rarely nearly glabrous, 5-8 mm. Stamens
more than 100. Disk hairy with 5 conspicuous
glands. Ovary with same indument as calyx,
11/,-2'/, mm high; style pilose towards the conical
base, about as long as the corolla. Fruit glabrous or
tomentellous, (obliquely) ovoid (or rarely narrowly
flask-shaped, pear-shaped or globular), more or less
narrowed towards the apex, 8-25 by 5-20 mm;
stone with c. 5(-10) ridges. Seeds curved, with
curved embryo.

Distr. Throughout Malesia, except New Guinea.

KEY TO THE VARIETIES

1. Twigs mostly glabrous. Leaves 7-23 cm long.
Fruit 8—15(—20) by 5-10 mm.
a. var. odoratissima

FLORA MALESIANA

[ser. I, vol. 8?

1. Twigs mostly patently pilose, pubescent or
tomentose. Leaves 15-27 cm long. Fruit 17-25
DYAU2—20 IMI. Asuras aes ree b. var. wenzelii

a. var. odoratissima. — Cf. Noort. Leid. Bot. Ser. 1

(1975) 247. — Dicalyx odoratissimus BL. Bijdr.

(1826) 1116. — Dicalyx aluminosus (non Lour.)

BL. /.c. 1117, p.p. — S. ciliata PReEsL, Rel. Haenk.

2 (1831) 61; F.-ViLL. Nov. App. (1880) 127. —

S. patens PRESL, Rel. Haenk. 2 (1831) 61; F.-VILL.

Nov. App. (1880) 127; BRAND, Pfi. R. Heft 6 (1901)

34, incl. var. ciliata BRAND, /.c. 35; Philip. J. Sc. 3

(1908) Bot. 4, incl. f. ciliata BRAND, l.c. 5 et f. elmeri

BRAND, /.c. 4; MERR. En. Philip. 3 (1923) 301. —

S. repandula Miq. FI. Ind. Bat. Suppl. 1 (1861) 474.

— S. racemosa (non Roxs.) F.-VILL. Nov. App.

(1880) 127. — S. spicata (non Roxs.) F.-VILL. Lc.

127; VIDAL, Sinopsis Atlas (1883) t. 64. — S. vil-

larii VIDAL, Rev. Pl. Vasc. Filip. (1886) 178, excl.

syn. Guettarda polyandra BLANCO, nom. illeg. —

S. pseudospicata VIDAL, I.c. 179. — Pygeum grandi-

florum KinG, J. As. Soc. Beng. 66, ii (1897) 228;

cf. KALKMAN, Blumea 13 (1965) 107. — S. alumi-

nosa BRAND, Pfi. R. Heft 6 (1901) 35. — S. poly-
andra sens. BRAND, l.c. 36, quoad descr. et syn.

Vidal. — S. floridissima BRAND, l.c. 35; Philip. J.

Sc. 3 (1908) Bot. 5; ibid., 7 (1912) Bot. 32; MERR.

En. Philip. 3 (1923) 298. — S. elmeri BRAND in

Perkins, Fragm. Fl. Philip. (1904) 36. — S. pul-

verulenta KING & GAMBLE, J. As. Soc. Beng. 74, ii

(1906) 234; Rip. Fl. Mal. Pen. 2 (1923) 300; Burk.

Dict. 2 (1935) 112. — S. floridissima BRAND var.

serrata BRAND, Philip. J. Sc. 4 (1909) Bot. 108. —

S. pulgarensis ELMER, Leafli. Philip. Bot. 5 (1913)

1841; MerR. En. Philip. 3 (1923) 302. — S. apoen-

sis ELMER, Leafli. Philip. Bot. 7 (1914) 2319; Merr.

En. Philip. 3 (1923) 297. — S. megabotrys MERR.

Philip. J. Sc. 9 (1914) Bot. 383; En. Philip. 3 (1923)

300. — S. dagamensis BRAND in Fedde, Rep. 14

(1916) 324; Merr. En. Philip. 3 (1923) 298. —

S. salix BRAND in Fedde, Rep. 14 (1916) 325;

Merr. En. Philip. 3 (1923) 302. — S. acuminatis-

sima Merk. Philip. J. Sc. 11 (June 1916) Bot. 31;

En. Philip. 3 (1923) 296. — Pygeum viride BAKER

f. J. Bot. 62 (1924) Suppl. 34; cf. KALKMAN,

Blumea 13 (1965) 107. — S. bulusanensis ELMER,

Leafl. Philip. Bot. 10 (1939) 3792, nom. illeg., angl.

— S. verdifolia ELMrr, I.c. 3793, nom. illeg., angl.

—Fig. 7.

Tree up to 30m, 50cm @. Twigs, petioles and
underside of leaves mostly glabrous, sometimes
however tomentellous, tomentose, or pubescent.
Leaves 7-23 by 2-12 cm, in watersprouts up to
40 cm. Fruit with thin, fleshy mesocarp, 8-15 by
5-10 mm, ovoid, or up to 20 mm, flask-shaped.
The stone with low ridges.

Distr. Throughout Malesia,
Guinea.

Ecol. Primary and secondary rain-forest, not
rarely along river-banks, on sandy river alluvium,
in Borneo also on brown sandy soils, black soils,
loam and sandstone, from sea-level to 2500 m. Fi.
Febr.—Nov., fr. Aug.—March. Flowers are noted to
be fragrant.

Uses. Dayak people extract salt from wood ash.

As in other species the bark is used for dyeing
purposes and Heyne /.c. even says that for the
purpose of obtaining bark and leaves the species is
planted by the Sundanese at Tjiamis.

except New

1977]

SYMPLOCACEAE (Nooteboom)

267

The tree is mainly useful for the inner bark which

is commonly sold in the medicinal market in West
Java as kayu or kulit sériawan. Decoctions are used
against sprue-like diseases; also pounded bark is
applied to the gums and young leaves are some-
times eaten or applied externally on mouth and
nose. Obat sériawan is even officially recognized in
the Dutch pharmacopeia.

Vern. Sumatra: sarigintung, Karo, tjirupago
uding, Simalur; Java: ki njatu, ki sariawan, ki
sériawan, S; Borneo: lisang, Kinabatangan, Dusun
lang., margaram, Sangkulirang I.; Bali: udu;
Talaud: labah.

b. var. wenzelii (MERR.) Noort. Leid. Bot. Ser. 1
(1975) 248. — S. wenzelii MERR. Philip. J. Sc. 10
(1915) Bot. 282; En. Philip. 3 (1923) 302. —
S. trichophlebia MeRR. Un. Cal. Publ. Bot. 15
(1929) 248. — Fig. 7.

Tree up to 26 m, 50 cm @. Twigs usually patently
pilose, pubescent or tomentose. Leaves mostly
densely pubescent, 15-27 by 12-20 cm; ridges on
the stone up to 4mm.

Distr. Malesia: Borneo (Sarawak and Kaliman-
tan), Philippines (Leyte, once).

Ecol. Primary and secondary rain-forest in the
lowland and hills in a variety of conditions: sandy
ridges and slopes, calcareous loam, dark red soil,
and black soil, near streams. F/. (March) June-
Dec., fr. (Febr._May) July. Obviously mature
fruits are often noted pale green or white.

Note. Size and shape of leaves are very variable
in S. odoratissima; var. wenzelii possesses the larger
and most hairy leaves. The flowers are exactly
matching those of var. odoratissima and with
collections without fruit it is not always possible to
decide to which variety they belong.

42. Symplocos is CLARKE, Fl. Br. Ind. 3
(1882) 579; K. & G. J. As. Soc. Beng. 74, ii (1906)
243; Rip. Fl. Mal. Pen. 2 (1923) 305; Noor. Leid.
Bot. Ser. 1 (1975) 249, f. 4a-e. — Eugeniodes
ophirense O. K. Rev. Gen. Pl. 2 (1891) 975. —
Fig. 4a-e.

For further synonyms see under the infraspecific
taxa.

Shrub or tree to 18 m high and 50 cm @. Twigs
glabrous, or sometimes the youngest parts appres-
sedly pubescent. Leaves glabrous, except some-
times the very youngest, cuneate or rounded to
acuminate, with cuneate base and entire, glandular
crenulate to denticulate or serrate margin, elliptic
to ovate or obovate, 5-22 by 1'/,-7 cm; nerves
4-13(-16) pairs, anastomosing or meeting in an
intramarginal vein; petiole 2-10(-20) mm. Flowers
in a short raceme, a 3—5-branched panicle of
racemes or a — of 1-3(-6 in Sumatra) cm,
rarely only 1-3 flowers together; axis appressedly
oneag to minutely puberulous or nearly gla-

. Bracts and bracteoles caducous or persistent,
with same indument as axis, '/,-1'/,(-3) mm and
slightly shorter than that respectively. Calyx with
same indument as ovary or less hairy, '/,~1 (2'/,-3
in - cumingiana var. pachyphylla) mm long.
Corolla 2-5 mm. Stamens , but more than 75
in var. pachyphylla. Disk glabrous to shortly pilose,

dular. Ovary mostly with same indument as
inflorescence axis, or densely aepreneds pubes-
cent, rarely glabrous, c. 1'/, mm high (2'/, mm in

var. pachyphylla); style glabrous to pilose, 3—5(-8)
mm. Fruit ampulliform, with long neck, to ovoid,
rarely ellipsoid or cylindrical; stone with coarse
surface, low lengthwise ridges, or high, interrupted
ridges and then with hollow base, filled with fleshy
mesocarp. Seed 1, embryo obscurely S-shaped,
curved with an angle of c. 90° + halfway its length,
or twice screw-like curved.

Distr. Malesia: Central West Sumatra (incl.
Lingga Is.), Malay Peninsula, Borneo, Celebes, and
throughout the Philippines.

KEY TO THE INFRASPECIFIC TAXA

1. Twigs densely appressedly pubescent or tomen-
tose . 1. ssp. ophirensis b. var. densireticulata
1. Twigs glabrous or sparsely fine-hairy.

2. Fruit ampulliform, with long neck. Ovary
1'/,mm_ high. Calyx lobes c. !/;mm long.
Corolla 2-3 mm. Disk globose or annular,
shortly pilose 2. ssp. perakensis

3. Terminal buds glabrous. Secondary veins
forming a rather coarse reticulation with the
slightly less prominent veins. Inflorescence a
many-flowered panicle of racemes, 1-4 cm
long. Style shortly pilose for its whole length.

c. var. perakensis

3. Terminal buds glabrous. Secondary veins
prominent, forming a fine reticulation with the
faintly prominent tertiary veins. Inflorescence
a 1-3-flowered raceme, up to lcm. Style
pilose only towards its base.

d. var. lingaensis

3. Terminal buds pubescent. Secondary and
tertiary veins much prominent, forming a fine
reticulation with the often also prominent
quaternary veins. Style glabrous.

e. var. sumatrana

2. Fruit ovoid, ellipsoid, or cylindrical. Ovary
1'/,(-2'/,) mm_ high. Calyx lobes '/,-1(-1'/2)
mm long. Corolla 3-5 mm. Disk 5-glandular,
glabrous or sparsely hairy.

4. Fruit ovoid to cylindrical; stone with shallow
lengthwise grooves. Seed ovoid, with small,
nearly straight embryo. Disk sparsely hairy.
Reticulation beneath very dense.

1. ssp. ophirensis a. var. ophirensis

4. Fruit ovoid, ellipsoid, or rarely cylindrical;
stone with high, interrupted ridges. Seeds
ovoid to horse-shoe-shaped, embryo curved,
or twice screw-like curved. Disk glabrous,
rarely with some hairs. Reticulation beneath
either very fine or coarse 3. ssp. cu

5. Inflorescence a raceme. Ovary 1'/; mm.
Calyx lobes '/,-1 mm. Corolla 3-4'/, mm.
Stamens 20-60 ; f. var. cumingiana

5. Inflorescence a spike. Ovary 2'/, mm, Calyx
lobes c. 1'/, mm. Corolla c. 5 mm. Stamens
more than 75 g. var, pachyphylla

1. ssp. ophirensis. — Cf. Noor. Leid. Bot. Ser. 1
(1975) 252. — S. ophirensis CLarke, Fl. Br. Ind. 3
(1882) 579.

For the description see the species.

a. var. ophirensis. — Fig. 7.

Shrub 1'/, m, or small tree to 6 m, Leaves acu-
minate or rounded, (narrowly) elliptic, 5*/,-9"/,
(-11'/,) by 1'/,-4'/,(-6) cm; nerves 5-6 pairs,

ee ee

268

FLORA MALESIANA

[ser. I, vol. 8?

meeting in a looped intramarginal vein; petiole
only with faint ridges towards the blade. Racemes
up to 10 mm, from the axils of the upper leaves or
from wood. Bracts caducous or persistent. Pedicels
1-3 mm. Calyx lobes '/,-1 mm. Corolla 3-5 mm.
Stamens 25-60. Disk usually sparsely hairy. Ovary
11/,mm high; style glabrous, 3'/,-5 mm. Fruit
ovoid to cylindrical, 6-12 by 4-5 mm. Seed ovoid,
with small, nearly straight embryo.

Distr. Malesia: Malay Peninsula (Perak,
Selangor, Johore).

Ecol. Montane forest, bush-like, on granite,
1200-1500 m. F/. July-Sept., fr. Aug. Young leaves
black purple.

b. var. densireticulata Noor. Leid. Bot. Ser. 1 (1975)
252:

Small, bushy treelet, 2-4 m. Twigs (appressedly)
pubescent to tomentose. Leaves cuneate to acumi-
nate with cuneate to cordate base, 31/,-11 by
11/,-41/, cm; nerves 6-9 pairs, anastomosing or
meeting in an intramarginal vein; petiole 2-9 mm.
Flowers in a short raceme to c. 3 cm; axis pubes-
cent. Bracts and bracteoles pubescent, soon
caducous, 2 and | mm long respectively. Calyx
pubescent, 1—1'/, mm long, the lobes + triangu-
lar, c. 1 mm. Corolla 2—2'/, mm. Stamens delicate,
c. 40. Disk inconspicuous, pilose. Ovary pubescent,
1 mm high; style glabrous, c. 2 mm. Fruit pubes-
cent, ellipsoid, 5-8 by 4-5 mm; stone smooth.
Seed not seen.

Distr. Malesia: Malay Peninsula (Pahang:
Cameron Highlands) and S. Celebes, in both areas
2 collections each.

Ecol. Montane forest, 1400-2500 m. F/. Sept.

2. ssp. perakensis (K. & G.) Noort. Leid. Bot.
Ser. 1 (1975) 254. — S. perakensis KING & GAMBLE,
J. As. Soc. Beng. 74, ii (1906) 241; Rip. FI. Mal.
Pen. 2 (1923) 304; Burk. Dict. (1935) 2114. —
S. caudata (non WALL. ex G. Don) RIDL. FI. Mal.
Pen. 2 (1923) 304.

Tree to 18 m high and 50cm @. Leaves faintly
acuminate to caudate with cuneate base, (narrowly)
elliptic, 5-12 by 2-4'/, cm; nerves 4~7 pairs, except
in var. sumatrana meeting in a looped intramarginal
vein; petiole 3-9 mm, not winged. Flowers in a
(basally) 3-5-branched very slender panicle of
racemes, a raceme, or in var. lingaensis only 1-3
flowers in each inflorescence. Bracts and bracteoles
persistent, minute. Pedicels 1-4 mm. Calyx divided
into semiorbicular c. !/, mm long lobes. Corolla
2-3 mm. Stamens 30-50. Disk shortly pilose. Ovary
1/,-1mm_ high; style pilose to glabrous. Fruit
ampulliform, c. 7 by 5 mm, with long beak; stone
with coarse surface, the inner wall of the stone
following the grooved surface of the deeply rumi-
nate cerebrum-like seed; embryo curved with an
angle of not yet 90°.

Distr. Malesia: Sumatra, Malay Peninsula, and
the Philippines.

c. var. perakensis. — Cf. Noort. Leid. Bot. Ser. 1
(1975) 255, f. 4a-c. — S. fragrans ELmer, Leafl.
Philip. Bot. 2 (1908) 508; BRAND, Philip. J. Sc. 7
(1912) Bot. 33; Merr. En. Philip. 3 (1923) 299. —
Fig. 4a—c, 7.

Leaves 5-11 by 2-4'/,cm; petiole 3-6 mm.
Flowers in a many-flowered panicle of racemes of

1-4 cm. Calyx and ovary appressedly pubescent;
style shortly pilose for its whole length.

Distr. Malesia: Malay Peninsula and the Philip-
pines (Negros, once).

Ecol. Primary lowland and montane forest,
hillsides, bamboo forest, 60-1500 m. Fi. April-
July, Sept., fr. Nov.

d. var. lingaensis Noor. Leid. Bot. Ser. 1 (1975) 255.

Leaves narrowly elliptic with caudate apex,
7-12 by 2-3'/, cm; petiole c. 5mm. Flowers in a
1-3-flowered raceme to 1cm. Calyx and ovary
minutely puberulous; style pilose only towards its
base. Fruit unknown.

Distr. Malesia: Sumatra (Lingga Arch.), Only
known from the type.

. var. sumatrana Noort. Leid. Bot. Ser. 1 (1975)
56.

Leaves faintly acuminate, narrowly elliptic,
6-10 by 2-3 cm; nerves 4-5 pairs; petiole 4-9 mm.
Flowers in a lax panicle or raceme of 1*/,-6cm.
Calyx and ovary minutely appressedly pubescent;
style glabrous. Fruit not known.

Distr. Malesia: Central West Sumatra.

Ecol. Montane forest, 900-1300 m.

3. ssp. cumingiana (BRAND) Noor. Leid. Bot. Ser.
1 (1975) 253. — S. cumingiana BRAND, Pfi. R. Heft
6 (1901) 58.

Shrub or small tree to 6m. Leaves + elliptic,
6-22 by 3-7 cm; nerves 6—13(-16) pairs; petiole
1-10(—20) mm, narrowly winged, except to its very
base. Flowers in a 3(—5) cm long often branched
raceme or spike. Pedicels 0-3 mm. Calyx 4/,-1
(-11/,)mm long. Corolla 3-5 mm. Disk 5-glan-
dular, glabrous, rarely with some hairs. Ovary
11/,(—2!/,) mm high. Fruit ovoid, ellipsoid or rarely
cylindrical, 5-12 by 3-8 mm; stone with high,
interrupted ridges which often protrude from the
base, enclosing some fleshy mesocarp. Seed ovoid
to horse-shoe-shaped, embryo curved with an angle
of about 90° to twice screw-like curved.

Distr. Malesia: Borneo, Philippines
Celebes.

f. var. cumingiana. — Cf. Noor. Leid. Bot. Ser. 1
(1975) 253, f. 4d-e, pl. 20a-e. — S. cumingiana
BRAND, Pfi. R. Heft 6 (1901) 58; Philip. J. Sc. 3
(1908) Bot. 8; ibid. 7 (1912) Bot. 34; MerRR. En.
Philip. 3 (1923) 297; H. HEINE, Pfi. Samml. Clemens
Kinabalu (1953) 87. — S. curtiflora ELMER, Leafl.
Philip. Bot. 2 (1908) 509; Merr. En. Philip. 3
(1923) 298. — S. angularis ELMER, Leafi. Philip.
Bot. 2 (1908) 510. — S. purpurascens BRAND, Philip.
J. Sc. 7 (1912) Bot. 33; MerRR. En. Philip. 3 (1923)
302. — S. minutiflora ELMER, Leafl. Philip. Bot. 7
(1914) 2320; Merr. En. Philip. 3 (1923) 300. —
S. agusanensis ELMER, Leafi. Philip. Bot. 7 (1914)
2321. — S. elliptifolia Merr. Philip. J. Se. 12
(1917) Bot. 292; En. Philip. 3 (1923) 298. — S.
brachybotrys MeErRR. Philip. J. Sc. 14 (1919) 447,
non Merr. 1917; En. Philip. 3 (1923) 297. —
S. ilocana Merk. Philip. J. Sc. 35 (1928) 7. — Fig.
4d-e, 7.

Shrub 1!/,m or small tree to 12 m, once even
30 m and 50cm @. Leaves + elliptic, 6-18 by
3-7 cm; nerves 6-13(-16) pairs, usually meeting
in a looped intramarginal vein; petiole 3—10(—15)

and

aia ol

1977]

SYMPLOCACEAE (Nooteboom)

269

mm. Racemes to 3(—5) cm long. Bracts and brac-
teoles usually very small, caducous or persistent.
Pedicels 1-3 mm. Calyx !/,-1mm, pubescent.
Corolla 3-4'/, mm. Stamens 20-60. Ovary 1!/, mm
high. Fruit 5-12 by 3-7 mm, ripe purple-blue.

Distr. Malesia: Borneo, Philippines, Celebes.

Ecol. Mostly in the mountain forest, on hillsides
in oak-Podocarpus forest, largely between 1000 and
3000 m, but on Mt Kinabalu once found as high as
3700 m (sterile), and once collected in lowland
Dipterocarp forest at 300 m in the Sierra Madre
Mts (Luzon), a very common species in the
Philippines. Flowers (once) noted to be faintly
fragrant. Fi. May—Dec., fr. March—Oct.

g. var. pachyphylla (MeRR.) Noor. Leid. Bot. Ser. 1
(1975) 254. — S. pachyphylla MeErr. Philip. J. Sc.
10 (1915) Bot. 283. — Fig. 7.

Small tree, 6m. Leaves 10-20 by 6-8!/, cm;
nerves c. 10 pairs; petiole 10-20 mm. Flowers in a
spike. Bracts and bracteoles appressedly pubescent,
2'/, and 3 mm long respectively. Calyx densely
appressedly pubescent, divided into c. 1/, mm long
lobes. Corolla c. 5 mm. Stamens more than 75, up
to 9mm. Disk glabrous, 5-glandular. Ovary
glabrous, 2'/, mm high; style glabrous, c. 8 mm.
Fruit ovoid, c. 10 by 6-8 mm, the stone as in var.
cumingiana, but several ridges totally lacking in the
upper half, c. 7 by 5-6 mm. Seed ovoid or curved,
and then as the embryo with an angle of about 90°
beneath the middle.

Distr. Malesia: Philippines (Leyte and Minda-
nao), 2 collections.

Ecol. Hill forest, c. 500 m. Fi. Sept.

43. Symplocos paucistaminea F.v.M. & F. M.
BaiLey, 3rd Suppl. Syn. Queensl. FI. (1890) 46;
F. M. BarLey, Queensl. FI. 3 (1900) 967; Noor.
Leid. Bot. Ser. 1 (1975) 262. — Fig. 7.

Tree 18 m, 45cm @. Twigs densely spreadingly
pubescent to tomentose. Leaves acuminate, with
acute to rounded base and dentate margin, sparsely
pubescent above and beneath, elliptic to obovate,
8-20 by 3-8 cm; nerves 7-12 pairs, meeting in a
looped intramarginal vein; petiole 5-10 mm.
Flowers in a basally branched spike to 5 cm long,
becoming longer in fruit; axis sparsely brown hairy.
Bracts and bracteoles persistent, spreadingly
hairy, c. 2 and c. 1'/,mm respectively. Calyx
divided into glabrous c. 1 mm long lobes, the lobes
tomentose. Corolla c. 2'/, mm. Stamens c. 10 to 60.
Disk glabrous or pilose. Ovary glabrous, c. 3/4 mm
high; style glabrous, c. 1'/, mm. Fruit ampulliform,
c. 6 by 4 mm, stone ampulliform with globose,
lengthwise grooved belly and narrow cylindrical
neck, 1-celled. Seed 1, filling the whole stone, with
the embryo twice curved.

Distr. Queensland and Malesia (New Guinea:
Milne Bay Distr.: Mt Suckling, two collections).

Ecol. Lowland rain-forest at 360 m. FI. July.

4. on pop polyandra (BLANCO) BrRANp, Pfl.
R. Heft 6 (1901) 436, quoad syn. Blanco, excl. descr.
et stirp., Merr. Sp. Blanc. (1918) 304; En. Philip. 3

1923) 301; Sreen. Bull. Bot. Gard. Btzg III, 12

1932) 170, f. 5; Noor. Leid. Bot. Ser. 1 (1975)
264. — Guettarda polyandra BLANCO, FI. be ed.
2 (1845) 500; ed. 3 (1879) 126. — Carlea oblongi-

folia PResL, Epim. Bot. (1851) 216. — Baranda
angatensis LLANOS, Mem. Acad. Cienc. Madrid 3,
2 (1857) 502. — S. oblongifolia RoLFE, J. Bot. 23
(1885) 214; VipAL, Phan. Cuming. Philip. (1885)
124; Rev. Pl. Vasc. Filip. (1886) 178; BRAND, Pfl.
R. Heft 6 (1901) 55; HALL. f. Beih. Bot. Centralbl.
39 B (1921) 94. — S. superba BRAND, Pfl. R. Heft 6
(1901) 55. — Fig. 7.

Tree up to 30 m, 50 cm @, rarely a shrub. Bark
dark, cracked. Twigs puberulous, glabrescent,
tapering off towards the apex, thick, at least 5 mm
@ beneath the leaves and there usually with many
pulvinate leaf-scars. Leaves crowded towards the
end of the twigs, rounded or cuneate-obtuse at the
apex, with cuneate, attenuate base and entire,
revolute margin, glabrous (except in innovations
and then puberulous), narrowly elliptic to obovate,
9-22 by 2'/,-7(-9) cm; nerves 11-15 pairs; petiole
2-4 cm. Many spikes from old wood beneath the
leaves, axis densely rusty appressedly puberulous,
glabrescent, 4-15 cm long. Bracts and bracteoles
with same indument, persistent under the fruit,
11/,-2mm_ long. Calyx with same indument,
becoming glabrous towards the apex, 2-3 mm, the
lobes c. 2mm long. Corolla 8-10 mm. Stamens 50
to more than 100. Disk glabrous, annular, and
then surrounding a lower, rarely shortly pilose
receptacle, or low pulvinate, only surrounding the
glabrous, 7-9 mm long style. Ovary with same
indument as calyx, c. 2mm high. Fruit ellipsoid,
c. 10 by 7 mm in vivo; stone rather smooth, with
few shallow lengthwise grooves, 8-10 by 4-5 mm
(s.s. the whole fruit as big as the stone), 3-celled.
Seed 1 in each cell with straight embryo.

Distr. Malesia: Borneo and adjacent islands
(Natuna, Banka, Billiton, Karimata, St. Barbe),
Philippines (throughout), and SW. Celebes
(Makassar: Baleh Angien, once).

Ecol. Secondary and primary forest, almost
always on sandstone, granite, kerangas, sandy
flats, more rarely on sandy loam, at low altitude,
below c. 300m, once found in montane forest
(Luzon: Sierra Madre) at 1000-1100 m in low,
mixed, primary rain-forest (JAcoss 7840). Fi.
Sept._March, fr. Febr.-June (July—Oct.). The
flowers are faintly fragrant, especially at night.

Vern. Bungur, dutat, Banka; sudjéng, Natuna;
Borneo: merbryot, Sarawak, beluno-beluno, salam-
buno, temasuk jantan, Sandakan; Philippines:
ditaman, rapo-rdpo, Tag., balakbaék, balakbakan,
bangkunai, mankénai, P.Bis., buli-bili, malabili,
ribili, Pang., dilangi-bdaka, Sbl.

45. Symplocos pulvinata Noor. Leid. Bot. Ser. |
(1975) 269. — Fig. 7.

Sparsely foliaged tree, 12-18 m high. Twigs thick,
at least 5mm. Leaves coriaceous, glabrous, acute
or faintly acuminate with cuneate base and glan-
dular crenate or dentate margin, obovate, 12-21 by
4'/,-10'/, cm; nerves 8-12 pairs; petiole stout,
1'/,-2'/, cm. Spike glabrous, c. 3.cm. Bracts and
bracteoles probably persistent, glabrous, 5-7 and
c. 4mm long respectively (older flowers often
fallen including bracts and bracteoles, leaving
conspicuous pulvinate light coloured scars on the
dark axis). Calyx glabrous, c. 3 mm, divided in 5,
2-2'/, mm long lobes. In some flowers corolla and
stamens absent or obsolete, in other flowers

270

FLORA MALESIANA

[ser. I, vol. 8?

corolla 5mm, 3(—4)-lobed and stamens 20-35.
Disk glabrous. Ovary glabrous, oblique, 1—-1'/, mm
at one, c. 2mm at the other side; style glabrous,
6 mm. Fruit ovoid, deeply violet, c. 13 by 6-8 mm;
stone with rather high lengthwise ridges in the
basal and low ridges in the apical half, in the middle
a deep transverse groove, l-celled. Seed 1,
uncinately curved towards the base with curved
embryo.

Distr. Malesia: East New Guinea (Koitaki and
Normanby I.), 2 collections.

Ecol. Under open canopy of tall forest, 450-
825 m. Fi. Febr.

46. Symplocos pyriflora Ript. J. Fed. Mal. St.
Mus. 6 (1915) 159; Fl. Mal. Pen. 2 (1923) 307. —
S. bakeri SYMINGTON, J. Mal. Br. R. As. Soc. 14
(1936) 356, t. xx. — Fig. 7.

Shrub or small to medium-sized tree. Twigs
often stout, glabrous. Leaves glabrous, mostly
faintly acuminate with cuneate or rounded base
and undulate to crenate margin, elliptic, 5-15 by
2-23/, cm; nerves 9-14 pairs, meeting in an intra-
marginal vein 2-4 mm from the margin; petiole
stout, 3-10 mm. Flowers in a subterminal, rarely
terminal, raceme or panicle of racemes; axis
pubescent to glabrous. Bracts and bracteoles
glabrous, soon caducous, c. 8 and c. 5 mm long
respectively. Pedicel at most 3 mm. Calyx glabrous,
3-5 mm, sometimes becoming symmetric by
tearing; lobes 2-3 mm, becoming longer by tearing
apart. Corolla 8-10 mm. Stamens c. 100 or more.
Disk 5-glandular, included the conical style base
glabrous or soft hairy. Ovary glabrous, 11/,-2 mm
high; style glabrous, c. 5 mm. Fruit ellipsoid, c. 15
by 8mm; stone smooth or with faint ridges,
1-celled. Seeds not seen, but probably with straight
embryo.

Distr. Malesia: Malay Peninsula (Pahang: G.
Tahan; Kuantan: G. Tapis), two collections.

Ecol. Montane rain-forest, 1400-1650 m. Fi.
Ju

ne.
Note. Closely allied to S. nivea, see there.

47. Symplocos robinsonii RipL. J. Fed. Mal. St.
Mus. 8 (1917) 60; Noor. Leid. Bot. Ser. 1 (1975)
276.

Twigs tomentose, dark brown pubescent or
(sparsely) appressedly pubescent or puberulous,
glabrescent. Leaves sparsely long pubescent,
appressedly fine dark-pilose or sparsely appres-
sedly minutely pilose, glabrescent beneath, acute
or acuminate with acute base and dentate, denticu-
late or crenulate margin, narrowly to broadly
elliptic, 3-91/, by 11/,-4cm; nerves 7-14 pairs.
Flowers in an often branched raceme to 1, 2 or 4cm;
axis pubescent or appressedly puberulous. Bracts
and bracteoles caducous, with same indument as
axis, 1-2 and 3/, to 11/, mm respectively. Pedicel to
2 or 3(—4) mm long. Calyx pubescent to puberulous,
often less hairy than ovary, 1-2 mm, the lobes
1/,-11/, mm. Corolla 4-5 mm. Stamens 25-55. Disk
with some hairs or shortly pilose, often the indu-
ment hardly visible. Ovary with same indument as
calyx or more hairy; style glabrous, or the base
shortly pilose. Fruit ellipsoid, 7-10 by 3-6 mm;
stone inconspicuously lengthwise grooved, 3-celled.
Seeds 1-3, straight with straight embryo.

Distr. Malesia: Sumatra.

KEY TO THE VARIETIES

1. Twigs tomentose
1. Twigs not tomentose.
2. Inflorescence to 4 cm long. Leaves 3-6 by 2-4
cm (index 11/,—21/,) . b. var. latifolia
2. Inflorescence 1—2 cm long.
3. Twigs densely dark-brown pubescent. Leaves
4-6!/, by 11/,—31/, cm (index 13/,—23/4).
c. var. pilosa
3. Twigs sparsely appressed-pubescent or pube-
rulous. Leaves 5-91/, by 2-3cm (index
21/,—31/5) cee d. var. angustifolia

a. var. robinsonii

a. var. robinsonii. — Cf. Noor. Leid. Bot. Ser. 1
(1975) 277. — Fig. 7.

Twigs tomentose. Leaves sparsely long-pubes-
cent, especially on midrib and nerves beneath, +
elliptic, 41/,-9 by 2-3%/,cm; nerves 7-9 pairs;
petiole 7-10mm. Raceme to 2cm, axis rusty
pubescent. Bracts and bracteoles appressedly
pubescent, 11/, and 1 mm long respectively. Pedicel
to 3mm. Calyx appressedly pubescent, c. 1 mm,
the + ovate lobes 3/,-1 mm long. Corolla c. 4 mm.
Stamens 25-40. Disk with some hairs. Ovary ap-
pressedly pubescent, c. 11/,mm high; style with
conical base, glabrous. Fruit 7 by 3mm; stone
3-celled.

Distr.
Kerintji).

Ecol. Gleichenia woodland in mountain forest,
2200-2500 m. Fi. May, Aug.

Malesia: Sumatra (Westcoast: G.

b. var. latifolia Noort. Leid. Bot. Ser. 1 (1975) 277.
— Fig. 7.

Treelet 6 m. Twigs (sparsely) appressedly pubes-
cent, glabrescent. Leaves sparsely minutely appres-
sedly pilose beneath, especially on midrib and
nerves, or glabrous, shortly acuminate, 3-6 by
2-4 cm; nerves 7-8 pairs; petiole 4-7 mm. Raceme
branched, to 4cm; axis (sparsely) appressedly
puberulous. Bracts and bracteoles ovate, 1-11/,
mm. Pedicel to 3(-4) mm. Calyx sparsely appres-
sedly puberulous, 11/,-2 mm long, the + semi-
orbicular lobes 1—11/, mm long. Corolla c. 5 mm.
Stamens 35-55. Disk with the conical style base
pilose. Ovary with same indument as calyx, c. 11/4
mm high; style glabrous, c. 4mm. Fruit c. 10 by
6 mm, blue-black. Seeds 1-3.

Distr. Malesia: northern half of Sumatra (Gajo
Lands: G. Losir; Westcoast: G. Kerintji).

Ecol. In dense ericoid shrub-forest, 2000-3000
(—3400) m. F/. May—Aug. Flowers scentless.

c. var. pilosa Noor. Leid. Bot. Ser. 1 (1975) 278.

Twigs densely dark brown pubescent. Leaves
appressedly fine dark pilose beneath, especially on
midrib and nerves, acute to acuminate, + elliptic,
4-6!/, by 1'/,-3'/, cm; nerves 7-10 pairs; petiole
5-7 mm. Raceme to 1 cm; axis appressedly brown
pubescent. Bracts and bracteoles with same
indument, 11/,-2 and 1-1'/,mm_ respectively.
Pedicel to 2 mm. Calyx sparsely fine puberulous,
c. 11/, mm, the lobes semi-elliptic to ovate, c. 1 mm
long. Corolla 4-5mm. Stamens 30-45. Disk
minutely pilose, hairs sometimes very inconspi-
cuous. Ovary appressedly fine puberulous, 11/,—
1!/, mm high; style glabrous, 4-5 mm.

——

1977]

SYMPLOCACEAE (Nooteboom)

271

Distr. Malesia: Sumatra (Westcoast: G. Merapi
and G. Singalang).

Ecol. Subalpine mountain forest, between lava
boulders, 2500-2800 m. F/. May-June.

d. var. angustifolia Noort. Leid. Bot. Ser. 1 (1975)
278.

Twigs sparsely appressedly pubescent or puberu-
lous. Leaves sparsely appressedly minutely pilose
beneath, acuminate, 5—9'/, by 2-3 cm; nerves 9-14
pairs; petiole 5-12mm. Raceme to 2cm; axis
minutely appressedly puberulous. Bracts and
bracteoles with same indument, ovate, c. 1 and
3/,mm respectively. Pedicel to 3 mm. Calyx less
hairy than ovary, c. 1 mm long, the lobes ovate,
1/,-5/, mm. Corolla c. 4mm. Stamens c. 35. Disk
minutely pilose. Style glabrous.

Distr. Malesia: Sumatra (Westcoast: G. Ophir
= G. Talakmau).

Ecol. Subalpine mountain forest, 1900-2700 m.
Fl. May.

48. Symplocos rubiginosa Wat. (Cat. 1831,
n. 4432, nomen) ex DC. Prod. 8 (1844) 257; Mia.
FI. Ind. Bat. 1, 2 (1859) 466; CLARKE, FI. Br. Ind. 3
(1882) 580; BrANpb, Pfil. R. Heft 6 (1901) 53;
K. & G. J. As. Soc. Beng. 74, ii (1906) 247; RIpL.
Fl. Mal. Pen. 2 (1923) 306; Noor. Leid. Bot. Ser. 1
(1975) 279. — Lodhra rubiginosa Miers, J. Linn.
Soc. Bot. 17 (1879) 299. — Fig. 7.

Shrub, or tree to 30 m high and 50 cm @. Twigs
tomentose, pubescent, tomentellous or glabrous,
rather thick (3-5 mm). Leaves sparsely appressedly
pilose to more or less densely patently soft-villous
beneath, especially on midrib and nerves, usually
abruptly acuminate with cuneate base and finely to
rather coarsely dentate margin, narrowly elliptic
to obovate, 15-45 by 5*/,-17 cm; nerves 12-17
pairs; petiole thickened, 10-25 mm. Flowers in a
spike from wood beneath or between the leaves;
in bud the inflorescence has the appearance of a
short cone; axis pubescent to tomentellous,
1-5S(-8) cm. Bracts and bracteoles caducous as
soon as the flower matures, ovate, boat-shaped,
densely silky-pubescent, 3-5 and 2—3 mm respec-
tively. Calyx appressedly puberulous to silky
pubescent, often symmetrically torn, 1'/,-3 mm,
the lobes '/,-2 mm. Corolla sparsely (minutely) stiff
hairy towards the outer base, 4-5 mm. Stamens
60-100. Disk glabrous or sparsely pilose. Ovary
pubescent to tomentellous or with same indument
as calyx, 1-2 mm high; style glabrous or pilose,
sometimes with thick conical pilose base. Fruit
blue in vivo, ovoid to ellipsoid, sparsely short pilose
to glabrous, 8-10 by 5-8 mm; stone lengthwise
grooved, at one side with a deep transverse con-
striction at '/, from the base. Seed 1, once or twice
and then S-shaped curved due to the constriction
of the stone.

Distr. Malesia: Sumatra, Malay Peninsula, and
Borneo (rare in Kalimantan).

Ecol. Both in the lowland and in the hills, from
sea-level to 1800 m, in arm and secondary
mixed rain-forest, not rarely in Dipterocarp forest,
along streamsides, on kerangas, in bertam
(Eugeissona) ridge forest. Fi. Oct.-Dec. (once
April), fr. Jan.-Dec. Fruit remain white for a long
time, then turn through red to light blue when ripe.

Uses. The wood is very hard and used for house-
building (BurK. Dict. 1935, 2115).

Vern. Sumatra: /empaong kantjil, Palemb.;
Malaya: pemasa, Sakai lang.; Borneo: smuak,
Sarawak, Land-Dayak.

49. Symplocos salicioides Noort. Leid. Bot. Ser. 1
(1975) 280.

Shrub 2 m, with pubescent twigs. Leaves faintly
acuminate to sharply acute, with cuneate to round-
ed base, pubescent beneath, narrowly elliptic,
3*/.-7 by 3/s-1'/,cm; nerves 6-8 pairs, rather
inconspicuous, meeting in an intramarginal looped
vein; petiole 3-4 mm. Spike 1-flowered. Bracts and
bracteoles pubescent, 2 and 1 mm long respectively.
Calyx densely pubescent, divided into 1-1!/,mm
long triangular lobes. Corolla 2-2'/, mm. Stamens
15-20. Disk with the conical style base softly long-
hairy. Ovary with same indument as calyx, 1'/, mm
high; style hairy for its lower half, c. 2 mm long.
Fruit long ellipsoid, pubescent, 13 by 5 mm, only
seen immature.

Distr. Malesia: East New Guinea (Sepik area,
once).

Ecol. Lowland rain-forest, 1000 m.

50. Symplocos sumatrana BRAND, Pfi. R. Heft 6
— 62; Noor. Leid. Bot. Ser. 1 (1975) 283. —
if. Te
Treelet 3 m. Twigs densely patently red-brown
long-hairy or tomentose. Leaves softly pilose
beneath, acuminate with rounded base and denticu-
late margin, narrowly elliptic to ovate, 6-14 by
2-4cm; nerves 7-15 pairs, meeting in a looped
much prominent intramarginal vein; petiole
5-7 mm. Flowers in a spike or raceme of 2-4 cm;
axis brown tomentose or spreadingly hairy. Bracts
and bracteoles soon caducous, the first not seen, the
latter appressedly long-hairy, c. 2'/;mm long.
Calyx divided into 5 appressedly pilose semi-
elliptic 2 mm long lobes. Corolla c. 5 mm. Stamens
45-70. Disk pulvinate, pilose. Ovary sericeous,
c. 1'/, mm high; style with some hairs in the lower
half, 2-5 mm. Fruit ellipsoid, hairy, c. 10 by 6 mm;
stone lengthwise ribbed, 3-celled, 1, 2 or 3 cells
fertile. Seed straight with straight embryo.
Distr. Malesia: Sumatra (Gajo Lands: Mt
Kemiri; Westcoast: Mt Singalang), 2 collections.
Ecol. Ericoid, elfin and subalpine mossy forest,
2700-3000 m. Fi. March, June-July. Flowers
fragrant.

51. Symplocos sumuntia BucH.-HAm. ex D. Don,
Prod. Fl. Nepal. (1825) 145; Clarke, FI. Br. Ind. 3
(1882) 578; Noor. Leid. Bot. Ser. 1 (1975) 284,
with full synonymy. — Fig. 7.

Low shrub to medium-sized tree. Twigs glabrous
or nearly so, dark-coloured. Leaves glabrous,
acuminate to caudate with attenuate base and
glandular dentate margin, + elliptic, 2-10 by
/,-4'/, cm; nerves 5-8 pairs, meeting in an intra-
marginal vein; petiole 2-10 mm. Raceme few to
many-flowered, 1-6 cm long; axis from nearly
glabrous to pilose or pubescent. Bracts at base of
pedicel, with the bracteoles soon caducous, ap-
pressedly hairy, 2-5 and '/,-4 mm long respectively.
Pedicel '/,-13 mm. Calyx glabrous to (sparsely)
appressedly hairy, divided into '/, to 1'/,; mm om
lobes. Corolla 4-8 mm. Stamens 25-40. Dis

272

FLORA MALESIANA

[ser. I, vol. 8?

glabrous. Ovary glabrous to shortly sparsely
appressedly hairy, 1-2 mm high; style glabrous,
2-11 mm. Fruit ovoid to ampulliform, 6-10 by
3-6 mm; stone shallowly (brain-like) grooved.
Seed curved, embryo once or twice curved.

Distr. Continental Asia (India, Burma, Thai-
land, Indo-China, China, Hong Kong, Hainan,
Formosa, Ryu Kyu Is., Japan, and Korea); in
Malesia: Malay Peninsula (Pahang: Cameron
Highlands and G. Tahan), 3 collections.

Ecol. Montane high forest, 1200-1500 m. FI.
Aug.—Oct.

52. Symplocos trichomarginalis Noor. Leid. Bot.
Ser. 1 (1975) 287. — Fig. 7.

Shrub 1-4m. Twigs often zigzag, appressedly
brown-pilose. Leaves alternate, sparsely appres-
sedly pilose beneath, the midrib and the finely
glandular-dentate recurved margin beneath
conspicuously densely appressedly brown-pilose,
acuminate with cuneate to rounded base, elliptic,
2-31/, by 11/,-13/,cm; nerves 5-7 pairs; petiole
2-4 mm. Flowers solitary, often several brown-
pilose bracts indicating the derivation from a more-
flowered inflorescence, the 2 uppermost bracts
3-5 by 1-3 mm, persistent. Pedicel from twig to
flower up to 1 cm. Calyx loosely appressedly pilose,
divided into the narrowly elliptic, acute, c. 3 mm
long lobes. Corolla c. 4mm. Stamens c. 50. Disk
glabrous. Ovary with same indument as calyx, c.
1 mm high; style glabrous, c. 5 mm. Fruit sparsely
pilose, ellipsoid to ovoid, green to deep indigo
when ripe, 8-9 by c. 4 mm; stone narrowly ovoid,
muricate with shallow lengthwise grooves. Seed 1,
embryo straight (only young seeds seen).

Distr. Malesia: Borneo (Sabah: Mt Kinabalu).

Ecol. Open places and forest edges, 1500-2400
m. Fl. May, fr. April.

Note. In habit similar to S. zizyphoides, but
differing in the veins being obscure and in the long-
pilose calyx with narrow triangular lobes being
longer than those of that species.

53. Symplocos tricoccata Noort. Leid. Bot. Ser. 1
(1975) 288. — Fig. 7.

Shrub 3 m to small tree to 10 m high, 15cm @.
Twigs glabrous. Leaves glabrous, yellowish or
olive-grey or water-green, sometimes glossy
beneath, acuminate, with acute base and dentate to
denticulate margin, + elliptic, 7-29 by 4-91/, cm;
nerves 5-10 pairs, meeting in an intramarginal
vein; petiole 5-15 mm. Flowers in a fascicle or very
short spike; axis glabrous, to 5mm long. Bracts
and bracteoles soon caducous, c. 11/, mm. Pedicel
0-1 mm. Calyx glabrous or with some hairs, c.
2mm long, the lobes 1-13/, mm. Corolla 5-8 mm.
Stamens 40 to more than 100. Disk 5-glandular,
the conical style base with some hairs to softly
short-pilose. Ovary glabrous, c. 2 mm high; style
glabrous to 7mm. Fruit narrowly obliquely
ellipsoid, 12-16 by 4-6 mm, + triangular in CS,
3-celled, each cell circular in CS; stone 3-lobed in
CS, endocarp thin, woody. Seed cylindrical,
with straight embryo.

Distr. Malesia: Borneo (Sarawak, Sabah, and
Kalimantan).

Ecol. Lowland and montane primary rain-
forest, near streams, on hillsides, in low undulating
flat country, on rocky soil, also in Dipterocarp

forest, 30-2100 m. Fl. Aug.—Nov., Febr., fr. Febr.-
June, Sept. Fruits often recorded to be whitish,
through purple to blue when ripe.

Uses. In Sarawak the wood is said to be used
for knife handles.

Vern. Borneo: atup, Sarawak, Kenyah lang.

54. Symplocos trisepala Merr. Philip. J. Sc. 12
(1917) Bot. 291; En. Philip. 3 (1923) 302; Noor.
Leid. Bot. Ser. 1 (1975) 289.

Twigs glabrous, but sparsely long-pilose in
innovations. Leaves sparsely appressedly pilose on
the midrib beneath, faintly acuminate, with
rounded or subcordate base and glandular denticu-
late margin, + elliptic, 5-9 by 23/,-5 cm; nerves
7-9 pairs, meeting in an intramarginal vein; petiole
15-25 mm. Spike to 1'/, cm; axis glabrous. Bracts
and bracteoles persistent, glabrous, ciliate, 3-5 mm.
Calyx glabrous, divided into three 2'/,-3 mm long
semi-elliptic rounded lobes. Corolla 5-6 mm.
Stamens 40-70. Disk glabrous, but style base hairy.
Ovary glabrous, 1mm _ high; style glabrous.
Fruits not known.

Distr. Malesia: Philippines (Luzon: Mt Umin-
gan, Nueva Ecija), only the type.

Ecol. Montane rain-forest, at least 400 m. FI.
Aug.—Sept.

55. Symplocos verticillifolia Noort. Leid. Bot. Ser. 1
(1975) 290. — Fig. 7, 20.

Treelet 7-9 m, 20cm @. Twigs hirsute, glabres-
cent. Leaves in whorls of 4 or 5, sparsely long-
pilose beneath, acuminate with cuneate base and
glandular denticulate margin, obovate, 6'/,-11 by
21/,-5 cm; nerves 6-9 pairs, meeting in an intra-
marginal vein; petiole 8-10mm. Flowers in a
reduced axillary fascicle-like spike; axis glabrous,
c. 3mm long. Bracts and bracteoles’ persistent
under the flower, 8-10 and c. 4 mm long respec-
tively. Calyx divided into unequal narrowly
triangular appressedly long-hairy 2-4mm _ long
lobes. Stamens 70 to more than 100. Disk pilose.
Ovary glabrous. Fruit ellipsoid to cylindrical,
immature whitish, 10-12 by c. 5mm; stone with
shallow lengthwise grooves, cylindrical, a little
swollen towards both ends, 3—-celled but mostly only
one cell fertile. Seed mostly 1, straight with straight
embryo.

Distr. Malesia: Philippines (Samar: Mt Can-
sayao).

Ecol. Lowland Dipterocarp forest, 200 m. Fr.
April.

56. Symplocos vidalii ROLFE, Kew Bull. (1912) 157;
Me_rr. En. Philip. 3 (1923) 302; Noor. Leid. Bot.
Ser. 1 (1975) 290. — S. luzoniensis (non ROLFE)
BRAND, Pfl. R. Heft 6 (1901) 61, pro descr. et
specim. Vidal 2141. — S. cagayanensis BRAND,
Philip. J. Sc. 7 (1912) Bot. 35; MERR. En. Philip.
3 (1923) 297. — Fig. 7.

Twigs villous to tomentose. Leaves patently soft
pilose beneath, acuminate with acute to rounded
base and recurved, entire to denticulate margin, +
elliptic, 23/,-8 by 1-3cm; nerves 7-10 pairs;
petiole 5-7 mm. Flowers in a lax raceme to 5 cm;
axis villous. Bracts and bracteoles linear, villous, at
least the latter persistent under the fruit, 2-3 and
11/,-2!/, mm long respectively. Pedicel 1-2 mm.
Calyx (appressedly) pilose, wholly divided into the

eee

1977]

SYMPLOCACEAE (Nooteboom)

273

Fig. 20. Symplocos verticillifolia Noort. a. Habit, in fruit, x 2/3, b. endocarp, c. ditto in CS, both x 3
(Sutit 14397).

narrowly triangular acute 1—11/, mm long lobes.
Corolla 2'/,-3 mm. Stamens 17-30. Disk glabrous.
Ovary with same indument as calyx, 1-1'/, mm
high. Fruit cylindrical, c. 10 by 3 mm; stone shal-
lowly lengthwise ribbed, 1-celled. Seed 1, straight
with straight embryo.

Distr. Malesia: Philippines (Luzon: Rizal and
Nueva Ecija Prov.).

Ecol. Rain-forest at low and medium altitude.
Fl. Febr., fr. April.

57. Symplocos whitfordii BRAND, Philip. J. Sc. 3
(1908) Bot. 8; Merr. En. Philip. 3 (1923) 302;
Noort. Leid. Bot. Ser. 1 (1975) 292. — Fig. 7.

Small tree, 6-10 m, 30cm @, sometimes fasti-
giate. Twigs glabrous. Leaves glabrous, acuminate
with acute, attenuate base and crenate margin, +
elliptic, 2—5*/, by 1—2'/,(-3) cm; nerves 5-9 pairs;
petiole 2-9 mm. Raceme basally branched; axis
glabrous or sparsely (long-)pilose 1'/,-3'/, cm.
Bracts and bracteoles persistent, glabrous or
sparsely pilose on the midrib, 3-8 and 1'/,-3 mm
long respectively. Pedicel (O-—)'/,-2'/,mm _ long.
Calyx glabrous, 1'/,-2'/, mm long, the lobes ovate,
acute, I'/,-2 mm. Corolla 5-7 mm. Stamens stout,
20-30. Disk glabrous. Ovary glabrous, 1'/,-3 mm
high; style glabrous. Fruit ovoid, 5-7 by 3-5 mm;
stone ampulliform, the belly irregularly grooved.
Seed |, i ne embryo U-shaped.

Distr. Malesia; Philippines (Luzon: Mt Pulog;
Negros).

Ecol. Montane rain-forest, also in mossy forest,
1600-2450 m. Fil. Jan.—-April, fr. Febr., May.
Flowers recorded as scentless, the white corolla
outside and apically blue violet tinged.

58. Symplocos zizyphoides STapr, Trans. Linn.
Soc. Bot. 4 (1894) 205; BRAND, Pfl. R. Heft 6
(1901) 65; Merr. En. Born. (1921) 488; Noor.
Leid. Bot. Ser. 1 (1975) 293. — S. clementis MERR.
J. Str. Br. R. As. Soc. n. 76 (1917) 111; En. Born.
(1921) 486. — Fig. 7.

Small shrub, '/, m, to treelet to 4(—10) m high.
Twigs appressedly brown-pubescent, often dis-
tinctly zigzag. Leaves alternate, olive-yellow
beneath and dark brown to green above when dry,
glabrous above, nearly glabrous to appressedly
fine-pilose beneath, faintly acuminate with rounded
to cuneate base and sharply glandular dentate
margin, ovate to elliptic, 2'/,-5'/, by 1-2'/, cm;
nerves 5-8 pairs; petiole 1-2 mm. Flowers solitary
and pedicels to 12 mm, or flowers up to 3 or 4
together in a raceme and then with very short
pedicel, except sometimes the uppermost flower;
axis, pedicels, the c. 4mm long bracts and the
2-3mm_ long bracteoles appressedly brown-
pubescent. Calyx less hairy than ovary, c. 2mm
long, the lobes 1-1'/,mm. Corolla 4-6 mm.
Stamens 40 to more than 100. Disk glabrous or
with some minute hairs. Ovary acpeeanedy
pubescent, 1—1'/, mm high; style glabrous or wit
some hairs, gradually thickened towards its base,
4-5mm. Fruit purple to blackish when ripe,
ellipsoid to ovoid, sometimes a little curved, 10-12
by 5-6 mm. Seed 1, straight with straight embryo.

Distr. Malesia; Borneo (Sabah: Mt Kinabalu).

Ecol, Subalpine shrub forest and open places,
between granite rocks and on ridges, 2400-3700 m.
Fl. Jan.-May, Oct., fr. Jan., March, July.

274 FLORA MALESIANA {ser. I, vol. 8?

Dubious

Symplocos aprilis BRAND, Bot. Jahrb. 54 (1916) 221. — Type: LEDERMANN 7559 (Bt), New Guinea,

Kaiser Wilhelmsland.
Symplocos argenna BRAND, Bot. Jahrb. 54 (1916) 223. — Type: LEDERMANN 11173, 11376 (BT), East

New Guinea, Hunsteinspitze.
Symplocos imperialis BRAND, Philip. J. Sc. 4 (1909) Bot. 109; Merr. En Philip. 2 (1923) 299. — Type:

BS 4133 FEénrx, Philippines, Babuyanes Is.

Symplocos ledermannii BRAND, Bot. Jahrb. 54 (1916) 218. — Syntypes: LEDERMANN 11901, 11925,
11977, 11980, 12107, 12118 (Bt), East New Guinea, Station Schraderberg.

Symplocos leucocarpa BRAND, Bot. Jahrb. 54 (1916) 221. — Syntypes: LEDERMANN 11031, 12430,

12683 (BT), East New Guinea, Hunsteinspitze.
Symplocos lilacina BRAND, Bot. Jahrb. 54 (1916) 223. — Type: LEDERMANN 11771 (Bf), East New

Guinea, Schraderberg.
Symplocos oranjeensis BRAND in Fedde, Rep. 26 (1929) 172. — Type: VERSTEEG 2481, New Guinea,

Oranje Mts.
Excluded

Symplocos atrocyanea ELMER (Philippines, ELMER 14679), nom. in sched. = Mastixia pentandra BL. ssp.
Philippinensis (WANG.) MATTHEW (Cornaceae).

Incompletely known taxa

A number of Malesian specimens which are represented by incomplete material, but possibly represent
new taxa, are listed by NOOTEBOoM in Leid. Bot. Ser. 1 (1975) 296.

———

LENTIBULARIACEAE (P. Taylor, Kew)

A small family of annual or perennial herbs, all of which are variously adapted for
the capture and digestion of small animals (insects, Crustacea, etc.). Only one genus
(Utricularia) occurs in Malesia.

The family is cosmopolitan, including arctic regions, but is more or less absent
from Polynesia. It includes 4 genera with c. 250 spp.

The largest and most widely spread is the cosmopolitan genus Utricularia L. with
c. 180 spp., almost half of which occur in the New World, the rest being more or
less equally distributed between tropical Africa, Asia, and Australia, with a few in
the north temperate zone, 22 spp. occurring in Malesia.

Pinguicula L., with some 50 spp., has a curious distribution, with a few circum-
boreal species and concentrations in the Mediterranean region and in North, but
especially in Central and in South America, as far south as Patagonia.

Genlisea ST.HIL., with c. 16 spp., is confined to the tropics of South America and
Africa.

Polypompholyx LEHM., with 2 spp., occurs only in Australia.

All genera are associated with damp or wet habitats and most frequently found
on sterile mineral soils where they are often associated with other carnivorous
plants (as e.g. Drosera).

Unfortunately no fossils are known with certainty, apart from some Quaternary
pollen.

The affinities of the family have been the subject of considerable discussion and
opinions are divided between a relationship with Scrophulariaceae and Primulaceae.
The combination of free basal (or free central) placentation, a spurred personate
corolla (the spur is always present but occasionally reduced), two stamens and the
carnivorous habit is diagnostic for the family. In favour of affinity with Scrophu-
lariaceae are the morphology of the corolla, the structure and number of the sta-
mens, the bilobed stigma, and such cytological evidence as is available. The pollen
of Lentibulariaceae is similar to that of both of the families in question. The placen-
tation (and no doubt the mode of dehiscence of the probably most derived aquatic
European species, i.e. those most usually studied) is certainly the reason for a sug-
gested alliance with Primulaceae but the two families have little else in common.
The transition from axile to free central (or basal) placentation by the loss of the
septum is quite feasible and the mode of dehiscence of at least what are presumably
the most primitive Utricularia species could support such a hypothesis.

Within the family the combination of two-lobed calyx and trap structure is
diagnostic for the genus Utricularia. Polypompholyx is very close to Utricularia but
with 4 calyx lobes in two whorls. Genlisea and Pinguicula both have true leaves and
a 5-lobed calyx, the traps of the former genus being extremely complex but quite
different from those of Utricularia. Genlisea has also a unique type of fruit dehis-
cence — likening the fruit to a globe it splits at the equator and at least partially at
both tropics. Pinguicula has an apparently much less complex trapping mechanism
consisting of two types of superficial glands on the leaves while the dehiscence is
constantly valvate. Theories have been advanced as to how the various trapping
mechanisms could be derived one from the other but they are on the whole uncon-
vincing.

(275)

276 FLORA MALESIANA [ser. I, vol. 82

UTRICULARIA

LINNE, Gen. Pl. ed. 5 (1754) 11; Sp. Pl. (1753) 18; A. DC. in DC. Prod. 8 (1844) 3;
B. & H. Gen. Pl. 2 (1876) 987; KAMIENSKI in E. & P. Nat. Pfl. Fam. 4, 3b (1895)
119; P. TAYLOR, Kew Bull. 18 (1964) 1; Mem. N.Y. Bot. Gard. 17, 1 (1967) 206;
Komiya, J. Jap. Bot. 48, 5 (1973) 149. — Polypompholyx (non LEHM.) BENS. in Mart.
Fl. Bras. 10 (1847) 251; Linnaea 2 (1847) 447; Grises. Cat. Pl. Cub. (1866) 162;
PELLEGR. Bull. Soc. Bot. Fr. 60 (1914) 514; ibid. 61 (1914) 20; PERRIER, Mém. Inst.
Sci. Madag. sér. B. 5 (1955) 199; in Humbert, Fl. Madag., Lentib. (1955) 19. —
Biovularia KAMIENSKI, Zap. Novoross. Obtsch. Est. 12 (1890) 204; in E. & P. Nat.
Pfl. Fam. 4, 3b (1895) 122; Bot. Jahrb. 33 (1902) 113; BARNH. Mem. N.Y. Bot.
Gard. 6 (1915) 58; MELCHIOR in Engl. Syll. Pfl. ed. 12, 2 (1964) 467. — Fig. 1-26.

Annual or perennial aquatic terrestrial or epiphytic herbs always of damp places,
without true roots or leaves but with stems modified in various ways to function as
rhizoids, stolons and foliar organs, all species bearing small complex bladder-like
traps for the capture and digestion of small aquatic organisms. /nflorescence
racemose, peduncled, usually simple, bracteate; sterile bracts (scales) often present
on the peduncle and sometimes also on the inflorescence axis; two bracteoles often
present, almost always at the base of the pedicel, usually free, rarely + connate
with the bract. Bracts very varied, basifixed, medifixed or variously produced below
the point of attachment. Calyx 2-lobed, usually + accrescent, the lobes + equal
or variously dissimilar, usually free, sometimes + connate at the base. Corolla
bilabiate, yellow, various shades of violet or purple, white or rarely blue or red;
upper lip entire or 2- or more-lobed; lower lip with an entire or 2—5-lobed limb, a
+ raised, often gibbous palate and a usually subulate or conical spur, in a few
species reduced to a short sac. Stamens 2 inserted at the base of the upper lip;
filaments usually short, linear, often curved and often + flattened and dilated
above; anthers dorsifixed, + ellipsoid, thecae + confluent. Ovary globose or ovoid,
ovules 2-many on a free basal or free central + fleshy placenta; style usually short;
stigma bilabiate, the lower lip usually much larger. Capsule globose or ovoid,
dehiscing very variously by longitudinal slits, dorsiventral or rarely lateral valves,
pores or circumscissile or rarely indehiscent. Seeds 1-many, very variously shaped
and sculptured.

Distr. Cosmopolitan but mostly in the tropical zone. About 180 spp., almost half of which occur in the
New World, the rest more or less equally distributed between tropical Africa, Asia and Australia with a few
in the north temperate zone; in Malesia 22 spp.

The geographical relationships of the Malesian spp. are of some interest. Twelve species are more or less
widespread throughout tropical Asia and Australia and four of these occur also in tropical Africa. U.
subulata is widespread in tropical America and Africa, apparently absent from India, but present in
Thailand, Malaya and Borneo. U. pulchra, which is allied to the very widespread U. striatula, appears to
be endemic in New Guinea while U. salwinensis of the same affinity is known only from the Gajo moun-
tains of North Sumatra and from SW. China (Yunnan). U. vitellina is apparently local-endemic in Malaya
while the allied U. involvens is known from that country, adjacent Burma, Thailand, and N. Australia.
U. heterosepala, a slightly anomalous species in the same group (which is predominantly Asian but with
representatives in tropical Africa and to a lesser extent in America) appears to be endemic in the Philip-
pines. The circumboreal species U. minor occurs at high altitudes in New Guinea and U. australis, which is
widely distributed in the Old World north temperate zone, occurs, mostly at high altitudes, in a number of
places in Malesia; it is known also in the mountains of tropical Africa and at lower altitudes in SE.
Australia. Two species known otherwise only from northern Australia occur in SE. New Guinea:
U. chrysantha and U. muelleri. One strange apparent absence from the Malesian region is U. stellaris
which is known from tropical Africa and Asia as far as Indo-China and reappears in northern Australia.
It is included in the key to the species as it seems very probable that it does occur in the area.

1977] LENTIBULARIACEAE (Taylor) 277
SS SS ee ce nc eee ene ek

Ecol. Marshes, wet grassfields, swamps, swamp-forest, streams and rivers and open damp sandy

a a few species epiphytic among moss on trees (and rocks), occurring in Malesia from sea-level to
m.

Pollination. The flowers often secrete nectar and in some species are fragrant. Pollination by Diptera and
Hymenoptera has been observed and the flowers are sometimes visited by Lepidoptera. However, self
pollination is probably usual and cleistogamous forms are frequent and in some species inflorescences
normally bear both cleistogamous and chasmogamous flowers.

Dispersal. Dispersal over short distances can easily take place in aquatic species by floating of entire
plants or parts thereof, or by dispersal of buds (turions), according to RIDLEY (Disp. 1930).

Seeds are mostly very small and sometimes winged and therefore perhaps sometimes dispersed by wind,
although gravity is probably the most normal agent. In some of the aquatic species the seeds do not float,
or they do so only for a time (RIDLEY, Disp. 1930, 220). A few (not Mal. spp.) growing in swiftly flowing
water have seeds with a mucilaginous testa and in the epiphytic species the seeds are either very small
(orchid-like) or winged or (U. striatula) beset with glochidiate processes. Fig. 13t.

The occurrence of some species which are epiphytic among moss on tree-trunks in dense primary rain-
— ‘ogg there is hardly any wind might point to very short-distance dispersal by ants or other insects.

ig. 14.

In open terrain the seed qualities would point to wider exozoic dispersal by migrating waterfowl and
waders for aquatic species, and by wind. This might induce dispersal enthusiasts to explain the enormous
disjunct gaps in the range of U. minor between Burma and New Guinea, and that of U. stellaris between
Indo-China and Australia by erratic long-distance dispersal.

However, several terrestrial species show similar wide disjunctions, e.g. U. salwinensis between Yunnan
and North Sumatra, U. scandens and U. limosa between the Malay Peninsula and New Guinea, and U.
baouleensis between Luzon and Java. Though the present revision is based upon some 2000 collections,
the latter two species may have escaped attention of collectors in intermediate stations. However, the
disjunct range of the subalpine U. salwinensis is certainly a real gap, as high mountains are at present
absent between Yunnan and N. Sumatra. A similar disjunction is found in the ranges of other high moun-
tain plants, such as for example Swertia bimaculata and Viola biflora which are certainly not overlooked.

It should be admitted, though, that Utricularia must often have escaped attention of collectors, especially
in seasonal areas where flowering is of short duration and ephemeral. On the other hand in a thoroughly
explored island as Java, U. baouleensis is known only from Madura I. in one collection. This leads to the
conclusion that it is most unlikely that the gaps mentioned above will be reasonably filled by later
exploration, especially these of the high altitude species. This argument is strengthened by the fact that
these disjunctions are by no means unique: the gap of U. salwinensis is matched by that of Viola biflora,
Hedyotis verticillaris, etc., the gap of U. minor by that of Drosera rotundifolia and several Carices, but also
by that found in Fagoideae. Even an extraordinary range as that of U. livida, which is found in East Africa
and Madagascar but also in Mexico, is + matched by a few other taxa or affinities with similar disjunction,
in tropical America, e.g. tribe Ravenalae (Musaceae) and Rheedia (Guttiferae).

Then there are some Indo-Australian species showing a huge disjunction: U. involvens, Burma, Thai-
a ga and N. Australia, and a closely related one, U. odorata, Thailand, Indo-China and N.

ustralia.

It gives thought to the assumption that these disjunctions cannot simply be explained by erratic long-
distance dispersal. Also the occurrence of three local-endemic species makes such a correlation with
dispersal capacity highly dubious and does not plead for easy dispersal. Neither does the fact that the
ecology of many Utricularia spp. is very wide; they are not particular to soil, many are found in the tropics
under both everwet and seasonal climatic conditions, and a fair number have a very large altitudinal
range.

On the other hand it must be realized that the very widely distributed U. australis, which ranges all over
the Old World with isolated sporadic localities on the southern hemisphere, is not known to produce
fruit and seed, which forces to assume dispersal of small particles of its vegetative parts by migrating
birds. This implies that such parts should be capable to withstand desiccation which will certainly happen
during such migratory flight. Experiments could add some evidence. It is e.g. shown by V. A. WAGER
(Trans. R. Soc. S. Afr. 16, 1928, 204, pl. 24) that U. australis (under the erroneous name U. stellaris)
forms resting buds towards the end of the season which may carry the plant over until the following spring.
These resting buds are not damaged by exposure to drought; buds taken from a herbarium specimen six
months old put into an aquarium slowly swelled and developed into healthy plants.

Chromosomes. Relatively few (about 15°) of the species of Utricularia have been examined; the
chromosomes are apparently small and not easily observed. Basic numbers of x = 7 and 9 seem to pre-
dominate but x = 6, 8, 10, 11 and 15 are recorded. An American species, U. inflata WALT., has 2n = 18
and 36, the latter being morphologically gigantic whereas the closely allied U. radiata SMALL has 2n = 28.
The common Australian species U. dichotoma LaBiLt. has 2n = 28 while conversely a morphologically
small variant of this, U. uniflora R.Br., has 2n = 56. Cf. J. Caspar in Fedde, Rep. 86 (1975) 211-232.

Morph. The most remarkable feature of the genus are the traps. They are minute vesicles provided
with an apical orifice at the ventral side. The narrow opening leading to the water-filled cavity is formed by
a ventral lip, and a dorsal valve which enables the prey to enter, but prevents it from ha Ne in which it
is also hampered 1 cer meee papillae of striking structure. The inner wall of the trap is densely glandular-
papillose and exudes proteolytic enzymes. j iN BD ;

functioning of the trap, by the opening of the valve, is caused by irritation of the sensitive hairs on
this lid by which small crustacea or other matter is ‘sucked’ in. See E. Mert (Flora: Allg. Bot. Z. 115,

278 FLORA MALESIANA [ser. I, vol. 8?

1922, 59-74) extracted by JACOBSON in Trop. Natuur 17 (1928) 107-113, 4 fig., in Dutch, and especially
F. E. Ltoyp, The Carnivorous Plants (1942) 233-270; furthermore the excellent survey by Y. HESLOP-
HARRISON (Endeavour 35, 1975, 114-122).

Palyn. Pollen grains are tricolporate or stephanocolporate with up to 28 colpi. Tetrads occur in two
species of Utricularia. Size ranges from 17 um in U. neottioides to 51 ym in U. humboldtii. Shape varies
from oblate to prolate. Sculpture is generally smooth in Utricularia or finely reticulate as in Pinguicula
(ERDTMAN, Pollen morphology and plant taxonomy. Angiosperms. Almquist & Wiksell, Stockholm,
1952, 233-234).

In Utricularia HUYNH (Etude de la morphologie du pollen du genre Utricularia L., Pollen et Spores 10,
1968, 11-55) has described a considerable amount of pollen-morphological variability. Tricolporate types
with ‘long or very short colpi and stephanocolporate types with equatorially fused endoapertures occur.
In some types the apocolpia are dissected by interconnections between colpi.

A remarkable similarity exists between the stephanocolporate grains in Utricularia and the pollen of
Polygalaceae. — J. MULLER.

Notes. The study of Utricularia has always been hampered by badly collected and inadequate material.
Of the aquatic species the vegetative parts should be suitably supported by a (rigid) sheet of paper and
thus be raised out of the water in their natural position and then dried as rapidly as possible. Additional
inflorescences and infructescences should be added. Terrestrial species have often not very obvious vegeta-
tive parts which are usually beneath or in the substrate. They must be carefully ‘unearthed’, or dried with
the adhering mud.

Specimens in liquid are of course excellent. FAA is not so suitable; the best solution is a mixture of
50-55 % alcohol, 40% water and 5-10% glycerine; the latter may be omitted or even added later; it is
important that the receptacle is entirely filled with liquid.

It is important to take many flowers but see that also fruit and seed are represented.

As two or more terrestrial species are frequently growing together, with their vegetative parts intimately
mixed in the substrate, care is needed in collecting. It is important to note the flower colour; this is in
several species very variable.

Hardly any Utricularia spp. have been collected in the Lesser Sunda Islands. Though this archipelago is
subject to a dry season, collecting at the the end of the wet season in rice-fields, shallow swamps and damp
grassland may yield interesting results, as several species have been found in the adjacent Madura &
Kangean Islands which have the same ‘climatic regime.

KEY TO THE SPECIES

1. Foliar organs conspicuous and numerous at anthesis, divided into narrowly linear to capillary segments,
the ultimate segments bearing apical and often lateral solitary or fasciculate setulae.
2. Turions (winter buds) of tightly clustered modified foliar segments present at the apex of some of the

stolons.
3. Turions glabrous or almost so. Ultimate foliar segments with minute apical but with or without
microscopic lateral setulae . 22. U. minor

3. Turions densely setulose. Ultimate foliar segments with aa numerous lateral setulae.

21. U. australis
2. Turions not present.
4. Primary foliar segments 3 or more semi-verticillate on the stolons usually with hyaline or foliose
setulose stipule-like segments at the base. Scales on peduncle absent.
5. Peduncle with a whorl of inflated ellipsoid floats some distance above the base. Basal segments of
foliar organs hyaline.

6. Floats shortly stipitate with capillary foliar segments at the base. Capsule much longer than the
calyx lobes. Seeds thinly lenticular with a single marginal wing . . . . 20. U. muelleri
6. Floats sessile without foliar segments at the base. Capsule shorter than the calyx. Seeds prismatic,
narrowly winged on the angles. Africa, Madagascar, SE. Asia, Australia. Cf P. TAYLoR, Kew

Bull. 18 (1964) 189, f. 77 6-11, 79; Aston, Aquat. Pl. Austr. (1973) f.39a-j . U. stellaris L. 7;
5. Peduncle without floats or with a whorl of narrowly fusiform floats at the base. Basal segments of
TOMATIOLSANSMONOSC Pasay atettajes eka) ce Pleo sd, eee nc OS Oe wp es, ee ee 19. U. aurea

4, Primary foliar segments 1 or 2 usually without stipule-like segments at the base. Scales (or at least
One) present on the peduncle.

7. Foliar organs less than 10 mm long with few (2-10) ultimate segments with few or no lateral setulae.

8. Ultimate segments distinctly flattened with microscopic or no lateral setulae. Bracts and scales

auriculate, the scales few but dispersed through the length of the peduncle. Spur of the corolla very

short, saccate. Capsule circumscissile. Seeds prismatic . . 22. U. minor

8. Ultimate segments terete, sometimes with a few lateral setulae. Bracts and scales not auriculate,

scales usually only one near the middle of the peduncle. Spur of the corolla narrowly conical.

Capsule laterally bivalvate. Seeds lenticular, winged ps is Pe et . . 18. U. exoleta

7. Foliar organs more than 20 mm long with very numerous ultimate segments ‘bearing + numerous

lateral setulae.

9. Ultimate segments terete. Traps absent from the lowermost part of the foliar organs. Bracts and

scales medifixed, the scales disposed through the Pai of the oes Corolla mauve or white.

Seeds with a broad dentate WIR e). .kuetas ATs : ; sos 6 ws vit. UU pete

ee

1977]

LENTIBULARIACEAE (Taylor)

279

9. Ultimate segments distinctly flattened. Traps present in the lowermost part of the foliar organs
and with rudimentary traps at the very base. Bracts and scales basifixed, slightly auriculate, the
scales few and usually present only in the upper part of the peduncle. Corolla yellow.

21. U. australis

1. Foliar organs simple, orbicular to linear, never setulose, often not present or conspicuous at anthesis.
10. Bracts medifixed or produced below the point of attachment.

11. Bracteoles present.

12. Calyx lobes approximately equal in size. Foliar organs linear to narrowly obovate, usually not

conspicuous at anthesis

11. U. caerulea

12. Calyx lobes very unequal in size, the upper much larger. Foliar organs reniform to obovate, usually

present and + conspicuous at anthesis.

13. Corolla lower lip 3-lobed, the lateral lobes obliquely emarginate, spur c. 2 mm long. Seeds ovoid,

densely echinate

14. U. salwinensis

13. Corolla lower lip + regularly 5-lobed, spur at least 5 mm long.
14. Spur over 10 mm long, usually longer than and widely diverging from the lower lip of thecorolla.

Flowers 1 or 2. Peduncle up to 6 cm long, relatively stout

13. U. pulchra

14. Spur c. 5 mm long, usually about as long as and + parallel with the lower lip of the corolla.
Flowers up to 10. Peduncle up to 15 cm long, very slender. Seeds pyriform, glochidiate.

11. Bracteoles absent.

12. U. striatula

15. Lower lip of corolla deeply 2-lobed, corolla pink or white. Pedicel minutely papillose. Seeds

globose with distinct coarse isodiametric reticulation

15. U. limosa

ie Y Oba te ed a Wee tee Gey oes

15. Lower lip of corolla deeply 3-lobed, corolla yellow. Pedicel smooth. Seeds ellipsoid with indistinct

coarse very elongate reticulation
10. Bracts basifixed.

16. Bracteoles absent. Lower calyx lobe much longer than upper in fruit

Pe en 6 Cae SR eee a ei nd, ey ae Rene el ae

16. U. subulata
6. U. heterosepala

16. Bracteoles present. Calyx lobes + equal or upper longer.
17. Bracteoles not much narrower (at least half as wide) than the bract.

18. Calyx, peduncle and pedicels + densely covered with often long septate hairs
18. Calyx glabrous. Peduncle glabrous or with a few short septate hairs

10. U. hirta
9. U. minutissima

17. Bracteoles much narrower (less than one quarter as wide) than the bract.

19. Lower lip of corolla deeply 4-lobed. Corolla yellow

8. U. cirysantha

19. Lower lip of corolla entire, emarginate or + 3-lobed.

20 Corolla yellow.
21. Peduncle twining.

22. Corolla (11-)15-20 mm long, upper lip orbicular, much wider than the calyx lobes.

22. Corolla 5-10 mm long, upper lip oblong, narrower than the calyx lobes

21. Peduncle erect.

23. Lower lip of corolla 3-lobed, corolla 15-22 mm long

2. U. involvens
. 5. U. scandens

1. U. vitellina

ae ak eee ee ee ae

23. Lower lip of corolla entire or emarginate, corolla 6-10 mm. Pedicels strongly recurved in

eee Corey ate ee Oe eee ee
20. Corolla mauve or white.

24. Peduncle twining. Pedicels strongly reflexed in fruit.
24. Peduncle erect. Pedicels erect or spreadinginfruit. ...........

1. Utricularia vitellina Ripv. Fl. Mal. Pen. 2 (1923)
492; Spare, Mal. Nat. J. 1 (1940) 89. — U. aurea
(non Lour.) Rip. J. Fed. Mal. St. Mus. 6 (1915)
165. — Fig. 1.

Terrestrial. Rhizoids capillary, simple. Stolons
capillary, sparsely branched. Foliar organs present
at anthesis, sparsely rosulate, narrowly linear,
membranous, 1-2cm long, 0.5-1mm_ wide,
l-nerved, apex rounded. Traps few on the stolons
and foliar Sepnes, globose, 0.5-1 mm long, shortly
stalked, mouth basal, upper lip with 2 short subu-
late reflexed appendages. Inflorescence erect,
2.5-5 cm long; peduncle filiform, terete, glabrous;
flowers 1-2; scales few in the lower third of the
peduncle, the lowermost usually at its very base,
similar to the bracts ; bracts basifixed, ovate-deltoid,
acute, c. 1.5 mm long; bracteoles narrowly linear,
acute, slightly shorter than the bract; pedicels

ing, usually curved, 4-8 mm long, dorsi-
ventrally fattened, very narrowly winged. Calyx
lobes unequal, upper narrowly ovate, c. 4.5 mm
long, apex obtuse, lower similar but c. 3 mm long,

eS g) eeeen=® Cea ee Lhe ee eee

3. U. bifida

7. U. baouleensis
4. U. uliginosa

apex obscurely bidentate. Corolla yellow streaked
with brown, 15-22 mm long, upper lip obovate-
oblong to + orbicular, slightly longer than the
upper calyx lobe, apex rounded or emarginate,
lower lip much larger, up to 10mm wide, +
orbicular, apex distinctly 3-lobed, palate scarcely
prominent, spur subulate, about as long as the
lower lip but widely diverging from it, usually
strongly curved. Filaments filiform, + straight,
c. | mm long, anther thecae distinct. Ovary ovoid,
style short, distinct, stigma lower lip orbicular,
upper obsolete. Capsule and seeds not known,

Distr. Malesia; Malay Peninsula (Pahang: G.
Tahan and G. Kerbau only).

Ecol, Peaty streambanks in moss, c. 1500-2100
m. Fl. Jan.-July.

2. Utricularia involvens Ript, J. Bot. 33 (1895) 11;
Pratn, J. As, Soc, Beng. 74, ii (1905) 371; Riou. Fl.
Mal. Pen. 2 (1923) 493, f. 121; Spare, Mal. Nat,
J, 1 (1940) 89. — Fig. 3.

Terrestrial. Rhizoids few, capillary, basally

280

Fig. 1. Utricularia vitellina RipL. a. Flowering
plant, x 4, b. trap, <x 24, c. bract & bracteoles,
x 12, d. corolla, the two lips from inside, x 4, e.
Stameniee elon wpIStiln, elie.) evcalyx: <6) (all
after RIDLEY, type, except b SPARE S4/41).

thickened, with numerous papillose branches
0.5-1 mm long. Stolons few, capillary, branched.
Foliar organs usually conspicuous at anthesis,
rosulate and on the stolons, obovate to narrowly
oblong, membranous, up to 2.5 cm long, 2-4 mm
wide, multinerved, apex rounded. Traps on the
vegetative organs, globose, 0.5—1 mm long, shortly
stalked, mouth basal, upper lip with 2 simple
subulate appendages, lower lip with 1 short obtuse
appendage. Inflorescence twining, up to 30cm
long; peduncle filiform, terete, glabrous; flowers
2-6, distant ; scales numerous, similar to the bracts;
bracts basifixed, ovate, acute, 2-3mm _ long;
bracteoles subulate, shorter than the bract; pedi-
cels erect or spreading, filiform, 10-15 mm long,
narrowly winged. Calyx lobes subequal, ovate to

FLORA MALESIANA

[ser. I, vol. 82

Fig. 2. Habit of some Utricularia species. a. U.
baouleensis A.CHEV. trailing along a sedge, b.
U. bifida L., c. U. aurea Lour. (inflorescence only),
d. U. uliginosa VAHL and d’. detail of vegetative
parts, e. U. striatula J.SM. with cleistogamous
flowers, f. U. exoleta R.Br. All x 2/3.

1977]

Fig. 3. Utricularia involvens RIDL. a. Trap, x 24,

b. bract & bracteoles, x 12, c. corolla, the two lips

from inside, x 2, d. pistil, x 12,e.stamen, x 12,

f. fruiting calyx, x 2, g. dehisced capsule, x 4, h.

seed, x 24 (a, b, h Dinc Hou 783, the others after
RIDLEY, type).

broadly ovate, 4-5 mm long at anthesis, up to
7mm long in fruit, upper slightly larger, apex
obtuse or subacute, lower shortly bidentate.
Corolla yellow, 11-20 mm long, upper lip orbicular,
longer than the upper calyx lobe, 7-12 mm wide,
apex rounded, lower lip + orbicular, up to 15 mm
wide, apex emarginate, palate very conspicuously
raised, gibbous, extending almost to the apex of
lower lip, spur subulate, straight or slightly curved,
about as long as but widely diverging from the
lower lip. Filaments linear, curved, c. 1.5 mm long,
anther thecae subdistinct. Ovary ovoid, style rela-
tively long, stigma lower lip orbicular, deflexed,
upper lip obsolete. Capsule broadly ovoid, dorsi-
ventrally compressed, uniformly membranous,
4-5 mm long, dehiscing by dorsal and ventral
longitudinal slits. Seeds numerous, ovoid, c.0.4 mm
long, testa conspicuously coarsely reticulate,
reticulations elongate.

Distr. S. Burma (Tenasserim), Thailand, N.
Australia, and Malesia: Malay Peninsula (Kedah
Peak, G. Jerai).

Ecol. Damp grassy places and along creeks,
900-1000 m in Malaya but at low altitude in
Thailand and Australia. F/. April-Jan.

3. Utricularia bifida Linné, Sp. Pl. (1753) 18;
Oxtver, J. Linn. Soc. Lond. Bot. 3 (1859) 182;
Crarke, Fl. Br. Ind. 4 (1884) 332; Rip. Trans,
Linn. Soc. Lond. II, Bot. 3 (1893) 327; Fl. Mal.
Pen. 2 a 492; Merr. En. Philip. 3 (1923) 466;
Peciecr. FI. Gén. L-C. 4 (1930) 482; Sreen.
Arch, Hydrobiol. Suppl. 11 (1932) 331, f. 8 1;

LENTIBULARIACEAE (Taylor)

281

HAND.-MAzz. Symb. Sin. 7 (1936) 872; Back. &
BAKH. f. Fl. Java 2 (1965) 518; P. TAYLor, Dansk
Bot. Ark. 23 (1968) 529. — U. recurva Lour. FI.
Coch. (1790) 26. — U. humilis VAHL, Enum. 1
(1804) 203. — U. ramosa VauHt, l.c. 204. —
U. antirrhinoides WALL. Cat. (1829) n. 1498,

nomen. — U. wallichiana BENS. Bot. Zeit. 3 (1845)
213. — U. brevicaulis BENJ. Linnaea 20 (1847) 303.
— U. sumatrana Mia. FI. Ind. Bat. 2 (1859) 998,
p.p.; Suppl. 1 (1860) 246. — U. biflora Hayata, J.
Coll. Sc. Imp. Un. Tokyo 30 (1911) 210; Ic. Pl.
Formos. 2 (1913) 125. — Fig. 2b, 4.

Fig. 4. Utricularia bifida L. a. Foliar organ, = 6, b.
trap, x 24, c. bract & bracteoles, < 15, d. flower,
x 6, e. upper lip of corolla, * 6, f. stamen, x 12,
g. pistil, x 12, h. fruiting calyx, = 6, i. capsule,
x 6,j. seed, x 45, k. ery x 75 (all after LARSEN

Terrestrial. Rhizoids few, capillary, basally
thickened, with numerous papillose branches
0.5-1 mm long. Stolons few, capillary, branched,
Foliar organs usually +. conspicuous at anthesis on
the stolons, narrowly linear, membranous, 1-2 cm
long, up to | mm wide, I-nerved, apex rounded,
Traps on the vegetative organs, globose, 0,6-1 mm
long, stalked, mouth basal, upper lip with 2 simple
subulate appendages, lower lip with a + well
developed obtuse swelling at the distal end of the
stalk, Inflorescence erect, 5-20 cm oy peduncle
filiform, terete, glabrous above, minutely glandular

282

FLORA MALESIANA

[ser. I, vol. 8?

below; flowers 1-10, distant; scales few, similar to
the bracts; bracts basifixed, broadly ovate-oblong,
obtuse, c. 1 mm long; bracteoles linear-subulate,
c. 0.5 mm long; pedicels erect at anthesis strongly
recurved in fruit, filiform, distinctly winged,
2-4mm long. Calyx lobes subequal, the upper
slightly larger, broadly ovate, base + broadly
connate and decurrent, apex obtuse, c. 3 mm long
at anthesis, up to 6mm long in fruit. Corolla
yellow, 6-10 mm long, upper lip narrowly oblong,
1-2 mm wide, apex rounded, slightly longer than
upper calyx lobe, lower lip + orbicular, up to
4mm wide, apex rounded, palate conspicuously
raised, gibbous, spur subulate, acute, curved, longer
than and widely diverging from the lower lip.
Filaments oblong, straight, c. 1 mm long, c. 0.5 mm
wide, anther thecae subdistinct. Ovary ovoid, style
short but distinct, stigma lower lip quadrate, de-
flexed, upper lip much shorter, entire or bidenticu-
late. Capsule broadly elliptic, dorsiventrally com-
pressed, uniformly membranous, 2.5—3 mm long,
dehiscing by a single ventral longitudinal slit.
Seeds numerous, + ovoid, c. 0.4mm long, testa
rugose, reticulate, reticulations relatively large,
elongate.

Distr. India to China and Japan, Indo-China,
Malesia to northern Australia; in Malesia not
recorded from the Lesser Sunda Is. (but present in
Kangean Is.) or Moluccas, but common elsewhere.

Ecol. Swamps and marshes, sometimes as a
weed in rice-fields, on moist sandy plains, floating
in lakes, in Sphagnum swamps, on edge of Mela-
leuca swamp and in moist Eucalypt savannahs,
mainly at low altitude, but up to at least 2000 m (in
Java and New Guinea). F/. Jan.—Dec.

Vern. Malaya: bunga janggut kéli, b. kéning
layah, M; New Guinea: ararébo, pékatorro,
Kapauko lang.

4. Utricularia uliginosa VAHL, Enum. 1 (1804) 203;
SANTAPAU, J. Bomb. Nat. Hist. Soc. 49 (1950) 217;
P. TayLor, Dansk Bot. Ark. 23 (1968) 532. —
U. cyanea R.Br. Prod. Nov. Holl. (1810) 431;
GUILLAUMIN, FI. Nouv.-Caléd. (1948) 319. —
U. affinis WiGuT, Ic. (1850) t. 1580; Mia. Fl. Ind.
Bat. Suppl. 1 (1860) 246; CLARKE, Fl. Br. Ind. 4
(1884) 330; TRIMEN, Handb. FI. Ceyl. 3 (1895) 269;
PELLEGR. FI. Gén. I.-C. 4 (1930) 479; STEEN. Arch.
Hydrobiol. Suppl. 11 (1932) 333, f. 8 III; Back. &
BAKH. f. Fl. Java 2 (1965) 518. — U. griffithii
Wiaurt, Ic. (1850) t. 1576; Mia. Fl. Ind. Bat. 2
(1859) 999; RipL. Fl. Mal. Pen. 2 (1923) 492;
SparE, Mal. Nat. J. 1 (1940) 89. — U. affinis var.
griffithii (WIGHT) OLIveR, J. Linn. Soc. Lond. Bot.
3 (1859) 179; CLarRKE, FI. Br. Ind. 4 (1884) 331;
Ripe. J. Str’ Br._R. As: Soc: 77: 33 (L900)0019;
PrRAIN, J. As. Soc. Beng. 74, ii (1905) 370; MERR.
En. Born. (1921) 537. — Fig. 2d, 5.

Terrestrial. Rhizoids few capillary, basally
thickened, with numerous papillose branches
c. 1 mm long. Stolons few, capillary, branched, up
to 6 cm long or more. Foliar organs often not con-
spicuous at anthesis, on the stolons, ovate to linear,
membranous, up to 4cm long, 1.5-6 mm wide,
multinerved, apex obtuse to subacute. Traps on the
stolons and foliar organs, globose, 1-2 mm long,
shortly stalked, mouth basal, upper lip with 2
simple subulate appendages. Inflorescence erect,
up to 30cm long; peduncle filiform, terete, glab-

Fig. 5. Utricularia uliginosa VAHL. a. Small, young,

foliar organ, x 6, b. part of fully developed foliar

organ, venation, x 4, c. trap, x 12, d. bract &

bracteoles, x 12, e. flower, x 6, f. fruiting calyx,

x 6, g. margin of this, x 24, h. capsule, x 6, i.
seed, x 24 (all after SINCLAIR 10157).

rous; flowers 2-10, rather distant; scales few,
similar to the bracts; bracts basifixed, ovate,
acuminate, 2—-3mm long, 3-nerved; bracteoles
subulate, about half as long as the bract, 1-nerved;
pedicels erect 1.5-2 mm long at anthesis, often
spreading and up to 3 mm long in fruit, filiform
distinctly winged. Calyx lobes subequal, very
broadly ovate to almost orbicular, c. 2.5 mm long
at anthesis, up to 5mm long in fruit, surface
minutely papillose, margin usually minutely
denticulate, upper lobe slightly larger, apex very
shortly acuminate, lower with apex shortly bifid.
Corolla blue, violet, mauve or white, 3-7 mm long,
upper lip + orbicular scarcely longer than upper
calyx lobe, apex rounded, emarginate or + bifid,
lower lip larger, + orbicular, apex rounded, entire
or + obscurely 3-crenate, palate raised, gibbous,
spur conical-subulate, acute, curved or +straight,
longer than and widely diverging from the lower
lip. Filaments linear, curved, c. 1.5 mm long, anther
thecae distinct. Ovary ovoid, style short but distinct,
stigma lower lip quadrate, upper lip obsolete.
Capsule broadly ellipsoid, dorsiventrally com-
pressed, 2-4mm long uniformly membranous,
dehiscing by dorsal and ventral longitudinal slits.
Seeds numerous, + globose or very broadly
ellipsoid, c. 0.3-0.4 mm long, testa thin, conspi-
cuously reticulate, reticulations + regularly
hexagonal isodiametric or very slightly elongate,
hilum not prominent.

Distr. India to Japan and Australia, also in New
Caledonia; in Malesia: Sumatra (also Banka &
Riouw Is.), Malaya, Java, Borneo, and New
Guinea.

1977]

LENTIBULARIACEAE (Taylor)

283

eee eeeeFFFFFsFsmmsMFsFhFsFseseF

Ecol. Swamps, streamsides and wet sandy
places, abandoned mining-land, in Melaleuca
savannahs, in siil grass-wastes, both under ever-
wet and seasonal climatic conditions, from sea-
level to 2100 m (in New Guinea). F/. Jan.—Dec.

5. Utricularia scandens BEN). Linnaea 20 (1847)
309; P. TAYLor, Dansk Bot. Ark. 23 (1968) 531. —
U. volubilis WIGHT ex BENJ. Linnaea 20 (1847) 309,
non R. Br. 1810, pro syn. — U. wallichii Wicut,
Hook. J. Bot. Kew Misc. 1 (1849) 372. — U.
wallichiana WIGHT, Ic. 4 (1850) t. 1572, non BEN’.
1845; Oxtver, J. Linn. Soc. Lond. Bot. 3 (1859)
182; CLARKE, FI. Br. Ind. 4 (1884) 332; PraIn, J.
As. Soc. Beng. 74, ii (1905) 371; Riv. Fl. Mal.
Pen. 2 (1923) 493; GAmBLE, FI. Madras 2 (1924)
982; PELLEGR. FI. Gén. I.-C. 4 (1930) 484; SPARE,

Fig. 6. Utricularia scandens Bens. a.
uncle with rhizoids, foliar organ and traps, * 4,

Base of

. trap, x 30, c. bracteoles, x 15, d.
calyx, < 6, ¢. a large flower, x 4, f. two small
flowers, x 4, g. upper lip, x 6, h. stamens, = 15, i.

istil, x 15, j. fruit concealed by calyx, = 4, k.
ruiting calyx, x 6, 1. seeds, x 45 (all after LARSEN
6195, except a and ¢ > seme & TAYLOR

Mal. Nat. J. 1 (1940) 89. — U. gibbsiae Starr, Fl.
Trop. Afr. 4 (1906) 574. — Polypompholyx
madecassa PERRIER, Mém. Inst. Sci. Madag. sér. B,
5 (1955) 199; in Humbert, Fl. Madag., Lentib.
(1955) 19. — U. scandens ssp. scandens P. TAYLOR in
Hutch. & Dalz. Fl. W. Trop. Afr. ed. 2, 2 (1963)
378; Kew Bull. 18 (1964) 46. — Fig. 6.

Terrestrial. Rhizoids numerous, capillary, with
papillose branches c. 0.5mm long. Stolons few
capillary branched. Foliar organs on the stolons,
linear, membranous, up to I cm long, c. 1mm
wide, 1-nerved, apex rounded or subacute. Traps on
the vegetative organs, globose, 0.6—-1 mm long,
shortly stalked, mouth basal, upper lip with 2
simple subulate recurved appendages, lower lip
with 1 shorter truncate or shortly bifid usually
apically glandular appendage. Inflorescence twin-
ing, 3-35cm long; peduncle filiform, terete,
glabrous; flowers 1-8, distant, with 1 or 2 sterile
bracts on the internodes; scales few, similar to the
bracts; bracts broadly ovate-deltoid, acute or
acuminate, 1-1.5 mm long, nerveless; bracteoles
linear-lanceolate, about as long as the bract,
nerveless ; pedicels erect, filiform, narrowly winged,
about as long as the flowering calyx. Calyx lobes
subequal, ovate or broadly ovate, 2.5-3 mm long
at anthesis, up to 5 mm long in fruit, upper slightly
larger, apex shortly acuminate, lower with apex
shortly bi- or tridentate. Corolla yellow, 5-10 mm
long, upper lip oblong, usually shorter and narrow-
er than upper calyx lobe, apex rounded, entire or
emarginate or + bifid, lower lip larger, + orbicu-
lar, apex rounded, entire or 2—3-crenate, palate +
raised, smooth or 2—4-gibbous, spur subulate,
acute, curved, longer than and widely diverging
from the lower lip. Filaments linear, + straight,
anther thecae + confluent. Ovary ovoid, style
short, indistinct, stigma lower lip semi-orbicular,
upper lip similar, smaller. Capsule oblong-ovoid,
dorsiventrally compressed, membranous, 2-2.5
mm long, dehiscing by a single ventral longitudinal
marginally thickened slit. Seeds numerous, broadly
ellipsoid or ovoid, c. 0.2mm long, testa thin,
smooth, reticulate, reticulations elongate, hilum
lateral, prominent.

Distr. Tropical Africa, South Africa, Madagas-
car, India, Indo-China, through Malesia to N.
Australia; in Malesia: a single record from Malay
Peninsula (G. Ledang) and more widespread in
New Guinea and the adjacent Aru Is.

Ecol. Damp places twining up Mab
Malaya at 1200 m, ascending in New Guinea from
the lowland to 2700 m. Fi. April-Aug.

6. Utricularia heterosepala Bens. Linnaea 20 (1847)
310; Mia. Fl. Ind. Bat. 2 (1859) 1000; VIDAL, Rev.
Pl. Vasc. Filip. (1886) 200; Merr. En. Philip. 3
(1923) 466. — rig. .

Terrestrial. izoids few, capillary, basally
thickened, with numerous papillose branches
c. 0.5 mm long. Stolons numerous br pee much
branched and mat-forming, up to 15cm long or
more. Foliar organs very numerous and conspi-
cuous at anthesis, on the stolons, narrowly ovate or
elliptic, membranous, 1-2 cm long, 2-3 mm wide,
apex rounded, multinerved. Traps numerous on
the vegetative organs, globose, 1-2 mm_ long,
shortly stalked, mouth basal, upper lip with 2 long
simple subulate appendages, lower lip sometimes

284

FLORA MALESIANA

[ser. I, vol. 8?

b

Fig. 7. Utricularia heterosepala BeENJ. a. Foliar
organ with trap, b. bract, c. flower, d. upper lip, all
x 6, e. glands inside spur, x 45, f. stamen, x 12,
g. pistil, x 12, h. calyx with dehisced fruit, a
thickened area bordering the cleft, x 6, i. seed,
« 24 (a—b, h-i ELMER 13127, c-g MERRILL 2085).

with a small + obscure lamelliform projection at
the distal end of the stalk. Inflorescence erect,
4-15 cm long; peduncle filiform, terete, glabrous;
flowers 3-12, distant; scales few, similar to the
bracts; bracts basifixed, ovate-deltoid, acute, 2-3
mm long, 3-nerved; bracteoles absent; pedicels
erect, filiform, very narrowly winged. Calyx lobes
subequal at anthesis, very unequal in fruit, narrow-
ly ovate, both c. 4mm long at anthesis, apex of
upper acute, of lower obscurely bidentate, lower
conspicuously accrescent and up to 6 mm long and
2.5mm wide in fruit. Corolla pink or white,
8-11 mm long, upper lip narrowly oblong or
oblong-obovate, about 1'/, times as long as upper
lobe, apex rounded or truncate, lower lip larger, +
orbicular, apex rounded, entire or emarginate,
palate conspicuously raised, gibbous, spur subulate,
acute, curved, longer than and widely diverging
from the lower lip. Filaments linear, straight,
c. 1mm long, anther thecae + confluent. Ovary
ovoid, style short, stigma lower lip quadrate,

deflexed, upper lip much smaller, rounded.
Capsule very narrowly ovoid, membranous, c. 3
mm long, dehiscing by a ventral longitudinal mar-
ginally thickened slit. Seeds few, globose, c. 0.5 mm
long, testa thin, reticulate, reticulations distinct,
isodiametric or slightly elongate.

Distr. Malesia: Philippines (Palawan, Luzon,
and Sibuyan).

Ecol. In wet places on stones in the forest and
along streams and creeks at medium and low
altitudes (MERRILL). F/. Febr.-May.

Note. Specimens of this very distinct species
(ELMER 13127, Palawan) were dist. ibuted bearing
the manuscript name U. e/meri Starr. I can find no
evidence of this name ever being published.

7. Utricularia baouleensis A. CHEV. Bull. Soc. Bot.
Fr., Mém. 8 (1912, ante Sept. 21); P. TAYLOR in
Hutch. & Dalz. Fl. W. Trop. Afr. ed. 2, 2 (1963)
378; Kew Bull. 18 (1964) 69; Back. & BAKH. f. FI.
Java 2 (1965) 518. — U. scandens (non BENJ.)
Ouiver, J. Linn: Soc. Lond. Bot. 3 (1859) 181;
CLARKE, FI. Br. Ind. 4 (1884) 332. — U. tenerrima
Merr. Philip. J. Sc. 7 (1912, Sept. 30) Bot. 247;
Fl. Manila (1912) 433; En. Philip. 3 (1913) 467;

Fig. 8. Utricularia baouleensis A.CHEV. a. Foliar

organs, x 4, b. trap, x 15, c. bract & one (of two)

bracteoles, x 15, d. calyx, x 6, e. flower, x 6,

f. upper lip of corolla, x 6, g. stamens, x 15, h.

pistil, x 15,h. fruiting calyx, x 6,i. calyx, flattened,

x 6, j. capsule, x 6, k. two seeds, x 45 (all after
FAULKNER 269A).

1977]

LENTIBULARIACEAE (Taylor)

285

Back. Onkr. Suiker. (1931) 635; BAKH. f. in Back.
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 194, p. 3. —
Fig. 2a, 8.

Terrestrial. Rhizoids few, capillary, with numer-
ous papillose branches c. 0.5 mm long. Sto/ons few,
capillary branched, up to 5 cm long. Foliar organs
usually not conspicuous at anthesis, on the stolons,
linear, membranous, up to 3 cm long, 0.4-1 mm
wide, apex acute or rounded, I-nerved. Traps few
on the stolons and foliar organs, globose, 0.8-
1.2 mm long, shortly stalked, mouth basal, upper
lip with 2 filiform-subulate sparsely branched
appendages. Inflorescence twining, up to 20cm
long; peduncle capillary, terete, glabrous; flowers
2-5, very distant; scales few, similar to the bracts;
bracts ovate to ovate-oblong, c. 1.2mm long,
nerveless, apex obtuse to shortly acuminate;
bracteoles linear-lanceolate, acute, + straight,
about half as long as the bract, nerveless; pedicels
erect at anthesis, sharply reflexed in fruit, about as
long as the calyx, filiform, dorsiventrally flattened
and narrowly winged. Calyx lobes subequal, ovate,
obtuse to subacute, c. 2mm long at anthesis,
3.54 mm long in fruit. Corolla pale blue or mauve,
3-4 mm long, upper lip oblong, apex truncate,
slightly longer than the upper calyx lobe, lower lip
larger, + orbicular, apex obscurely 3-crenate,
palate scarcely raised, spur narrowly conical,
obtuse, longer than and + diverging from the
lower lip. Filaments linear, + straight, anthers
c. 0.3 mm long, thecae subdistinct. Ovary ovoid,
style short, distinct, stigma lower lip orbicular,
upper much shorter, truncate. Capsule broadly
ovoid, dorsiventrally compressed, membranous,
dehiscing by a longitudinal ventral marginally
thickened slit. Seeds numerous, ovoid or ellipsoid,
c. 0.3mm long, testa loose, corky, distinctly
reticulate, reticulations elongate.

Distr. Scattered in tropical Africa from Mali to
Mozambique, in Madagascar, India, China
(Hainan), Thailand, and Malesia to Queensland;
in Malesia: E. Java (Madura: Lampek), Philip-
pines (Luzon).

Ecol. Damp places twining up grasses, in
Madura abundant in rice-fields. F/. March, Sept.

Note. This extremely widespread and distinct
but very inconspicuous species is probably much
commoner than the few specimens seen (less than
20) would suggest. Despite the wide geographical
range the Queensland examples (and all from inter-
mediate localities) are identical in every respect
with those from West Africa.

8. Utricularia chrysantha R. Br. Prod. Nov. Holl.
(1810) 432; Bru. Fl. Austr. 4 (1869) 527; F. M.
BAILEY, Queens!. Fl. 4 (1901) 1127; Ewart &
Davies, Fl. North. Terr. (1917) 249: SPECHT,
Arnhem Land Exped. 3 (1958) 301. — Fig. 9.
Terrestrial. Rhizoids numerous, capillary, with
numerous short ron branches. Stolons few,
capillary, sparsely branched. Foliar organs not
crore conspicuous at anthesis, on the stolons,
-prnarcle | linear, up to 3 cm long and 0.7 mm wide,
. Traps few on the stolons and foliar
organs, obliquely ovoid, 0.4-0.7 mm long, sessile,
mouth lateral, upper lip with a prominent dorsi-
ventrally flatte , apically rounded aepentens
fringed with long unicellular setae, lower lip with a
shorter appendage fringed with shorter setae.

Fig. 9. Utricularia chrysantha R.Br. a. Foliar
organ, x 6, b. part of rhizoid, x 6, c. trap, lateral
view, X 24, d. ditto, dorsal view, x 24, e. fruit in
calyx, x 6, f. bract, bracteoles & (on right) scale,
Xl Des flower with upper lip half bent back,
x 4, h. stamen, x 12, i. pistil, x 12, j. capsule,
ventral view, X 6, k. seed, x 75 (a-d, f-j ADAMS
1737, e GEORGE 12231, k PULLEN 7136).

Inflorescence erect, up to 55cm long (-63 fide
PULLEN); peduncle filiform, terete, glabrous
throughout or minutely papillose at the base;
flowers 1-15, distant; scales numerous, especially
below, similar to the bracts; bracts basifixed, ovate-
deltoid, 1-1.5 mm long, base shortly auriculate,
apex acute or acuminate, 1—3-nerved; bracteoles
linear-subulate, acute, 0.6-1 mm long; pedicels
erect, c. 1.5 mm long, terete. Calyx lobes unequal,
upper broadly ovate, cucullate, c. 2.5 mm long,
apex rounded, scarcely accrescent, nerves c. 9,
plicate, lower narrowly ovate, about as long, apex
emarginate, 5-nerved. Corolla bright yellow with an
orange palate and externally tinged with reddish
brown, c. 10 mm long, upper lip + orbicular, apex
retuse, longer than upper calyx lobe, lower 4
much larger, + orbicular in outline, apex + dee
4-lobed, palate prominent, bigibbous, spur su
late, curved, acute, about as long as the lower lip
and diverging from it at an angle of c. 90°. Filaments
linear, c. 1.5mm long, anthers c. 1 mm po
c. 0.3 mm wide, thecae distinct. Ovary ovoid, style
short but distinct, stigma lower lip orbicular,
upper minute, deltoid. Capsule globose, c. 2mm
long, firm and relatively thick, dehiscing by a ven-
tral narrowly linear lanceolate pore. Seeds numer-

286

ous, obovoid, c. 0.25 mm long, hilum terminal,
distinct, testa thin, reticulate, reticulations slightly
elongate.

Distr. Australia (northern Ww. Australia,
Northern Territory and Queensland) and Malesia:
New Guinea (Papua).

Ecol. Sandy open heathland and in Melaleuca-
Acacia savanna, at low altitude. F/. June—Aug.

Note. This well known Australian species has
been discovered (as has U. muelleri) in a number of
localities in Papua relatively recently. It is possible
that other smaller and less conspicuous Australian
species may yet be found in the same region.

9. Utricularia minutissima VAHL, Enum. 1 (1804)
204; Oxiver, J. Linn. Soc. Lond. Bot. 3 (1859)
190; Ripv. Trans. Linn. Soc. Lond. II, Bot. 3
(1892) 327; J. Bot. 33 (1895) 11; PRAIN, J. As. Soc.
Beng. 74, ii (1905) 371; RIDL. Fl. Mal. Pen. 2 (1923)
492; HEND. J. Mal. Br. R. As. Soc. 17 (1939) 60;
SpaRE, Mal. Nat. 7. 2 (1940) 89; P. TAYLOR,
Dansk Bot. Ark. 23 (1968) 531. — U. pygmaea
R. Br. Prod. (1810) 432; non Bru. Fl. Austr. 4
(1869) 526, quae est U. exoleta. — U. capillacea
(non WILLD.) WALL. Cat. (1832) n. 6399; OLIVER,
J. Linn. Soc. Lond. Bot. 3 (1859) 184; THw. En.
Pl. Zeyl. (1860) 171; TRIMEN, Handb. FI. Ceyl. 3
(1895) 270. — U. siamensis OSTENF. in Fedde,
Rep. 2 (1906) 68; PELLEGR. Fl. Gén. I.-C. 4 (1930)
480, — U. nipponica MAKINO, Bot. Mag. Tokyo
20 (1906) 95; OHwI, Fl. Japan (1965) 814. —
U. nigricaulis Rwt. J. Linn. Soc. Lond. Bot. 38
(1908) 317; J. Fed. Mal. St. Mus. 6 (1915) 164;
Fl. Mal. Pen. 2 (1923) 493; SYMINGTON, J. Mal.
Br. R. As. Soc. 14 (1936) 357. — U. calliphysa
SraPF in Gibbs, J. Linn. Soc. Lond. Bot. 42 (1914)
115; Merr. En. Born. (1921) 537; SPECHT, Arnhem
Land Exped. 3 (1958) 300. — U. brevilabris LACE,
Kew Bull. (1915) 404; PELLEGR. Fl. Gén. I.-C. 4
(1930) 481, incl. var. parviflora. — U. lilliput
PELLEGR. Bull. Mus. Nat. Hist. Paris 26 (1920)
181; Fl. Gén: L.-C. 4 (1930) 482; SUBRAMANYAM &
BALAKRISHNAN, Bull. Bot. Surv. India 2 (1960) 347.
__ U. evrardii PELLEGR. Fl. Gén. I.-C. 4 (1930) 476.
— U. barnesii LLOYD, The Carnivorous Plants
(1942) 232, nomen. — Fig. 10.

Terrestrial. Rhizoids few, capillary, simple.
Stolons capillary, sparsely branched. Foliar organs
sparsely rosulate and on the stolons, narrowly
linear to narrowly obovate-spathulate, up to
2cm by 0.5-1 mm, apex obtuse, 1-nerved. Traps
numerous on the vegetative organs, broadly ovoid,
c. 0.2 mm long, stalked, mouth lateral, upper lip
with a solitary multicellular subulate appendage,
lower lip with radiating rows of basally connate
obliquely gland-tipped processes. Inflorescence
erect, 3-12 cm long, glabrous or with a few short
septate hairs on the peduncle; peduncle terete,
filiform; flowers 1-10, distant; scales numerous,
similar to the bracts; bracts basifixed, narrowly
ovate, acute, 0.8—1 mm long; bracteoles similar or
less acute; additional bracts subtending dormant
(inflorescence branch) buds usually present on
inflorescence axis a short distance above each or
most of those subtending flowers; pedicels erect,
capillary terete, c. 1 mm long. Calyx lobes sub-
equal, broadly ovate, 1-2 mm long, apex of upper
obtuse, of lower emarginate, nerves obscure, not
raised. Corolla mauve or white, 2.5-7 mm long,

FLORA MALESIANA

[ser. I, vol. 8?

upper lip narrowly oblong, much longer than upper
calyx lobe, apex emarginate or rounded, lower lip
larger, + orbicular, obscurely to distinctly 3-lobed,
palate raised, spur subulate, obtuse, straight,
usually much longer than and parallel with the
lower lip. Filaments linear curved, anther thecae +
confluent. Ovary ovoid, style short, stigma lower
lip orbicular, upper lip much smaller, deltoid.

Utricularia minutissima VAHL. a. Stolon

Fig. 10.
with traps and foliar organ, * 12, b. trap, x 75,
c. ditto, dorsal view, x 75, d. bract & bracteoles,
x 12, e. flower, lateral view, x 6, f. ditto, frontal
view, X 6, g. pistil, x 12, h. stamen, x 124.
capsule, asymmetric in lateral view, x 12, re
fruiting calyx, < 12, k. dehisced capsule, x 12,
1. seeds, <X 45, m. dehisced capsule, 12,
flowering (left) and fruiting calyces, < 6, O. i
x 45 (a-c MEUER 26395, d_lJ. RAYNAL 17295, m-o
MEER 26395).

1977]

LENTIBULARIACEAE (Taylor)

287

SS ee SSSSSSSSSSSSSSSSm—MFhFsFeseFFFeFeseses

Capsule obliquely oblong-ovoid, 1.5-2 mm long,
membranous in texture, translucent, dehiscing by a
longitudinal ventral slit which is marginally scarcely
thickened. Seeds few globose, 0.17-0.2 mm long,
testa smooth, reticulate, reticulations relatively
large, + isodiametric.

Distr. India to China and Japan, Indo-China,
Malesia, and Australia (Northern Territory and
Queensland); in Malesia: Sumatra, Malaya,
Borneo, Philippines, and New Guinea.

Ecol. Damp open sandy or rocky places and
peaty swamps, also found on wet limestone, mostly
at low altitude but ascending to 2100m on G.
Tahan (Malaya). F/. March—Dec.

Note. This species is very variable in overall and
in flower size, and the degree to which the lower
corolla lip is lobed. The peduncle varies from quite
glabrous to sparsely hairy but the hairs when
present do not extend to the calyx as in U. hirta.
The ‘rigid patent black bristly hairs’ mentioned by
PRAIN and often present are in fact not hairs but
fungal growths.

10. Utricularia hirta KLEIN ex Link, Jahrb. 1, 3
(1820) 55; Oxtver, J. Linn. Soc. Lond. Bot. 3
(1859) 183; CrLarke, FI. Br. Ind. 4 (1884) 332;

Fig. 11. Utricularia hirta KLein ex Link. a. Trap,
* 75, b. part of hairy peduncle, ~ 24, c. flowering
calyx (left) and fruiting calyx, d. flower, e. ditto,
lateral view, all « 6, f. bract & bracteole, g. stamen,
h. pistil, all x 12, i. calyx with fruit, » 6, j. capsule,
thickened area shaded, ~ 6, k. seed, » 45 (all after
SMITINAND & WARNCKE s.n., except e G. TAYLOR
sn).

PELLEGR. FI. Gén. I.-C. 4 (1930) 479, incl. var.
elongata; P. TAYLOR, Dansk Bot. Ark. 23 (1968)
531. — U. capillacea (non WiLLD.) VAHL, Enum. 1
(1804) 204. — U. setacea (non MICHX.) WALL. Cat.
(1832) n. 6398. — Fig. 11.

Terrestrial. Rhizoids few, capillary, simple.
Stolons capillary, sparsely branched. Foliar organs
sparsely rosulate and on the stolons, narrowly
obovate-spathulate, up to 2 cm by 0.5-1 mm, apex
obtuse, 1-nerved. Traps numerous on the vegetative
organs, broadly ovoid, c. 0.2 mm long, stalked,
mouth lateral, upper lip with a solitary multicel-
lular subulate appendage, lower lip with radiating
rows of basally connate obliquely gland-tipped
processes. Inflorescence erect, 8-30cm long, +
densely covered throughout (including the calyx
and at least the spur of the corolla) with short to
long septate hairs; peduncle filiform, terete;
flowers 1-5, distant,; scales numerous, similar to
the bracts; bracts basifixed, narrowly ovate, acute,
1-1.2 mm long; bracteoles slightly narrower and
longer; additional bracts subtending dormant
(inflorescence branch) buds usually present on
inflorescence axis a short or very short distance
above each or most of those subtending flowers;
pedicels erect, capillary, terete, 1-2 mm long.
Calyx lobes subequal, broadly ovate, 2—3.5 mm
long, upper slightly smaller, apex obtuse, nerves
raised, lower relatively broader, apex emarginate
or obscurely tridentate. Corolla mauve or white,
8-10 mm long, upper lip narrowly oblong, much
longer than upper calyx lobe, apex + emarginate,
lower lip larger, + orbicular, distinctly 3-lobed,
palate raised, spur subulate, acute apically curved,
usually longer than and + parallel with the lower
lip. Filaments linear, curved, anther thecae dis-
tinct. Ovary ovoid, style short but distinct, stigma
lower lip orbicular, upper lip much smaller, del-
toid. Capsule oblong-ovoid to globose, 1.5—2.5 mm
long, firm in texture, opaque, dehiscing by a longi-
tudinal ventral marginally thickened slit. Seeds
few, + globose, c. 0.25 mm long, testa smooth,
reticulate, reticulations relatively large, slightly
elongate.

Distr. India, Indo-China, Thailand and
Malesia: Borneo (Bako National Park).

Ecol. Damp open sandy or muddy places at low
altitude. F/. Febr., July.

Note. Very similar to U. minutissima (with
which it sometimes occurs) but distinguished by its
larger size, larger flowers and hairy calyx.

11. Utricularia caerulea Linné, Sp. Pl. (1753) 18;
Burm. f. Fl. Ind. (1768) 11; Wiaur, Ic. (1850)
t. 1583; GamBLe, FI. Madras 2 (1924) 983;
SANTAPAU, J. Bomb. Nat. Hist. Soc. 49 (1950) 217;
P. Taytor, Dansk Bot. Ark. 23 (1968) 530. —
U. nivea VAHL, Enum. 1 (1804) 203; Wiont, Ic.
(1850) t. 1582; Oxtver, J. Linn. Soc. Lond. Bot. 3
(1859) 186; Merr. Fl. Manila (1912) 433; Philip.
J. Sc. 7 (1912) Bot. 247; En. Philip. 3 (1923) 466;
Back. Onkr. Suiker. (1931) 636; BAKH. /. in Back.
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 194, p. 3;
Back. & BAKH. f. Fl. Java 2 (1965) 518. — U.
baueri R. Br. Prod. Nov. Holl. (1810) 431; Bru.
Fl. Austr. 4 (1869) 531; Specur, Arnhem Land
Exped. 3 (1958) 300. — U. complanata WALL. Cat.
(1829) n. 1497, nomen, — U. racemosa WALL, ex
Wa cp. in Meyen, Observ. Bot. Nov. Acta 19

288

FLORA MALESIANA

[ser. I, vol. 8?

(1843) 401; A. DC. in DC. Prod. 8 (1844) 21;
WicGut, Ic. (1850) t. 1584(1); OLiver, J. Linn. Soc.
Lond. Bot. 3 (1859) 186; CLARKE, FI. Br. Ind. 4
(1884) 333; Forses & Hems-. J. Linn. Soc. Lond.
Bot. 26 (1890) 224; Rip. Trans. Linn. Soc. Lond.
II, Bot. 3 (1893) 327; J. Str. Br. R. As. Soc. n. 33
(1900) 119; ibid. n. 59 (1911) 144; STAPF in Gibbs,
Arfak (1917) 180; Merr. Lingn. Sc. J. 5 (1927) 167;
MeEtcuior, Bot. Jahrb. 62 (1929) 383; PELLEGR.
Fl. Gén. I.-C. 4 (1930) 485; HAND.-MAzz. Symb.
Sin. 7 (1936) 873; Merr. J. Arn. Arb. 18 (1937) 73;
SPARE, Mal. Nat. J. 1 (1940) 90. — U. filicaulis
WALL. (Cat. 1829, n. 1501) ex A. DC. in DC.
Prod. 8 (1844) 21; OLiver, J. Linn. Soc. Lond. Bot.
3 (1859) 186; Mia. Fl. Ind. Bat. 2 (1859) 999;
PELLEGR. Fl. Gén. I.-C. 4 (1930) 486, incl. var.
papillosa. — U. bifida (non L.) MACRAE ex A. DC.
in DC. Prod. 8 (1844) 21, pro syn. — U. squamosa
BENJ. Bot. Zeit. 3 (1845) 212. — U. rosea EDGEw.
Proc. Linn. Soc. Lond. 1 (1847) 352. — U. purpurea
(non WALT.) WILLD. ex BENJ. Linnaea 20 (1847)
309, pro syn. — U. paucifolia BENS. l.c. 309. —
U. obtusiloba BeENs. I.c. 312. — U. albiflora (non
R. Br.) GrirF. Notul. 4 (1854) 168. — U. racemosa
var. filicaulis CLARKE, Fl. Br. Ind. 4 (1884) 333;
Boerv. Hand. 2, 2 (1899) genus 559; PRAIN, J. As.
Soc. Beng. 74, ii (1905) 373; Merr. En. Born.
(1921) 538. — U. campestris MiQ. ex CLARKE,
Fl. Br. Ind. 4 (1884) 333, pro syn. — U. warburgii
GoeBEL, Ann. Jard. Bot. Btzg 9 (1891) 22. —
U. ophirensis Rv. J. Bot. 33 (1895) 10; PRAIN, J.
As. Soc. Beng. 74, ii (1905) 373; Ript. Fl. Mal.
Pen. 2 (1923) 495, f. 122; PELLEGR. FI. Gén. I.-C. 4
(1930) 487. — U. cavalerii STAPF, Kew Bull. (1910)
195. — U. sootepensis CratB, Kew Bull. (1911)
430; PeLttecr. Fl. Gén. L-C. 4 (1930) 487. —
U. kerrii Crates, Kew Bull. (1911) 429; PELLEGR.
Fl. Gén. I.-C. 4 (1930) 485. — U. charnleyensis
FITZGERALD, J. R. Soc. W. Austr. 3 (1918) 207. —
U. albina Rw. Fl. Mal. Pen. 2 (1923) 493; HEND.
Gard. Bull. S. S. 4 (1928) 295. — U. roseo-purpurea
STAPF ex GAMBLE, Fl. Madras 2 (1924) 983. —
Fig. 12.

Terrestrial. Rhizoids few to very numerous,
capillary, simple. Stolons capillary, sparsely
branched. Foliar organs not always conspicuous at
anthesis, rosulate and on the stolons, narrowly
obovate-spathulate, up to 7 mm long and 1.5 mm
wide, apex rounded, 1I-nerved. Traps rather few on
the vegetative organs, + dimorphic, ovoid, shortly
stalked, mouth terminal, oblique, the larger traps
up to 1.5 mm long including a long carinate beak
on the upper lip, the smaller traps about half as
long with a relatively shorter beak, mouth and
beak of both types densely stipitate-glandular.
Inflorescence erect, 5-30 cm long; peduncle filiform
to relatively stout, 0.3-1.5mm _ thick, terete,
glabrous; flowers 1-20 or more very variably dis-
posed, distant to quite densely congested and sub-
capitate; scales numerous, similar to the bracts;
bracts medifixed, variably attached above or below
the middle, narrowly rhombic, acuminate at both
extremities, 2-2.5 mm long; bracteoles similar but
often shorter and narrower; pedicels erect at
anthesis, spreading or reflexed in fruit, filiform,
terete, 0.5-1mm long. Calyx lobes unequal,
usually minutely papillose, upper ovate-oblong,
cucullate, 2-3 mm long, apex rounded, lower shor-
ter, transversely elliptic with conspicuously

inrolled margins. Corolla pink, mauve, purple or
blue, often white and sometimes ? yellow, 4-10 mm
long, externally minutely papillose, upper lip
narrowly ovate-oblong, longer than upper calyx
lobe, apex rounded or truncate, lower lip larger, +
orbicular, apex retuse, palate raised, + conspi-
cuously transversely crested, spur narrowly conical
or cylindrical from a broader conical base, straight
or curved, usually longer than and + parallel
with the lower lip. Filaments filiform, straight,
c. 1mm long, anther thecae distinct, minutely
papillose. Ovary ovoid, style variably in length,
usually distinct, stigma lower lip orbicular, upper

Fig. 12. Utricularia caerulea L. a. Foliar organ,
x 6, b. large and small trap, x 24, c. bract &
bracteoles, all peltately attached, x 6, detail of
surface papillae, x 24, d. calyx in flower, x 6,
detail of surface papillae, x 24, e. flower, lateral
view, and upper lip from front, f. flower, front view,
g. small flower, all x 6, h. stamen, x 12, i. pistil,
x 12, j. calyx with fruit, x 6, k. fruit exposed,
x 6, 1. seed, x 45, with detail of testa enlarged,
75 (all after J. RAYNAL 17069, large form, except
a-b, d, g LARSEN 5160, small form).

1977]

lip minute, deltoid. Capsule globose, c. 2 mm long,
firm, opaque, dehiscing by a ventral longitudinal
slit. Seeds numerous, obliquely oblong-ellipsoid,
c. 0.3mm long, testa thin, smooth to distinctly
papillose, obscurely reticulate, reticulations elon-

gate.

Distr. India to China and Japan and Australia;
in Malesia: not recorded from the Lesser Sunda Is.
(but present in Kangean Is.) or Celebes, but other-
wise widespread and common.

Ecol. Damp or wet open situations on sand
(also on kerangas and in heath-forest) or mud both
under everwet and seasonal climatic conditions,
from sea-level to 2700 m. Fi. Jan.—Dec.

Note. This is an excessively variable plant in
which the extreme forms, often of quite distinct
appearance, are linked by intermediates.

12. Utricularia striatula J.SmM. in Rees, Cyclop. 37
(1819) m. 17; Forses & HeEms-. J. Linn. Soc. Lond.
Bot. 26 (1890) 224; GoeseL, Ann. Jard. Bot. Btzg 9
(1890) 68; Rpt. Fl. Mal. Pen. 2 (1923) 495;
GAMBLE, FI. Madras 2 (1924) 983; PELLEGR. FI.
Gén. I.-C. 4 (1930) 474; HAND.-MAzz. Symb. Sin.
7 (1936) 872; Spare, Mal. Nat. J. 1 (1940) 88;
BaKuH. f. in Back. Bekn. Fl. Java (em. ed.) 8 (1949)
fam. 194, p. 2; SANTAPAU, J. Bomb. Nat. Hist. Soc.
49 (1950) 220; P. TaAyLor in Hutch. & Dalz. Fl. W.
Trop. Afr. ed. 2, 2 (1963) 378; Kew Bull. 18 (1964)
91; Back. & Baku. f. Fl. Java 2 (1965) 517; P.
TayLor, Dansk Bot. Ark. 23 (1968) 532. — U.
pusilla (non VAHL) Grau. Cat. Pl. Bombay (1839)
165. — U. orbiculata WALL. (Cat. 1829, n. 1500)
ex A. DC. in DC. Prod. 8 (1844) 18; Oxiver, J.
Linn. Soc. Lond. Bot. 3 (1859) 187; CLARKE, FI.
Br. Ind. 4 (1884) 334; Strapr, Trans. Linn. Soc.
Lond. II, Bot. 4 (1894) 211; Trimen, Handb. FI.
Ceyl. 3 (1895) 271; Ript. J. Str. Br. R. As. Soc.
n. 59 (1911) 144; Merr. En. Born. (1921) 538; En.
Philip. 3 (1923) 467; Steen. Arch. Hydrobiol.
Suppl. 11 (1932) 333, f. 8 IV; H. J. LAM, Blumea 5
(1945) 582. — U. rosulata Bens. Linnaea 20 (1847)
310; Mia. Fl. Ind. Bat. 2 (1859) 1000; Merr. En.
Philip. 3 (1923) 189. — U. glochidiata Wicut, Ic.
(1850) t. 1581. — U. harlandii OLIver ex Btu. FI.
Hongk. (1861) 257; Forpes & Hems-. J. Linn. Soc.
Lond. Bot. 26 (1890) 223. — U. anthropophora
Riwi. J. Fed. Mal. St. Mus. 6 (1915) 165. —
U. striatula var. minor Rip. Trans. Linn. Soc.
Lond. II, Bot. 9 (1916) 122; Metcuior, Bot. Jahrb.
62 (1929) 384. — Fig. 2e, 13-14.

Epiphytic or terrestrial. Rhizoids few, capillary,
simple. Stolons capillary, branched. Foliar organs
numerous and conspicuous at anthesis, rosulate
and on the stolons, obovate, orbicular or reniform
with a + distinct pseudopetiole, membranous,
3-10 mm long, up to 6 mm wide, with numerous
dichotomously branched nerves. Traps numerous
on the stolons, obliquely globose or ovoid,
0.6-0.8 mm long, long stalked, mouth lateral,

lip projecting with 2 divergent stipitate-
glandular appendages. Inflorescence erect, 1-15 cm
g; peduncle filiform, glabrous, usually less
than 0.3 mm thick; flowers 1-10, distant; scales
few, similar to the bracts; bracts medifixed, lanceo-
late, apex obtuse or acute, base truncate to acute;
bracteoles similar; pedicels spreading at anthesis,
+ reflexed in fruit, capillary, usually less than
0.2 mm thick. Calyx lobes very unequal, papillose,

LENTIBULARIACEAE (Taylor)

289

upper orbicular-obcordate, 1.5-2.5 mm long at
anthesis, apex emarginate, lower very much smaller,
ovate-oblong, apex truncate or retuse. Corolla
white, pink or mauve with a yellow spot on the
palate, 3-10 mm long, upper lip minute, semi-
orbicular, apex bidentate, very much shorter than
the upper calyx lobe, lower lip orbicular or trans-
versely elliptic, 3-10 mm wide, apex + distinctly
3-S-lobed, palate slightly raised, spur subulate,

” a
G00 KO

Fig. 13. Utricularia striatula J.SM. a. Foliar organ,
x 6, b-c. trap, lateral & ventral views, x 24, d.
insertion of pedicel, with bract & bracteoles in situ,
= 6,e. the latter enlarged, = 12, f. calyx in flower,
x 6, g. flower calyx lobe bent back to show upper
corolla lip, x 5, h. ditto, in lateral view, * 5, i.
upper & lower lip flattened, « 6, j-k. two flowers,
x 5, l-o. four ditto, x 5, p. stamens, x 12, q.
pistil, x 12, r. dehisced fruit, < 6, s. fruit in lateral
view, x 6, t. seed, * 45 (a~c, g-h, r-t ALLEN 5.7.,
Borneo, f, /-q J. wer ie , i-k CRUTTWELL

290

FLORA MALESIANA

[ser. I, vol. 8?

Fig. 14. Utricularia striatula J.SM. Epiphytic on a
tree trunk in primary forest on sandstone at Lubuk
Bangko, 20 km E of Pajakumbuh, Central West
Sumatra, Febr. 2, 1957. Photogr. W. MEIER. Note
orbicular foliar organs at right margin.

usually curved, 1-4 mm long, shorter than and +
parallel with or somewhat divergent from the lower
lip. Filaments filiform, c. 0.6mm long, anther
thecae subdistinct. Ovary globose, style short but
distinct, stigma lower lip semi-orbicular, upper
obsolete. Capsule + globose, obliquely dorsiven-
trally compressed, membranous, obscurely ven-
trally keeled, dehiscing by a ventral longitudinal
slit. Seeds few, pyriform or obovoid, c. 0.25 mm
long, hilum terminal, prominent, testa smooth,
bearing, especially distally, numerous unicellular
apically glochidiate processes.

Distr. Tropical Africa (but apparently absent
from Madagascar), widespread from India to
China, Indo-China and throughout Malesia.

Ecol. Rocks or trees or less commonly damp
soil among moss in somewhat shady permanently

moist conditions, on mossy tree trunks, on stones
in rivers, near waterfalls, on wet talus, from sea-
level up to 3300 m. Fi. Jan.—Dec.

Vern. Banka: kakrak, rumput kitjekar, M; New
Guinea: romaripi, Orne lang., Wantipi.

13. Utricularia pulchra P. TAYLOR, sp. nov. —
Fig. 15.

Affinis U. striatulae J.Sm. sed floribus duplo
majoribus, paucioribus in inflorescentia crassiore
congestis differt. — Type: New Guinea, Irian Jaya,
Tembagapura (southern slopes of Mt Carstensz),
alt. 2400 m, 30 April 1973, RAYNAL 17440 (K, P).
— U. striatula (non J.SM.) RIDL. Trans. Linn. Soc.

Fig. 15. Utricularia pulchra P. TAyvor. a. Habit of
vegetative parts, < 4, b-c. trap, lateral & dorsal
views, < 24, d. bract & bracteoles, x 8, e. their
insertion, x 6, f. calyx, x 4, g. (large) flower, x 3,
h. upper lip, ventral & lateral views, = 6, detail of
hairs, x 24,i. stamens, x 12, j. pistil with circular
adnation to calyx at base, « 12 (all after J. RAYNAL
17440, type).

1977]

LENTIBULARIACEAE (Taylor)

291

Lond. II, Bot. 9 (1916) 122; MELCHIOR, Bot. Jahrb.
62 (1929) 384.

Terrestrial. Rhizoids few, capillary, simple.
Stolons few, capillary, up to 5cm long. Foliar organs
present at anthesis, rosulate and on the stolons,
reniform, fleshy, + pseudopetiolate, 2-4 mm long.
Traps numerous on the stolons, globose, 0.6—-1 mm
long, stalked, mouth lateral, upper lip projecting,
with 2 short divergent stipitate-glandular appen-
dages. Inflorescence erect, 4-6 cm long; peduncle
filiform, glabrous, 0.5—0.7 mm thick; flowers 1-3,
congested; scales 1 or 0, similar to the bracts;
bracts medifixed, ovate, 1.5-2mm long, apex
obtuse, base truncate or bidentate; bracteoles

imilar; pedicels erect, filiform, dorsiventrally
flattened, 4-5 mm long, c. 0.5mm wide. Calyx
lobes very unequal, papillose, upper orbicular or
broadly reniform, apex rounded or retuse, 3-4 mm
long, 3.5—-5 mm wide, lower much smaller, orbicu-
lar or broadly ovate, 2-2.5mm long. Corolla
mauve or violet with a yellow spot on the palate,
17-20 mm long, upper lip about as long as upper
calyx lobe, oblong, cucullate, apex emarginate,
lower lip flabellate, 5-lobed, up to 20 mm wide,
palate slightly raised, spur subulate, + straight,
10-12 mm long, as long as or longer than the lower
lip and diverging from it at a very obtuse angle.
Filaments linear, c. 1.2mm long, anther thecae
distinct. Ovary obliquely ovoid, c. 1.5 mm long,
style very short, stigma lower lip semi-oribicular,
upper lip obsolete. Capsule and seeds not known.

Distr. Malesia: New Guinea (West New
Guinea: Mt Carstensz; T.N.G., Sepik Distr.:
Sirius Plateau).

Ecol. Wet cliffs and damp leached sand or
among moss, 2400-3000 m. Fi. March-April, Oct.

Note. U. pulchra is very close to U. striatula but
differs constantly in its fewer, much larger flowers
and its shorter stouter habit.

14. Utricularia salwinensis HAND.-MaAzz. Symb.
Sin. 7 (1936) 873. — Fig. 16.

Terrestrial. Rhizoids few, capillary, simple.
Stolons capillary, sparsely branched, up to 5cm
long. Foliar organs present at anthesis, sparsely
rosulate and on the stolons, flabellate, cuneate or
obovate-spathulate to orbicular, with a distinct
sey esi 1.5-3.5mm wide, up to 2.5cm
ong. Traps rather few on the stolons, ovoid, long
stalked, c. 0.8 mm long, mouth lateral, upper lip
with a relatively large flabellate appendage c. 1 mm
long and 2 mm wide including c. 8 marginal subu-
late multicellular gland-tipped processes. /nflores-
cence erect, up to 8cm long; peduncle filiform,
terete, glabrous; flowers 1-3, + distant; scales
0 or 1 in the upper part of the peduncle, similar to
the bracts; bracts medifixed, ovate, membranous,
sparsely glandular, 1.2-1.5 mm long, apex acute,
base truncate, apiculate; bracteoles similar but
base obliquely truncate; pedicels erect at anthesis,
strongly recurved in fruit, 24mm long. Calyx
lobes very unequal, papillose, upper + orbicular,
c. 1.5mm long at anthesis, up to 3 mm long in
fruit, apex emarginate, lower much smaller, ovate-
a c. 1mm long, apex truncate or retuse.
Corolla white (in Mal.) with a yellow (fide De
WiLpe) or brown (fide VAN STEENIS) spot on the

te or pink (in Yunnan, fide HANDEL-
), 46mm long, upper lip transversely

Fig. 16. Utricularia salwinensis HAND.-MAZZ. a.
Rosette of foliar organs, b. stolon with foliar
organ, branch and trap, b’. lamina of two foliar
organs, all x 3,c. trap, x 12, d. bract (right) &
bracteole, x 12, e. flower, x 6, f. corolla, upper &
lower lip laid out, * 6, g. stamen, x 24, h. pistil,
= 24, i. dehisced fruit in calyx, j. ditto, lower calyx

lip removed, k. capsule in LS, seeds removed, all
x 6,1. seed, x 24 (a VAN STEENIS 8598, the others
after pe WiLpe 15258).

oblong, cucullate, apex emarginate, much shorter
than upper calyx lobe, lower lip flabellate, 4-5 mm
wide, 3-lobed, midlobe quadrate or semi-orbicular,
side lobes smaller, obliquely and rte pie un-
equally 2-lobed, palate scarcely raised, shortly
fimbriate, spur narrowly cylindrical, apex obtuse,
much shorter than the lower lip. Filaments filiform,
c. 1 mm long, anther thecae distinct. Ovary glo-
bose, c. 0.6 mm long, style very short, sti lower
lip orbicular, upper -* minute, deltoid. Capsule

obose, obliquely rsiventrally compressed
c. 2mm long, dehiscing by a longitudinal ventral
slit. Seeds few, ovoid,.c. 0.5mm long, shortly
densely echinate.

292

FLORA MALESIANA

[ser. I, vol. 8?

Distr. China (Yunnan) and Malesia: N, Suma-
tra (Gajo Lands: Mts Goh Lembuh, Kemiri,
Bandahara, Mamas, and Losir).

Ecol. Moist sandy rocky or mossy places in
open or low scrub, 2500-3300 m. Fl. Febr.—May.

Note. The only difference between the Yunnan
and Sumatra plants is the recorded corolla colour
and this is not considered significant. Similarly dis-
junct distribution of mountain plants are not un-
known.

15. Utricularia limosa R. Br. Prod. Nov. Holl.
(1810) 432; Bru. Fl. Austr. 4 (1869) 531; SPECHT,
Arnhem Land Exped. 3 (1958) 302. — U. verticil-
lata BENJ. Linnaea 20 (1847) 312; OLiver, J.
Linn. Soc. Lond. Bot. 3 (1859) 184; Mia. FI. Ind.
Bat. 2 (1859) 1000; CLARKE, Fl. Br. Ind. 4 (1884)
333; RipL. Trans. Linn. Soc. Lond. II, Bot. 3
(1893) 327; Boerv. Handl. 2, 2 (1899) genus 559;
PRAIN, J. As. Soc. Beng. 74, ii (1905) 372; Rmw-. J.
Str. Br. R. As. Soc. n. 59 (1911) 144; Fl. Mal. Pen.
2 (1923) 493; PELLEGR. FI. Gén. I.-C. 4 (1930) 484;
Spare, Mal. Nat. J. 1 (1940) 90. — U. bifida (non
L.) Wicut, Ic. (1850) t. 1584(2). — U. biloba (non
R. Br.) P. TAyLor, Dansk Bot. Ark. 23 (1968)
529. — Fig. 17.

Terrestrial. Rhizoids few, filiform, basally
thickened, with numerous, often geminate, papil-
lose branches c. 0.5mm long. Stolons capillary,
branched. Foliar organs often not present or
conspicuous at anthesis, on the stolons, narrowly

N
N
N
\

SS

Fig. 17. Utricularia limosa R.Br. a—b. Traps, large
& small, x 12, c. bract, d. scales, mid and lower
(small one), e. fruiting calyx, all « 6, f-g. flower, in
two views, < 6, h. stamen, x 12, i. pistil, x 12, j.
capsule, x 6, and enlarged surface of pedicel,
x 24, k. capsule, ventral view, thickened area
shaded, x 6,1. seed, x 45 (a-e, k Must 1004, the
others after J. RAYNAL 17296).

linear, 1.5—2.5 cm long, c. 0.3 mm wide, 1-nerved,
apex acute. Traps few on the stolons and foliar
organs, obliquely ovoid, 0.5-1 mm long, stalked,
stalk 2-3 times as long as trap, mouth lateral,
upper lip with 2 long densely hairy setiform appen-
dages. Inflorescence erect, up to 25cm long;
peduncle filiform, glabrous or basally glandular,
0.5-0.8 mm thick; flowers 2-10, rather distant;
scales few, similar to the bracts; bracts medifixed,
narrowly elliptic, 1.5-2 mm long, apex acute, base
acute or obliquely truncate; bracteoles absent;
pedicels filiform, slightly dorsiventrally flattened,
minutely papillose, distinctively curved, erect at
base and apex, spreading in the middle part, up to
6mm long in fruit. Calyx lobes unequal, upper
transversely elliptic, apex rounded, up to 1.5mm
long in fruit, lower obovate-oblong, apex rounded
or obscurely crenulate, up to 2 mm long in fruit.
Corolla violet, pink or white, 4.5-6 mm long, upper
lip broadly ovate, about twice as long as upper
calyx lobe, apex rounded ; lower lip larger, quadrate,
bilobed to half its length, lobes ovate-oblong,
curved upwards, palate raised, spur narrowly cylin-
drical from a conical base, apex obtuse, somewhat
longer than lower lip. Filaments linear, strongly
curved, flattened and apically dilated, c. 0.6mm
long, anthers c. 0.3mm long, thecae confluent.
Ovary globose, style short, stigma lower lip
orbicular, upper lip obsolete. Capsule globose,
c. 2mm _ long, membranous, dehiscing by a
longitudinal ventral marginally thickened slit,
the thickened area before dehiscence apically
acute, broader and emarginate at the base. Seeds
numerous, globose or slightly angular, c. 0.25 mm
long, testa thin, coarsely reticulate, reticulation +
isodiametric.

Distr. Indo-China, Thailand, and Australia; in
Malesia: Malay Peninsula and New Guinea.

Ecol. Seasonally flooded grassland at low alti-
tude. F/. Febr._May, Sept.

Norte. U. limosa belongs to a small, very dis-
tinct group of species, the others in the group
being apparently confined to Australia.

16. Utricularia subulata LINNE, Sp. Pl. (1753) 18; P.
TAYLOR, Dansk Bot. Ark. 23 (1968) 532. — For
full synonymy, which is large and exclusively
American and African, see P. TAYLOR, Kew Bull.
18 (1964) 81-83 and Mem. N.Y. Bot. Gard. 17
(1967) 223. — Fig. 18.

Terrestrial. Rhizoids few, capillary, basally
thickened and rigid, with numerous papillose
branches c. 0.5 mm long. Stolons capillary, much
branched. Foliar organs often not conspicuous at
anthesis, narrowly linear, 1-2 cm long, c. 0.5 mm
wide, apex acute, l-nerved. Traps very numerous
on the vegetative organs, ovoid, stalked, 0.2-0.5
mm long, mouth lateral, upper lip projecting with
2 spreading curved subulate sparsely branched
appendages. Inflorescence erect, up to 25 cm long;
flowers 1-25, moderately distant; peduncle
capillary, glabrous above, usually minutely papil-
lose below; scales few, similar to the bracts but
narrower and acuminate at base and apex, often
papillose; bracts medifixed, peltate, broadly elliptic
to orbicular, membranous, 0.75—1 mm long, obtuse
or subacute at base and apex; bracteoles absent;
pedicels ascending, capillary, terete, 2-10 mm long.
Calyx lobes subequal, broadly ovate to orbicular,

,
J

1977]

Fig. 18. Utricularia subulata L. a. Base of peduncle
and vegetative parts, x 6, b. trap, « 45, c. smooth
peduncle with lower scale, d. ditto, with upper
scale, both x 12, e. peduncle near base with
papillae and teeth on margin of scale, x 15, with
details, x 75, f. bract, flattened, x 12, g. calyx in
flower, h-i. corolla, j. ditto, small size, k. ditto,
adaxial aspect, |. fruiting pedicel, m. capsule, ven-
tral view, all x 6, n. two seeds, x 45 (a, j, k
LARSEN & WARNCKE 13, b-d, f Jacoss 5494, e, i
Kerr 14338, g, h FRANCK s.n., 1, m MILNE-
REDHEAD & TAYLOR ae n MELVILLE & HOOKER
3).

c. 1 mm long at anthesis, slightly accrescent, apex
rounded or truncate. Corolla yellow, 6-10 mm long,
upper lip broadly ovate to orbicular, 2—3 times as
long as upper calyx lobe, apex rounded, lower lip
larger, + flabellate, deeply 3-lobed, midlobe
larger than side lobes, apex of lobes rounded to
truncate, palate much raised, bigibbous, spur
subulate from a short conical base, parallel with
and about as long as lower lip, apex usually ob-
tuse, sometimes acute or 2—4-denticulate. Filaments
filiform, curved, anther thecae confluent. Ovary
globose, style very short, stigma lower lip orbicular,
upper lip obsolete. Capsule globose, 1—1.5 mm
long, membranous, dehiscing by a small ventral
ovate pore. Seeds numerous, ovate, 0.2-0.25 mm
testa thin, obscurel prycoerreggar striate.

istr. Eastern North America, throughout

Central and South America to Argentina, through-

LENTIBULARIACEAE (Taylor)

293

out tropical Africa and in S. Africa and Madagas-
car, also in Portugal (possibly introduced); in
Thailand and in Malesia: Malaya (Selangor) and
Borneo (Bako National Park, Kalabit Highlands,
and once in Central Borneo).

Ecol. Damp open sandy or boggy ground at low
altitude. F/. Sept.

Note. It is curious that this species, which is
widespread and abundant in the New World,
Africa and Madagascar, should never have been
found in India. There can be no doubt that the
Malesian plant is the same as the African and
American ones.

17. Utricularia punctata WALL. (Cat. 1829, n. 2121)
ex A. DC. in DC. Prod. 8 (1844) 5; WiaGur, Ic.
(1850) t. 1570; OLtver, J. Linn. Soc. Lond. Bot. 3
(1859) 175; Mia. Suppl. 1 (1860) 246; Kurz, Nat.
Tijd. N. I. 27 (1864) 213; Rip. Trans. Linn. Soc.
Lond. II, Bot. 3 (1893) 327; Boeri. Handl. 2, 2
(1899) genus 559; PRAIN, J. As. Soc. Beng. 74, ii
(1905) 369; Merr. En. Born. (1921) 538; Ript. FI.
Mal. Pen. 2 (1923) 491; SPARE, Mal. Nat. J. 1
(1940) 88; P. TayLor, Dansk Bot. Ark. 23 (1968)
531, excl. syn. U. rogersiana. — U. fluitans RID . J.
Str. Br. R. As. Soc. 1. 61 (1912) 32; Fl. Mal. Pen. 2
(1923) 491 ; Disp. (1930) 180, sphalm. ‘fluviatilis’. —
U. aurea (non Lour.) P. TAYLor, Dansk Bot. Ark.
23 (1968) 529, quoad syn. U. fluitans. — Fig. 19.
Aquatic. Rhizoids apparently absent. Stolons
filiform, terete, sparsely branched, up to 20cm
long, 0.5-1_mm thick, glabrous. Foliar organs
numerous on the stolons, 2-6 cm long, the primary
segment divided in 2 or 3 just above the base, then
repeatedly divided into very numerous segments,
the secondary pinnae + reflexed, the lowermost
somewhat remote from the base which may or may
not be provided with sparse stipule-like segments
resembling the ultimate segments, the latter capil-
lary, terete, sparsely minutely setulose; the primary
segments of the foliar organ basal to the inflores-
cence often longer, inflated, + fusiform and bear-
ing fewer shorter ultimate segments. Traps not
numerous, lateral on the penultimate and ultimate
segments, broadly obliquely ovoid, 1-2 mm long,
shortly stalked, mouth lateral with short marginal
setae, upper lip with 2 long, branched setiform
appendages. Inflorescence erect, 8-30 cm long;
peduncle filiform, terete; flowers 6-12, + distant;
scales few, similar to the bracts; bracts medifixed,
peltate, inserted shortly above the base of the
pedicel, ovate, apex acute, base rounded, mem-
branous, c. 2 mm long, attached below the middle;
bracteoles absent; pedicels capillary, terete, erect
or ascending, c. 6mm long at anthesis, up to
15mm long in fruit. Calyx lobes subequal, the
lower slightly smaller, orbicular, membranous,
c. 1.5mm long, scarcely accrescent. Corolla lilac,
violet, pink or rarely white, with a yellow spot on
the palate, 6-10(—15) mm long, externally glabrous,
upper lip transversely elliptic or orbicular, cucul-
late, apex rounded, lower lip larger, transversely
oblong-elliptic, up to 12 mm wide, base auriculate,
apex rounded, lateral margins incurved, palate
raised, gibbous, spur conical, slightly curved, apex
obtuse, about as long as and + parallel with the
lower lip. Filaments falcate, considerably ex-
panded above, c. 1.5 mm long, anther thecae con-
fluent. Ovary ovoid, style about as long as ovary,

294

FLORA MALESIANA

[ser. I, vol. 8?

stigma lower lip orbicular, upper lip much smaller,
deltoid. Capsule ellipsoid, membranous, c. 3 mm
long, laterally bivalvate. Seeds few, lenticular,
1.5-2 mm wide, margin winged, deeply irregularly
dentate.

Fig. 19. Utricularia punctata WALL. ex A.DC. a.
Stolon with part of a branched foliar organ, at the
base with stipule-like basal segments, x 5, b. trap,
0245 ci) bract. in situ, <"4 do brace, 2 7/5, e:
flowering calyx, x 4, f. flower, lateral view, x 4,
detail of glands inside spur, x 75, g. corolla, front
view, < 4, h. stamen, x 15, i. pistil, x 15, detail
of glands on ovary x 75, j. fruiting calyx, x 6, k.
fruit, x 4, 1. placenta, x 4, m. dehisced fruit,
abaxial ventral view, x 6,n. seed, x 7?/, (all after
LARSEN 82, except b CASTLE 88, j, m FLENLEY 134).

Distr. Burma, Indo-China, Thailand; in
Malesia: Sumatra, Malaya, and Borneo.

Ecol. Still or slow flowing water in lakes, rivers
and swamps at low altitude. F/. Aug.—May.

Vern. Banka: kantur, kembang ganga, kramor,
M, used as fodder for pigs.

Note. U. punctata is anomalous in the ‘aquatic’
group of the genus (i.e. those with dissected setu-
lose foliar organs) in both the mode of dehiscence
of the fruit and the medifixed bracts. RIDLEY stated
that the corolla of his U. fluitans was 25 mm wide
but that preserved on the type specimen is less than
half this dimension.

18. Utricularia exoleta R. Br. Prod. Nov. Holl.
(1810) 430; Bru. FI. Austr. 4 (1869) 526; CLARKE,
Fl. Br. Ind. 4 (1884) 329; Forpes & HEmsL. J.
Linn. Soc. Lond. Bot. 26 (1890) 223; GOEBEL,
Ann. Jard. Bot. Btzg 9 (1890) 91; TRIMEN, Handb.
Fl. Ceyl. 3 (1895) 268; Riv. J. Str. Br. R. As. Soc.
n. 33 (1900) 119; Pratn, J. As. Soc. Beng. 74, ii
(1905) 368; Koorp. Exk. Fl. Java 3 (1912) 204;
Merk. En. Born. (1921) 537; En. Philip. 3 (1923)
466; Rip. Fl. Mal. Pen. 2 (1923) 491; PELLEGR.
Fl. Gén. I.-C. 4 (1930) 473; STEEN. Arch. Hydro-
biol. Suppl. 11 (1932) 331, f. 8 V; Miki, Bot. Mag.
Tokyo 49 (1935) 847; SPARE, Mal. Nat. J. 1 (1940)
88; BAKH. f. in Back. Bekn. Fl. Java (em. ed.) 8
(1949) fam. 194, p. 2; SANTAPAU, J. Bomb. Nat.
Hist. Soc. 49 (1950) 218. — U. diflora Roxs. Hort.
Beng. (1814) 4, nomen nudum. — U. biflora (non
LAMK) Roxs. FI. Ind. 1 (1820) 144. — U. diantha
Roxs. ex Roem. & SCHULTES, Mant. 1 (1822) 169;
Wicut, Hook. J. Bot. Kew Misc. 1 (1849) 372; Ic.
(1850) t. 1569; Mig. Fl. Ind. Bat. 2 (1859) 999;
Oxtver, J. Linn. Soc. Lond. Bot. 3 (1859) 176. —
U. roxburghii SPRENG. Syst. 1 (1825) 52. — U.
pauciflora BL. Bijdr. (1826) 739; Miq. Fl. Ind. Bat.
2 (1859) 999. — U. ambigua A. DC. in DC. Prod.
8 (1844) 9. — U. pterosperma EDGEw. Proc. Linn.
Soc. Lond. 1 (1847) 352. — U. conferta Hassk. Nat.
Tijd. N. I. 10 (1855) 90 (Retzia 1 (1855) 90); Mia.
Fl. Ind. Bat. 2 (1859) 998. — U. sumatrana Mia.
FI. Ind. Bat. 2 (1859) 998, p.p.; Suppl. 1 (1860) 246.
— U. saharunporensis RoyLe ex OLtver, J. Linn.
Soc. Lond. Bot. 3 (1859) 176, pro syn. — U.
elegans WALL. (Cat. 1829, n. 1502) ex OLIver, J.
Linn. Soc. Lond. Bot. 3 (1859) 176, pro syn. —
U. gracilis Lepr. ex OLIveR, J. Linn. Soc. Lond.
Bot. 9 (1865) 147, pro syn. — U. amphibia WELW.
ex KAMIENSKI, Bot. Jahrb. 33 (1902) 112, pro syn.
— U. exoleta var. lusitanica KAMIENSKI, Bot. Jahrb.
33 (1902) 112. — U. riccioides A. CHEV. Bull. Soc.
Bot. Fr. Mém. 8 (1912) 187. — U. nagurai MAKINO,
Bot. Mag. Tokyo 27 (1913) 59. — U. gibba L. ssp.
exoleta (R. Br.) P. TAYLOR, Mitt. Bot. Staatssamml.
Miinchen 4 (1961) 101; in Hutch. & Dalz. Fl. W.
Trop. Afr. ed. 2, 2 (1963) 381; Kew Bull. 18 (1964)
204; Dansk Bot. Ark. 23 (1968) 530; BAcK. &
BAKH. f. Fl. Java 2 (1965) 51. — Fig. 2f, 20.
Aquatic. Rhizoids few, filiform with short botry-
form branches. Sto/ons filiform, terete, up to 20 cm
long or more, up to 1 mm thick, much branched
and often mat-forming. Foliar organs numerous on
the stolons, up to 15 mm long, sparsely dichoto-
mously divided, ultimate segments few, capillary,
terete, glabrous or very sparsely setulose. Traps
numerous, lateral on the foliar segments, obliquely
ovoid, stalked, 1-1.5mm long, mouth lateral,

1977]

LENTIBULARIACEAE (Taylor)

295

Fig. 20. Utricularia exoleta R.BR. a—b. Foliar organs
on stolon, x 4, c. bract in situ, x 6, d. ditto,
flattened, x 15, e-g. flower, in adaxial, lateral and
frontal views respectively, x 4, h. glands on spur
outside, x 75,i. ditto inside, x 75,j.stamen, x 15,
k. pistil, x 15, 1. fruiting calyx, x 6, m. fruit, x 6,
n. ditto, dehisced, x 6, o—p. seeds, x 15 (a, e-i
Jones 310, b-d, j-] Jones 386, m—n, p ADAMES 201,
o JORDAN 2096).

sometimes with short + th grat setae, upper lip
with 2 long, much branched setiform appendages.
Inflorescence erect, 2-15cm long, solitary or
fasciculate; peduncle filiform, terete, glabrous;
scales usually 1 near the middle of the peduncle,
similar to the bracts; bracts basifixed, trans-
versely oblong or semi-orbicular, c. 1 mm long,
pirat ren or obscurely crenulate; bracteoles
a ; pedicels filiform, terete, suberect, 2-12 mm
long. Calyx lobes subequal, + orbicular, apex
rounded. mad apace 4-8 mm long, upper lip
orbicular or broadly ovate, about twice as long as
upper calyx lobe, 3-4 mm wide, apex rounded or
truncate, lower lip similar, palate raised, gibbous,
spur narrowly cylindrical from a conical base,
straight, apex obtuse, slightly longer than and +

lel with the lower lip. Filaments linear,
curved, anther thecae + confluent. Ovary globose,
style short but distinct, stigma lower lip semi-
orbicular, upper lip much shorter or obsolete.
Capsule glo , 2-3 mm long, firm in texture,
laterally bivalvate. Seeds few, lenticular, 1-1.6 mm

wide, with a broad irregular corky wing, testa
smooth or slightly verrucose, hilum prominent.

Distr. Tropical Africa, Portugal, India to China
and Japan and N. Australia; in Malesia common
and widespread but not recorded from the Lesser
Sunda Is.

Ecol. Shallow still water in lakes, swamps and
marshes, also in Pandanus swamp forest, largely at
low altitude, occasionally ascending to 1600, and
even to 2100 m. Fi. Febr.—March, July—Dec.

Vern. W. Java: lukut tjai, S; W. Borneo: veen-
eoblg Pontianak; New Guinea: keenapidu, Enga
ang.

Fig. 21. Utricularia aurea Lour. a. Foliar organ
with traps, x 4, b. foliose stipule-like segments at

base of foliar organs, = 6, c. small and large trap,
x 6, d. bract, flattened, = 6, e¢. flowering calyx
x 4, f. flower in lateral view, = 4, g. ditto, frontal
view, x 4,h. stamen, = 12, i. pistil, x 12, j. young
fruit, with swollen pedicel, * 4, k. circumscissile
upper half of dehisced capsule, x 4, 1. seed, left to
right in abaxial, adaxial view, = 12, and in section,
x 24 (a, d-e,g, j/-k Larsen & WARNCKE 1663, b-c,
f, h-i Darsysuire 605, / HAVILAND 2177A).

296

19. Utricularia aurea Lour. Fl. Coch. (1780) 26;
Merr. Trans. Am. Phil. Soc. 24 (1935) 356; HAND.-
Mazz. Symb. Sin. 7 (1936) 874; STEEN. Trop.
Natuur Jub. no. (1936) 123, f. 20; BAcK. & BAKH.
f. Fl. Java 2 (1965) 517; P. TAYLor, Dansk Bot.
Ark. 23 (1968) 529, excl. syn. U. fluitans. — U. vul-
garis LINNE, Sp. Pl. (1753) 18, partim quoad ref. FI.
Zeyl. — U. flexuosa VAHL, Enum. 1 (1804) 198;
BL. Bijdr. (1826) 739; OLtver, J. Linn. Soc. Lond.
Bot. 3 (1859) 175, excl. syn. U. ramosa; BTH. FI.
Austr. 4 (1869) 525, excl. syn. U. australis; CLARKE,
Fl. Br. Ind. 4 (1884) 329; Forses & HEMSL. J. Linn.
Soc. Lond. Bot. 26 (1890) 223; RipL. Trans. Linn.
Soc. Lond. II, Bot. 3 (1893) 327; TRIMEN, Handb.
Fl. Ceyl. 3 (1895) 267; Boerv. Handl. 2, 2 (1899)
560; Ript. J. Str. Br. R. As. Soc. n. 33 (1900) 119;
PRAIN, J. As. Soc. Beng. 74, ii (1905) 368; RIDL.

S. V7]
—— wa
Vio 4

am \

FLORA MALESIANA

RNY, / BAN = 0.

EA

[ser. I, vol. 8?

J. Str. Br. R. As. Soc. n. 59 (1911) 144; Koorp. Exk.
Fl. Java 3 (1912) 204; Merr. FI. Manila (1912)
432; Botp. Zakfl. (1916) 126; RipL. Fl. Mal. Pen.
2 (1923) 491; Merr. En. Philip. 3 (1923) 466;
Lingn. Sc. J. 5 (1927) 167; JAcoBson, Trop. Natuur
17 (1928) 112, fig.; HEND. Gard. Bull. S. S. 4 (1928)
295; MELCHIOR, Bot. Jahrb. 62 (1929) 385;
PELLEGR. FI. Gén. I.-C. 4 (1930) 471; BAck. Onkr.
Suiker. (1931) 633; SANDs, Mal. Agric. J. 21 (1933)
175; Spare, Mal. Nat. J. 1 (1940) 88; SANTAPAU,
J. Bomb. Nat. Hist. Soc. 49 (1950) 218; HEND. Mal.
Nat. J. 6 (1950) 336. — U. fasciculata ROxB.
(Hort. Beng. 1814, 4, nomen) FI. Ind. 1 (1820) 143;
WALL. Cat. (1829) n. 1568; Miq. Fl. Ind. Bat. 2
(1859) 997; Suppl. 1 (1860) 246. — U. confervifolia
Jacks. ex D. Don, Prod. FI. Nepal. (1825) 84. —
U. flexuosa var. blumei A. DC. in DC. Prod. 8

YY ee

a
’
bg

a
Ne

7 / Kr Y
=4@\ P

Fig. 22. Utricularia aurea Lour. Habit, about nat. size, a fruit in natural poise separately (DOCTERS VAN
LEEUWEN 2281).

1977]

LENTIBULARIACEAE (Taylor)

297

Fig. 23. Utricularia aurea Lour. Form with conspicuous inflated rhizoids from the base of the peduncle,
slightly enlarged. In clay ditch in seasonal savanna, Indramayu, W. Java, March 1936 (VAN STEENIS s.71.).

(1844) 24. — U. inaequalis BEN). Linnaea 20 (1847)
304. — U. calumpitensis LLANOs, Fragm. (1851) 11;
Merr. Sp. Blanc. (1918) 351. — U. extensa HANCE
in Walp. Ann. 3 (1852) 3. — U. reclinata HAssk.
Versl. Med. Kon. Ak. Wet. A’dam 4 (1855) 161;
Retzia 1 (1855) 92; ibid. 10 (1856) 92; BoeRL.
Handl. 2, 2 (1899) genus 559. — U. blumei (A.
DC.) Mia. Fl. Ind. Bat. 2 (1859) 997. — U. vulgaris
var. pilosa MAKINO, Bot. Mag. Tokyo 9 (1895)
111. — U. pilosa (MAKINO) MAKINO, Bot. Mag.
Tokyo 11 (1897) 70. — Fig. 2c, 21-23.

Aquatic. Rhizoids usually present, verticillate at
or near the base of the peduncle, fusiform, inflated,
2-6 cm long, 1-3 mm thick (fig. 23), with filiform
branches bearing botryform clusters of ellipsoid
——— 0.1-0.2 mm_long. Stolons filiform to
relatively thick, up to 50 cm long, terete, branched,

brous or + densely covered with short simple

irs. Foliar organs numerous and conspicuous
2-6 cm long, primary segments 3-4, semi-verticil-
late, filiform or sometimes thick and inflated, each
pinnately repeatedly divided from near the base
into numerous segments, ultimate segments capil-
lary, terete, setulose; stipule-like clusters of short
capillary setulose segments usually present at the
base of the primary segments. 7raps usually
numerous, lateral on the foliar segments and some-
times also in the angle between segments, er oe
ovoid, shortly stalked, 1-4 mm long, mouth lateral,
upper lip usually with 2 short sparsely branched
setiform = Acme or without appendages.
Inflorescence erect, 5~25 cm long; peduncle filiform,
terete, glabrous; flowers 5-10, initially congested

becoming + distant; scales always absent; bracts
basifixed, + orbicular, 1.5-2mm long, apex
rounded or subacute; bracteoles absent; pedicels
filiform, dorsiventrally flattened, elliptic in cross
section, 4-20 mm long, erect at anthesis, usually
sharply reflexed and thickened, especially apically,
in fruit. Calyx lobes subequal, the upper slightly
longer, ovate, 3-4mm long at anthesis, rather
fleshy, apex obtuse, margins strongly incurved, very
accrescent, up to 9mm long and spreading or
reflexed in fruit. Corolla yellow, externally glab-
rous or sparsely hairy, 10-15 mm long, upper lip
broadly ovate, about twice as long as the upper
calyx lobe, apex rounded, lower lip larger, trans-
versely elliptic, apex rounded, entire or retuse
palate raised, gibbous, spur cylindrical from a coni-
cal base, often constricted at the middle, apex
obtuse, slightly shorter than and + parallel with
the lower lip. Filaments linear, expanded above,
c. 2mm long, anther thecae confluent. Ovary
lobose, glandular, style relatively long, stigma
inet lip orbicular, margin hyaline, ciliate, upper
lip much shorter, margin glabrous. Capsule
dolce up to 5 mm long, relatively thick and fleshy,
circumscissile, style usually beds! enlarged and
elongated, often equalling or exceeding the capsule,
Seeds numerous, lenticular-prismatic, 5-angled,
1.5-2 mm wide, very narrowly winged on all the
angles, testa thin, obscurely reticulate, reticulations
slightly elongate.
Distr. India to China and Japan to Australia,
throughout Malesia, common and bg
Ecol. Deep or shallow still water in lakes and

298

FLORA MALESIANA

[ser. I, vol. 8?

rice-fields, also in pandan swamp forest at low
and medium altitude, ascending to 1200 and occa-
sionally to 1500 m. F/. Jan.—Dec.

Vern. Sumatra: giamon, M, Enggano; E.
Borneo: klanibu udang, Kutei; Philippines:
bagingan gadudugge, bagiw di udongo, If.; New
Guinea: pehra pehra, Matapaili lang.

Note. This very common species is very variable
in size. The inflated rhizoids at the base of the
peduncle (fig. 23) are sometimes very conspicuous
but often absent.

20. Utricularia muelleri KAMIENSKI, Ber. Deut. Bot.
Ges. 12 (1894) 5. — U. stellaris (non L. f.) BTH. FI.
Austr. 4 (1869) 525, partim quoad spec. Gulf of
Carpentaria, F.v.MUELLER. — U. inflexa Forsk.
var. stellaris (L. f.) P. TAYLor, Kew Bull. 18 (1964)
191, partim quoad syn. U. muelleri excl. ref. Bot.
Jahrb. 33 (1902) 108. — Fig. 24.

Aquatic. Rhizoids apparently absent. Stolons
filiform, terete, glabrous, up to 60cm long, un-
branched. Foliar organs numerous and conspi-
cuous, 2—3 cm long, primary segments 3 or more,
semi-verticillate, filiform, each dichotomously
repeatedly divided from near the base into numer-
ous segments, ultimate segments capillary, terete,
minutely setulose; stipule-like clusters of short
hyaline flattened densely setulose segments usually
present at the base of the primary segments. Traps
usually numerous, lateral on the intermediate
segments, obliquely ovoid, 1-3 mm long, shortly
stalked, mouth lateral, apparently without appen-
dages. Inflorescence erect, 4-15 cm long, peduncle
filiform, terete with a whorl of 4-6 spongy floats
approximately midway, floats ellipsoid, 10-20 mm
long, shortly stipitate, with capillary foliar seg-
ments at the apex and laterally and more numerous
at the base; flowers 2-14, rather congested; scales
absent; bracts transversely elliptic when flattened,
cucullate, amplexicaule, hyaline, nerveless, 2.5-3
mm long, apex + deeply denticulate-laciniate;
bracteoles absent. Calyx lobes unequal, connate,
scarcely accrescent, upper broadly ovate, c. 2mm
long, cucullate, apex rounded, lower about as long
but broader + orbicular, apex emarginate. Corolla
yellow with pink nerves on the palate and spur,
6-9 mm long, glabrous, upper lip oblong, cucullate,
much longer than upper calyx lobe, lower lip
shorter, transversely oblong, apex emarginate,
palate raised, gibbous, spur broadly conical,
slightly curved, apex rounded, about as long as
lower lip. Filaments falcate, c. 2mm long, some-
what expanded above, anther thecae subdistinct.
Ovary globose, style distinct, stigma lower lip
quadrate, reflexed, upper lip very short, emarginate.
Capsule globose, 34mm long, circumscissile.
Seeds few, thinly lenticular, 1.5-2 mm @ including
a distinct narrow hyaline wing, testa thin with
small conspicuous reticulation.

Distr. N. Australia (Northern Territory and
Queensland) and Malesia: New Guinea (Papua).

Ecol. Shallow water in lowland Melaleuca
swamp forest. F/. Aug.

Note. U. muelleri is superficially very similar to
the more widespread U. stellaris L. f. which occurs
from tropical Africa to Indo-China and reappears
in northern Australia, but is apparently absent
from Malesia. For differences see the key to the

species.

=

Fig. 24. Utricularia muelleri KAMIENSKI. a. Whorl
of five floats showing stipites and segments at or
near their bases (+ ‘air shoot’), x 5, b. hyaline
stipule-like segments from base of primary seg-
ments, < 12, c. secondary to ultimate segments of
foliar organs with traps, x 5, d. apex of foliar
segments on floats, x 24, e. part of deflorated
infiorescence, x 4, f. bract, flattened, x 6, g.
flowering calyx, 6, h. corolla, front view, x 4,
and below it spur apex, with internal sessile and
external stipitate glands, x 24, i. stamen, x 12, j.
pistil, x 12, k. capsule, with circumscissile line
for later dehiscence, x 5,1. seed, x 12, and below
it in section filled with embryo, x 24, m. testa,
x 24, n. apex of air shoot (a—c, n ByRNES 2381, dj
Latz 3697, k PARKER 117, /-m PULLEN 7058).

a

1977]

LENTIBULARIACEAE (Taylor)

299

a eeeeEeEeeEeSSSSSSSSSSSSSSSSSSSSSSSSeeEeEeeeee

21. Utricularia australis R. Br. Prod. Nov. Holl.
(1810) 430; A. DC. in DC. Prod. 8 (1844) 6; P.
TAYLOR in Tutin et a/. Flora Europaea 3 (1972) 297.
— U. neglecta LEHM. Pugillus Plantarum | (1828)
38. — U. sacciformis BEN. Linnaea 20 (1897) 302.
— U. flexuosa (non VAHL) Bru. FI. Austr. 4 (1869)
525; CLARKE, Fl. Br. Ind. 4 (1884) 329, partim et
quoad syn. U. australis R. Br. — U. incerta
KAMIENSKI, Bot. Jahrb. 33 (1902) 111. — U. japoni-
ca MAKINO, Bot. Mag. Tokyo 28 (1914) 28. —
U. stellaris (non L. f.) WAGER, Trans. R. Soc. S.
Afr. 16 (1928) 204. — U. vulgaris (non L.) P.
TAYLOR, Kew Bull. 18 (1964) 171. — Fig. 25.
Aquatic. Rhizoids 2-4 near base of peduncle,
capillary, 1-2 cm long with a few short botryform
segments. Stolons filiform, up to 50cm long,
terete, branched, glabrous. Foliar organs numer-
ous, 1.5—4 cm long, primary segments 2, filiform or
sometimes + inflated, each repeatedly pinnately

Fig. 25. Utricularia australis R.Br. a. Foliar organ,
at the base with two rudimentary traps, some traps
not drawn indicated by ‘t’, « 5, b. trap, x 12, c.
scale, * 4, d. insertion of flower (left stalk), with
bract in situ, * 4, e. bract, flattened, = 4, f.

flowering calyx, lowest margin hyaline, * 4, g.

flower, front view, spur separately drawn, = 4,

h. spur, lateral view (the punctate glands are in-

ternal), x 4, i. stamen, 6, j. pistil, * 6 (all after
ASTON 1799).

divided from very near the base into numerous
segments, ultimate segments capillary, distinctly
flattened, marginally setulose. Globose or ovoid
turions (winter buds) 0.5—-1.5 cm long of reduced
densely setulose foliar segments often present at
apex of stolons or stolon branches. Traps usually
numerous, lateral on the segments and usually also
at the base of the primary segments, obliquely
ovoid, 1-2 mm long, shortly stalked, mouth lateral,
with simple marginal setae, upper lip with two long
branched setiform appendages. Inflorescence erect,
up to 40cm long; peduncle at first straight
becoming flexuous post anthesis, relatively stout,
1-2 mm thick terete, glabrous; flowers 4-10,
initially congested becoming + distant; scales 2-3
always present in the upper half of the peduncle,
similar to the bracts; bracts basifixed, + orbicular,
3-5 mm long, base auriculate, apex rounded or
obscurely tridentate; bracteoles, absent; pedicels
filiform, 1.5-2.5cm long, erect at anthesis, later
+ spreading. Calyx lobes subequal, ovate-oblong,
c. 3mm long, apex of upper rounded, of lower
emarginate. Corolla yellow, c. 15 mm long, exter-
nally glabrous, upper lip ovate to orbicular, 2-3
times as long as upper calyx lobe, lower lip much
larger, transversely elliptic, up to 16mm wide,
apex rounded or retuse, palate raised, gibbous,
spur cylindrical or broadly conical, obtuse, slightly
curved, shorter than the lower lip. Filaments linear,
curved, anther thecae confluent. Ovary globose,
style distinct, about as long as ovary, stigma lower
lip semi-orbicular, margin denticulate, upper lip
very short. Capsule and seeds not known.

Distr. W. Europe to China and Japan, tropical
and S. Africa, India (south to Ceylon) to SE.
Australia; in Malesia: Sumatra, Java, Philippines,
and New Guinea.

Ecol. Pools in swamps, in Malesia at high alti-
tudes, 1300-2500 m, often not flowering. F/. April-
July, Nov.

Note. This widespread temperate Eurasian
species apparently never sets seed and its some-
what sporadic occurrence, mostly at high altitude,
in the African and Asian tropics is probably due
to transmission of small particles of its vegetative
parts by migrating birds. It has been much con-
fused with U. aurea but is easily distinguished by its
1 or 2 (not 3 or more) primary foliar segments
without basal stipule-like segments and when in
flower by the presence of scales on the peduncle.

22. Utricularia minor LINNé, Sp. Pl. (1753) 18; A.
DC. in DC. Prod. 8 (1844) 7; OLtver, J. Linn. Soc.
Lond. Bot. 3 (1859) 176; P. TAYLOR in Tutin et al.
Fl. Europaea 3 (1972) 296. — U. rogersiana LACE,
Kew Bull. (1915) 405. — U. minor var. multispinosa
Mik1, Bot. Mag. Tokyo 48 (1934) 337. — U. multi-
spinosa (Mik1) Miki, Water Phaner. Jap. (1937)
109; Onwi, Fl. Japan (1965) 815. — U. punctata
(non WALL. ex DC.) P. TAyLor, Dansk Bot. Ark.
23 (1968) 531, partim, quoad syn. — Fig. 26.
Aquatic. Rhizoids not clearly differentiated.
Stolons filiform, terete, up to 20cm long, 0.1-
0.3mm _ thick, sparsely branched, glabrous, +
dimorphic, some green, suspended or floating,
others without chlorophyll and buried in the sub-
strate. Foliar organs numerous, polymorphic, +
circular in outline, 0.2-1.5 cm long, + palmately
divided into rather few segments, the ultimate

300 FLORA MALESIANA [ser. I, vol. 8?

segments flattened, capillary to narrowly linear,
0.1-0.5 mm wide, the margin sometimes sparsely
denticulate but not or only microscopically
setulose, the apex acute with or without a micro-
scopic seta. Globose turions (winter buds) 1.5-4
mm @ of reduced glabrous foliar segments often
present at apex of stolons or stolon branches.
Traps rather few on the narrower foliar segments,
usually absent on the broader ones and most
numerous on reduced segments on the subter-
ranean stolons, obliquely ovoid, shortly stalked,
0.8-2.5mm long, mouth lateral with marginal
simple hairs, upper lip with two long much
branched setiform appendages. Inflorescence erect,
5-20 cm long; peduncle filiform, terete, straight;
flowers 2-8 somewhat distant, scales 2-4, +
equally spaced on the peduncle, similar to the
bracts; bracts basifixed, broadly ovate-deltoid
1.5-2 mm long, 1-nerved, apex obtuse, base con-
spicuously auriculate; bracteoles absent; pedicels
erect at anthesis, spreading and apically reflexing
in fruit, filiform, terete, 4-7 mm long. Calyx lobes
subequal, the upper somewhat larger, broadly
ovate, c. 2 mm long, apex obtuse, cucullate. Corolla
pale yellow, 8-10 mm long, upper lip ovate, apex
obtuse, lower lip larger, obovate, apex retuse,
palate raised, elongate, distally narrowed, spur
saccate, c. 1.5mm long, internally densely glan-
dular. Filaments linear, curved, dilated above,
anther thecae confluent. Ovary globose, style
distinct, stigma lower lip orbicular, reflexed, upper
lip narrowly deltoid, both. fimbriate. Capsule
globose, c. 3mm_ long, circumscissile. Seeds
lenticular-prismatic, c. 0.6mm @, very narrowly
winged on the angles.

Distr. Circumboreal, extending southwards
into the Himalaya, Burma, and Malesia: New
Guinea (Mt Hagen area).

Ecol. Shallow water in high mountain swamps,
2500-3660 m.

Note. The three Malesian specimens seen are
without inflorescences but there can be little doubt
that they are the same as the circumboreal plant.
The small, almost glabrous, turions distinguish it
from all other species in the genus. Like U.
al hd it is probably distributed by migratory

irds.

Fig. 26. Utricularia minor L. a. Narrow foliar
segments, from stolon, with one trap, 6, b.
ditto, apex of segment, x 12, c. narrow foliar
segments, x 5, d. apex, x 12, e. broad foliar
segments, with stolon, x 6, e’. apex of broad
segment, x 12,f. turion, 6, g. scale in situ, x pe
h-h’. bract in situ, and flattened, x 6, i. flowering
calyx, x 6, j. flower, lateral view, x 4, k. ditto,
upper and lower lip flattened, and below it lateral
view of spur, x 5,1. stamen, x 12, m. pistil, x 12,
with flattened lower lip of stigma, < 12, n. fruiting
calyx, x 6, 0. fruit on pedicel, x 6, p. capsule,
dehisced, x 6, q. seed, from lateral (lat), apical (ap)
and basal (bas) views, x 12(a—b, e-f ANU 667, c-d,
g, n-q P. TAYLOR s.n., h-m R. B. DRUMMOND .n.).

Doubtful

Lemnopsis mnioides Z1PPEL, Flora 12 (1829) i, 285, nomen; Alg. Konst- & Letterbode 1 (1829) 297, nomen,
is according to HALLIER f. Med. Rijksherb. 1 (1910) 40, cf. Fl. Males. I, 4 (1949) 68 = Utricularia orbicu-
lata WALL. = U. striatula SM.

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#. @

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 REQUESTS AND RELATED CORRESPONDENCE

SHOULD BE DIRECTED EXCLUSIVELY TO

SIJTHOFF & NOORDHOFF INTERNATIONAL PUBLISHERS

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Series I - Spermatophyta (Flowering Plants)

Cyclopaedia of collectors & collections. 1950. pp. clii + 639. Out of
print. Available in microfiche.

Malesian vegetation (In preparation)
Malesian plant geography (In preparation)

General chapters and revisions. 1948-1954. pp. ccix + 631
Bibliography, specific delimitation & revisions. 1955-1958. pp.
cecxlii + 596

Systematic revisions. Parts 1-6. 1960-1972. pp. 20 + 1023. Out of print.
Available in microfiche.

Systematic revisions. Parts 1-4. 1971-1976. pp. 18 + 876
Cyclopaedia of Collectors, Suppl. 2. Systematic revisions. Part 1. 1974.
Systematic revisions. Part 2. 1977.

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