REINWARDTIA
Published by Herbarium Bogoriense, Kebun Raya Indonesia
Volume 2, P a r t 2, pp. 185-224 (1953)
THE GENUS TETRACERA (DILLENIACEAE) IN THE
EASTERN OLD WORLD
(With Plate 1)
R . D . HOOGLAND*
SUMMARY
1. An account of the genus Tetracera L. in the eastern Old World (Asia, Malaysia,
Australia, New Caledonia) is given. The Malaysian species have already been in-
cluded in the revision of the Dilleniaceae in "Flora malesiana" (I 4: 141-149. 1951).
2. The main part of the present paper consists of a key to the species, followed
by a systematic treatment of the 15 species admitted for the region.
3. Latin diagnoses are given for three new subspecies under Tetracera asiatica
(Lour.) Hoogl. and two new varieties under Tetracera nordtiana F. Muell.
4. A number of species are reduced either to the rank of variety or to synonymy.
5. Distribution-maps are provided for the species of which a relatively large;
number of specimens has been studied.
INTRODUCTION.—The present paper forms an extension of my revision
of the genus Tetracera L. in "Flora malesiana" (I 4: 141-149. 1951) and
the revision now covers, besides Malaysia, continental Asia (Ceylon and
Hainan included), Australia, and New Caledonia.
The study was made possible by the loan of specimens from several
herbaria, which have been indicated in the citation of type specimens by
the abbreviations proposed by Lanjouw [in- Reg. Veg. 2 (Ind. Herb. 1):
106-117. 1952] as follows:
A Arnold Arboretum Herbarium, Harvard University, Jamaica Plain, Mass.
BM Department of Botany, British Museum (Natural History), London.
BO Herbarium Bogoriense, Bogor, Java,
BR Herbier du Jardin Botanique de l'£tat, Bruxelles.
BRT Botanical Museum and Herbarium, Brisbane.
BRSL Botanical Institute, Wroclaw (formerly Breslau).
C Universitetets Botaniske Museum, Copenhagen.
CAL The Indian Botanic Garden and Herbarium, Calcutta,
CGE Botanical Museum and Herbarium of the University, Cambridge (England).
DD Forest Research Institute Herbarium, Dehra Dun.
E Herbarium of the Royal Botanic Garden, Edinburgh.
FI Erbario del Instituto Botanieo dell' University, Firenze.
G Institut de Botanique Systematique de l'Universite, Geneve.
GH Gray Herbarium, Harvard University, (Jambridge, Mass.
•Botanist , Flora Malesiana Foundation, now C. S. I. R.O., Canberra, A. C.T.,
Australia.
-; [REINWARDTIA, Vol. 2, P a r t 1 was issued September 12, 1952]
REINWARDTIA [VOL. 2
GL Botanical Department Herbarium, Glasgow.
GRO Botanisch Laboratorium der Rijksuniversiteit, Afdeling Plantensystematiek,
Groningen.
I F I Herbarium of the Imperial Forestry Institute, Oxford.
K The Herbarium, Royal Botanic Gardens, Kew, Surrey.
L RIjksherbarium, Leiden.
LINN Herbarium of the Linnean Society of London, London.
M ' Botanische Staatssammlung, Miinehen.
MEL Botanic Gardens and National Herbarium, Melbourne.
MICH Herbarium of the University of Michigan, Ann Arbor, Mich.
MO Herbarium of the Missouri Botanical Garden, St. Louis, Mo.
NY Herbarium of the New York Botanical Garden, New York, N. Y.
OXF Oxford University Herbarium, Oxford.
P Laboratoire de Phanerogamie du Museum National d'Histoire Naturelle,
Paris.
5 Naturhistoriska Riksmuseet, Stockholm.
SING Herbarium of the Botanic Gardens, Singapore.
U Botanisch Museum en Herbarium, Utrecht.
UC University of California Herbarium, Berkeley, Calif.
U P S Botaniska Museet, Uppsala.
US United States National Herbarium, Washington, D. C.
I wish to express here my indebtedness to the Directors and Keepers
of these herbaria for their valuable help in putting their specimens at my
disposal.
The detailed citation of specimens examined has been omitted and
in its place distribution-maps have been prepared for the species of which
a relatively large number of specimens have been studied. All specimens
studied by the author have been provided with an identification-label.
TETRACERA L.
Tetracem L., Sp. PI. 533. 1753; Gen. PI., 5th Ed., 237. 1754; D C , Syst. 1; 397.
1818; Prod. 1: 67. 1824; Roxb., F l . ind., ed. Carey, 2: 646. 1832; Hook. f. & Thorns.,
PI. ind. 1: 62. 1855; Miq., Fl. Ind. bat. 1 (2): 8. 1859; Benth. & Hook, f., Gen. PI. 1:
12. 1882; Drury, Handb. Ind. Fl. 1: 9. 1864; Miq. in Ann. Mus. bot. Lugd. Bat. 4:
74. 1868; Hook. f. & Thorns, in Fl. Br. Ind. 1: 81. 1872; Kurz, For. Fl. Br. Burma 1:
22. 1877; Eichl., Blutendiagr. 2: 251. 1878; Martelli in Becc, Malesia 3: 150. 188G;
King in J. As. Soc. Beng. 58 (2) : 362. 1889; K. Schum. & Hollr., PI. Kais. Wilhelmsl.
47. 1889; Boer]., Handl. 1 (1) : 6. 1890; Trim., Handb. Fl. Ceyl. 1: 6. 189S; Gilg in Engl.
6 Prantl, Nat. PflFam. 3, 6: 110. 1895; Pritzel in Bot. Jb. 24: 352. 1897; F. M. Bail.,
Queensl. Fl. 1: 8. 1899; Ridl. in J. S t r . B r . R. A. S. 33: 37. 1900; K. Schurn. & L a u t ,
Fl. deut. Schutzgeb. Sudsee 444. 1901; Brand., Ind. Trees 4. 1906; Fin. & Gagnen.-
in Fl. gen. Ind.-Ch. 1: 12. 1907; Back., Fl. Batavia 1: 3. 1907; Schoolfl. J a v a 8.
1911; Koord., ExkFl. J a v a ' 2 : 600. 1912; Ridl. in Saraw. Mus. J. 1: 68. 1913; Merr.,
Bibl. En. Born. PI. 281. 1921; Ridl., Fl. Mai. Pen. 1: 4. 1922; Drels in Bot. J b . 57:
439. 1922; Merr., En. Philip, fl. PI. 3; 58. 1923; Craib, Fl. siam. En. 1: 19. 1925; Gilg
& Werderm. in Engl. & Prantl, Nat. PflFam., 2nd Ed., 2 1 : 16. 1925; Back., B e t a . PI.
J a v a (Nooduitg.) 4 (Fam. 80) : 1. 1942; Hoogl. in PL mal. I 4: 141. 1951.
1853] HOOGLAND: Tetmcera 187
Delima L., Gen. PI., 5th Ed., 231. 1754; Syst., 10th Ed., 1076. 1759; D C , Syst.
1: 397. 1818; Prod. 1: 67. 1824; Hook. f. & Thorns., Fl. ind. 1: 61. 1855; Miq., Fl.
Intl. bat. 1 (2); 7. 1859; Benth. & Hook, t., Gen. PI. 1: 12. 1862; Triana & Planch.
in Ann. Sci. nat., Bot. IV 17: 20. 1862; Drury, Handb. Ind. Fl. 1: 11. 1864; Miq.
in Ann. Mus. bot. Lugd. Bat. 4: 73. 1868; Hance in 3. of Bot. 7: 115. 1869; Hook. f.
& Thorns, in Fl. Br. Ind. t : 31. 1872; Kurz, For. Fl. Br. Burma 1: 22. 1877; King in
J. As. Soc. Beng. 38 (2): 361. 1889; BoerL, Handl. 1 (1) > 6. 1890; Trim., Handb. Fl.
Ceyl. 1: 5. 1893; Brand., Ind. Trees 5. 1906; Ridl. in Saraw. Mus. J. 1: 58. 1613; Fl.
Mai. Pen. 1: 3. 1922; Burk., Diet. econ. Prod. Mai. Pen. 776. 1935.
Korosvel Adans., Fam. des PI. 2: 442. 1763.
Assa Houtt., Nat. Hist. 5: 275. 1776; Christm. & Panz., PflSyst. 1: 40. 1779.
Enryandra Foist., Char. Gen 81. 1776.
Wahlbomia Thunb. in Vet. Akad. Handl., Stockh., 215. 1790; Eafin., Sylva tellur.
105. 1838 ("Valbomia").
Roehlingia Dennst., Schluess. Hort. malab. 31. 1818.
Bleiastis Rafin., Sylva tellur. 165. 1838.
Leontoglossttm Hance in Walp., Ann. 2: 18. 1859.
Delimopsis Miq., Fl. Ind. bat. 1 (2): 9. 1859.
T Y P E SPECIES.—Te t ra ce ra: T. volubilis L., 1753, I.e. (Central America).—
Delima: D . sarmentosa L . , 1 7 5 9 , I.e. r = T . scandens ( L ) . M e r r . — Korosvel:
"Korosvel. Herm. Zeyl. 19." 1717 = T. asiatica (Lour.) Hoogl. subsp. zcylanica Hoogl.
— Ass a: A. indica Houtt. ex Christm. & Panz., 1779, Lc. — T. indica (Houtt. ex Christm.
& Panz.) Merr. — Euryandra: E. scandens Forst., 1776, I.e. = T. euvyandra
Vahl. — Wahlbom i n : W. indica Thunb., 1790, I.e. = T. indica (Houtt. ex Christm.
& P a n z . ) M e r r . — Roehlingia: R. suaveolens D e n n s t . , 1 8 1 8 , I.e. = T. akara
( B u r m . f.) M e r r . — E l e i a s t i s : E . laevis ( V a h l ) R a f i n . , 1 8 3 8 , I.e. = T . indica
( H o u t t . ex C h r i s t m . & P a n z . ) M e r r . — Leontoylussu m: L. scabrum H a n c e , 1 8 5 1 ,
Lc. = T. asiatica ( L o u r . ) H o o g l . s u b s p . asidtica. — Delimopsis: D. hirsnta M i q . ,
1859, I.e. = T. askitica (Lour.) Hoogl. subsp. sumatrana Hoogl.
Shrubs, sometimes straggling, or lianas, with flexuous branches;
older branches with flaky bark with longitudinal fissures. Leaves spirally
arranged, simple, petiolate, without stipules; base decurrent; margin
manifestly dentate, most distinctly so in upper half of leaf, with teeth
at apex of nerves to entire; leaves penninerved, often scabrid on one
or both sides; petiole short, slightly channelled above. Inflorescences few-
to many-flowered panicles, terminal or axillary, often with bracts; peduncle
distinct or not. Flowers fragrant, actinomorphic, bisexual. Sepals 4—6,
rarely up to 15, imbricate, persistent, often reflexed when flowering
and fruiting, circular to oval with rounded apex and base and usually
ciliate margin. Petals 3—5, caducous, obovate-spathulate with slightly
emarginate apex, narrowed towards the base, whitish, often slightly
reddish. Stamens ~ (about 60—500) ; filament thin; anther with broad-
ened connective; thecae divergent towards the base, touching each other
at the apex or more or less separated (connective in the latter case often
emargrinate between thecae), usually lateral, rarely more or less on outer
side of connective, opening with longitudinal slit. Carpels 4—1, free,
each with short style ending in hardly differentiated stigma; placentae
marginal, adaxial, each with single row of 1—10 ovules. Fruits coriaceous
1 8 8 R E I N W A R D T I A [VOL. 2
capsules, opening with longitudinal slits along ventral and dorsal suture
into two valves, with short beak, one- to few-seeded. Seeds glossy dark-
brown to black with abundant endosperm and microscopically small
embryo (cf. Pritzel, 1897, I.e.), arillate; aril fleshy, fimbriate or laciniate
at margin for 1/3 to nearly the whole length.
HISTORY OF THE GENUS. — The genus Tetracera was founded by
Linnaeus (1753, 1754) on a single species from Central America. The
genus Delima, already mentioned by Linnaeus in 1748 (Fl. zeyl. 92) was
not included in the first edition of "Species Plantarum" (1753), though
it was included in the fifth edition of "Genera Plantarum" (1754); a
single species was described in the tenth edition of "Systema Naturae"
(1759). Triana & Planchon (1862) were the first to reduce Delima to Tetra-
cera, as the only differential character (the single carpel, to which in some
flowers a second is added) does not justify its distinction as a separate
genus. Nevertheless, it was still treated as such by Ridley (1922) and,
following him, by Burkill (1935).
The other generic names, proposed for species from the region con-
cerned, have hardly been taken up after their first publication.
RELATIONSHIPS.—The genus Tetracera was included by Gilg & Wer-
dermann (1925) in the tribus Tetracereae; in my opinion it is justifiable
to raise this tribus to the rank of a subfamily {cf. Blumea 7: 7. 1952).
Related genera within this subfamily are found only in the New World
(Davilla, Guratella, Doliocarpus). The subfamily is primarily character-
ized by the structure of the stamens (broadened and, sometimes, thickened
connective).
Within the genus, Gilg & Werdermann distinguished three sections.
The first of these, Empedoclea (St. Hi].) Gilg, is found only in the New
World (Brazil). The other two sections agree with the former genera Tetra-
cera L. s.s. (section "Eutetracera" Gilg) and Delima L. [section Delima
(L.) Gilg.]. As far as the species revised here are concerned, these sections
do not form natural taxa; I can not judge about the New World species.
According to the principal characters of these two sections (the number
of carpels) the first four species of the present revision would belong to
Delima. For Tetracera, scandens and T. asiatica I am not able to indicate
any relationship with other species; T. glaberrima, is most closely related
to T. akara,. and T. maingayi to T. fagifolia, thus both to species with a
greater number of carpels.
DISTRIBUTION.—The genus is pantropical with the exception of the
Pacific Islands east of New Britain and New Caledonia. The species them-
selves all have a more or less restricted area. None of those considered
here has been found outside the region concerned.
1953] HOOOLAND: Tetracera 189
All species occur in the lowland, rarely above 600m altitude; the
highest collection known to me is from about 1500 m (T. asiatica, Cam-
bodia). No general habitat can be indicated for the genus, some species
occurring in forests, others in scrub or even more open places.
VERNACULAR NAMES.—The Malay name mempelas (ampalas, ampelas,
empelas, mumplas) is in general use in western Malaysia, often with epi-
thets which are not specific. Akar (= root or liana) is found in some
names. The Sundanese name is assahan.
USES.—The scabrid leaves of some species are used as sandpaper;
hence the Malay name for sandpaper has been derived from the plant
name: mempelas. The stems are sometimes used as cordage.
NOTE.—The generic description above has been based on the species
studied, thus exclusive of the African and American species. The following
additions to the description are taken from the generic description by
Gilg & Werdermann: Sometimes trees. Sepals 3—15, "usually 5." Petals
1—6, "usually 5." Filaments very rarely more or less highly united to
fascicles. "Anthers usually extrorse, very rarely introrse." The parts
between inverted commas certainly do not hold for the species studied
here; the other parts are additions outside the variability of the genus
in the region treated in the present revision.
KEY TO THE EASTERN OLD WORLD SPECIES OF TETRACERA
1. Carpels in most flowers solitary, but often in some flowers of the same plant 2
carpels.
2. Carpels and capsules glabrous or with minute scales.
',i. Sepals 4; inflorescences up to 5-flowered, usually axillary; flowers about
2.5 cm across d. T. glaberrima
3. Sepals 5; inflorescences lo- and mure-flowered, terminal; flowers about 1—1.5
cm across.
4. Sepals glabrous inside 2. T. asiatica
4. Sepals sericeous inside 4. T. maingayi
2. Carpels and capsules hirsute 1. T, scandens
1. Carpels in all flowers 2—4.
5, Carpels and capsules hirsute all over their whole surface.
6. Indument of carpels consisting of rather thin villous hairs, caducous. Species
from W Malaysia 14. T. arborescens
G. Indument of carpels consisting of rather rigid, straight hairs, persistent.
Species from E Malaysia.
7. Inflorescenses 2—4-flowered; hairs of the carpels about 2 mm long.
5. T. lanuginosa
7. Inflorescenses 15—50-flowered; hairs of the carpels about 0.5 mm long.
6. T. uordtiana
5. Carpels and capsules glabrous, with minute scales, or with few strigose hairs
on the back only.
190 REINWARDT1A [VOL. 2
8. Sepals 4; inflorescenses few- (up to 12-) flowered, terminal or axillary, without
leaves in the basal p a r t ; flowers about 2.5—3 cm across.
9. Sepals glabrous inside 1. T. indica
9. Sepals sericeous inside 8. T, akara
8. Sepals 5—6; inflorescenses many- (15- and more-, rarely less-) flowered, ter-
minal, often with small leaves in basal part, sometimes on short axillary
few-leaved branch; flowers about 0,8—2.5 cm across.
10. Sepals all glabrous inside.
11. Branches of inflorescenses strigose 9. T. loureiri
11. Branches of inflorescenses with single appressed to distant hairs as
well as stellate groups of shorter hairs 12. T. korthalsii
10. Sepals, at least 3 inner ones, sparsely to densely sericeous inside.
12. Two outermost sepals glabrous inside 13. 7". macrophylla
12, All sepals sericeous inside.
13. Stamens extrorse; apex of leaves usually rounded; flowers r a t h e r
large (15—20 mm across) 11. T. curyandra
13. Stamens latrorse; apex of leaves usually obtuse to acute; flowers
r a t h e r small (8—15mm across).
14. Younger sterile branches absolutely glabrous. 10. T. daemeliana
14. Younger sterile branches always more or leas hairy.
15. Younger branches villous; leaves immediately below the
inflorescences small (about 4 x 3 cm), obovate; sepals sub-
equal 14. T. arborescens
15. Younger branches strigose; leaves immediately below the
inflorescences larger, elliptic to oblong; two outer sepals
distinctly smaller than inner ones. . . 15. T. fagifolia
1. TETRACERA SCANDENS (L.) Merr. — Fig. 1
[Fmtis urens aepera Rumph., Herb. amb. 5: 13 pi. 9*1 1747.]
Tragia scandens L. in Stickm., Herb. amb. 18. 1754; Amoen. acad 4: 128. 1759.
Delima sarmentosa L-, Syst., 10th Ed., 1076. 1759; Miq., Fl. Ind. bat. 1 (2): 7.
1859; Vill., Nov. App. 2. 1880; Ridl., Fl. Mai. Pen. 1: 3. 1922.
Tetracera, sarmodosa (L.) Vahl, Symb. bot. 3: 70. 1794; Roxb., Fl. ind., ed. Carey,
2: 645. 1832, p.p.; Blanco, Fl. Filip., 2nd Ed., 320. 1845; 3d Ed., 2: 227. 1878; Merr. w
Govt Lab. Philip. Publ. 27: 15. 1905; Hunter (ed. by Ridl.I hi J. Str. Br. R. A. S. 5 3 :
97. 1909; Merr., Fl. Manila 331. 1912.
Delima hebecarpa D C , Syst. 1: 407. 1818; Deless., Ic. sel. PL 1: pi 72* 1821; D C ,
Prod. 1: 70. 1824.
Delima intermedia Bl.f Bijdr. 1: 4. 1825; Schlecht. in Linnaea 1: 492. 182G; Hassk.,
PI. jav. r a r . 176. 1848.
Delima tripetala Bl. ex Spreng., Syst veg. 2: 597. 1825; G. Don, Gen. Hist, dichl.
PI. 1: 71. 1831.
Delima frangulaefolia Presl, Rel. Haenk. 2: 73. 1835—6; Vill., Nov. App. 2. 1880.
Delima aspera Blanco, Fl. Filip. 429. 1837; 2nd Ed., 299. 1845; 3d Ed., 2: 191
pi. 190** 1878.
Tetracera monocurpa Blanco, Fl. Filip. 459. 1837.
An asterisk indicates an uncoloured illustration; a double asterisk a coloured one.
1953] HoodLAND: Tetracera 191
Delima sarmentosa var. hebecarpa (DC.) Hook. f. & Thorns., F l . ind. 1: 61. 1855;
Miq., PI. Ind. bat. Suppl. 618. I860; in Ann. Mus. bot. Lugd. Bat. 4: 73. 1868; Hook. f.
& Thorns, in PL Br. Ind. 1: 31. 1872; Kurz in J. As. Soc. Beng. 43 (2) : 45. 1874; King
in J. As. Soc. Beng. 58 (2) : 362. 1899.
Delima sarmentosa var, jl Miq., Fl. Ind. bat. 1 (2) : 7. 1859 (= Delima hebecarpa
D C ) .
Tetracera sarmentosa var. hebecarpa (DC.) Martelli in Becc, Malesia 3: 150.
1886; Vid., Rev. PI. Vase. Pilip. 36. 1886; Pin. & Gagnep. « Bull. Soc. bot. F r , Mem.
4: 4. 1906; in F l . gdn. Ind.-Ch. 1: 16. 1907.
Tetracera hebecarpa (DC.) Boerl., Cat. Hort. bot. bogor. 3. 1899; Back., Fl.
Batavia 1: 4. 1907; Schoolil. J a v a 8. 1911; Koord., EjtkFl. J a v a 2: 600. 1912; Ciaib,
Fl. siam. En. 1: 19. 1925.
Tetracera xcaudens (L.) Merr., Int. Rumph. Herb. amb. 365. 1917; Brown, Minor
Prod. Philip. For. 3: 59. 1921; Merr., Bibl. En. Born. PL 382. 1921; En. Philip, fl.
PI. 3: 59. 1923; Back., Bekn. Fl. J a v a (Nooduitg.) 4 (Fam. 80): 2. 1942; Henders.,
Mai. wild Flow. 1: 19. 1949; Hoogl. in Fl. mal. I 4: 143. 1951.
Tetracera vnlubilis Auct, (non L.); Merr., Spec, blanc. 362, 1918, in error, T.
scandens being intended.
Tetracera scandens var. hebecarpa (DC.) Heyne, Nutt. PL Ned. Ind., 2nd Ed.,
1070. 1927.
Delima scandens Burk., Diet. econ. Prod. Mal. Pen. 1: 776. 1935, doubtlessly
based on Tragia scandens L., though not expressly stated.
TYPES.—Tragia scandenx: Rumphius, I.e. pi. !}. — Delima sarmentosa: lectotype
in LINN. — Delima hebecarpa: La Haye s.n., J a v a ; lectotype in G (original of
Delessert's figure, 1821). — Delima intermedia: Blume s.n., J a v a ; leetotype in L.
— Delima tripetala: J a v a ; unknown to me. — Delima fra,ngnlaefolia: Haenke s.n.,
Luzon; holotype in Herb. Prague, not seen. — Delima as]tera & Tetracera monocarpa:
Blanco s.n., Malinta, Philippines; probably lost.
Small shrub (up to 2 m high) or climbing or creeping woody vine
(up to 30m long), much-branching; trunk up to 16cm thick; branches
strigose, glabrescent, younger ones light brown, older ones with light
grey bark. Leaves oblong to obovate, (3.5—)6—15(—20) X (1.5—)3—7
(—9) cm, with (6—)10—14(—20) nerves on either side; apex rounded
to obtuse; base obtuse; margin entire to distinctly dentate, most so in
saplings; nerves slightly curving upward, ending in apex of teeth; leaves
above sometimes slightly glossy, sparsely strigose to glabrous on inter-
venium, sparsely pubescent on midrib, beneath dull, sparsely pubescent to
glabrous on intervenium, strigose on nerves and midrib, without or with
slightly to distinctly developed hairy domatia in axil of nerves, particu-
larly upper ones, scabrid on both sides; petiole (4—)6—12(—I5) mm,
sparsely pubescent above, strigose beneath. Inflorescences terminal, up
to 40 x 20 cm, up to 200-flowered, often in basal part with 1—5 small
leaves; branches strigose, extreme ones most densely so; bracts usually
caducous, lanceolate, 3 x 1 mm, acute at apex, attached with broad base,
glabrous above, strigose beneath. Flowers 6—8 mm across; pedicel 2—6
mm, rather densely strigose, without or with 1—2 bracteoles; bracteoles
1 x 0.5 mm. Sepals 4, on same plant in about 5% of flowers 5, reflexed,
about 3 x 2 mm, scabrid, slightly strigose outside, smooth, glabi'ous
R E I N W A R D T I A [VOL. 2
inside, ciliate at margin. Petals 3, 3—5 X 2—3.5 mm, white, yellowish
white, or reddish white. Stamens about 65—80, 3 mm long, white, with
thecae touching each other at apex. Carpels 1(—2), densely hairy with
rigid, appressed, 0.4—0.7 mm long hairs, ovoid, 0.75—1 X 0.5—0.75 mm
with 3 mm long style, with about 10 ovules. F r u i t s ovoid, about 1 0 x 6 mm,
acute with 1—3 mm long beak, with 0.5—2 mm long rigid, appressed
haira, reddish brown, glossy, 1(—2)-seeded. Seeds ovoid, about 4 x 3 mm,
(flossy black; aril 2—3 mm long, scarlet, fimbriate for 3/4 - 9/10 of its
length.
DISTRIBUTION.—Southern China (Yunnan), Burma (Arracan, Tenas-
serim), Peninsular Siam, Indo-China (Cambodia, Cochinchina), Nicobar
Islands, Sumatra, Malay Peninsula, Banka, Java, Borneo (only Kuching
and British North Borneo), Philippines, Celebes, Kangean Islands, and
Lesser Sunda Islands (Bali, Sumbawa, Flores).
ECOLOGY.—Common at low altitudes (up to 500 m), in Yunnan at
950 m; in thickets, scrub, hedges, secondary forests, open primary forests,
and teak-forests (Central & East Java), on moist to rather dry soil, often
on riverbanks.
VERNACULAR NAMES.—Siam: kapot (Prachuap), pet kai (Songkla),
pot (Puket), pot kai (Patalung). S u m a t r a : akar ampala (Priaman),
akar (am) pelas (Palembang), ampelas padang (Bengkalis), baik sipi hendak.
1953] HoogLiAND: Tetracera 193
hasahan (Lampung), galinggin (Asahan), ompe (Atjeh). M a l a y P e n i n -
a u l a : akar ampelas hari betina, ampelas hari (Malacca), akar ampelas
tikus (Pahang), akar ampelas puteh, ampelas tikus (Sungei Ujong State),
akar pelah (Trengganu), ampelas (Kuala Lumpur, Pahang, P e n a n g ) ,
ampelas hari betina (fide Ridley), ampelas kasap (Taiping), ampelas
putih (fide Ridley), ampelas rimau (Kuala L u m p u r ) . J a v a : (akar)
ampelas hari betina, (akar) ampelas ojod, (akar) ampelas putih, (akar)
ampelas tikus (Malay), areuj ki assahan, areuj ki assahan lalaki, kaju
as(s)ahan, ki asahan, asahan areuj (Sundanese), bo, debo, dembo, kroko,
kroko ojod, ojal, roko, rokokan, singaran (Javanese). B o r n e o : agupit,
kerubkerub (Bajau), akar ampelas (Kudat, Membakut), akar pampan
(Kinabatangan), ampelas (Bajau, Bingawa, Papan, Tawao), ampelas akar
(Bukit P a d a n g ) , kariskaris, panpan (Sungei). P h i l i p p i n e s : anggi'-
git (Tagb.), dangilian (Bag.), eses na bagin, malbastigbalang, tagbalang
(Tag.), malakatmon (Pamp., Tag.), pakiling (Sbl.). C e l e b e s : lumpiwi
apaelae (Galiraeng). K a n g e a n: ampelas. B a l i : bun api-api.
U S E S . — T h e leaves are used for polishing wood and (fide Rumphius)
metal. The stems may be used as cordage. The medical use is unimportant
(cf. Burkill, 1935).
The glabrous-fruited form, in the present paper considered to re-
present a distinct species, Tetracera, asiatiea (Lour.) Hoogl., was formerly
included in the present species.
The genus Delima was first proposed by Linnaeus in his " F l o r a
zeylanica." The species which occurs in Ceylon is not the present one, but
Tetracera asiatiea (Lour.) Hoogl. The specimen in the Linnaean her-
barium, however, is a fruiting specimen of the present species of unknown
provenance. This specimen is considered to represent the lectotype of
Delima sarmentosa L.
Rumphius' illustration is not very good. After comparison of his
description and plate with the present species it is however safe to
identify Rumphius' plant with the present species rather than with
Tetracera nordtiana F. Muell., the only species which is known from
Amboina. Tragia scandens L. antedates Delima sarmentosa L., which was
erroneously omitted from the first edition of the "Species plantarum."
2. TETRACERA ASIATICA (Lour.) Hoogl. — Fig. 2
[Fructus indicus savuieutosus Burm., Thes. zeyl. 101. 1737.]
{Delima. L., Fl. zeyl. 92. 1748.]
Scauieria asiatica Lour., Fl. cochinch. 341. 1790.
Leontogloesum ecabrum Hance in Walp., Ann. 2: 18. 1851.
Delima sarmentosa var. glabra Hook. f. & Thorns., Fl. ind. 1: 61. 1855; in Fl.
Br. Ind. 1: 31. 1872; King in J. As. Soc. Beng. 58 (2) : 362. 1889.
Detimopsis kirsuta Miq.., Fl. Ind. bat. 1 (2i : 10. 1859; ibid. Suiipl. 152, 618. 1860.
Delivia sarmentosa f. liirsutio-r Miq. hi Ann. Mus. bot. Lugd. Bat. f: 73. 1868.
Tetracera hirsuta (Miq.) Boerl., Cat. Hort. bot. bogcr. 3. 189S.
194 RE1NWARDT1A |v0L. 2
Tetracera sarmentosa var. hirsuta (Miq.) Fin. & Gagnep. lit Bull. Soc. bot. Fr.,
Mem. 4: 4. 1906.
Tetracera levinei MeiT. in Philip. J. Sci., Bot. 8: 147. 1918.
Tetracera asiatica (Lour.) Hoogl. in Fl. mal. I 4: 143. 1951.
Delima sarmentosa Auct. (nov L.); Burm. f., Fl. ind. 122 pi. 37." 1768; Gaertn.,
Fruct. Sem. PI. 2: 112 pi. 106.'" 1791; DC, Syst. 1; 407. 1818; Lara., Illustr. 3: pi.
1,75* 1823; DC, Prod. 1: 69.1824; Hook, in Curt., Bot. Mag. 58: t. Sons.*" 1831; Drury,
Handb. Ind. Fl. 1: 12. 1864; Schnizl., Iconosx 3: vl- 177." 1843-70; Kurz, For. Fl. Br.
Burma 1: 22. 1877; Trim., Handb. Fl. Ceyl. 1; 5. 1893; Gamble, List Darjeeling Distr.
Beng. 1. 1896; Man. Ind. Timb., 2nd Ed., 3. 1902; Prain, Beng. PI. 195. 1903; Brand.,
Ind. Trees 5. 1906; Parkinson, For. Fl. Andam. Isl. 72. 1923; Kanjilal, Kanjilal, & Das,
Fl. Assam 1: 10. 1934.
Tetracera sarmentosa Auct. [von (L.) Vabl]; Roxb., Fl. ind., ed. Carey, 2; 645.
1832, p.p.; Hanee in J. Linn. Soc, Bot. 8; 99. 1873; Forb. & Hemsl. in J. Linn. Soc,
Bot. 23; 22. 1886; Fin. & Gagnep. in Bull. Soc. bot. Fr., Mem. 4: 3. 1906, p.p.; in Fl.
gen. Ind.-Ch. 1; 15. 1907, p.p.; Crev. & Petel. in Bull. econ. Ind.-Ch. II 32: 19. 1929;
Cowan & Cowan, Trees N. Bens. 8. 1929; Crook, Fl. PI. Hongkong, Ran.—Meliac.
22." 1930; Gagnep. in Fl. gen. Ind.-Ch., Suppl. 1; 20. 1938.
Tetracera scandens Auct. [non (L.| Merr.]; Merr. in Lingn. Sci. J. 5: 128.
1927; Alston in Trim., Handb. Fl. Ceyl. 8 (Suppl.) : 3. 1931; Merr. in Trans. Am. phil.
Soc. II 24 (2): 264. 1935; Masamune, Fl. kainant. 205. 1943.
TYPES.—Seguieria asiatica: d'Alleizette s.»., Tonkin, near Jen Hay, April 1908; '
neotype in L. — Leontoglossum scabnim: Seemann 2461, China; holotype in BM.
— Delima sarmentosa var. glabra: Wallich 6632, Sylhet, February 1829; lectotype in
K, isotypes in BM, CAL, CGE, G. — Delimopsis kirsuta: "Davilla hirsuta T.'& B.";
holotype in U, isotype in L; probably from cultivation in the Botanic Garden, Bogor,
or from the Lampung, Sumatra. — Delima sarmentosa I. hirsntior: is based on the
preceding. — Tetracera levinei: Levine 1794, Canton & vicinity, October 22, 1927;
holotype in A, isotypes in E, GH, MO.
Small shrub (up to 3 m high) or climbing or creeping liana (up to 6 m
long), much-branching; branches scabrid, sparsely strigose with 1 mm
long hairs with or without sparsely to profusely distributed, small, diver-
gent tufts of 3—12, shorter (0.3—0.5 mm) hairs or hirsute to sparsely
so with 1 mm long hairs and also profusely distributed, small, divergent
tufts of 3—12, shorter hairs, glabrescent, older branches smooth, brown
or reddish brown. Leaves oblong, (3—)5—11(—15) x (1.5—)2—5(—7.5)
cm, with (5—)10—14(—20) nerves on either side; apex obtuse to acute,
sometimes rounded; base acute; margin slightly emarginate at end of
nerves or entire to distinctly dentate; nerves ending mueronately in
margin, or in apex of teeth where present; leaves above deep lustrous
green, more or less shining, sparsely strigose with 1 mm long hairs on
intervenium, glabrous to hirsute with up to 2 mm long hairs, particularly
in basal part, on nerves and midrib, beneath light green, glabrous to
sparsely stellate-hairy with some solitary hairs on intervenium, sparsely
strigose with or without rather sparsely distributed tufts of 3—8, short
hairs, or hirsute with 1.5—2.5 mm long hairs as well as rather sparsely
distributed tufts of short haira on nerves and midrib; petiole 5—10(—15)
mm, strigose or hirsute and with tufts of shorter hairs beneath, glabrous
1953] HOOOLAND: Tetracera 195
to hirsute above. Inflorescences terminal, 10—25 x 5—15 cm, 30—150-
flowered, often with 1—4 small leaves in basal part; branches scabrid,
sparsely strigose or strigose with (sometimes profusely distributed) tufts
of smaller hairs, or hirsute with tufts of small hairs; bracts usually
caducous, lanceolate, 3 x 1 mm, acuminate at apex. Flowers 7—10 mm
across; pedicel 1—5 mm, the indument like that of extreme branches
of inflorescences, without or with 1—2 bracteoles; braeteoles 1 x 0.5
mm. Sepals 5, reflexed, 2 outermost ones about 2 x 1.5 mm, 3 inner ones
about 4 x 3 mm, scabrid, sparsely hirsute with 0.2—0.4 mm long hairs
outside, smooth, glabrous inside. Petals 3, 3—4 X 2—3 mm, white,
yellowish white, or greenish white. Stamens about 100—125, 3—4 mm
long, whitish yellow, the thecae slightly separated at apex. Carpels
1(—2), glabrous, 0.75—1 x 0.5—0.75 mm with 0.3 mm long style, with
about 10—12 ovules. Fruits ovoid, 6—10 x 4—6 mm, acute with 2—5
mm long beak, glabrous, purplish, shining, 1(—2)-seeded. Seeds ovoid,
4 x 3mm, glossy black; aril up to 5mm long, fimbriate for 1/2—2/3 of
its length with rather broad fringes.
Subsp. ASIATICA.
Segitieria asiatica Lour. — Leontoglossnm scabrum Hance. — Tetracera levinci
,Merr. — Tetracera asiatica (Lour.) Hoogl.
Branches sparsely strigose with 1 mm long hairs. Leaves glabrous
on intervenium, sparsely strigose on nerves and midrib beneath; petiole
strigose beneath. Branches of inflorescences sparsely strigose.
Subsp. andamanica Hoogl., subsp. nov.
Tetracera asiatica subsp. andamanica Hoogl. in Fl. mal. I 4: 144. 1951, cum
desev. angl.
Delima sarmcntosa var. fflabra Hook. f. & Thorns., p.p., lectotypo incluso.
TYPE.—Dr. King's Collector 337, Port Bhing, Andamans, July (5, 1884; holotype
in L, isotypes in CAL, G, K.
Ramis novellis et ramis inflorescentiarum strigosis et pilis stellato-
fasciculatis; foliis nervis facie inferiore strigosis, intervenio glabris.
Branches sparsely strigose together with sparsely to profusely
distributed small divergent tufts of about 3—12, shorter (0.3—0.5 mm
long) hairs. Leaves glabrous on intervenium, sparsely strigose on nerves
and midrib beneath; petiole strigose beneath. Branches of inflorescences
sparsely strigose to strigose, with also on thicker branches, very sparsely
distributed, and on extreme branches, up to densely distributed tufts
of smaller hairs.
Subsp. SUMATRANA Hoogl.
Tetracera asiatica subsp. sumatrana Hoogl. in Fl. mal. I 1: 144. 1951.
Delimopsis hirsnta Miq. — Delima sarnientoea f. hirsutior Miq. — Tetracera
hirsuta (Miq.) Boerl. — Tetracera sarmentosa var. hirsata (Miq.) Fin. & Gagnep.,
quoad typ.
TVPE.—Same as for Delimopsis hirsuta Miq.
190 R E I N W A R D T 1 A [VOL. 2
Branches hirsute to sparsely so, with in addition profusely distri-
buted, small, divergent tufts of about 3—12, shorter hairs. Leaves with
sparsely distributed tufts of 3—8, divergent hairs of 0.3—0.5 mm, as
well as with a smaller number of identical, but solitary hairs on inter-
venium, on nerves and midrib beneath hirsute with 1.5—2.5 mm long
hairs as well as (particularly on the sides) rather sparsely distributed
tufts of hairs like those on intervenium; petiole hirsute beneath, together
with tufts of shorter hairs like those on midrib, hirsute above. Branches
of inflorescences hirsute with 1.5—2.5 mm long hairs as well as tufts of
smaller hairs, the latter on extreme branches up to densely distributed.
Subsp. zeylanica Hoogl., subsp. nov.
[Fructus indicus sarmentosus Burm.] — [Delima L.]
TYPE.—Anonymus G9, Ceylon; holotype in L.
Ramis novellis et rwtnis inflorescentiarum strigosis et pilis stelluto-
fascicvlatis; foliis nervis facie inferiore strigosis et pilis stellato-fascicu-
latis, intervenio facie inferiore pilis stellato-fasciculatis.
Branches sparsely strigose as well as with sparsely to profusely
distributed, small, divergent tufts of about 3—12, shorter (0.3—0.5 mm
long) hairs. Leaves with sparsely distributed tufts of 3—8 divergent
hairs of 0.3—0.5 mm, as well as with smaller number of identical, but
solitary hairs on intervenium, sparsely strigose with about 1 mm long
hairs, particularly on central part, as well as rather sparsely distributed,
small tufts like those on intervenium, particularly on sides, on nerves
and midrib, beneath. Branches of inflorescences sparsely strigose, with
also rather densely to densely distributed tufts of smaller hairs.
DISTRIBUTION.—Ceylon and from Assam and South China to Borneo
and South Sumatra. Subspecies asiatica in South China (Kwangsi,
Kwangtung, and Hainan), Indo-China, and Siam; subspecies andamanica
in Darjeeling, Assam, Bengal, Andaman Islands, South Burma (Tenas-
serim), and Malay Peninsula (Perak, Selangor); subspecies sumatrana
in Sumatra, the Malay Peninsula, and Borneo; subspecies zeylanica in
Ceylon. The subspecies are geographically isolated except subspecies
andamanica and subspecies sumatrana, which are both found in the
Malay Peninsula.
ECOLOGY.—Subspecies asiatica along roadsides, in scrub, hedges,
thickets, open forests, and, rarely, in woods; generally at low altitudes,
up to 1500 m (Cambodia) ; flowering from March to November, fruiting
from May to February. Subspecies andamanica in evergreen forests
(Assam) and "hill jungle" (Andamans), up to 650m altitude; flowering
from May to October, fruiting from June to February. Subspecies
sumatrana in primary forest, up to 1300 m (Atjeh). Subspecies zeylanica
rare in woods at low altitude; flowering in August and September.
VERNACULAR NAMES.—I n d i a: ou-lota, oua-lota, panilewa (Ass.),
bau-taruk (Daff.) , samphot-rikang (Mik.) , aithlang shrui (Kuki) , hruisen
(Tipp.) , tiegdi-douka (Cach.) (fide Kanjilal, Kanjilal, & Das). C h i n a :
sahan yau ma (Cantonese), shap ip t'ang, sut t'ang (Hainan). I n d o -
C h i n a : cay chay chiu (Annamese: Vinh), cay giay chiu (Annamese: Ba
1953] HOOGLAND: Tctracera
ngoi), chong kho (Tonkin, Yen Vuc), dak kuon (fide Gagn'epain), day chiu
(Annamese: Quang T r i and N h a t r a n g ) , day sanh (Mg, Rhanh Hoa), gay-
sac, gay-trieu (Tonkin), giay chieu (Annamese: Bien Hoa and Vinh Yen),
giay chiu (Annamese: Hue and Thua Thien), sang at (Moi, Quang T r i ) ,
vie chieu (Annamese: Tourane). C e y l o n : corose-wel, korasawel, korese
wel, korosse (Singhalese). M a l a y P e n i n s u l a : memplas rimau (Ke-
maman ; noted under subspecies sumatrana).
PIG. 2. Tetrucet-a aBialica- (Lour.) Hoogl.: ssp. aeiatica ( •) , ssp.
andamavica Hoogl. ( + ) , ssp. mntatrava Hoogl. ( x ) , ssp.
zeylctniea Hoogl. ( o ) .
USES.—The stems of subspecies asiatica are used for binding pur-
poses in Indo-China, its leaves are used for cleaning tin-ware in China.
Subspecies sumatrana is used in the Malay Peninsula for polishing wood.
I have preferred to give the infraspecific taxa the rank of subspecies
rather than of varieties for the following reasons: (i) the marked geo-
graphical distribution of each, centering in the Malay Peninsula, where
three of the subspecies occur; (ii) the absence of intermediate forms; (iii)
in addition to the differences in the composition of the indument the
subspecies show a markedly different habit of the leaves.
198 R E I N W A R D T I A [VOL. 2
Merrill (1935, I.e.), in his discussion of the binomials proposed by
Loureiro, gives under Tetracera scandens (from which I have separated
the present species) the following of Loureiro's names as synonyms:
Actaea aspera, Calligonum asperum, and Seguieria asiatica. Of these
Actaea aspera and Cailigonum asperum are excluded here on account of
the fruit, said to be a berry in both of them. For Cailigonum asperum,
Loureiro expressly stated the difference from Seguieria asiatica and
Delima sarmentosa Burm. f. (Fl. ind.) : "Minime vero ad istas aceedit
ratione fructus"; for Seguieria asiatica the fruit is indicated to be a
capsule. The axillary flowers in spicate racemes and the quadrifid calyx
and corolla in Actaea aspera, the hispid fruit and deeply bipartite stigma
in Cailigonum asperum can only give more support to my opinion. The
only point likely to raise doubt as to Seguieria asiatica is the absence of
a corolla, which is, however, considering the caducous corolla of the
Dilleniaceae, not a point to which great value can be attached.
The species is here separated from Tetracera scandens (L.) Merr.
(formerly usually known as Delima, or Tetracera, sarmentosa), in which
it has been currently included, on account of the following distinctive
characters: (i) its glabrous carpel and fruit (T. scandens, hirsute) and (ii)
its pentamerous calyx (T. scandens, tetramerous or in a few flowers on
the same plant pentamerous). Tetracera scandens has the centre of its
distribution in Malaysia, T. asiatica in continental Asia.
From Borneo the species is known only from sterile specimens,
having slightly larger leaves than the specimens of the same subspecies
from Sumatra. The specimens from Korthals are provided with the
manuscript name "Tetracera setigera Khs," mentioned by Miquel {in Ann.
Mus. bot. Lugd. Bat. 4: 75. 1868) under T. scaberrima Miq.
3. TETRACERA GLABERRIMA Martelli
Tetracera glaberrima Martelli m Becc, Malesia 3: 150. 1886; Boerl., Cat. Hort.
bot. bogor. 3. 1899; Merr., Bibl. En. Born. PI. 381. 1921; Hoogl. in Fl. mal. I 4: 148.
1951.
TYPE.—Beccari Piante Bornensi 298, Kuching, Sarawak, August 1865; bolotype
in F l , isotypes in BM, G, K, M, NY, S.
Scandent shrub; branches glabrous, younger ones brown, older ones
greyish white. Leaves elliptic to obovate, (2.5—)5—9(—15) X (1.5—)2—4
(—0) cm, with (8—)5—7(—9) nerves on either side; apex distinctly
acuminate; base acute; margin entire; nerves slightly curving upward,
ending in margin mucronately; leaves glossy, glabrous above, less glossy,
glabrous on intervenium, sparsely strigose along midrib and nerves
beneath, smooth on both sides; petiole 3—8 mm, glabrous. Inflorescences
axillary, up to 4 x 2.5 cm, 1—-5-, usually 3-flowered; peduncle 0.5—1 cm,
1953] HOOGLAND: Tetracera 199
like branches sparsely strigose, without bracts. Flowers about 2.5 cm
across; pedicel 7—12 mm, sparsely strigose to glabrous, without or with
1—2 minute bracteoles. Sepals 4, 8—10 x 5—7 mm, glabrous outside,
glabrous to sparsely sericeous inside, not ciliate at margin. Petals 4,
10—12 x 6—8 mm, white. Stamens about 100, 5 mm long, the thecae touch-
ing each other at apex. Carpels 1(—2), glabrous, 1.5 x 1 mm with 0.5—4
mm long style, with about 10 ovules. Fruits with 1(—2) capsules devel-
oped in each flower; capsules ovoid, 15 X 10mm, acute with 2—3mm
long, slightly lateral beak, glabrous, glossy, 1(—3)-seeded. Seeds ovoid,
glossy black, 3—4 x 2—3 mm; aril 5 mm long, laciniate with broad slips
to about 1/2 of its length.
DISTRIBUTION.—Borneo (Kuching), once collected; formerly cultivat-
ed in the Botanic Garden of Bogor from unknown provenance.
VERNACULAR NAME.—In the Botanic Garden, Bogor, the Sundanese
name aroy kiasahan was noted.
The species is closely related to T. akara (Burm. f.) Merr. The charac-
ters in common are the sericeous inner side of the sepals, the structure
of the inflorescences, the tetramerous calyx, and the large flowers; the
differences are the less dense sericeous indument on the inner side of the
sepals, the slightly broader leaves, and the solitary carpels.
4. TETRACERA MAINGAYI Hoogl.
Delimit laeuis Maing. IT King in J. As. Soc. Beng. 5S (2) : 362. 1889, nun Tetracera
hevis Vahl.
Tetracera muingayi Hocgl. in Fl. mal. I 4: 144. 1951 (= Delima laevis Maing.
ex King).
Tetracera borncenaw Auct. (noil Miq.l; Kidl., Fl. Mal. Pen. 1 : 8. 1922.
Tetracera. sumatrana Auct. (non Miq.l ; Ridl., Fl. Mal. Pen. 1: <>. 1922, p.p.
T Y P E — M a i n g a y 1570 (Kew Distribution 10), Malacca, April 10, 1867; holotype
in CAL, isotype in K.
Scandent shrub; branches strigose-hirsute with solitary hairs and
hairs in divergent tufts of 2—5 each, glabrescent, younger ones scabrid,
later smooth. Leaves oblong, (3.5—)7.5—15(—20) x (1.7—)3—6(—9) cm,
with 8—11 nerves on either side; apex acute, somewhat acuminate;
base rounded to obtuse; margin entire; nerves curving upward, not reach-
ing margin; leaves shining, glabrous or sparsely strigose only on base
of midrib above, dull, hirsute with divergent tufts of 2—5, 0.2—0.4 mm
long hairs to glabrous on intervenium, rather densely strigose-hirsute
with 0.75—1.5 mm long hairs on midrib and nerves beneath, smooth on
both sides; petiole 10—20 mm, slightly winged, 2—3 mm broad, glabrous
to sparsely strigose along the middle above, glabrous to strigose-hirsute
. beneath. Inflorescences terminal, 15—20 X 10—20 cm, up to 250-flower-
ed, often with 1—3 small leaves in basal part; branches strigose-hirsute
like younger sterile branches, slightly scabrid; bracts caducous, lance-
olate, 3—6 X 1—2 mm, acute at apex, attached with broad base, glabrous
above, strigose beneath. Flowers 12—15 mm across; pedicel 1.5—4mm,
strigose-hirsute like branches of inflorescences, without bracteoles. Sepals
2 0 0 R E I N W A R D T I A [VOL. 2
5, 2 outermost ones 3.5 x 3 mm, 3 inner ones 5 X 3.5—4.5 mm, all scahrid,
2 outermost ones sparsely strigose in central part, 3 inner ones completely
glabrous outside, all with smooth inner surface, densely sericeous except
for 0.5—1 mm broad glabrous margin. Petals 3, about 7 x 4 mm. Stamens
about 110, 2.5 mm long, with thecae distinctly separated at apex, con-
nective not emarginate between thecae. Carpels 1, glabrous, 1.8 x 1.4 mm,
gradually tapering into 1.2 mm long style, with about 6 ovules. F r u i t s
oblong, 8—12 X 3—4 mm, acute with 2—3 mm long beak, glabrous, dull,
1—2-seeded. Seeds unknown to me.
DISTRIBUTION.—Malay Peninsula (Penang, Malacca, Selangor), ?Borneo.
ECOLOGY.—Climber in low altitude (up to 200 m) forests.
VERNACULAR NAMES.—M a l a y P e n i n s u l a : akar mempelas (Se-
langor), akar mempelas betina (Alvins 1066, without locality). B o r n e o :
a k a r amplas.
The species is more closely related to Tetracera fagifolia Bl. than to
the other species with single carpels.
The Borneo record is uncertain. The only specimen bears the indica-
tions : "Borneo, no 1703, Remow, no information" (SING).
5. TETRACERA LANUGINOSA Diels
Tetracera laimginosa Diels in Bot. Jb. 57: 439. 1922.
TYPE.—Ledermann 8586, April River, Sepik Region, New Guinea, September 8.
1912; leetotype (isotype) in BM, isotype in K.
Scandent s h r u b ; branches hirsute with single, up to 2 mm long and
stellately grouped, about 0.1 mm long hairs, slightly scabrid. Leaves
elliptic, 5—6 x 3—3.8 cm, with 9—12 nerves on either side; apex and
base rounded; margin e n t i r e ; nerves curving upward, ending close to or in
m a r g i n ; leaves slightly shining, r a t h e r sparsely hirsute with thin, r a t h e r
rigid, up to 2 mm long hairs above, the under surface dull, sligthly more
densely hirsute t h a n above, with stellate groups of about 0.1 mm long hairs,
mainly on n e r v e s ; petiole 10—15 mm, long-hirsute with single hairs above,
hirsute with long and stellate groups of very short hairs beneath. In-
florescences axillary, 2—4-flowered; branches hirsute like younger sterile
branches; bracts lanceolate, about 4 x 2 mm, glabrous above, hirsute
beneath with long and stellate groups of small hairs. Flowers about 18 mm
across (or more?). Sepals 5, 2 outermost ones 4 x 4 mm, 3 innermost ones
6 x 7 mm, lanuginose outside with in addition stellate groups of small
hairs, glabrous to very sparsely and shortly strigose inside. Petals 3.
9 x 6 mm, on outer side on central p a r t with stellate groups of hairs as
well as a few lanuginose hairs. Stamens about 500, 2.5—3 mm long, the
thecae strongly separated at apex. Carpels 2—3, densely covered, mainly
in basal part, with 2 mm long, r a t h e r thin, ferrugineous hairs, each carpel
1.5 x 1 mm with 2 mm long style. F r u i t s unknown.
DISTRIBUTION.—New Guinea (April River, Sepik region), only knowr.
from the type collection.
ECOLOGY.—In primary forest, at 50—100 m altitude.
1863] HOOGLABD: Tetracera 201
In the original description Diels indicates that the number of sepals
is seven to eight. The only flowerbud I studied had five sepals and three
petals, the latter not much differing from the first. The fact that the
petals, which are somewhat larger than the innermost sepals, are slightly
hairy outside may have confused Diels. In some other species, e.g. T. fagi-
folia Bl., the differences between the innermost sepals and the petals also
hardly exist, except that there the indument on the inner and outer side
is found only in the sepals.
6. TETRACERA NORDTIANA F. Muell. — Fig. 3
Tetracera nordtiana F. Muell., Fragm. 5: 1. 1865; F. M. Bail., Synops. Queensl.
PI, 3. 1883; K. Schum. & Hollr., Fl, Kais. Wilhelmal. 47. 1889; F. M. Bail., Queensl. Fl.
1: 9. 1899; K. Schum. & Laut., Fl. deut. Sehutzgeb, Siidsee 444. 1901; F. M. Bail.,
Compr. Cat. Queensl. PI. 18 f. 3." 1909; Hoogl. in Fl. mal. 1 4 : 144. 1951.
Tetracera wuthiana F. Muell., Fragm. 10: 49. 1876; F. M. Bail., Synops. Queensl.
Fl. 3. 1883; Queensl. Fl. 1: 10. 1899; Compr. Cat. Queensl. PI. 18. 1909; in Dept Agric.
Brisb. Bot. Bull. 18: 3. 1916.
Tetracera everillii F. Muell., Descr. Notes Papuan PI. 7: 25. 1886.
Tetracera moluccana Martelli HI Becc, Malesia 3: 153. 1886; Kaneh. & Hatus.
in Bot. Mag., Tokyo 57: 63. 1943.
Tetracera coivleyana F. M. Bail, in Dept Agric. Brisb. Bot. Bull. 5: 7. 1892;
Queensl. Fl. 1: 9. 1899; Compr. Cat. Queensl. Fl. 18 /. Sbis.' 1909.
Tetracera boerhgei Merr., Int. Rumph. Herb, ami), 366. 1917.
Tetracera floribunda Diels in Bot. Jb. 57: 440. 1922.
Tetracera pilophylla Diels in Bot. Jb. 57: 440. 1922.
Tetracera euryandra Auct. (noM Vahl) ; Eoxb., Fl. ind., ed. Carey, 2: 646. 1832.
Tetracera volubilis Auct. (no« L.); Kendle in J. of Bot. 59 (Suppl.) : 2. 1923.
TYPES.—Tetracera nordtiana: Dallachy s.n., Rockinghams Bay, Meunga Creek,
January 4, 1846; lectotype in MEL, isotype in BM. — Tetracera wuthiana: Dallachy
s.n., Rockingham's Bay, November 21, 1865; lectotype in MEL, istotypes in BM, BRI,
K. — Tetracera everillii: Bauerlen 472, Fly River (Branch), October 1885; holotype
in MEL, isotype in BRI. — Tetracera inohiccana: Beceari s.n., Piante delle Molucche,
Amboina, 1873; holotype in F l . — Tetracera cowleyana: Cowley s.n., Cairns, Queens-
land (Cook Distr.); holotype in BRI, isotype in MEL. — Tetracera boerlagei: Ro-
binson Plantae Rumphianae amboinenses 485, Amboina, July—November 1913;
isotypes in BO, K, L. — Tetracera floribunda: Ledermann 10723a, Malu, Sepik Region,
January 1913; probably lost. — Tetracera pilophylla: Ledermann 8937, Etappenberg,
Sepik Region, October 2, 1912; isotypes in K, SING.
Shrub or large climber (up to 10 m long) ; branches often slightly
scabrid, strigose to hirsute, sometimes together with stellate groups of
very short hairs, glabrescent, older branches smooth. Leaves elliptic or
ovate to oblong or lanceolate, (3—) 5—10(—15) X (2—) 3—5(—7) cm, with
(6—)12—16(—24) nerves on either side; apex and base rounded to acute,
margin entire to distinctly dentate; nerves curving upward, ending in
margin, or in apex of teeth where present; leaves above glabrous or hirsute
with single hairs or shortly hirsute with stellate groups of hairs together
with longer solitary hairs on intervenium, nerves, and midrib, beneath
2 0 2 R E I N W A R D T 1 A [VOL. 2
glabrous or hirsute with solitary hairs or shortly hirsute with stellate
groups of hairs, together with longer solitary hairs on intervenium, sparse-
ly strigose to strigose or strigose-hirsute or hirsute on nerves and midrib;
petiole 5—15 ram, as to indument like leaves. Inflorescences terminal,
often on short axillary few-leaved branches, 5—15(—30) x 3—6(—15)
cm, 15—50-flowered; branches as to indument like younger sterile
branches; bracts partly caducous, lanceolate, 2 x 0.5 mm, acute at apex.
Flowers 6—10 mm across; pedicel 0.5—2 mm, as to indument like extreme
branches of inflorescences. Sepals 4—5, reflexed, 2 outermost ones 1.5—2
x 1.5—2 mm, 2—3 inner ones 3—4.5 X 2—3.5 mm, densely and shortly
stellate-hairy or strigose or densely villous outside, glabrous or, rarely,
slightly sericeous in central part inside, ciliate at margin. Petals 3, 5—6
X 3—4 mm, white. Stamens about 140, 4—5 mm long; thecae strongly
separated at apex; connective slightly to deeply emarginate between thecae.
Carpels 2—4, usually 3, densely covered with rather rigid, about 0.5 mm
long hairs, ovoid, 2 X 1.5 mm with 1—2 mm long style, with about 10
ovules. Fruits with 2—3(—4) capsules developed in each flower; capsules
ovoid, 5—8 X 3—5 mm, rounded to acute with ]—2 mm long beak at. apex,
rather rigidly hairy, slightly glossy, usually 1-seeded. Seeds ovoid, about
3 X 2.5 mm, glossy black; aril about 5 mm long, red or crimson, laciniate
at margin to 2/3i—4/5, of its length with rather broad slips.
Var. NORDTIANA.
Tctracera nordtiana F. Muell. var. nordthina; Hoogl. in Fl. mal. I 4: 145. 1951.
Tetraccra nordtiatia F. Muell. — Tctracera pilophylla Diels.
Younger branches strigose or strigose-hirsute with up to 1 mm long
hairs together with stellate groups of much shorter, 0.1—0.2 mm long
hairs, sparsely to profusely distributed (up to nearly covering whole
surface). Leaves sparsely to rather densely covered with stellate groups
of 8—15 very short, 0.1—0.2 mm long hairs (in transitional 'forms to
variety molnccana groups of 2—6 hairs of 0.2—0.5 mm) together with
0.4—0.8 mm long single hairs above, intervenium of lower surface as
upper, on nerves strigose with up to 1 mm long hairs. Sepals densely cov-
ered with stellate groups of very short hairs accompagnied by fewer
solitary 0.2—0.5 mm long hairs outside, glabrous inside. Capsules rather
small, about 5 x 3 mm.
Var. WUTHIANA (F. Muell.) Hoogl.
Tetracera nordtiana var. wutkiana (F. Muell.) Hoogl. in Fl. mal. I 4: 145. 1951.
Tetraccra wuthiana F. Muell.
Younger branches strigose with up to 1.5 mm long hairs. Leaves
glabrous above, beneath glabrous on intervenium, sparsely strigose on
nerves and midrib. Sepals strigose with 0.2—0.5 mm long h a i r s outside,
glabrous inside. Capsules r a t h e r small, about 5 x 3 mm.
Var. louisiadica Hoogl., var. nov.
Tetracera nordtiana var. louisiadica Hoogl. in Fl. mal. I 4: 145. 1951, cum descr.
angl
1953] HOOGLAND: Tetruceru 203
TYPE.—MacGregor s.n., Joannet Island, Louisiades, 1888; holotype in MEL,
isotype in L.
Ramis inflorescentiarum et ramis novellis hirsutis et villosis; sepalis
intus glabris, extus villosis; foliis facie superiore intervenio sparsim hir-
sutis, nervis hirsutis, facie inferiore intervenio villosis, nervis strigosis.
Younger branches strigose-hirsute with up to 0.8 mm long hairs and
a rather dense, closely appressed indument of shorter, villose hairs. Leaves
above hirsute with up to 0.5 mm long hairs, moat densely so on nerves and
midrib, beneath densely shortly villose on intervenium, strigose-hirsute
on nerves and midrib. Sepals densely villose outside, glabrous inside.
Capsules rather small, about 5 x 3 mm.
Var. MO1UCCANA (Martelli) Hoogl.
Tetracera nordtiana var. mohiccaim (Martelli) Hoogl, in Fl, mal. I 4: 145. 1951.
Tetracera moluccana Martelli. — Tetraceru cowleyana F. II. Bail. — Tetracera
boerlagei Merr. — fTetracera floribuvda Diels.
Younger branches strigose to hirsute with up to 1.5 mm long hairs.
Leaves sparsely hirsute with rather rigid, up to 1.5 mm long hairs on
intervenium, slightly more densely so on nerves and midrib above, beneath
glabrous to hirsute with up to 1.2 mm long hairs on intervenium, strigose
to hirsute with up to 1.5 mm long hairs on nerves and midrib. Sepals
strigose with up to 0.7 mm long hairs, solitary or in groups of 2—5, out-
side, glabrous inside. Capsules rather large, about 8 x 5 mm.
Var. EVERILLII (F. Muell.) Hoogl.
Tetracera nordtiana var. crcrillii (F. Muell.) Hoogl. in Fl. mal. I 4: 145. 1951.
Tetracera everillii F. Muell.
Younger branches hirsute with up to 2 mm long hairs as well as a
closely appressed indument of shorter, villose hairs. Leaves hirsute with
up to 3 mm long hairs on both sides, slightly more densely so on midrib
beneath. Sepals densely villose outside, glabrous inside. Capsules rather
large, about 8 x 5 mm.
Var. celebica Hoogl., var. nov.
Tetracera nordtiana var. celebica Hoogl. in Fl. mal. 1 4: 145. 1951, cum deser. angl.
TYPE.—Elbert 3S82, Kabaena Island, SE Celebes, October 26, 1909; holotype in L.
Ramis inflorescentiarum et ramis novellis dense hirsutis; sepalis intus
partim sparsim senceis, extus strigoso-hirsutis; foliis hirsutis ambis
faciebus.
Younger branches rather densely hirsute with up to 0.7 mm long
hairs. Leaves hirsute with up to 1 mm long hairs on both sides, most
densely so on midrib beneath. Sepals strigose-hirsute with 0.1—0.3 mm
long hairs, partly in groups of 2—5, outside, slightly sericeous in central
part inside. Capsules unknown.
DISTRIBUTION.—From south-eastern Celebes (Kabaena Island) east-
ward to the Louisiades and. north-eastern Queensland; variety nordtiana
204 R E I N W A R D T I A [VOL. 2
in New Guinea and Queensland; variety moluccana in the Moluccas, Am
Islands, New Guinea, and Queensland; variety wuthiana in Queensland
and a transitional form to variety nordtimm in SE New Guinea (Koitaki);
variety everillii in SE New Guinea (Fly River); variety celebica in
Kabaena Island (SE Celebes); variety louisiadica in the Louisiades. A
single sterile collection is known from New Britain (Kew Herbarium) ;
it probably belongs to variety moluccana.
FIG. 3. Tetracera nordtiana F. Muell.: var. n o r d t i a n a (•) , vai. wutniana
(F. Muell.) Hoogl. (X), var. everillii (F. Muell.) Hoogl. ( + ), var. mo-
luccana (Martelli) Hoogl, (c), var. celebica Hoogl. (•) , var. louisiadica
Hoogl. (T .
ECOLOGY.—Climber in rain forest, from sea-level to about 1000 in
altitude.
VERNACULAR NAMES.—A m b o i n a: talir hassat, hassat, and hassat
cotel. Probably also gumi uccu in T e r n a t e (fide Rumphius).
The species is remarkably polymorphic, but the varieties as distin-
guished here mainly on the character of the indument do not appear to
deserve higher taxonomic rank, many intermediate forms occurring. Most
typical variety nordtiana has an indument of stellate groups of very
short hairs on intervenium and young branches; this typical form I have
seen only from Australia. Most of the New Guinea collections which I
refer to this variety have the hairs in the stellate groups longer and in
smaller groups, and often mixed. with single hairs as in variety moluccana.
Tetracera pilophylla, of which I have seen only an infructescence (and
no leaves), may come nearest to the most typical variety nordtiana. Carr
12832 (Koitaki, Territory of Papua) represents an intermediate form
between variety nordtiana and variety wuthiana. In variety celebica an
1963] HOOOLAND: Tetracera 205
additional character is found in the indument on the inner side of the
sepals. Though this character is often of specific value in Tetracera, I
prefer to keep the specimen within T. nordtiana in view of the great
variability of the species, in which variety celebica is inserted easily, and
the imperfect condition in which it is known.
Variety nordtiana and variety wuthiana have in general smaller,
elliptic, variety moluccana larger, oblong, leaves. Variety louisiadica in
this respect is nearest to variety nordtiana, but is only known from a
single collection; variety everittii and variety celebica, both also known only
from single collections, are nearest to variety moluccana. However, this
character of the leaf is not constant and, as in the case of the indument,
many intermediates occur.
The varietal status of T. floribunda Diels cannot be decided with
certainty from the description.
Funis urens glabra Rumph. (Herb. amb. 5: 12. 1747) may be this
species as has been suggested by Merrill. The description, however, is
very defective.
7. TETRACERA INDICA (Houtt. ex Christm. & Panz.) Merr. — Plate 1, Fig. 4
Assa Houtt., Nat. Hist. 5: 275 pi. 21 f. 1." 1776.
Asm indica Houtt. ex Christm. & Panz,, PflSyst. 4: 40, pi. 26 f. 1." 1779.
Wahlbomia in&iea Thunb. in Vet. Akad. Handl., Stockh. 216 pi. 9* 1790; Lam..
Illustr. 3: pi 485.* 182S.
Assa exotica Gmel., Syst. 839. 1791.
Tetracera laevis Vahl, Symb. bot. 3: 71. 1794.
Tetracera assa D C , Syst. 1; 402. 1818; Prod. 1: 68. 1824; Hook. f. & Thorns.,
PI. ind. 1: 63. 1855; Miq., Fl. Ind. bat. 1 (2): 8. 1859; Drury, Handb. Ind. Fl. 1: 9.
1864; Miq. in Ann. Mus. bot. Lugd. Eat 4: 74. 1868; Hook, f. & Thorns, in Fl. Br. Ind.
1: 31. 1872; Kurz in J. As. Soc. Beng. 43 (2): 45. 1874; For. Fl. Br. Burma 1: 22.
1877; King in J. As. Soc. Beng. 58 (2): 362. 1889; Brand., Ind. Trees 5. 1906; Fin. &
Gagnep. in Bull. Soc. bot. Fr., Mem. 4: 3. 1906; in Fl. gen. Ind.-Ch. 1: 14.* 1907; Back.,
Fl. Batavia 1; 3. 1907; Schoolfl. Java 9. 1911; Koord., ExkFl. Java 2: 600. 1912; Ridl.,
Fl. Mai. Pen. 1: 5. 1922; Craib, Fl. siam. En. 1: 19. 1925; Gagnep. in Fl. gen. Ind.-Ch.,
Suppl. 1: 18. 1938.
Tetracera wahlbomia D C , Syst. 1; 403. 1818; Prod, t: 68. 1824.
Tetracera malabarica Lam., Illustr. 3: 32 pi. isr, f. 1." 1823.
Tetracera dichotoma BL, BJjdr. 1; 4. 1825.
Tetracera gracilis Bl., Bijdr. 1: 4. 1825; Miq., Fl. Ind. bat, 1 (2): 9. 1859.
Tetracera trigyna Roxb., Fl. ind., ed. Carey, 2: 645. 1832; Hunter (ed. by Ridl.)
in J. Str. Br. R. A. S. 53: 98. 1909.
Eleiastis laevis (Vahl) Eafin., Sylva tellur. 165. 1838.
Tetracera indica (Houtt. ex Christm. & Panz.) Merr., Int. Rumph. Herb. amb.
367. 1917; Back. & Sloot., Theeonkr. no. 174.* 1924; Heyne, Nutt. PL Ned. Ind.,
2nd Ed., 1070. 1927; Burk., Diet. econ. Prod. Mai. Pen. 2143. 1935; Back., Bekn. Fl.
206 R E I N W A H D T I A [VOL. 2
Java (Nooduitg.) 4 (Fam. 80) : 3. 1942; Henders., Mai. wild Flow. 1: 20 /. s.* 1949;
Hoogl. in Fl. mal. I 4: 146 /. 1.' 1951.
Assa tetragynia Houtt. ex Stapf, Ind. lond. 1: 331. 1929; misinterpretation ol'
text of Houttuyn, 1776, I.e.
Eugenia malaccensis Auet. (non L.); Bm-m. f., Fl. ind. 114. 1768; cf. Steenis.
in Bull. bot. Gard. Buitenz. Ill 18: 459. 1950.
TYPES.—Assa, Assa itidica, Assa exotica, Tetracera assa, Assa tetragynia: Her-
barium Houttuyn, Planta Ind. Orient, Java; holotype in L (original of Houttuyn's
figure). — Wahlbomia indica, Tetracera wahibomia; Herbarium Thunberg; holotype
in UPS. — Tetracera luevis: Herbarium Vahl, dedit Lamarck; holotype in C. — Tetra-
cera malabarica: herbarium Lamarck; holotype in P (original of Lamarck's figure).
— Tetracera dichotoma: Blume s.n., Java; holotype in L. — Tetracera gracilis: Blume
1638, Java; holotype in L. - Tetracera trigyna: Wallich 6629A (Herbarium Rox-
burgh) ; lectotype in K, isotype in BR.
Small shrub, up to 2 m high, or small liana, up to 6 m long, creeping
on the ground or clambering upwards over other plants, much branching,
younger branches strigose, reddish green, smooth, older ones glabrous
with greyish brown bark. Leaves elliptic to oblong or obovate, (3.5—)6—
10(—20) x (1.5—)3—B(—9) cm, with (7—)9—11(—15) nerves on either
side; apex obtuse to acute, not or only slightly acuminate; base acute;
margin entire to, usually, more or less dentate; nerves slightly curving
upward, ending mucronateiy in apex of teeth; leaves above often glossy,
glabrous on intervenium, sparsely pubescent on midrib, beneath dull, slight-
ly pubescent to glabrous on intervenium, rather densely to very sparsely
strigose on midrib and nerves, without or with only very inconspicuous
hairy domatia,- on both sides smooth; petiole (4—)6—10(—15) mm,
sparsely pubescent above, strigose beneath. Inflorescences terminal,
sometimes (particularly on climbing plants) on short lateral few-leaved
branches, up to 8 X 6 cm, (2—) 4—7(:—12) -flowered; peduncle 0.5—2 cm,
like branches strigose to strigose-hirsute; bracts lanceolate, 2.5—12 X 1—5
mm, acute at apex, attached with broad base, glabrous above, strigose
beneath. Flowers 2.5—3cm across; pedicel 8—15mm, strigose, hairs
patent near apex, most densely hairy immediately below flower, without
or with 1—3 bracteoles; bracteoles lanceolate, 3—4 X 0.8—1.2 mm. Sepals
4, 8—10 x 7—9 mm, glabrous on both sides, yellowish green or slightly
tinged with red. Petals (3—)4(—5), 12—15 x 6—8 mm, reddish white.
Stamens about 375, 6—8 mm long, red, white at base, with thecae touching
each other at apex. Carpels 3—4, glabrous except for few 0.5 mm long,
appressed, stiff hairs on back, 1.5 x l m m with 5 mm long style, with
about 10—15 ovules. Fruits with (1—)2—3(—4) capsules developed in
each flower; capsules almost globular, about 10 mm in diameter with
2—6 mm long beak, glabrous, glossy, red or reddish brown, 1—7-, usually
2-, seeded. Seeds ovoid, 3—4 X 2—3 mm, glossy black; aril 8—10 mm long,
bright red, fimbriate nearly to its base.
DISTRIBUTION.—Assam, Bengal (Chittagong), Burma (Pegu), Siam,
Indo-China (Cambodia, Cochinchina), China (Fukien), Sumatra, Malay
Peninsula, Banka, Java, Madura, Kangean.
1958] HOOGLAND: Tetracera 207
ECOLOGY.—Small shrub in open places, e.g. recently .abandoned fields;
low liana, climbing over low shrubs, in brushwood and open forests,
common in the Malay Peninsula and Java, less common in South Indo-
China and Sumatra, rarely
collected in the other parts
of its area; at low alti-
tudes, up to 600 m.
VERNACULAR NAMES.—
S i a m; pot luan (Song-
kla), pot lun (Krabi), yan
pad lun (Trang). I n t l o -
C h i n a : cay giay chieu,
cheait betao (fide Gagne-
pain), chon que (Moi;
Bien HoA). S u m a t r a :
aplas kedjong (Djambi),
baih siepiek, baih sipiek
suloh, sipik suluh, wajit
sipit (Lampung), djelati
(Palembang), memplas ga-
dja (East Coast). M a l a y
P e n i n s u l a : akar pulas
puyio, ma ampalasu akar
(Malacca), ampelas lichin
(fide Burkill), ampelas
mihsak (Trengganu), am-
pelas minyak, ampelas
payah (fide Ridley), ka-
lintat niamok (Singapore).
B a n k a: akar tempelas.
J a v a : akar mempelas
(tempelas), empelas (mempelas) akar (Malay), (areuj) ki as(s)han, (kaju)
asahan (Sundanese), bo (Javanese). K a n g e a n : buko-buko.
USES.-—The stems may be used as cordage. The leaves, though smooth
on both sides, are said to be used as sandpaper (Burkill, 1935). The medical
use is unimportant (Burkill, 1935).
The type specimen of Assa indica Houtt. ex Christm. & Panz., present
in the Rijksherbarium, Leyden, agrees with the figure of Houttuyn, the
latter being the reverse image of the actual specimen. This is perhaps
the first specimen known from Ilouttuyn's collection which could be iden-
tified as his with certainty (cf. Merrill in J. Arn. Arb. 19: 291. 1938).
Tetraceru hygrophila Kurz (nomen nudum) is represented in the
Calcutta and Kew herbaria by sterile specimens (Kurz 1809 from Pegu)
probably belonging to the present species. The only Bengal record is a
small-leaved, sterile specimen collected by Hooker f. & Thomson (K), the
Fie. 1. Trtracera indica (Houtt. ex Christm.
& Panz.) Merr.
2 0 8 R E I N W A R D T I A [VOL. 2
only Assam record a specimen in the Pierre herbarium (P), communicated
by T. Anderson. The only Chinese record is based on a specimen in the
Arnold Arboretum herbarium (H. H. Chung 7051, Fukien, without exact
locality).
8. TETRACERA AKARA (Burm. f.) Merr. — Fig. 5
[Akdra-Patsjoti Rheede, Hort. malab. 5: 15 pi. 8* 1085.]
Calophyllmn akara Burm. f., PI. ind. 121. 1768.
Roehlingia suaveolens Dennst., Schliiss. Hort. ind. malab. 31. 1818.
Tetracera rkeedii D C , Syst. 1: 402. 1818; Prod. 1: 68. 1824; Wight & Am.,
Prod. PI. Pen. Ind. or. 1: 5. 1834; Wight, Ic. PI. Ind. or. 1: pi. 70." 1838; Drury, Hanilb.
Ind. PI. 1: 9. 1864.
Tetracera sericea BL, Bijdr. 1: 3. 1825; Miq., Fl. Ind. bat. 1 (2): 9. 1859.
Tetracera axillaris Martelli in Bece., Malesia 3: 151. 1886; Merr., Bibl. En.
Born. PI. 381. 1921.
Tetracera assa "vai1." Ridl. in J. Str. Br. R. A. S. 33: 37. 1900.
Tetracera sylvestris Ridl. in J. Str. Br. R. A. S. 54: 8. 1910; Fl. Mai. Pen. 1: 6.
1922.
Tetracera akara (Burm. I.) Merr. in Philip. J. Sci. 19: 366. 1921; Hoogl. in Fl.
mal. I 4: 146. 1951.
Tetracera laevis Auet. (noil Vahl) ; D C , Syat. 1: 401. 1818; Prod. 1: 68. 1824;
Hook. !. &. Thorns., Fl. ind. 1: 62. 1855; Miq. m Ann. Mus. bot. Lugd. Bat. 4: 74.
1868; Hook. i. & Thorns, in Fl. Br. Ind. 1: 31. 1872; Trim., Handb. Fl. Ceyl. 1: 6. 1893;
Brand., Ind. Trees 5. 1906; Fin. & Gagnep. in Bull. Soc. bot. Fr., Mem. 4: 3. 1906, p.p.;
Back., Schoolfl. J a v a 9. 1911; Gamble, Fl. Pres. Madras 1: 7. 1915; Back., B e t a . Fl.
J a v a (Nooduitg.) 4 (Fam. 80) : 2. 1942.
Tetracera assa Auct. (nc-n DC.); Hassk., PI. jav. r a r . 177. 1848.
TYPES.—Calophyllnm akara, Roehlingia suaveolens, Tetracera rheedii: Akdra-
Patsjoti Rheede, I.e., pi. 8. — Tetracera sericea: Blume 835, Gunung Seribu, J a v a ;
holotype in L. — Tetracera axillaris: Beccari Piante Bornensi 2844, Gunung Wah,
Sarawak, November 1866; lectotype in F l , isotype in K. — Tetracera sylvestris:
Ridley 6179, Garden Jungle, Singapore, April 1894; lectotype in SING, isotypes in
BM, CAL.
High climbing- or creeping liana, up to 25 m long, with bright brown
trunk, up to 6 cm thick, much-branching; younger branches strigose,
older ones glabrous with bright brown to whitish bark. Leaves oblong
to lanceolate, (5—)g—13(—22) x (1.5—)3.5—6(—10.5) cm, with (5—)
6—8(—10) nerves on either side; apex distinctly acuminate; base acute;
margin entire to slightly undulate or dentate; nerves curving upward
along margin, sometimes with vein towards margin, ending mucronately
in apex of teeth; leaves above glossy bright green, glabrous on inter-
venium, glabrous to slightly pubescent on nerves, beneath dull, glabrous on
intervenium, sparsely strigose on nerves, without or with only slightly
developed hairy domatia, on both sides smooth or, rarely, slightly scabrid;
petiole (3—)5—7(—10) mm, glabrous to slightly pubescent above, sparsely
strigose beneath. Inflorescences terminal or axillary, up to 8 by 6 cm, (2—)
5—8(—12)-flowered; peduncle 0.5—1.5(—3.0) cm, like branches sparsely
1953] HOOGLAND: Tetracera 209
hirsute to glabrous; bracts lanceolate, about 2 x 1 mm, acute at apex,
attached with broad base, glabrous to slightly hirsute above, sparsely to
densely strigose-hirsute beneath. Flowers 2.5—3.0 cm across; pedicel
10—25 mm, sparsely hirsute to glabrous, without or with 1—2 bracte-
oles; bracteoles lanceolate, about 1 X 0.5 mm. Sepals 4, 8—10 X 6—8 mm,
often reflexed in fruit, green or reddish green, glabrous outside, densely
whitish or yellowish sericeous except 1—2 mm broad glabrous margin
inside, ciliate at margin. Petals 3—4, 12—15 x 6—8 mm, white or
greenish white. Stamens about 230, 7—8 mm long, yellowish white with
grey tips, the thecae touching each other at apex. Carpels 3(—4), glabrous,
1.5 X 1 mm with 5 mm long style, with about 10 ovules. Fruits with (1—)
2—3 capsules developed in each flower; capsules almost globular, about
10 mm in diameter with 1—3 mm long beak, glabrous, glossy, 1—2-
seeded. Seeds ovoid, 2—4 x 1—3mm, glossy black; aril 6mm long,
fimbriate for 1/2—3/4 of its length.
DISTRIBUTION.—Deccan Peninsula (Malabar, Travancore), Ceylon,
Indo-China (Cambodia), Sumatra, Malay Peninsula, West Java, Borneo,
and Celebes.
FIG. 5. Tetmcerci akara (Burm. f.) Merr.
ECOLOGY.—Climber in lowland forests, up to 750 m altitude.
VERNACULAR N A M E S . — I n d i a : akar a patsjoti (Malabar), tilo sameno
(Brachm.). C e y l o n : et-korasa-wel. S u m a t r a : daun amplas (Pa-
lembang). M a l a y P e n i n s u l a : akar rusa-rusa, mumplas rimba (Ma-
lacca). J a v a : aroj pengasaman (Sundanese). B o r n e o : daun ampelas
(Pulau Kuwala).
U S E S . — T h e leaves are said to be used as sandpaper, though the sur-
face is hardly rough (Polak 2120, Borneo). The stems are used for ropes.
210 R E I H W A R D T I A [VOL. 2
This species is most commonly known under the name Tetracera
laevis Vahl which, however, is identical with T. indica (Houtt. ex Christm.
& Panz.) Merr., as is evident from the type specimen in the Copenhagen
herbarium.
Tetracera axillaris Martelli represents a small-leaved form, which
has been collected only in Borneo. I do not think it to be of any taxonomic
value.
The only specimen from Indo-China (Pierre 763, Samrongtong
Province, Cambodia) much resembles T. indica; the inner side of the
sepals has the sericeous indument as is found in the present species,
though it is distinctly less dense than in all other specimens. The identity
of this specimen appears somewhat dubious.
The species is closely related to T. indica; the main difference is the
indument on the inner side of the sepals. With exclusion of the specimen
just cited this character is very clear and sharp. In addition the acuminate
apex of the leaf and the generally more glossy and darker appearance of
the upper side of the leaves are usually characteristic of 2'. akara. The
latter is found almost exclusively in forests, T. indica mostly in much more
open, low vegetation.
9. TETKACERA LOUEEIRI (Fin. & Gagnep.) Pierre ex Craib. — Fig. 6
Tetracera assa var. hnreiri Fin. & Gagnep. in Bull. Soc. hot. F i \ , Mem. 4: 3.
1906.
Tetracera sarmentosa var. loitreiri (Fin. & Gagnep.) Fin. & Gagnep. in PI. gen.
Ind.-Ch. 1: 16. 1907; Craib in Aherd. Univ, Studies 57: 4. 1912.
Tetracera fragrana Ridl. m J. Str. Br. R. A. S. 5 9 : 62. 1911; Fl. Mai. Pen. I : 6.
1922; lion Wildem. & Dur. 1899.
Tetracera loureir! (Fin. & Gagnep.) Pierre M Craib in Kew Bull. 1922: 1(55;
Craib, Fl. siam. En. 1: 20. 1925; Gagnep. in Fl. gen. Ind.-Ch., Suppl. 1: 28. 1938;
Hoogl. in Fl. mal. I 4; 147. 1951.
Tetracera kampotensifi Gagnep. in Not. ayst. 6: 40. 1987; in Fl. gen. Ind.-Ch.,
Suppl. 1: 18/..?. * 1938.
TYPES.—Tetracera assa var. hnreiri: Zimmermann 74, Bangkok, Siam, 1899;
lectotype in P, isotypes in BM, BO, BR, G, K, L, U. — Tetracera fragravs: Ridley
15183, Chupcng, Perlis; leetotype in SING, isotype in K. — Tetracera kampotensis:
Chevalier 31763, Kampot, Cambodia, March 18—19, 1914 (collector: F l e u r y ) ; holo-
type in P.
Woody vine with flexuous branches; branches sparsely strigose,
glabrescent, sometimes slightly scabrid. Leaves elliptic to oblong, (1.5—)
6—10(—13) x ( 1 _ ) 3 _ 5 ( _ 7 ) cm, with (3—)8—12(—15) nerves'on either
side; apex rounded to obtuse; base obtuse to acute; margin entire to
distinctly dentate; nerves slightly curving upward, ending in margin or
in apex of teeth; leaves above glossy, glabrous except the sparsely strigose
1953] HOOGLAND: Tetracera
basal Vs of the midrib, beneath dull, glabrous on intervenium, glabrous to
sparsely strigose on nerves, sparsely strigose on midrib, without or
with only inconspicuous hairy domatia, on both sides smooth or only
slightly scabrid, with many small circular spots; petiole (4—)7—10mm,
glabrous to sparsely strigose above, sparsely strigose beneath. Inflores-
cences terminal, 10—20 x 5—10 cm, about 40—80-flowered, often with
1—2(—4) leaves in basal part; branches sparsely strigose, extreme ones
most densely so; bracts lanceolate, 4—10 x 1—3 mm, acute at apex, attach-
ed with broad base, glabrous above, sparsely strigose on midrib beneath,
sparsely ciliate at margin. Flowers 8—11mm across; pedicel 3—7mm.
sparsely strigose, without or with 1—3 bracteoles; bracteoles lanceolate,
2—3 X 0.5—1 mm, glabrous, ciliate at margin. Sepals 5, 2 outermost ones
4—5 x 3—3.6 mm, 3 innermost ones 6—-7 x 4—5 mm, glabrous, slightly
scabrid outside, glabrous, smooth inside, ciliate at margin. Petals 3, 5 x
2.5 mm, white or pink. Stamens about 150, 4—5 mm long, the theeae
distinctly separated to nearly touching each other at apex; connective
only slightly emarginate between theeae. Carpels (2—)3, glabrous, 1.5 X
1 mm with 2.5 mm long style, each with about 8—16 ovules. Fruits with
1—3 capsules developed in each flower; capsules ovoid, 7—8 x 5—6 mm,"
acute with 2 mm long beak, glabrous,
glossy, 1—2-seeded. Seeds globular,
2—3 mm in diameter, glossy dark
brown; aril asymmetric, on one side
5—6, on other 2—3 mm long, lacin-
iate to1/8—1/4 of its length.
DISTRIBUTION.—Around the Gulf
of Siam, on the west coast of the Ma-
lay Peninsula from Kedah to Setul,
on the east coast from Singora north-
ward, in Indo-China on the east
coast north to Bleu Hoa Province.
ECOLOGY.—Climber in open for-
ests, scrubs, and hedges, from sea-
level up to 400 m altitude.
VERNACULAR NAMES.—S i a m: baw
rakhon (Koh Chang), lin rat (Lam
Nang Eong), rot sukon (Singora),
tana kim (Chantaburi), thao kapot bai suam, thao harakon, thao orakon
(Prachuap), yan pot (Nakawn Sritamarat). I n d o - C h i n a : bay giay
omen, giay chien (Annamese: Bien Hoa), chait betao (Moi: Bien Hoa),
dak kuon (fide Gagnepain), day chien (Cochinchina), giey no nuoc hon
(Cambodia).
The species is not closely related to T. korthalsii Miq., as might be
suggested by its position in the key. In addition to the character given
in the key, the species differ in the following characters: T. loureiri has
generally narrower and thicker leaves, it has often in a number of flowers
only two carpels, it has small circular spots on both sides of the leaves,
FIG. 6. Tetracera loureiri (Fin. &
Gagnep.) Pierre ex Craib.
2 1 2 R E I N W A R D T I A [VOL. 2
which are not found in T. korthalsii, the thecae are much less separated
at the apex in T. loureiri than in T. korthalsii. The latter is related to
T. fagifolia, Bl., whereas there are no species closely related to T. loureiri.
10. TETRACERA DAEMELIANA F. Muell.
Tetracera daemeliana F. Muell., Fragm. 5: 191. 1865; F. M. Bail., Synops.
Queensl. Fl. S. 188S; Queensl. Fl. 1: 10. 1899; Compr. Cat. Queensl. PI. 18. 1909; White
in Contr. Arn. Arb. 4: 71. 193.3.
TYPE.—Daemel s.n.t Cape York, October; holotype.in MEL, isotype in BRI.
Large woody vines, much branching; branches glabrous, smooth.
Leaves subcoriaceous, oblong to narrowly obovate, (6—)14—20(—23) x
(2.5—)5—7(—8) cm, with (8—)10—14 nerves on either side; apex acute
to obtuse; base acute; margin entire; nerves curving upward, each anas-
tomosing with the next one through a secondary nerve; leaves more or less
shining, glabrous above, dull, glabrous beneath, smooth on both sides;
petiole 6—20 mm, winged, 2—4 mm broad, glabrous. Inflorescences
terminal, up to 20 x 10 cm, up to 200-flowered, often with 1—2 small
leaves in basal part; branches, particularly extreme ones, sparsely
strigose; bracts ovate, 2.5 X 1.5 mm, broadly acute at apex, attached by
broad, rounded base, glabrous above, strigose beneath. Flowers 12—15
mm across; pedicel 2—3 mm, sparsely strigose, without or with 1—3
minute bracteoles. Sepals 5—6, 1—2 outermost ones 4—5 x 3—4 mm,
inner ones about 6 X 4.5 mm, very sparsely strigose to glabrous, slightly
scabrid outside, sparsely to rather densely short-sericeous, smooth
inside, ciliate at margin. Petals 3, 6—7 x 3.5—4 mm, rather thick, green-
ish white or cream-coloured. Stamens about 110—150, 5—6 mm long,
the thecae manifestly separated at apex; connective slightly emarginate
between thecae. Carpels 3, glabrous except for some 0.2 mm long stiff
hairs on the back, each carpel 1.5 x 1.5mm; style 2"mm long; ovules
about 10. Fruits with 2—3 capsules developed in each flower; capsules
ovoid, 1 0 x 6 mm, acute with 1 mm long beak, glabrous, dull, 1-seeded.
Seeds ovoid, 4 x 3 mm, glossy black; aril unknown to me.
DISTRIBUTION.—Queensland (Cape York Peninsula: Etty Bay, John-
stone River, Daintree River, Barron River).
ECOLOGY.—Climber in rainforests and scrub, at low altitudes; col-
lected in flower in October, November, and December, in fruit in March.
11. TETRACERA EURYANDRA Vahl
Euryandra aeandem Forst., Char. Gen. 41 pi. J,l". 1776.
Tetracera em-yandra. Vahl, Symb. bot. 3: 71. 1794; D C , Syst. 1: 402. 1818;
Deless., Ic. sel. PI. 1: pi. 70*. 1821; Lam., Illustr. 3: pi. i83*. 1823; D C , Prod. 1;
68. 1824; Labill., Sert. austr.-caled. 55 pi. 55". 1825; Montrous. in Mem. Acad.
Lyon 10: 175. 1860; Fin. & Gagnep. m Bull. Soc. bot. Fr., Mem. 4; 2. 1906, p.p.;
in Fl. gen. Ind.-Ch. 1: 13. 1907; Guill. in Ann. Mus. col. Marseille II 9: 93. 1911; in
Bull. Soc. bot. F r . 67: 47. 1920.
Tetracera billardieri Martelli in Becc, Malesia 3: 152. 1886.
1953] HOOGLAND: Tetraccra 213
Tetraeera seandens (Foist.) Gilg & Werderm. in Engl. & P r a n t l , Nat. PflFam.,
2nd Ed., 2 1 : 18. 1925; DSnik. in Vierteljahrsschr. naturf. Ges. Zurich 78: 206.
1933; Guill., Fl. Nouv.-Cal. 213. 1948; won (L.) Merr. 1917.
TYPES:—Ewryandra, seandens, Tetraeera euryandra: Forster 137, 228, New
Caledonia; lectotype in BM, isotypes in BM, S. — Tetraeera billardieri: De la Billar-
diere s.n., New Caledonia; lectotype in G.
Woody vine, much branching; branches strigose with 0.5 mm long
hairs, glabrescent. Leaves oval to elliptic, (3.5—)6—10(—15) x (2—)
3—5(—7) cm, with (6—)8—10(—12) nerves on either side; apex rounded,
sometimes slightly emarginate, to obtuse, often mucronate; base rounded
to obtuse; margin entire; nerves curving upward, each anastomosing with
the next one through a secondary nerve; leaves glossy, glabrous above,
slightly glossy, glabrous on intervenium, glabrous to sparsely strigose on
midrib and nerves beneath, smooth on both sides; petiole 10—20 mm,
slightly winged, about 1.5 mm broad, glabrous above, sparsely strigose
beneath, ciliate at margin, mainly in lower part. Inflorescences terminal,
5—15 X 3—9 cm, 20—125-fIowered, sometimes with 1(—2) small leaves in
basal part; branches strigose with 0.5mm long hairs; bracts caducous,
oblong, 6—10 X 2—4 mm, obtuse at apex, attached by broad, rounded base,
sparsely strigose above, strigose beneath. Flowers 15—20 mm across;
pedicel 1—3 mm, strigose, without or with 1—3 bracteoles; bracteoles
oblong, 2—4 x 1—2 mm. Sepals 5—6, 2 outermost ones about 5 x 4 mm,
inner ones about 1 0 x 8 mm, sparsely strigose, smooth outside, sparsely
sericeous with 0.2—0.3 mm long hairs inside, ciliate at margin. Petals
S, 9 X 5 mm, white. Stamens about 250, 4—6 mm long, the thecae on
outer side of broadened connective, extrorse, nearly touching each other
on outer side, distinctly separated on inner side, manifestly separated at
apex. Carpels 2—3, glabrous, 2 x 1,5 mm with 4 mm long style, ovules
about 10. Fruits with 2—3 capsules developed in each flower; capsules
ovoid, 10—12 x 8—10 mm, acute with 1—2 mm long beak, glabrous,
shining, 1—4-seeded. Seeds ovoid, flattened, 4 x 3.5 mm, 2.5 mm thick,
slightly glossy, black; aril 6—10 mm long, fimbriated to "/IO of its length.
DISTRIBUTION.—New Caledonia, all over the island.
ECOLOGY.—On plains, in open forests, on edge of forests, and in
riverside-scrub; from low altitude up to 700 m; collected in flower in
October to January, in fruit in October to April.
There is one collection from Indo-China in the Paris herbarium
(Spire 74, from Laos) which does not seem very reliable. The species was
included in the "Flore generale de l'lndo-Chine" on account of this spec-
imen. I think the record should be neglected, as from a geographical
point of view the occurence in Indo-China is improbable and an inter-
change of labels may be possible.
12. TETRACERA KORTHALSII Miq. — Fig. 7
Tetraeera korthalsii Miq. in Ann. Mus. bot. Lugd. Bat. 4: 75. 1868; Merr., .BibI,
En. Born. PI. 381. 1921; Hoori. in Fl. mal. I 4: 147. 1951.
214 REINWARDTIA [VOL. 2
Tetracera subrottmda Elm. in Leafl. Philip. Bot. 5: 1771. 1913; Mere., E n .
Philip, fl. PI. 3: 59. 1923; Elm. ex. Prain, Ind. kew. Suppl. 5: 257. 1921 C suhrotundata",
error for the preceding).
Tetracera clmeri Meir. in Univ. Calif. Publ. Bot. 15: 104. 1929.
TYPES:—Tetracera korthalsii: Korthals s.n., Tewe River, Borneo: lectotype in
L. — Tetracera siibrotundu: Elmer 13048, Puerta Prineesa, Palawan, April 1911;
lectotype (isotype) in L, isotypes in A, BM, BO, CAL,, E, P I , G, GH, K, MO,
K U, US. — Tetracera clmeri: Elmer 2137G, Tawao, British North Borneo, October
1922—March 1923; holotype in UC, isotypes in A, BM, BO, C, G, GH, K, L, MO, NY,
P. SING, U.
Large climbers or creepers, much-branching; branches scabrid, stri-
guse to hirsute with 1 mm long hairs, sometimes also small tufts of 0.2—
0.4 mm long hairs, glabrescent. Leaves oval to elliptic-oblong or obovate,
(3.5—) 6.5—17(—22) X (2.3—)4—8(—13) cm, with (6—)10—16(—20)
nerves on either side; apex rounded to acute, sometimes slightly acuminate;
base rounded to acute; margin slightly dentate; nerves slightly curving up-
ward, ending in apex of teeth; leaves above hirsute to sparsely strigose
to glabrous on intervenium, glabrous on nerves, glabrous on upper part
of midrib, short-hirsute to glabrous on basal part, beneath sparsely
hirsute to glabrous on intervenium, hirsute to strigose on nerves and
midrib, on both sides smooth to slightly scabrid; petiole (5—)8—20(—SO)
mm, slightly winged, short-hirsute to glabrous above, hirsute to strigose
beneath. Inflorescences terminal, 10—30(—100) X 6—20 cm, 40—200- and
more-flowered, often with 1—2 small leaves in basal part; branches densely
to slightly hirsute to strigose with solitary, 1 mm long hairs and small
fascicles of 0.2—0.4mm long hairs; bracts caducous, lanceolate, 3—5 x
1—2 mm, acute at apex, attached by broad base, glabrous above, strigose
to hirsute beneath. Flowers about 10mm across; pedicel 1.5—2.5mm,
as to indument like extreme branches of inflorescences, without or with
1—2 bracteoles; bracteoles caducous, lanceolate, 1.5 x 0.5 mm. Sepals
5{—6), 2 outermost ones 4 x 3 mm, 3(—4) inner ones 5—5.5 x 4—5 mm,
glabrous or sparsely strigose-hirsute with small fascicles of short hairs
on central part, scabrid outside, glabrous inside, not ciliate at margin.
Petals about 5 x 3.5 mm. Stamens about 125, 3.5—4 mm long, with
thecae strongly separated at apex. Carpels 3, glabrous or covered by
minute scales, ovoid. 1.3 X 1 mm with 1—2 mm long style, with about
9 ovules. Fruits with 1—3 capsules, developed in each flower; capsules
ovoid, 7 x 4 mm, acute with 1—2 mm long beak, glabrous, glossy, 1-seeded.
Seeds ovoid. 4.5 x 3.5 mm, glossy black; aril mainly one-sided, 2.5—5 mm
long, fimbriate with broad lobes to 1/3—1/2 of its length.
Var. KORTHALSII.
Tetracera korthalsii var. korthalsii; Hoogl. in Fl. mal. I 4: 147. 1951.
Leaves elliptic-oblong or obovate, (3.5—) 6.5—17(—20) x (2.3—) 4—
8(—16) cm; apex acute or slightly acuminate; base acute; petiole (5—)
g—20(—30) mm.
1953] HooiiLAND: Tetracera
Var. SUBROTUNDA (Elm.) Hoogl.
Tetracera korthalsii var. subrotunda (Elm.) HOOKI. in Fl. mal. I 4: 147. 1951.
Tetracera aubrotunda Elm. — Tetracera elmeri Merr.
Leaves broadly oval to nearly circular, (6.5—)8.5—22 x (4—)5.5—13
cm; apex and base rounded; petiole 12—30 mm.
DISTRIBUTION.—VariefJ kort-
halsii in Borneo (eastern half),
Celebes, and Moluccas (Taliabu
Island) ; variety subrotunda in
British North Borneo and Pala-
wan.
ECOLOGY.—Climber in forests
at low altitudes, up to 700 m.
VERNACULAR NAMES.—Empelas
(Malay, Sandakan), Pampad (Du-
sun, Sandakan).
USES.—The leaves are used
for polishing wood.
The differences between the
two varieties are found in the
shape and, to a lesser degree, in
the size of the leaves. Tetraceru
elmeri Merr. represents a hirsute
form.
The species is closely related to and much resembles T. fagifolia BL,
differing- from the latter by the absence of the sericeous indument on
the inner side of the sepals.
13. TETRACERA MACROPHYLLA Wall, ex Hook. 1 & Thorns.— Fig. 8
Tetracera macrocarpa Wall., Cat. no. 6628, 182S, nomen nudum.
Tetracera macrophylla Wall, ex Hook. f. & Thorns., Fl. ind. 1: 6.3. 1855; Miq.,
PI. Ind. bat. 1 (2): 8. 1859; Hook. f. & Thorns, in Fl. Br. Ind. 1: 32. 1872; King
in J. As. Soc. Beng. 58 (2): 363. 1889; Gamble, Man. Ind. Timb., 2nd Ed., 3. 1902;
Ridl., Fl. Mal. Pen. 1: 4. 1922; Hoogl. in Fl. mal. I 4: 147. 1951.
Tetracera scaberrima Miq., Fl. Ind. bat. 1 (2) : 9. 1859; in Ann. Mus. bot. Lugd.
Bat. 4: 75. 1868; Merr., Eibl. En. Born. PI. 382. 1921.
Tetracera teysmannii Mai-telli in Becc, filalesia 3: 150. 1886.
Tetracera radnla Martelli in Becc, Malesia 3: 153. 1886; Merr., Bibl. En. Born.
PI. 382. 1921; non Martius 1863.
Tetracera grandis King in J. As. Soc. Beng. 58 (2) : 363. 1889; i« Ann. roy. bot.
Card. Calc. 5: 115 pi. 119.* 1896; Eidl., Fl. Mal. Pen. 1: 4. 1922.
Tetracera havilavdii Ridl. in Kew Bull. 1912: 381; Merr., Bibl. En Born PI
381. 1921.
Tetracera scabricaulis Ridl. in Kew Bull. 1912: 381; Merr., Bibl En Born
PI. 382. 1921.
F I G . 7. Tetracera korthatiii Miq.:
var. korthalsii ( ), var. subrotunda
(Elm.) Hoogl. ( X ) .
216 REINWARDTIA [VOL. 2
TYPES.—Tctracera- macrophylta: Wallieh 6628, Singapore, 1822; lectotype in
K, isotypes in BM, CAL, CGE, G. — Tetracera scaberrima: Teysmann 452HB, Lubuk-
alung, Sumatra West Coast; holotype in U, isotypes in BO, L. — Telracera Uysmannii:
Beccari s.n, cultivated in the Botanie Garden, Bogor (Buitenzorg), from Banka;
holotype in FI. — Tetracera radula: Beccari Piante Bornensi 3448, Sungei Kantu,
Pontianak, May 1867; holotype in FI. — Tetraceta grandis: Seortechini g.n., Perak;
holotype in CAL, isotypes in BM, FI, K, SING, UC. —tetracera havilandii: Haviland
1811, near Kuching, Sarawak, October 28, 1892; holotype in K. — Tetracera scabricau-
lis: Creagh s.n,, Sandakan, British North Borneo, April 20, 1895; holotype in K.
Woody vine, up to 10 m long, or tree(?), with winding or straight
branches; branches strongly scabrid to smooth, densely hirsute with 1—
1.5 mm long single hairs and stellate groups of about 0.3 mm long hairs,
glabrescent. Leaves elliptic to oblong, (5—)8—15(—30) X (3—)5—10(—
17) cm, with 8—12(—17) nerves on either side; apex and base rounded
to obtuse; margin slightly emarginate at end of nerves, entire or slightly
dentate with nerves ending in apex of teeth; nerves slightly curving up-
ward, ending mucronately; upper surface of leaves often slightly glossy,
the intervenium rather densely hirsute with single, 1—2 mm long hairs
and, at extreme basal part only, few stellate groups of 0.3 mm long hairs to
nearly glabrous, midrib densely to slightly hirsute with both kinds of
hairs, under surface densely hirsute with both kinds of hairs to nearly
glabrous, margin ciliate with 1—2 mm long single hairs only near base,
very scabrid to smooth on both sides; petiole 15—30 mm, winged, 2—4 mm,
in leaves in base of inflorescences 5—8 mm, broad, slightly to densely
(most densely beneath) hirsute with both kinds of hairs, with single longer
hairs mainly along edge. Inflorescences terminal, 10—40 X 4—15 cm,
(20—) 50—125(—200)-flowered, often with 1—4 small leaves in basal
part; branches more or less densely velvety-hirsute like younger sterile
branches; bracts oblong, 5—10(—20) x 2—4(—8) mm, acute at apex,
attached by a broad base, densely hirsute with single hairs above, with
single and stellate groups of hairs beneath, ciliate at margin. Flowers
2—2.5 cm across; pedicel 4—20 mm, densely hirsute like extreme branches
of inflorescences, without or with 1—3 bracteoles; bracteoles oblong, 2—4
x 1—2 mm, acute at apex, attached by a broad base, as to indument like
bracts. Sepals 5—6, 2 outermost ones 8—9 x 7—8 mm in flower, 9—12
x 6—8 mm in fruit, inner ones 11—12 x 8—9 mm in flower, 12—15 x
7—9 mm in fruit, all more or less densely velvety, scabrid outside, 2 outer-
most ones glabrous, inner ones sericeous except 1—2 mm broad margin,
smooth inside, all ciliate at margin, greenish, turning to red in fruit. Petals
3, 8—11 x 5—7 mm, rather thick, not emarginate at apex, attached by
rounded, 1—1.5 mm broad base. Stamens about 375, 4—7 mm long, the
thecae distinctly separated at apex, connective strongly emarginate be-
tween thecae. Carpels 3—4, glabrous except few strigose hairs on back,
2 X 1.5 mm with 4 mm long style, with about 14 ovules. Fruits with 2—3
(—4) capsules developed in each flower; capsules ovoid, 8—10 X 6—8 mm,
acute with 2—3 mm long beak, glabrous, glossy brown, 1—2-seeded.
Seeds ovoid, 6—7 x 4—5mm, glossy black; aril mainly one-sided, 5—9
mm long, slightly laciniate for about 1/8 of its length.
1953] HOOGLAND: Tctraccra
DISTRIBUTION.—Sumatra, Malay Peninsula, Banka, Borneo.
ECOLOGY,—Climber in dry aa well as in swampy forests and thickets,
up to 300 m altitude. According to Ridley very common in the Malay
Peninsula, but rarely in flower.
F I G . 8. Tttmccnt macrophylla Wall, ex Hook. f. & Thorns.
VERNACULAR NAMES.—S u m a t r a: akar ampaleh riembu (West Coast).
M a l a y P e n i n s u l a : ampelas gajah (Alvins 600, without locality),
ampelas lidah kuching, ampelas rimbah (Malacca), ampelas rimau (fide
Ridley). B o r n e o : akar tembara (W. Kutei), ampalas (Sarawak).
USES.—The medical use is unimportant (cf. Burkill, 1935).
Within this species the size and shape of the leaves and their
scabridness is very variable. Some specimens from the Malay Peninsula
are nearly smooth, most specimens are very scabrid. Small-leaved speci-
mens are the type specimen of T. seabricaulis Ridl. and Forbes 3059 from
Sumatra. The type specimen of T. havilandii Ridl. is a short one-leaved
fruiting branch with a small, rather smooth leaf, possibly only part of an
infructescence.
The most typical feature of the species is the absence of an indument
on the inner side of the two outermost sepals, whereas the inner sepals
have a dense sericeous indument.
14. TETRACERA ARBORESCENS Jack. — Fig. 9
r e t r a c e r a arborescens Jack hi Mai. Misc. 1 (5) : 45. 1820; D C , Prod. 1: 69. 1824;
Miq., Fl. Ind. bat. 1 (2) : 9. 1859; Gage & Boik. in J. Str. Br. R. A. S. 7 3 : 242. 1916;
Hoogl. in Fl. Mai. I 4: 148. 1951; Meir. in J. Arn. Arb. 33: 248. 1952.
r e t r a c e r a laavigata Miq., Fl. Ind. bat. 1 (2): 8. 1859; in Ann. Mus. hot. Lugd.
Bat. 4: 74. 1868; Men-., Bibl. En. Born. PI. 382. 1921.
Tetracera subcordata Boerl., Cat. Hort. bot. bogor, 3. 1899.
218 R E I N W A R D T I A [VOL. 2
Tetracera lucidu Wall., Cat. no. G631. 1828, nomen nudum; e,c Ridl., Fl. Mai. Pen.
1: B. 1922.
Tetracera lucida var. lanugmosa Ridl., Fl. Mai. Pen. 1: 5. 1922.
Tetracera euryandra Auct. (non V a h l ) ; Hook. f. & Thorns., Fl. Ind. 1: 63. 1855;
Miq., PL Ind. bat. 1 (2): 8. 1859; in Ann. Mus. bot. Lugd. Bat. 4: 75. 1868; Hook,
f. & Thorns. « Fl. Br. Ind. 1: 32. 1872; King in J. As. Soc. Beng. 58 (2) : 362. 1889;
Fin. & Gagnep. in Bull. Soe. bot. Fr., Mem. 4: 2. 1906, p.p.; Back., Sehoolfl. J a v a 9.
1911.
TYPES:—Tetracera arboresceiis: Jack x.n., Tapanuli, W Sumatra. 1829 ( ? ) ; holo-
type in L. — Tetracera Uievigata: Teysmann 457HB, Sibogu, Sumatra West Coast;
holotype in U, isotypes in BO, CAL, P I , L, MEL. — Tetracera subcordatn: Boerlage
s.n., cultivated in Hortus Bogoriensis no. XI. A. 74, May 17, 1890; holotype in BO,
isotypes in CAL, K, L. — Tetracera lucida: Wallich 6631, Singapore, September 1822;
lectotype in K, isotypes in CAL, CGE. — Tetracera lucida var. lanuginosa: Kunstler
5579, Larut, Perak, February 1884; holotype in SING, isotypes in CAL, K.
Strong, woody climbers, up to 8 m long, or shrubs, or (?) small trees,
much-branching; branches villose to densely villose-floccose with rusty
brown tomentum, glabrescent. Leaves subcoriaceous, obovate to elliptic-
oblong, those below inflorescences distinctly smaller [(2—)3—4(—6) x
(1—)2—3(—4) cm] than those lower at branches [(4—)6—10(—15) x
(2—)3—5(—8) cm], all with (4—)5—6(—8) nerves on either side; apex
rounded or obtuse to acute or acuminate; base obtuse to acute; margin
entire; nerves curving upward, ending 0.5—1 mm from margin; leaves
above bright green, often slightly glossy, short-strigose to glabrous on
basal part of midrib, further on villose-floccose to glabrous, glabrescent,
beneath slightly to densely villose-floccose, glabrescent on intervenium,
strigose to glabrous on midrib and nerves beneath, smooth on both sides;
petiole 3—5(—8) mm, villous-floccose, glabrescent, short-strigose to gla-
brous on central part above, strigose beneath. Inflorescences terminal, up
to 15 x 6 cm, (6—)10—15-flowered, usually with some (1—7) small leaves
in the basal part to high up in the panicle; branches villous-floccose,
glabrescent, or hirsute, extreme ones most densely so; bracts caducous,
lanceolate, 3—6 X 1—2 mm, .acute at apex, attached by broad base.
Flowers about 15mm across; pedicel (2—)4—10mm, villous-floccose,
glabrescent, without braeteoles. Sepals 5—6, about 5 x 3 mm in flower,
7 x 3 mm in fruit, reddish yellow, outside sparsely to densely villous-
hirsute, mainly in central part, glabrescent in fruiting state, slightly
scabrid, inside densely sericeous except 0.5—1 mm broad glabrous margin,
densely ciliate at margin. Petals 3, 5 X 4 mm, white. Stamens about 110
—150, 4—5 mm long, the thecae slightly to manifestly separated at apex.
Carpels 3, glabrous to hirsute or villous, 1.25 x 0.75 mm; style 0.5—3 mm
long; ovules about 10—12. Fruits with 2—3 capsules developed in each
flower; capsules ovoid, 7 x 4 mm, acute with 2—3 mm long beak, glabrous
or, rarely, villose-floccose, glossy, 1-seeded. Seeds ovoid, 2—3 X 1—2 mm,
glossy black; aril 3—5 mm long, laciniate with broad lobes to about 1/2 of
its length.
DISTRIBUTION.—Sumatra (Tapanuli, East Coast), Malay Peninsula,
Banka, Billiton, Borneo, and ?Java (Korthals: Papandajan).
1058] HOOGLAND: Tetracera 219
F I G . 9. Tetracera urborescens Jack.
ECOLOGY.—In swampy forests, riverside shrubs, open jungle, and on
borders of woods, only at low elevations. Probably rarely in flower.
VERNACULAR NAMES.—S u m a t r a: andor ruhas igung (Tapanuli),
mohi-mohi (Sibolga). M a l a y P e n i n s u l a : akar mamplas paya (Ma-
lacca) . B a n k s : akar tamplas (tempelas). B i 11 i t o n: akar memplas.
The type specimen of
Jack has a label ''Tetracera
arborescens WJ. Tappanoo-
ly," probably in Jack's hand-
writing, and "Legit Jack
Sumatra Occident. 1829" in
Hasskarl's handwriting. The
original description is in-
sufficient for specific dis-
tinction and the binomial has
never been interpreted be-
fore.
The species is most close-
ly related to T. fagifolia Bl.,
but differs mainly in the shape and size of the leaves, in the venation, and
in the indument of the younger parts. The venation in T. arborescens is
reticulate, in T. fagifolia parallel, at about right angles to the nerves, but
as to this character intermediates are sometimes found.
The hairy-fruited form has on the carpels and capsules an indument
which agrees with the indument on the younger vegetative parts. In these
specimens the indument is generally denser than in the specimens with
glabrous carpels. The hairs are distinctly thinner than in those species,
where the carpels are constantly hairy {T. scandens (L.) Merr., T. lanu-
ginosa Diels, and T. nordtiana F. Muell.].
15. TBTEACERA FAGIFOLIA Bl. — Fig. 10
Tetracera fagifoliu Bl., Bijdr. 1: 4. 1825; Miq., PI. Ind. bat. 1 (2): 9. 1859;
in Ann. Mus. hot. Lugd. Bat. 4: 75. 1808; Ridl. in J. Str. Br. R. A. S. 54: 10. 1909; PI.
Mai. Pen. 1: 6. 1922; Hood, in Fl. mal. 1 4 : 148 /. 2." 1961.
Tetracera rigida Bl., Bijdr. 1: 4. 1825; Jliq., Fl. Ind. bat. 1 (2) : 9. 1859; Back.,
SchooHl. Java 9. 1811; Bekn. Fl. J a v a (NooduitE.) 4 (Fam. 80) : 2. 1942.
Tetracera blumei Walp., Rep. 1: 67. 1842.
Tetracera sumatrana Miq., Fl. Ind. bat. Suppl. 1 : 618, 619. 1861; Ridl., Fl. Mal.
Pen. 1: 6. 1922, p.p.
Tetracera fapifolia f. submtegeryima Miq. hi Ann. Mus. bot. Lup:d. Bat. 4: 75.
1868.
Tetracera borneensis Miq. in Ann. Mus. bot. Lugd. Bat. 4: 76. 1868; Merr., Bifal.
En. Born. PI. 381. 1921.
220 R E I N W A R D T I A [VOL. 2
Tetracera obovata Boerl., Cat. Hort. bot. bogor. 3. 1899.
Tetraccra philippinensis Merr. in Philip. J. Sci., Eot. 9: 375. 3 014; En. Philip,
fl. PI. 3 : 58. 1923.
Tetratiera obliquinervia Elm. in Leafl. Philip. Bot. 7: 2621. 1915.
TYPES:—Tetracera jagifolia, Tetracera blamei: Blume s.n., Bantam, Java; holo-
type in L. — Tetracera rigida, Tetracera fagifolia i. wibintegerrima (lectotypc) : Blume
1734, Java; lectotype in L. — Tetraccra suwatrana: Teysmann 4454HB, Natar, Lam-
pung, Sumatra, December 29, 1857; holotype in U, isotypes in BO, CAL, L. — Tetracera
borneensis: Korthals s.»-., Pulu Lanipei, Borneo; holotype in L, isotypc in U. — Tetra-
cera, pkilippineims: Wenzel 812, Leyte, June 2, 1914; isotypes in A, BM, G, MO, US.
— Tetracera obliquinervia: Elmer 13862, Cabadbaran, Agusan Province, Mindanao,
September 1912; isotypes in A, BM, BO, CAL, E, FI, G, GH, K, MO, NY, U.
Scandent shrubs, up to 14 m high, much-branching; branches sparse-
ly strigose, glabrescent, slightly scabrid. Leaves elliptic to lanceolate,
(2.4—)6—20(—30) X (1.9—)3—1C(—13) cm, with (6—) 10—16(—23)
nerves on either side; apex rounded to acute, sometimes slightly acuminate;
base rounded to acute, sometimes asymmetric; margin entire to slightly
dentate; nerves curving upward, either ending in margin or following
margin closely until next nerve, anastomosing with the latter either direct-
ly or through major venation; leaves above rather dark green, shining to
dull, smooth to very slightly scabrid, very sparsely hirsute to glabrous
on intervenium, densely short-hirsute to glabrous on nerves and midrib',
beneath bright green, dull, smooth, sparsely hirsute to glabrous on
intervenium, strigose to very sparsely so on nerves and midrib; petiole
7—20(—30) mm, winged, 2—4 mm broad, densely hirsute, mainly on
central part, to glabrous above, more or less strigose beneath, ciliate at
margin. Inflorescences terminal, often on rather short axillary few-
leaved branches, 15—40 x 8—25 cm, 40—250-flowered, often with 1(—3)
small leaves in basal part; branches scabrid, densely to slightly hirsute
with small tufts of stellate groups of 0.2—0.4 mm long hairs, in larger
inflorescences only on extreme branches, basal branches of larger inflo-
rescences slightly strigose with single 1 mm long hairs; bracts caducous,
lanceolate, 3—7 X 1—2 mm, acute at apex, attached by broad base. Flow-
ers 8—12 mm across; pedicel 1—5 mm long, hairy like extreme branches
of inflorescences. Sepals 5(—6), 2 outermost ones 4 x 4 mm, 3(—4) inner
ones 5.5—7 x 4.5—5.5 mm, hirsute with hairs solitary to arranged in
small groups to glabrous, scabrid outside, inside sparsely to densely
sericeous for basal part extending to 1/4—1/2 of length and 1/4.—1/2 of
breadth on outermost one, for 1/3 to nearly whole sepal on innermost
one, with gradually increasing sericeous surface between. Petals 3, 6 x 4
mm, white. Stamens about 80, 4.5—6 mm long, white, the thecae distinctly
separated at apex, connective slightly emarginate between thecae. Carpels
3, ovoid, 2 X 1.5 mm, acute at apex, tapering into 1—3 mm long style, with
about 10 ovules. Fruits ovoid, 5—8 x 4—6 mm, tapering into 1 3 mm
long beak, with 2—3 capsules developed in each flower; capsules ovoid,
5—8 x 4—6 mm, tapering into 1—3 mm long beak, glabrous, glossy,
1(—2)-seeded. Seeds ovoid, 5 X 3 mm; aril one-sided, up to 7 mm longj
divided to 1/4— 1/2 of its length with broad lobes.
1953] Hoora-AND: Tetracera 221
Var. PAGIFOLIA.
Tctracem fagifolia var. fagifolia; Hoogl. in Fl. mal. I i: 148 f.2.* 1951.
Tetracera fagifolia Bl. — Tetracera rigida Bl. — Tetracera blumei Walp. —
. Tetracera sumatrana Miq. — Tetracera fagifoUa f. subintege-rrima Miq. — Tetracera
obovata Boerl. — Tetracera philippineusis Merr. — Tetraccra obliqllincrvia Elm.
Leaves elliptic to oblong, length about 1.4—2.25 x breadth, (4—)
7_20(—30) x (2.3—)5—10(—13) cm, with (7—) 12—16(—23) nerves on
either side; margin entire to slightly dentate; nerves curving upward,
ending in margin,
Var. BORNEENSIS (Miq.) Hoogl.
Tetracera fagifolia var. borneensis (Miq.) Hoogl. in Fl. mal. I 4: 148. 1951.
Tetracera bomee-nsis Miq.
Leaves elliptic to lanceolate, length about 1.4—3.5 x breadth, (2.4—)
6—13(—18) X (1.9—)2.7—5.5(—6.5) cm, with (6—) 8—10(—14) nerves on
either side; margin entire; each nerve following the margin closely
until it meets the next one, anastomosing with the latter either directly
or through major venation.
F I G . 1 0 . Tetracera fagifoUa B l . : v a r . fagifoUa ( 9 ) , v a r .
borneensis (Miq.) Hoogl. ( + ).
DISTRIBUTION.—Variety fagifoUa in Sumatra (also Simalur and Sibe-
rut), Malay Peninsula (Johore only), West Java, Borneo, and Philippines;
variety borneensis in Sumatra, Malay Peninsula (Singapore only), Banka,
Borneo, and Celebes. Variety fagifoUa is the most common form in
Sumatra, variety horneensis in Borneo.
222 R E I N W A R D T I A [VOL. 2
ECOLOGY.—Climber in primary forest, scrub, or bamboo forest, variety
fagifolia only known from primary forest, 100—750 m altitude, variety
bomeensis on Mt. Kinabalu up to about 1300 m, but from here known
sterile only.
VERNACULAR NAMES.—S u m a t r a: alor ampaleh (Simalur), ampalu
riembu (Lampung), sapbet (Siberut) . J a v a : aroy (ki) assahan, kiassahan,
ki saun-(Sundanese). P h i l i p p i n e s : balau-balau (Mbo). All these ver-
nacular names have been noted with specimens belonging to variety
fagifolia.
The two varieties show a marked difference in general appearance.
Generally specimens can be easily located, but there is a relatively small
number of intermediate specimens.
D o u b t f u l s p e c i e s
TETRACERA TRIPETALA Turcz.
Tetracera tripetala, Turcz. in Bull. Sec. Nat. Mosc. 36 (1): 547. 1863.
This species was described from a specimen collected by Horsfield
in Java. It is impossible to identify the species from the description; most
probably it is identical with Tetracera fagifolia El., of which species a
specimen collected by Horsfield in Java is present in the Calcutta her-
barium.
E x c l u d e d a n d i n v a l i d l y p u b l i s h e d n a m e s
Actaea aspera Lour. (Fl. cochinch. 332. 1790) is considered a synonym
of Tetracera scandens (L.) Merr. by Merrill (A commentary on Loureiro's
"Flora Cochinchinensis" in Trans. Am. phil. Soc. N.S. 24: 264. 1935). It is
excluded here: see this paper under T. asiatica (Lour.) Hoogl.
Calligonum asperum Lour. (Fl. cochinch. 342. 1790) likewise cited
by Merrill (1. c.) as a synonym of Tetracera scandens (L.) Merr., is also
excluded here: see this paper under T. asiatica (Lour.) Hoogl.
Tetracera aspera (Lour.) Eaeusch. (Nomencl., 3rd Ed., 147. 1797) is
based on Calligonum asperum Lour. (1790; see above) and, therefore,
excluded here. The binomial, which was not included in the "Index
kewensis," antedates Tetracera aspera (Aubl.) Willd. (Sp. PI. 2: 1241.
1799; Tigarea aspera Aubl., PI. Gui. fr. 2: 918. 1775) by two years. The
correct name for this South American species most probably is Tetracera
tigarea DC. (Syst. 1: 403. 1818).
Delima piripu DC. (Syst. 1: 408. 1818), based on Piripn Rheede (Hort.
malab. 7: 101 pi. 54- 1688), is no Dilleniacea, but Polygonum chinense L.
(cf. Hook. f. & Thorns., Fl. Br. Ind. 1: 62. 1855).
1953] HOOOLAND: Tetracera 223
Trachytella, DC. (Syst. 1: 410. 1818) with two species:
Trachytella actaea DC. (I.e.), baaed on Actaea aspera Lour. (1790);
see above.
Trachytella calligonum DC. (I.e.), based on Calligonum asperum Lour.
(1790); see above.
Tetracera juncea Hort. Angl. ex Steud. (Nomencl., 2nd Ed., 2: G70.
1821), included in the "Index kewensis," was published as a nomen nudum
only; as origin Australia was indicated.
Tetracera heyneana Wall. (Cat. no. 6630. 1828), included in the "Index
kewensis," was published as a nomen nudum only; the specimen is an
Euphorbiaceous plant.
Traxilisa Rafin. (Sylva tellur. 161. 1838), with one species:
Traxilisa aspera (Lour.) Rafin. (op. cit. p. 162), based on Calligonum
asperum Lour. (1790) ; see above.
Tetracera hygrophila Kurz (For. Fl. Br. Burma 1: 22. 1877), included
in the "Index Kewensis," was published as a nomen nudum only; the
specimen in the Calcutta herbarium, which is sterile, probably represents
Tetracera indica (Houtt. ex Christm. & Panz.) Merr.
Tetracera borneensis Auct. (non Miq.) ; Rolfe (in J. of Bot. 23: 209.
1885; Vidal, Revis. PI. Filip. 36. 1886), described after the specimen.
Vidal 940 from Luzon, represents Dillenia luzoniensis (Vidal) Martelli ex
Dur. & Jacks.
Trachytella, aspera DC. was cited by Forb. & Hemsl. (in J. Linn. Soc,
Bot. 23: 22. 1886—8) as a synonym under Tetracera sarmentosa (L.) Vahl.
Tetracera corymbosa Lignier & Bey (in Bull. Soc. linn. Normandie
V 5: 163. 1902), included in "Index kewensis," was published as a nomen
nudum only; New Caledonia.
Tetracera graudiuscida F. Muell. & Tate ex Dur. & Jacks. (Ind. kew.
Suppl. 1: 422. 1906, error). The binomial intended is Teucrium grandius-
culumF. Muell. & Tate (TO Trans. Proc. roy. Soc. S. Austral. 13: 108. 1890).
INDEX
New names and the final members of new combinations are in bold face type.
Actaea aspera 198, 222.
Akara-Patsjoti 208.
Assa 187, 205, 206; exotica 205, 206;
Calophyllum akai'a 208.
Curatella 188.
Davilla 188; hirsuta 194.
indica 187, 205-207; tctragynia 206. Delima 187, 188, 193, 196; aspera 190,
Calligonum asperum 198, 222, 223. 191; frangulaefolia 190, 191; hebecar-
224 R E I N W A R D T I A [VOL. 2
pa 190, 191; intermedia 190, 191; laevis
199; piripu 222; sarmentosa Auct.
194; sarmentosa L. 187, ISO, 191, 193;
aarmentosa var. fi 191; sarmentosa var.
glabra 193-195; sarmentosa var. hebe-
carpa 191; sarmentosa /. hirsutior 193-
195; scandens 191; tripetala 190, 191.
Delimopsis 187; hirsuta 187, 193-195.
Doliocarpus 188.
Eleiastis 187; laevis 187, 205.
Eugenia malaecensis 20G.
E u r y a n d r a 187; scandens 187, 212, 213.
Fructus indicus sarmentosus 193, 196.
Punis urens aspera 190.
Funis urens glabra 205.
Korosvel 187.
Leontoglossum 187; scabrum 187, 193-195.
Roehlingia 187; suaveolens 187, 208.
Seguieria asiatiea 193-195, 198.
Tetracera 186, 188.
Sect. Delima 188; Empedoclea 188;
Eutstracera 188.
a k a r a 187, 188, 190, 199, 208-210;
arborescens 189, 190, 217-219; asiatiea
188, 189, 193-198; asiatiea sap. anda-
manica 195, 196; asiatiea ssp. asiatiea
187, 195-197; asiatiea sup. sumatrana
187, 195-197; asiatiea «sp. zeylanica
187, 196; aspera (Lour.) Raeusch. 222;
aspera (Aubl.) Willd. 222; assa Auct.
208; assa DC. 205; assa var. 208;
assa var. loureiri 210; axiltaris 208,
210; billardiei'i 212, 213; blumei 219-
221; boerlagei 201, 203; borneensis
Auct. 199, 223; borneensis Miq. 219-
221; eorymbosa 223; cowleyana 201,
203; daemeliana 190, 212; diehotoma
205, 206; elmeri 214, 215; euryandra
Auet. 201, 218; euryandra Vahl 187,
190, 212, 213; everillii 201, 203; fagifo-
lia 188, 190, 200, 212, 215, 2T9-222;
fagifolia var. borneensis 221, 222; fa-
gifolia var. fagifolia 221, 222; fagifo-
lia /. subintegerrima 219-221; floribun-
da 201, 205; fragrans 210; glaberrima
188, 189, 198, 199; gracilis 205, 206;
grandis 215, 216; grandiuscula 223;
havilandii 215-217; hebecarpa 191;
heyneana 223; hirsuta 193, 195; hygro-
• phila 207, 223; indica 187, 190, 205-208,
210; juneea 223; kampotensis 210;
korthalsii 190, 211-213, 214, 215; kort-
halsii var. korthalsii 214, 215; kort-
halsii var. subrotunda 215; laevigata
217, 218; laevis Auct. 208; laevis Vahl
199, 205, 206, 210; lanuginosa 189, 200,
201, 219; levinei 194, 195; loureiri
190, 210-212; lucida 218; lucida var.
lanuginosa 218; macrocarpa 215;
macrophylla 190, 215-217; maingayi
188, 189, 199, 200; malabarica 205, 206;
moluccana 201, 203; monocarpa 190,
191; nordtiana 189, 193, 201-205, 219;'
nordtiana var. celebica 203-205; nord-
tiana var. everillii 203, 204; nordtiana
var. louisiadtca 202, 204; nordtiana
var. moluccana 203-205; nordtiana i w .
nordtiana 202-205; nordtiana var. wu-
thiana 202-205; obliquinervia 220, 221;
obovata 220, 221; philippinensis 220,
221; pilophylla 201, 202; radula 215,
216; rheedii 208; rigida 219-221; sar-
mentosa Auct. 194; sarmentosa (L.)
Vahl 190; sarmentosa vcr. hebecarpa
191: sarmentosa var. hirsuta 194; sar-
mentosa var. loureiri 210; scaberrima
198, 215, 216; seabricaulis 215-217;
scandens Auct. 194; scandens (Forst.)
Gilg 213; seandens (L.) Merr. 187-190,
191-193, 198, 219; seandens var. hehe-
carpa 191; sericea 208; setigera 198;
subcordata 217, 218; subrotunda 214,
215; subrotundata 214; sumatrana
Auct. 199; sumatrana Miq. 219-221;
sylvestris 208; teysmannii 215, 216;
tigarea 222; trigyna 205, 206; tripe-
tala 222; volubilis Auet. 191, 201;
volubilis L. 187; wahlbomia 205;
wuthiana 201, 202.
Tigavea aspera 222.
Trachytella 223; actaea 223; aspera 223;
calligonum 223.
Tragia scandens 190, 191.
Traxitisa 223; aspera 223.
Valhomia 187.
Wahlbomia 187; indica 187, 20S, 206.
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224