Untitled




280
R E I N W A R D T I A

[VOL. 4

272; pteridifolia PresL 276; truneata
Willd. 261, 271.

Platyzoma R. Br. 257.
Polypodium dichotovvum Thunb. 261, 277;

glaucum, Thunb. 261; lineare Burm.
261, 275.

Subfam. Sticherioideae Nakai 260.
Sticherus Presl 258, 259, 260, 261, 266;

caudatus Copel. 270; habbemensis
Copel. 268; hirtus var. candidus Nakai
269, var. virescens Nakai 270; lami-
anus Copel. 269; myriapoda Nakai 271;
perpaleaceus Copel. 272; pinnatus Co-
pel. 267, 270; jmlcher Copel. 271.

Stromatopteris Mett. 257, 260, 261; mo-
niliformis Mett. 261.

Subfam. Stromatopteroideae Nakai 261.

ADDENDUM

In Gleichenia subgenus Diplopterygium the following species must

be placed:
Gleichenia deflexa Holtt., sp. nov.

Rami rhacheos 120 cm vel ultra longi, maximi 35 cm lati, rami mi-
nores (steriles) interdum 18 cm lati; pinnulae omnes angulo c. 75° deflexae;
pinnulae maximae 16—20 cm longae, 3.5—3.8 cm latae, costae 4—5 cm
inter se distantes (costae ramorum minorum sterilium 2.5—3.5 cm inter se
distantes) ; costulae 4—5 mm inter se distantes, medii rectangulariter
patentes, infimae interdum leviter deflexae; lamina tenuis, segmenta c.
20 paria basi leviter constricta (i.e., supra basin ampliata), ala angustissima
laminae conjuncta, segrnentum infimum foliolum disjunctum constatum;
venae utroque basi prominentes, cetera planae; sori sporangiis 3—5 con-
stati; f oliola stipulif ormia.ad 4 cm longa, bipinnatif ida, segmenta 1 mm lata.
Paleae apicis rhacheos c. 10 mm longae, haud 1 mm latae, pallidae, margine
brunneo setis brevibus obliquis nitidis praeditae; rami rhacheos costaeque
utroque primo paleis parvis et pilis stelliformibus coarctis paleis elongatis
angustis intermixtis vestiti, demum plus minusve glabri et verruculosi;
venae infra pilis stelliformibus pallidis adspersis praeditae.

NEW GUINEA. P a p u a . Fergusson Island, 800 m, climbing to 6 m in rather
open rain forest in steep ravine, Brass 27171 (L, typs). Normanby Island, Mt Pali-
nama, 850 m, scrambling to 7 m in tall mossy forest, Brass 25736 (L).

R E I N W A R D T I A

Published by Herbarium Bogoriense, Kebun Raya Indonesia
Volume 4, Part. 2, pp. 281-310 (1957)

THE GENUS FIRMIANA MARSILI (Sterculiaceae)

A. J. G. H. KOSTERMANS *

SUMMARY

1. Firmiana Marsili and Erythropsis Lindley are congeneric.
2. Firmiana Marsili links Sterculia L. and Scaphium Schott & Endl.
3. Eight species are recognized: Firmiana eolorata (Roxb.) R. Br.;
F. diversifolia A. Gray; F. fidgens (Wall, ex King) Corner; F. hainanensis
Kosterm.; F. major Hand.-Mazz.; F. papuana Mildbr.; F. philippinensis
Kosterm.; F. simplex (L.) W. F. Wight.
4. Seventeen binomials are referred to other genera.

While a paper on Firmiana was in the press, I received some addition-
al, important information from the Kew Herbarium and from Mr. J. E.
Dandy of the British Museum. These "Additional Notes" were placed at.
the end of the article.

In a subsequent letter that reached me when the article already had
been printed and issued**, the problem of Firmiana simplex was solved,
thanks to the tenacity of Mr. Dandy. Moreover, F. fulgens came into
flower in the Bogor Botanical Garden, which enabled me to add some
field notes and photographs.

We had in mind to incorporate the paper in "Reinwardtia" in its
original form, but due to the additions, mentioned above, it was considered
better that it should be rewritten to this new, separate publication.

F I R M I A N A * Marsili
Firmiana Marsili in Saggi Scientifici e Letterari dell' Accademia Padova 1:

106-116, tab. 1 & 2. 1786; Lamarck-Podret, Encycl. meth. 7: 432. 1806 (as a syn.
under Sterculia platanifolia L.f.; excl. syn. Culhamia Forsk.) ; Steudel (Firmiana
Medic.), Nomencl. 343. 1821; ed. 2, 1: 642. 1840 (as a syn. cf Sterculia L.) ; DC,
Prodr. 1: 481. 1824 (as a syn. of Sterculia L.); Bartling, Ordin. 340. 1831 (as a syn.
of Sterculia L.) ; Schott & Endlicher, Melet. bot. 33. 1832; Spach, Hist. Veg. phan.

' D. Sc.j Botanist, Forest Service of Indonesia; cooperator Herbarium Bogoriense.
* Published under the title "The Genus Firmiana Marsili" as Communication

no. 54̂  of the Forest Research Institute, Indonesia, in December 1956, p. 3-33.
*** Named for count K. J. von Firmian, born 1716 at Deutsch-Metz in Tirol, died

1782 Milan, an Austrian statesman who was Governor of Lombardy under Maria
Theresia.

— 281 —



282 R E I N W A R D T I A [VOL. 4

3: 516. 1834; Meissner, Gen. 31 (25). 1837; Reichenbach, Handb. 291. 1837; Nomencl.
202. 1841; R. Wight, Illustr. Ind. Bot. 1: 77. 1840; Endlicher, Gen. 994. 1840 (as a
submenus of Sterculia L.); Suppl. 4 (3): 61. 1857; Enchir. 516. 1841 (as a submenus
of Sterculia L.); R. Brown in Bennett & Brown, PL Jav. rar. 235. 1844; Walpers,
Rep. 5: 104. 1846; Lindley, Veg. Kingd., ed. 3: 362. 1853; Miquel. PI. Nederl. Ind. 1
(2) : 178. 1859; Bentham in Benth. & Hook, f., Gen. PI. 1: 218. 1867 (as a subgenus of
Sterculia L.); Baillon, Hist. PI. 4: 60. 1872 (as a section of Sterculia L.; excl. syn.
Carpophyllum Miq.); Pfeiffer, Nomencl. 1 (2): 1353. 1874; Masters in Hook, f., Fl.
Brit. Ind. 1: 359. 1874 (as a section of Sterculia L.); Pierre, Fl. for. Cochinch. 3,
fasc. 13: t. 198. 1889 (as a syn. of Sterculia L.); Schumann in Engl. & Prantl, Nat.
Pfl. Fam. 3 (6) : 97. 1895 (excl. syn. Hildegardia Schott & Endl. and Scaphium Schott &
Endl.); Koorders & Valeton in Meded. Lands PI. tuin Buitenz. 14: 160. 1895; de Dalla
Torre & Harms, Gen. 313. 1903; Hutchinson & Dalziel, Fl. W. Trop. Afr. 1: 251. 1928;
Alston in Trimen, Handb. Fl. Ceylon 6 (Suppl.) 31. 1931; Lemee, Diet, descr. 3: 126.
1931 (excl. syn.); Kanjilal & Das, Fl. Assam 1 (1) : 154. 1934; Ridley in Kew Bull.
1934: 214 (as a syn. of Erythropsis Endl.); Gamble, Fl. Madras 1: 106. 1935; Shun
Ching-Lee, For. Bot. China 824. 1935; Corner, Ways. Trees Mai. 1: 610. 1940; Little,
Checklist Trees U.S.A. 185. 1953; Kostermans in Bull. Jard. bot. Bruxelles 24: 335-338.
1954; Comm. For. Res. Inst. Bogor 54: 3 - 33. 1956. — Erythropsis Lindley in Brandes,
Quart. J. Sci. Lit. Arts 1: 112. 1827; in Bot. Reg. t. 1236. 1829; Veg. Kingd., ed. 3:
362. 1853 (as a syn. of Firmiana Marsigli); Schott & Endlicher, Melet. 33. 1832;
Spach, Hist. Veg. phan. 3: 517. 1834; Meissner, Gen. 31. 1837; R. Wight, Illustr.
Ind. Bot. 1: 77. 1840; Endlicher, Gen. 994. 1840 (as a subgenus of Sterculia L.),
Enchir. 516. 1841 (as a section of Sterculia L.); Suppl. 5: 61. 1850; Steudel, Nomencl.
ed. 2, 1: 597. 1840; Reichenbach, Nomencl. 202. 1841; Bentham in Benth. & Hook, f.,
Gen. 1: 218. 1867 (as a syn. of the subgenus Firmiana Mars.); Pfeiffer, Nomencl. 1
(2): 1259. 1874; Boerlage, Handl. Fl. Ned. Ind. 1: 123. 1890 (as a syn. of Sterculia
L.); Ridley, Fl. Mai. Pen. 1: 277. 1922; in Kew Bull. 1934: 214-217 (Erythropsis
Endl.); Lemee, Diet, descr. 3: 15. 1931 (excl. descript.); Burkill, Diet. econ. Prod.
Mai. Pen. 1: 950. 1935; Adelbert in Backer, Fl. Java, (Nooduitg.) 4b, Fam. 107: 25.
1944; Kostermans in Bull. Jard. bot. Bruxelles 24: 335-338. 1954. — Clompanus
[Rumphius] 0. Kuntze, Rev. 1: 77. 1891. — Karaka Rafinesque, Sylv. Tellur. 72.
1838; Merrill, Index Rafin. 167. 1949. (p.p.).

TYPE SPECIES.—Firmiana, simplex (L.) W. F. Wight.

Trees. Leaves often cordate, glabrous or with stellate hairs, with
long petioles. Flowers often in coralliform panicles, brightly coloured,
covered with stellate hairs, unisexual (perhaps sometimes polygamous);
calyx with a more or less developed tube with nectarial disc at base inside;
corolla none. Androgynophore growing out of the tube after anthesis;
androecium consisting of usually 10 shortly stipitate or sessile, 2-celled
anthers opening longitudinally; the filaments attached to the margin of
the sunken top of the androgynophore; ovaries 5 (enveloped by the
stamens), conglutinate; styles short, stigmas curved outwards. After
anthesis the ovaries soon separate and expand, opening in an early state
of development, one or two pairs of ovules at the margins. Male and
female flowers differ only slightly in size and in development of the

1957] A. J. G. H. KOSTERMANS : The genus Firmiana 283

androecium or gynaecium. Follicles papery, dispersed with the adhering
seeds.

DISTRIBUTION.—Asiatic continent, Malaysia, Pacific Islands.

The genus was originally included in the older genus Sterculia L.
(Sp. pi. 1007. 1753; Gen. pi., ed. 5: 438. 1754). Created as a separate genus
by Marsili (sometimes wrongly spelt Marsigli) in 1786, Firmiana has
been alternately included in Sterculia L. and reinstated again as a proper
genus. Endlicher (Gen. pi., I.e.) considered Firmiana Mars., like Erythrop-
sis Lindley, a subgenus of Sterculia L.; in his Suppl. (I.e.), he adopted
R. Brown's view in accepting it as a proper genus and dividing it into two
sections: Eufirmiana Endl. (type species: Sterculia platanifolia L.f.) and
Erythropsis (Lindl.) Endl. (type species: Sterculia colorata Roxb.), dif-
fering in the size of the flower tube in comparison to the lobes. This
subdivision was generally accepted, but not the status of Firmiana. —
R. Brown (1844) recognized Firmiana as a proper genus; Bentham (1867)
reduced the genus again to a subgenus, herein followed by Baillon (1872),
Masters (1874) and Brandis (1874). — Pierre (t. 198. 1889), although
enumerating the specimens under Sterculia L., emphatically stated that
Firmiana was a proper genus. Schumann (1893) reinstated Firmiana,
incorporating Hildegardia and Scaphium. into that genus.

Ridley (1922 and 1934) not only considered Firmiana a good genus
but also Erythropsis, a view formerly presented by Pierre.

Lemee (Diction, descript. 3: 15. 1931) gives a wrong description of
Erythropsis Lindl., which he confused with Pterygota.

The present author (1954) accepted Firmiana as a proper genus;
Erythropsis was considered to be congeneric with it.

Firmiana is closely related to Sterculia and may be differentiated
from that large genus only by its membranous fruit, which is dispersed
with the adhering seeds, whereas in Sterculia the follicles are leathery
and the seeds are dispersed separately; furthermore the inflorescence is
often different.

On the other hand Firmiana. is also related to Scaphium and Ptero-
cymbium, the difference being in the usually 2-seeded fruit (the seeds
are marginal) of the former, as contrasted with the basal, one-seeded
fruit of the latter. In this sense Scaphium and Pterocymbium are closer
to Hildegardia of which the fruit does not open at all and reminds one
of a samara, as found in Tariettia and Heritiera.

In flower characters the above-mentioned genera cannot be separated.
Saunders (in Ann. Bot. 45: 97-103. 1931), after examining the vas-

cular bundles, came to the conclusion that the calyx actually represents



284 R E I N W A R D T I A [VOL. 4

a combination of calyx and corolla. Her ingenious hypothesis that each
carpel represents actually1/2 -l-1/2 carpel, which would explain the peculiar
venation, is however not in agreement with the position of the ovules.

The position of the radicles, often serving to point out affinity of
genera, is so variable in Stercniliaceae that it is not of much use (see
Pierre). Culhamia Forskal (Fl. Aeg.-Arab. 96. 1775), formerly included
in Firmiana Mars., was relegated to Cola by Pfeiffer (Nomencl. 1: 289.
1870), although not definitely. Kuntze (Rev. 1: 78. 1891) remarks that
the shape of the leaf and the inflorescence, as described by Forskal,
point to Firmiana.

Rafinesque (Sylv. Tellur. 73. 1838, descr. op. cit. 158. 1838) coined
the name Culhamia triloba. Merrill (Index Rafin. 167. 1949) considers this
species to be identical with StercuUa setigera Delile.

In his lengthy article *, Marsili discussed the tree, flowers and fruit
of a specimen of F. platanifolia grown in the Botanical Garden of Padua.
On page 116 he gives a Latin diagnosis of the genus. The two drawings
are exceptionally good, the first one represents an inflorescence accom-
pained by drawings of male and female flowers, the second one depicts the
young and opened fruit.

In a note on page 116 he remarks that StercuUa platanifolia L.f.
(Suppl. Plantarum 423) is the same species. Linnaeus fil. received a speci-
men from Alstroemer, who in turn got it perhaps from Marsili; Alstroe-
mer brought it to Sweden. The contention of Linnaeus fil. that the flowers
are hermaphroditic, is, according to Marsili, attributable to the poor state
of the dried material available to Linnaeus fil.

Marsili apparently did not intend to describe a species or to procure
a specific epithet, but his drawings manifestly represent F. platanifolia
,and he mentioned StercuUa platanifolia as being identical with his plant;
hence I follow the current custom to attribute the combination to Marsili.

Much controversy exsists in the matter of the sexes. In a recent Flora
(Kanjilal) the flowers are considered hermaphroditic, although most
botanists are of the opinion that they are unisexual. In all species I could
examine there are male and female flowers in the same inflorescence.
In the male flowers we find minute, sterile ovaries. The anthers in the
female flower are not smaller (sometimes even larger) than those in the
male flower, but they do not open. I have not been able to make sure
whether their pollen are viable.

The number of anthers indicated by different authors varies between

* I obtained a photographic reproduction through the courtesy of Prof.
C. G. G. J. van Steenis.

Dr.

1957] A. J. G. H. KOSTERMANS: The genus Firmiana 285

10 and 30. I suppose that there are only 10 anthers, each with two cells,
As in other Sterculiaceae the anther cells are either parallel, which results
in a regular ring of anthers, or the cells are not at the same level and
often one is turned in another direction than the other, (upwards or
downwards), which results in an irregular globular clump of cells, some-
what resembling a knot of wool.

KEY TO THE SPECIES

la. Carpels and follicles densely pilose . 1 2
b. Carpels and follicles glabrous or nearly so 4

2a. Leaves entire 3
b. Leaves deeply lobed 6. F. simplex

3a. Lower leaf-surface glabrous 3. F. philippinensis
b. Lower leaf-surface densely pilose (matted) 7. F. papuana

4a. Calyx lobes very short, erect or incurved 5
b. Calyx lobes long, reflexed, lanceolate-oblong 6

5a. Lobes Vs of the tube length. Lower leaf surface softly pilose. Flower tube inside
and androgynophore glabrous 2. F. fulgens

b. Lobes % of the tube length. Lower leaf surface glabrous, but for the axils of the
main nerves. Flower tube inside and androgynophore sparsely pilose. 1. F. colorata

6a. Leaves lobed; lower surface grey, densely pilose (matted); follicle up to 15 cm
long. Leaf-base cordate 4. F. major

b. Leaves entire; lower leaf surface densely pilose (matted), follicles up to 7,5 cm
long. Leaf base truncate or subcordate 8. F. hainanensis

c. Leaves entire or lobed. Lower leaf surface practically glabrous. Follicles up to
7 cm long 5. F. diversifolia

1. FIRMIANA COLORATA (Roxb.) R. Br.—Fig. 1, 2, 3

Firmiana colorata (Roxb.) R. Brown in Bennett & Brown, PI. Jav. rar. 235.
1844 (quoad var. a ) ; Walpers, Rep. 5: 104. 1845-46, p.p.; Miquel, Fl. Ind. bat. 1 (2):
178. 1859 (excl. var. ft (?) et y et spec. Korthalsii) ; Thwaites, Enum. 29. 1864 (nomen);
Trimen, Fl. Ceylon 1: 166. 1893 (as a syn. of StercuUa colorata Roxb.); Masters in
Hook, f., Fl. Brit. India 1: 360. 1874 (as a syn. of StercuUa colorata Roxb.) ; King
in J. As. Soc. Bengal 60 (ii) : 71. 1892; K. Schumann in Engl. & Prantl, Nat. Pfl.
Fam. 3 (6): 97. 1893; Koorders & Valeton in Meded. PI. tuin Buitenzorg 14: 160.
1895 (excl. spec. Korthalsii); Atlas t. 406. 1914; Koorders, Exkurs. Fl. Java 2: 598.
1912; Koorders-Schumacher, Syst. Verz., Fam. 178: 20. 1913; Gagnepain in Lecomte,

1- gen. Indoch. 1: 459. 1911 (as a syn. of StercuUa colorata Roxb, excl. var.); Backer,
bcnoolfl. Java 1: 135. 1911 (as a syn. of StercuUa colorata Roxb.); Gamble, Fl.

1 S 1 T 1 : 1 0 7 ' 1 9 3 5 ; M e r r i U i n P h i l i P P - J - S c L * 4 : 2 4 6 - 1 9 1 9 ; Enum. Born. P L 380.
9^1 (quoad nomen tantum); in Lingnan Sci. J. 13: 63. 1934; Adelbert in Backer,

• Java, (Nooduitg.) 4b, Fam. 107: 25. 1944 (as a syn. of Erythropsis colorata (Roxb.)
irk.) ; Kosterm., Comm. For. Res. Inst. Bogor 54: 7-16, /. 1, 2, 3. 1956. — StercuUa

oiorata Roxburgh, PL Coromand. 1: 26. 1795 (quoad tab. 25, descript. excl.) ; Hort
eng. 5. 1814 (nomen); Fl. Ind. 3: 146. 1832 (reprint 3: 507. 1874), quoad nomen;

wumenow, Sp. pi. 2: 873. 1799; Lamarck-Poiret, Encycl. meth. 7: 432. 1860, p.p.;



286 REINWARDTIA [VOL. 4

Persoon, Synops. 2: 240. 1807; Steudel, Nomencl. 814. 1821; ed. 2, 2: 639. 1841; DC,
Prodr. 1: 483. 1824, p.p.; Sprengel, Syst. Veg. 3: 83. 1826, p.p.; G. Don, Gen. Hist.
1: 517. 1831; Schott & Endl., Melet. 33. 1832 (as a syn. of Erythropsis roxburghiana
Schott & Endl.); Spach, Hist, Veg. phan. 3: 516. 1834 (as a syn. of Erythropsis rox-
burghiana Schott & Endl.), p.p.; Wight & Arnott, Prodr. 63. 1834; in Hooker's Icon.
2: t. US. 1837; R. Brown in Bennett & Brown, PI. Jav. rar. 235. 1844 (as a syn. of
Firmiana colorata R. Br., quoad var. a ) ; Moritzi, Verz. 28. 1845-46 (nomen) ; Hasskarl
in Tijdschr. Nat. Gesch. 12: 115. 1845 (nomen); Dalzell & Gibson, Bombay Fl. 23.
1861 (n.v.); Thwaites, Enum. 29. 1864 (nomen); Bentham in Benth. & Hook., Gen.
pi. 1: 218. 1867; M. R. Brown, PI. Wild Fl. S. & W. India tl 8 (n.v.) ; Beddome, Fl.
sylv. 32. 1869 (n.v.); Pfeiffer, Nomencl. 1 (2): 1353. 1874; Masters in Hook, f., Fl.
Br. Ind. 1: 359. 1874; Brandis, For. Fl. 34, 1874 (excl. syn. Sterculia wallichii Falc.) ;
Ind. Trees 84, fig. 40, 190,6, p.p.; Kurz in J. Asiat. Soc. Bengal 43 (ii) : 117. 1874;
For. Fl. Burma 1: 138, 1877; Pierre, Fl. for. Cochinch. 3, fasc. 13: t. 199. 1889;
King in J. Asiat. Soc. Bengal 60 (ii) : 71-72. 1892; Watt, Diet. 6: 361. 1893 p.p.;
Talbot, Trees Bombay 22, 1894 (n.v.); For. Fl. Bombay 1: 141-142. 1909; Woodr. in
J. Bomb. Nat. 11: 129. 1897; Koorders & Valeton, I.e. 160. 1895 (as a syn. of Firmiana
colorata Mars.); Trimen, Handb. Fl. Ceylon 1: 166. 1893; id. 6: (Alston): 31. 1931;
Cooke, Fl. Bombay 1: 125. 1903; Backer, Schoolfl. Java 1: 135. 1911; Gagnepain in
Lecomte, Fl. gen. Indoch. 1: 459. 1911 (var. bracteosa Gagn. excl.); Tardieu-Blot,
Suppl. 1: 401. 1945; H. A. Clerx in Trop. Natuur 1: 10-13. 1912, cum fig.; Haines,
Fl. Bihar & Orissa 2: 76. 1921; Gamble, Man. Ind. Timb. 96. 1922; Ridley in Kew
Bull. 1934: 215 (as a syn. of Erythropsis colorata Ridley) ; Parkinson, For. Fl. Andam.
Isl. 101, fig. 16. 1923; Craib, Fl. Siam. Enum. 1: 166. 1925; Kanjilal, Kanjilal, &
Das, Fl. Assam 1 (1): 151. 1934; Gamble & Fischer, Fl. Madras 107. 1935; L. H.
Bailey, Stand. Cyclop. Hort. 3: 3239. 1947. — Erythropsis colorata (Roxb.) Burkill
in Gard. Bull. S.S. 5: 231. 1931; Ridley in Kew Bull. 1934: 215; Adelbert in Backer,
Fl. Java, (Nooduitg.) 4b, Fam. 107: 26. 1944; Mooney, Suppl. Haines, Bot. Bihar
& Orissa 28. 1950 (quoad nomen tantum). — Clompanus colorata (Roxb.) 0. Kuntze,
Rev. Gen. pi. 1: 78. 1891. — Karaka colorata, (Roxb.) Rafinesque, Fl. tellur. 72. 1838;
Merrill, Index Rafin. 167. 1949. — Erythropsis roxburghiana Schott & Endl., Melet.
33. 1832; Spach, I.e. 517, p.p.; Sprengel, Syst. veg. 3: 83. 1826; Spach, Hist. Veg.
phan. 3: 517. 1834; Steudel, Nomencl. ed. 2, 1: 597. 1840 et ed. 2, 2: 639. 1841 (as a
syn. of Sterculia colorata Roxb.) ; Voigt, Hort. Suburb. Calcut. 104. 1845; Masters,
I.e. (as a syn. of Sterculia colorata Roxb.); Koorders & Val., I.e. (as a syn. of Firmiana
colorata R. Br.). — Tab. 25 in Roxburgh, PI. Corom. 1.

Erythropsis colorata, var. bracteata (A. DC.) Ridley in Kew Bull. 1934 :215. —
Firmiana bracteata A. DC. in Bull. Herb. Boiss., ser. 2, 3: 369. 1903. — Balansa
3743 (G).

Sterculia rubicunda Wall., Catal. 1119 D, F, G, ex Masters in Hook, f., Fl. Brit.
Ind. 1: 360. 1874. — Wallich 1119 D, F. G. (K).

Sterculia fulgens (non Wall.) Kurz in J. Asiat. Soc. Beng. 43 (ii) : 117. 1874
(p.p.) ; For. Fl. Br. Burma 1: 139. 1877 (p.p.) ; King in J. Asiat. Soc. Beng. 60 (ii) :
72. 1891 (p.p., quoad specim.) ; Kostermans, Comm. For. Res. Inst. Bogor 54: 11.
1956 (p.p.). — Erythropsis fulgens (Wall, ex Mast.) Ridley, Fl. Mai. Pen. 1: 277.
1922 (p.p., quoad specim.) ; Burkill, Diet. econ. Prod. Mai. Pen. 1: 950. 1935 (p.p.,
quoad specim.). — Firmiana fulgens (Wall, ex King) Corner, Wayside Trees Mai.
1: 610. 1940 (p.p., quoad specim.).

1957] A. J. G. H. KOSTERMANS : The genus Firmiana 287

PIG. 1. Firminia colorata (Roxb.) R.Br. — Flowering branch (photo: Huysmans, Bogor).



288 R E I N W A R D T I A
[VOL. 4

FIG 2. Firmiana eolorata (Roxb.) R.Br. - Different leaves (X 0.5). - After living
material.

1957] A. J. G. H. KOSTBRMANS : The genus Firmiana 289

Tree up to 25 m high with smooth grey or grey-green bark. Buttres-
ses up to 2 m high, out 50 cm. Branches glossy dark brown. Branches
and the grey or greenish grey branchlets with many lenticels. Terminal
bud rather flat, surrounded by densely rusty pilose, ovate-triangular,
acute, deciduous bud-scales measuring 1-2 mm. Petiole 10—20 cm long,
glabrous, base pulvinate. Leaf-blade glabrous (axils of main nerves pilose
below; in very young leaves scattereed stellate hairs on lower surface),
variable in size and shape, entire or deeply incised, ovate, 6-25 cm long,
6—20 cm wide; top acuminate, base cordate or subcordate; palmately
veined; dark green on upper, paler green on lower surface; primary
veins slightly raised on upper, raised on lower surface, pale; main ribs 5,
rarely 7, arcuate; secondary nerves lax, at right angles from primary
nerves, pale; reticulation rather dense.

PIG. 3. Firmiana eolorata (Roxb.) R.Br. — (From left to right) Female flower
before anthesis; female flower after anthesis; male flower before anthesis; male
flower after anthesis; top of androgynophore of female flower before anthesis; young
follicle (below); young follicles; (below) top of male androgynophore (all parts slightly

less than natural size; tops of androgynophore x 3.3). — After living material.

Flowers in axillary panicles or racemes from the axils of previous
year's fallen leaves (as a rule they appear when the tree is bare) ; main
peduncle stout, up to 7 cm long, yellowish orange; ramifications slender,
short (up to 1 cm long), coralliform. Flowers 3 cm long, densely covered
with minute orange stellate hairs and scales; calyx funnel-shaped, some-
what inflated towards apex, slightly curved; teeth triangular, acute,
5—6 mm long, with a conspicuous, flattened, grey-pilose margin; inside of
calyx with sparse stellate hairs; above the base a ring of glistening, long
bundles of strigose hairs. Androgynophore in mature flowers extending
5 10 mm beyond calyx, covered with very small scale-like stellate hairs.
Anthers 10, two-celled; the cells curved; filaments very short, attached
to the rim of the shallow tube at the top of the androgynophore. Ovaries
5, flask-shaped (at base knee-like), glabrous, white (red towards the top)
later turning pale green, merging into the short, curved, red styles; stigma



290 REINWARDTIA [VOL. 4 1957] A. J. G. H. KOSTERMANS : The genus Firmiana 291

bent outside, peltate, pellucid, reddish. Follicles up to 7 cm long, mem-
branous, glabrous, veined, stipitate, opening at an early stage; ovules
2-4, ellipsoid, glossy, black, alternate at the margins of the follicle.

The flowers are hermaphroditic in appearance but one sex apparently
remains sterile. The male and female flowers occur on the same branch,
In the male ones the anthers are smaller than in the female ones, but in
the latter the anthers do not open.

The anthers of the male flower already open when it is still closed.
After dehiscence of the flower the androgynophore stretches. In this
stage the disc of anthers is about 1.5 to 2 mm in diameter; the pollen
is smooth; the antherial disc is always bent to one side at an angle of
almost 90°. In the female flower the antherial disc is about 1.5 to 2 times
as large as that in the male flower; it is not bent (or hardly so) ; the
anthers contain pollen, but do not open, even after the stretching of the
androgynophore.

The styles and stigmas gradually turn red during their development,
like the top of the ovary.

The male flowers develop earlier than the female ones.
Burkill's specimen 6990 is accompanied by a note, that in many

flowers the flower tube is partly ripped open from base to top; apparently
this is done by birds looking for honey.

DISTRIBUTION.—Indian Peninsula, Burma, Ceylon, Andamans, Siam,
Malay Peninsula, Sumatra; introduced in Java.

USE.—The inner bark yields a fibre; twigs and leaves are used as
cattle fodder in the Western part of the Indian Peninsula. The wood is
dingy greyish white in colour, very soft, marked with conspicuous medul-
lary rays. The roots (tubers) are eaten by the Lepchas.

ECOLOGY.—The tree is frequent in deciduous forests; it is bare in
winter (dry season) ; in W. Java it becomes partly or completely leafless
before coming into bloom (October, November). In Burma it flowers
in March-April (Kurz, Brandis). In Ceylon it flowers in June-July
(Trimen).

LOCAL NAMES (these are probably applicable both to F. colorata and
F. fulgens. — India: Bodula, Walena, Samarri (Hind) ; Nustruk (Ajmere) ;
Sisi (Kol., Sonth) ; Khowsey (Mar., Berar) ; Maraka, Karu boppaji
(Tel.) ; Samari (Oudh) ; Kowsey, Bharkoi, Bheckhol, Samarri, Walena,
Bhai-koi (Bomb.) ; Bodala, Bodal (Kumaon) ; Mutruk (Merwara) ; Mula
(Beng.) ; Pisi, Sisi (Kol.) ; Bolazong, Bolajun, Chengsu, Walgem (Caro) ;
Sito udal, Phirphiri omra (Nepal) ; Kanklyem (Lepcha) ; Jhari-udal, Jari
udal, Kath udal (Assam) ; Sangkru-aromy (Mik.) ; Bohog-odla (Mech.) ;
Dieng-symphlor, Dieng-Sangkhlor (Kli) ; Lersima (Kharna) ; Burma:

Wetshaw, yaseng-shaw; Andamans: Berda; Siam: Paw kao (Loas, Chieng-
mai) ; Karaka (Telingas) ex Roxb., Fl. Corom.

INDIA. C h i t t a g o n g : Kodala Hill near Chittagong, Apr., fl., fr., Badul Khan
496 (CAL); Manbhum, fl., Rev. J. Campbell s.n. (CAL). A s s a m : South Lushai,
hills near Fort Lungleh, Apr., fl., Gage 111 (CAL); Naga Hills, ster., Anon. s.n.
(CAL). S i k k i m: locality not indicated, ster., Anderson s.n. (CAL, L). S o u t h
I n d i a : Shevaron Hills, fr., Perrottet 34 (CAL); Travancore, Santhanpora, Dec,
ster., Meebold 565 — 13257 (CAL) ; ibid., Feb., fl., Bourdillon 91 (CAL) ; Harsleyhondo-
Chittoor Distr., May, fr., Fischer 4647 (CAL).

LOWER SIAM. Pingah, Dec, fl., Haniff & Nur 3940 (SING) . A n d a m a n I s . :
South Andaman, fl., Kurz s.n. (CAL) ; Southpoint, fl., Kurz s.n. (CAL.).

MALAYSIA. — M A L A Y P E N I N S U L A . P a h a n g : near Jerantut, Feb.,
fl., Corner s.n. (SING). P e r a k : fl., Ridley 14640 (CAL, SING); Ulu Kuching, on
river bank, Mar., fl., King's Coll. 8673 (CAL, SING); Kuala Lumpur, Batu Rd., Mar.,
fl., Foxworthy 1709 (SING); ibid., Feb., fl., Burkill 4415 (SING); July, fl., Curtis
232,2 (CAL, SING). P e n a n g : Govt Hill, Jan., in bud, Curtis 144 (SING); Head of
Waterfall Gardens, Mar., fl., Burkill 6990 (SING); Ayer hitam, alt. 70 m, ster.,
Nov., Haniff s.n. (SING) ; Penara Bukit, Feb., fl., For. Guard s.n. (SING). K e l a n -
t a n : Sg. Ketah, Gua Nanah, Feb., fl., Nur 12131 (SING). S e l a n g o r : Kepong
Plantations, May, fl., Hussin F.D. 15360 (SING). Locality not indicated, fl., Scor-
techini s.n. (CAL, SING). — S U M A T R A . Hoch Ankola, fl., Junghuhn s.n. (L);
Ranau Lake, alt. 600 m, fl., Forbes 2105 (L); R. Warkau, alt. 600 m, fl., Forbes 2106
(CAL). — J A V A . W e s t J a v a : no loc, fl., Kuhl & van Hasselt s.n. (L); no loc,
fl., Zollinger ad 494 (L) ; Bantam, fl., Spanoghe s.n. (L) ; Bandung, alt. 700 m, fl.,
Sep., Popta 866 (L); Southcoast Sukapura, low nits, fl., Junghuhn s.n. (L).

CULTIVATED. Hort. Calc, fr., von Mueller s.n. (L) ; fl., Pierre 2849a (P).
Hort. Bogor. sub XI C 105, fl., (SING).

The original publication of Sterculia colorata Roxburgh consists of
a description and a coloured drawing. The description, which is slightly
emendated in Roxburgh's Flora, indicates the leaves as being broader
than long, villose with soft lobes and downy petioles. This part of the
description certainly refers to the species, which in our paper is called
Firmiana fulgens. The description of the flowers is insufficient to' make
out, whether the flowers are those of F. fulgens or of F. colorata, as
nothing is said about the length of the lobes of the flower tube.

Roxburgh's plate 25 (which is preserved in the Kew Herbarium, no.
46), depicts without doubt F. colorata with tubular flowers with short
lobes; a leaf is added in outline, which also belongs to F. colorata in our
sense, as it is not broader than long. Moreover, in the figure the flowers
are orange; those of F. fulgens are white.

Consequently Roxburgh's description covers a mixture of two species.
In order to prevent the abolishing of a well-established name for

a plant, currently interpreted as F. colorata, I adopt here Roxburgh's



292 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. KOSTERMANS: The genus Finniana 293

plate 25 as the basis of his Sterculia colorata, with exclusion of the des-
cription (at any rate with exclusion of the description of the leaves; the
flower description is so incomplete, that it may cover both species).

Through the courtesy of the Kew Herbarium, I received photographic
reproductions of a sheet, marked Sterculia, colorata R. Br. with the number
1119, the specimen on which R. Brown based his variety a of Firmiana
colorata. It consists of a leafy branch; a branch with fruit and a branch
with young flowers. This specimen is certainly Firmiana. colorata in our
sense. A second sheet consists of 4 branches with inflorescences and a
couple of loose inflorescence-branches and two loose follicles. The sheet
bears four labels, all different. The right hand top label gives the follow-
ing: 1119 Sterculia colorata Roxb. and three different habitat indications,
which make it evident that there are three different specimens, one
collected in Silhet in 1823, one in Hundnar in 1825, and one from Mungger
Hills (marked Herb. Calcutta). The original labels from Hundnar and
Silhet are present. The fourth label bears the numbers 1119 D Sterculia
colorata Hb. Wight; 1119 G Sterculia colorata Herb. Roxburgh; 1119 F.
Sterculia rubicunda Hb. Hamilton (Mt. Mongher).

A third sheet, marked Sterculia rubicunda, Mungger 20 July 1811
and 5 April 1811 with an added number 1119a represents Firmiana colo-
rata.

Sterculia wallichii Falconer, enumerated by Brandis under F. colorata,
belongs certainly to F. fulgens. The specimen is conserved in the Kew
Herbarium and consists of inflorescence bearing branches only.

Korthals' specimen from Mt. Sakumbang in Borneo, is preserved in
Leiden. It represents a young twig of Scaphium macropodum. This spe-
cimen was cited by Miquel and by Merrill.

Ridley (Flora Mai. Pen., I.e.) under Erythropsis fulgens gives a
description of the species, called F. fulgens in our paper. All material
from the Malay Peninsula (autographed by Ridley), however, belongs to
F. colorata with glabrous leaves. I have not seen a single specimen of
F. fulgens from the Malay Peninsula. F. fulgens does not occur in Java.

Brown's third variety King (I.e.) under Sterculia fulgens enumerated
represented in the British Museum, belongs to F. colorata. Forbes 2105
from Sumatra. This specimen is manifestely F. colorata with glabrous
leaves, contrarily to King's description,. The same holds true for the
specimens Scortechini s.n. and King's Collector 8673 from the Malay
Peninsula, I think Masters was right in indicating that F. fulgens occurs
in the tropical Western Himalayan; the species, occuring in Malay, Su-
matra and Java is F. colorata, with glabrous leaves.

The specimen Wallich 1135 represents F. fulgens.

Corner's (I.e.) description of F. fulgens covers indeed that species;
but his specimens from the Malay Peninsula all represent F. colorata with
glabrous leaves.

2. FIRMIANA FULGENS (Wall, ex Mast.) Corner — Fig. 4, 5, 6

Firmiana fulgens (Wall, ex Mast.) Corner, Ways. Trees Mai. 1: 610. 1940 (p.p.,
quoad descr.). — Sterculia fulgens Wallich, Cat. 1135 ex Masters in Hook, f., Fl.
Br. Ind. 1: 360. 1874; Kurz in J. Asiat. Soc. Beng\ 43 (ii) : 117. 1875, et For. Fl. Br.
Burma 1: 139. 1877 (quoad nomen et descr.) ; King- in J. Asiat. Soc. Beng. 60 (ii) :
72. 1891 (quoad descr., p.p.) ; K. Schumann in Engl. & Pr., Nat. Pfl. Fam. 3 (6) :
97. 1895; Haines, Bot. Bihar & Orissa 2: 77. 1921 (as a syn. of S. pollens). —
Clompanus fulgens (Wall.) O. Kuntze, Rev. 1: 78. 1891. — Erythropsis fulgens (Wall.
ex Mast.) Ridl., Fl. Mai. Pen. 1: 277. 1922 (quoad descr., p.p.); in Kew Bull. 1934:
215; Burk., Diet. econ. Prod. Mai. Pen. 1: 950. 1935 (quoad nomen). — Firmiana
colorata var. ft fulgens R. Brown in Benn. & Br., PI. Jav. rar. 235. 1844. — Wallich
1135 (K).

Sterculia colorata Roxburgh, PL Corom. 1: 26. 1795 (quoad descr., tab. 25
exclud.); Fl. Ind. 3: 146. 1832, (repr. 3: 507. 1874). — Firmiana colorata (Roxb.)
R. Brown in Benn. & Br., PL Jav. rar. 235. 1844 (quoad var. jj) ; Walpers, Rep. 5:
104. 1846 (p.p.).

Sterculia wallichii Falconer (non G. Don) ex Brandis, For. Fl. NW. & C. Ind.
34. 1874. — Coll. unknown no 26 (K).

Firmiana pallens F. von Mueller in Viet. Nat. 3: 48. 1866 (Combination indicated,
bat not made); (Wall, ex King) Kostermans, Comm. For. Res. Inst. Bogor 54: 16.
1956. — Sterculia pallens Wallich ex Voigt, Hort. suburb. Calc. 105. 1845 (nomen);
King in J. Asiat. Soc. Beng. 60 (ii) : 73. 1891; Hochreut. in Bull. Inst. Btzg. 19: 22.
1904 (nomen); Brandis, Ind. Trees 84. 1906; Haines, Bot. Bihar & Orissa 2: 77.
1921. — Erythropsis pallens (Wall, ex Voigt) Ridl. in Kew Bull. 1934: 215. — Fal-
coner 289 (K).

Tree up to 20 m high and 1 m in diameter (Haines). Branches thick,
densely pilose (matted stellate hairs with short horizontal branches) with
large round leaf scars. Leaves palmately veined, chartaceous or papyra-
ceous, more or less reniform (broader than long), up to 30 X 25 cm,
shallowly 3—4-lobed, the lobes acutish, base cordate; upper surface soon
glabrous, lower surface laxly pilose (stellate hairs with long, more or less
erect branches) ; veins flat above, prominent below; the 3 main ribs with
a 4 pairs of arcuate secondary nerves; tertiary nerves lax, somewhat
prominent reticulation inconspicuous. Petioles up to 15 cm long, densely
pilose (stellate hairs with erect, short branches), base swollen.

Flowers in panicles up to 25 cm long; peduncle and branches stout.
flowers white or pale yellowish, the female ones slightly smaller and more
slender than the male ones,or te reverse (both in the same inflorescence

and flowering at the same time), subcampanulate, 1.5—2.5 cm long, den-
Selly pilose (stellate hairs with long, erect branches) ; lobes 4—6 mm long,



294 REINWARDTIA [VOL. 4

-.?-). . i . 3 . . . . -

1957] A. J. G. H. KOSTEKMANS : The genus Firmiana 295

F I G . 4. Firmiana fulgens (Wall, ex Mast.) Corner — Flowering branch. — After
H. B. XV J. B. IX 2.

F I G . 5 Firmiana fulgens (Wall, ex Mast.) Corner. — Leafy branch and Dart of
inflorescence (X 0.4); flowers (X 2). - After H.B. XV JB'IX\ (BO)



296 R E I N W A R D T I A [VOL. 4

FIG. 6. Firmiana fulgens (Wall, ex Mast.) Corner. — Flowering branch (X V2) ; two
types of flowers (nat. size; top of male androgynophore (X 5) ; the same but anthers

removed (X 5). — After H. B. XV J. B. IX 2.

fleshy, the inside with a few hairs (stellate hairs with shorter branches)
or glabrous; tube inside glabrous, the base with a fleshy, glossy,̂  dark
yellow part (nectary) ; immediately above this glossy part a broad ring ot
long, strigose hairs." Androgynophore at last exsert, glabrous. Male flowers
with large anther-cells (about 20) and undeveloped carpels; female ilowers
with smaller anthers and flask-shaped carpels, with a few, white stellate
hairs at the ventral suture; stigmas slender, recurved. Fruit glabrous,
not seen.

DISTRIBUTION. — Western Himalaya.

1957]

LOCAL NAMES.
ex Haines).

A. J. G. H. KOSTERMANS : The genus Firmiana 297

Khardala, N.W. provinces (ex Brandis) ; Phap (Th.,

The first description of Sterculia pallens Wallich should be attributed
to King (I.e.) who in a note under S. fulgens, differentiated it from the
latter species. No type specimen was indicated by King.

Ridley gave the first extensive description under Erythropsis pallens
Ridley and indicated Falconer 289 (K) as type specimen.

Roxburgh's description of Sterculia colorata in his PI. Coromand.
and in his Flora fits Firmiana pallens (lower leaf surface and petioles
pilose), but the typical funnel-shaped flower with long tube lobes is not
indicated. Roxburgh's plate 25 refers certainly to our Firmiana colorata.

Masters enumerated the specimens Falconer 289 and Strachey &
Winterbottom 2 (K), which are also mentioned by Ridley under Firmiana
pallens. Furthermore Masters cited: Wallich 1135 and 1119 2. The latter
is perhaps Wallich 1119/2, which represents F. colorata.

Kurz {in J. As. Soc. Beng. 43 (ii) : 117. 1875) under Sterculia fulgens,
already suggested, that Masters described actually Sterculia pallens, which
he considered specifically different.

King's and Kurz' description cover each other completely, but the
material, cited by King is certainly F. colorata. I do not know, why King
described the lower leaf surface as densely stellate pubescent, as the spe-
cimens, which he enumerated (King's Coll. 8673; Scortechini, and Forbes
2105) have completely glabrous leaves.

Sterculia wallichii Brandis is based on the specimen Falconer 289,
which is the type specimen of Erythropsis pallens Ridley. The sheet is
preserved in the Kew Herbarium and consists of flower-bearing branches
nly. The label bears the indication: Sterculia wallichii and the number 26,
nd the distributing label Falconer 289 from the Kew Herbarium.

The inside of the flower tube in F. fulgens is glabrous, like the andro-
gynophore; in F. colorata these parts are pilose.

W. HIMALAYA. Gurhwal, fl., Falconer 289 (BO, CAL, K, L, P) ; ibid., Herb.
Hook. f. & Thorns. 58045 (CAL) : ibid., King s.n. (CAL). Missoorie, 1000 m alt., Apr.,
fl-, Mackinnon s.n. (CAL); ibid., May, fr., Maekinnon s.n. (CAL). Rajpoor. fl., Anon.

(CAL). — CULTIVATED. Hort. Bogor sub XV J B IX 2. fl. * (e horto Saharum-
'°re). Hort. Calc. s.n. (BO, CAL); ibid., fl., Pierre 3780 (P).

* In flower in January and February 1957: flowers almost white; there are
Wo kinds of flowers, the male ones are smaller and more slender than the female
ones; both occur in the same inflorescence (see fig. 4, 6).



298 R E I N W A R D T I A
[VOL. 4

3. FIRMIANA PHILIPPINENSIS Kosterm.—Fig. 7

Firmiana philippinensis Kostermans, Comm. For. Res. Inst. Bogor 54: 19. 1956.
— Firmiana simplex (non W. F. Wight) Merrill in Philip. J. Sci., Bot. 13: 308. 1918;
Enum. Philip, fl. PL 3: 56. 1923. — Fenix 29858 (BO).

Tree. Branchlets thick, soft, filled with soft pith, glabrous, covered
with large leaf scars. Leaves entire, crowded near apex of branchlets, soon
glabrous, chartaceous, ovate, acute, base subcordate, 9 x 14 cm; nerves

FIG. 7. Firmiana philippinensis Kosterm. — Leafy_ branch and inflorescence (X 0.5);
flower (X 2.5). — After Femx 29858 (BO).

1957] A. J. G. H. KOSTERMANS: The genus Firmiana 299

palmate. Petioles 7 cm long, glabrous. Inflorescence and flowers as in
F. platanifolia.

LOCAL NAME.—Bitnong (Pang.), according to Merrill.

The leaves of this species are different from those of F. simplex.
As the follicles are unknown and the leaves of the type specimens are
rather young, these is a possibility that it represents Hildegardia merrittii
(Merr.) Kosterm. Of the latter species I have never seen a specimen.

PHILIPPINES. L u z o n . Prov. Pangasinan, Mt. San Isidro, Fenix 29858, fl.
Nov. (BO).

4. FIRMIANA MAJOR Hand.-Mazz.

Firmiana major Handel-Mazzetti in Anz. Akad. Wiss. Wien; Mat.-Nat. 60: 96.
1923. (1924); Ridley in Kew Bull. 1934: 214. (nomen; maior); Kosterm., Comm. For.
Res. Inst. Indon. 54: 20. 1956. — Sterculia platanifolia var. major W. W. Smith in
Notes roy. bot. Gard. Edinburgh 9: 130. 1916. — Hildegardia major (Hand.-Mazz.)
Kostermans in Bull. Jard. Bot. Bruxelles 24: 338. 1954. — G. Forest 10820 (E).

Small tree or shrub. Branchlets stout, glabrous, with large, round leaf
scars; apical buds with small, densely rusty pilose, ovate-lanceolate bud-
scales. Leaves crowded near apex of branchlets, palmately veined, sub-
reniform (broader than long), chartaceous, shallowly 3-lobed, the lobes
either acuminate or very shortly acute; base cordate; up to 30 X 25 cm;
upper surface glabrous, ribs rather flat; lower surface densely, woolly
pilose (stellate hairs with short ramifications) ; of the 7 main ribs, the
3 or 5 central ones have 3—4 pairs of secondary nerves, tertiary nerves
ladder-shaped, lax; upper leaf-surface slightly pitted (in dried condition)
under the lens. Petiole up to 20 cm long, very minutely grey woolly at base
or completely glabrous. Inflorescence 11 cm long, glabrous, but for the
ends of the branches. Pedicels 0.5—1.5 cm long, grey-woolly; flower tube
funnel-shaped, 3—4 mm long, woolly outside; inside with a dense layer of
long, strigose, glossy hairs, protruding from the tube; lobes lanceolate,
up to 1.5 cm long, reflexed, fleshy, inside glabrous, outside woolly.
Androgynophore glabrous; male (?) one with a pinhead-like top of 10—15
anther-cells. Infructescence with dense felt-like, minute, woolly tomentum
towards end of ramifications, glabrescent. Follicles 5, glabrous, up to
15 cm long and 6 cm wide (opened) with 3—5 subglobose, smooth seeds
(10~ 15 mm in diam.) ; androgynophore 1 cm long, glabrous; stalk of
follicles 2 cm, glabrous; pedicel 1 cm, densely pilose.

The species is manifestly related to F. simplex, from which it differs
by its glabrous, larger carpels and its larger flowers.

C H I N A . Yunnan, near Dali (Talifu), alt. 1750 m., May, fl., Handel-Mazzetti
1132 (A); in thickets on the Tong Shan in the Yangtze bend, lat. 27°20' N., alt. 3000 m,
f r-, Forest 10820 (A, E); locality not indicated, fr., Delavay s.n. ,(P).



300 REIN WARDTIA [VOL. 4

5. FIRMIANA DIVERSIFOLIA A. Gray.—Fig. 8

Firmiana diversifolia A. Gray, Botany U.S. Expl. Exped. 1: 185, t. 13. 1854;
Ridley in Kew Bull. 1934: 214 (nomen) ; A. C. Smith in J. Arnold Arb. 36: 283. 1955;
Kosterm., Comm. For. Res. Inst. Indon. 54: 21, /. 6. 1956. — Sterculia diversifolia
(non G. Don) Seemann, PL Vit. 23 (Viti, Append. 433) 1865 (nomen) ; Drake del
Castillo, Illustr. Flor. Ins. Pacif. 122. 1886. — Pickering s.n. (US).

FIG. 8. Firmiana diversifolia A. Gray. — Leaves and infructescence (x 0.5). — After
Pickering s.n. (P).

1957] A. J. G. H. KOSTEBMANS : The genus Firmiana 301

Tree of considerable size (Pickering'). Leaves ovate, palmately nerved,
entire or three-lobed (lobes short, triangular, acute or acuminate), 15—24 x
12—20 cm, base cordate, thinly chartaceous or membranaceous, glabrous
(or the lower surface and the branchlets etc, very minutely stellate pu-
berulent under the lens) ; upper surface smooth, nerves flat; lower surface
with 5 prominent main nerves, the 3 central ones with 3 pairs of lateral
nerves (arcuate and running out at margin) ; tertiary nerves lax, ladder-
shaped; veins inconspicuous. Petioles 10—12 mm long, rather slender,
glabrous, hardly swollen at base. Infructescence glabrous, 14 cm long
with few, distant, patent ramifications, up to 6 cm long. Pedicel densely
pilose, 5 mm long. Androgynophore 5—6 mm long-, glabrous, broadened
at apex into a pateriform cup. Stalks of follicles 5—10 mm long, almost
glabrous. Follicles oblong-lanceolate, up to 2 x 7 cm, glabrous or nearly
so, obtuse at both ends, abruptly tipped with a small mucroniform style.
Seeds globular, glossy, 6 mm in diameter.

The species differs from F. eoiorata because of the few hairs on leaves
and the lack of hairs in the axils of the main nerves. It is perhaps related
to F. papuana, which has the same tomentum on branchlets and petioles,
but differs by its cordate leaves and densely pilose fruit and lower leaf-
surface.

FEEJEE ISL. Ovalau, fr., Pickering s.n. (P, US.).

6. FIRMIANA SIMPLEX (L.) W. F. Wight—Fig. 9

Firmiana simplex (L.) W. F. Wight in U.S. Dept. Agr. Bur. PL Industr. Bull.
142: 67. 1909; F. N. Meyer in Bull. 204: 56. 1911; Rehder & Wilson in Sargent, PL
Wilson. 2 (2): 377. 1915; Arnold Arb. Exped. E. Asia 1917-18, t. N-691. 1920; Nyi
in Contr. Biol. Lab. Sci. Soc. China 5 (3): 17-20, fig. 1929; Sasaki, Catal. Herb.
Taihoku 343. 1930; Liu, Syst. bot. Fl. Fam. N. China 73. 1931; Yen in Bot. Gaz. 93:
206, 210. 1932; Chow, Fam. Trees Hopei (Fan Mem. Inst. Biol. Peiping, Handb. n. 4)
324. 1934; Shun Ching Lee, For. Bot. China 825. pi. 230, 1935; Kostermans in Bull.
Jard. Bot. Bruxelles 24: 335. 1954; Comm. For. Res. Inst. 54: 26. 1956; Petelot, I.e.
125 (as a syn. of Sterculia platanifolia L.f.) ; Chittenden, Diet. Gard. 2, 2: 824. 1956.
— Clompanus simplex (L.f.) O. Kuntze, Rev. 1: 77. 1891.

Firmiana platanifolia (L.f.) Marsili in Sagg. Sci. Lett. Ace. Pad. 1: 106-116.
1786; Schott & Endlicher, Melet. 33: 1832; Spach, Hist. Veg. phan. 3: 516. 1834,
Atlas, t. 69. 1846; Steudel, NomencL, ed. 2, 1: 642. 1840; id. 2 (2): 640. (as a syn.
of Sterculia platanifolia L.f.); R. Brown in Bennet & Brown, PL Jav. rar. 235. 1844;
Walpers, Rep. 5: 104. 1845-46; Voigt, Hort. Suburb. Calcut. 104. 1845; Seemann, Bot.
H.M.S. Herald 365. 1857; Miquel, Ann. Mus. Lugd. Bat. 3: 92. 1867; Prolus. 256.
K. Schumann in Engl. & Prantl, Nat. Pfl.Fam. 3 (6) : 97, fig. 49 B. 1895; Hemsley
in J. Linn. Soc. 23: 90. 1886 (as a syn. Sterculia platanifolia L.f.) ; Diels in Engl. bot.
Jahrb. 29: 470. 1900; Britton, N. Amer. Trees 694. 1908; Shirasawa, Icon. Ess. for.
J a p . 2: t. 51 f. 10-34. 1908; Makino, Illustir. FL Japan 969. 1924; Yushun kudo, etc.
(Usef. Trees of Japan), ed. 2: 310. 1930; Terasaki, Nippon Shokubutsu Zufu 636,



302 R E I N W A R D T I A [VOL. 4

1609. 1933; Small, Man. S. Eastern PI. 864. 1933 (cum fig.); Coker, Totten, Trees
S.E. Un. States 322. 1934; Burkill, Diet. econ. Prod. Mai. Pen. 1: 1019. 1935. —
Sterculia platanifolia L.f., Suppl. PL Syst. Veget. 423. 1781; Buchoz, Grand Jard. Univ.
t. 94. 1785; Marsili in Saggi Scient. e Lett. Acad. Padova 1: 116, t. 1 et 2 (in nota).
1786; Cavanilles, Diss. 5: 287, no. 417, t. 1U5; ibid. 6: 352. 1788; Vahl. Symb. 1: 80.
1790 (excl. Culhamia Porsk.); de Jussieu, Gen. (ed. Suteri) 309. 1791; Houttuyn,
Linn. PL Syst. 4: 152. 1773-83; Willdenow, Sp. PL 2 (2): 873. 1799 (excl. syn.
Culhamia Forsk.); Persoon, Syst. Veg. 910. 1791; Synops. 2: 240. 1807; Thunberg,
Icon. PL Japon, Decas IV: tab. 8. 1802; Lamarck-Poiret, Encycl. meth. 7: 431. 1806
(excl. Culhamia Forsk., diagn. p.p.) ; Steudel, Nomencl. 814. 1821 (exel. syn. Culhamia
Forsk.); DC, Prodr. 1: 483. 1824 (excl. Culhamia Forsk.); Sprengel, Syst. Veg.
3: 83. 1825 (excl. Culhamia Forsk.); G. Don, Gen. Hist. 1: 517. 1831 (excl. Culhamia
Forsk.) ; Schott & Endlicher, I.e. (as a syn. of Firmiana platanifolia R. Br.); End-
licher, Gen. 994. 1840; Suppl. 4 (3): 61. 1847 (under subgenus Eryt'firopsis of Fir-
miana); Loudon, Arb. Frut. Britt. 363. 1838; ed. 2, 1: 363, fig. 90. 1844; R. Brown,
I.e. (as a syn. of Firmiana platanifolia R. Br.) ; Walpers, Rep. 5: 104. 1845-46;
Bentham, PI. Hongkong. 36. 1861; Bentham in Benth. & Hook, f., Gen. PL 1: 218.
1867; Le Maout & Decaisne, Traite gen. Bot. 344. 1868; Le Monnier in Ann. Sci.
nat. Bot., ser. 5, 16: 260-61. 1872; Baillon, Hist. PL 4: 60, fig. 85, 86, 87. 1873;
Traite Bot. medic. 2: 792. 1844; Kurz in J. Bot. 11: 193. 1873; Franchet & Savatier,
Enum. PL Japan. 1: 65. 1875; Dai-Nohon Ju-Moku Thio Rio Yaku (Jap. Trees, Fig.
& Expl.) t. 85. 1878; Debeaux in Act. Soc. Linn. Bordeaux 30: 73. 1875; ibid. 33:
36. 1879 (PL Tientsin 13; Fl. Shanghai 21); Godfrin in Ann. Sci. nat. Bot., ser. 6,
19: 155, pi. 3. 1884; Saccardo in Inst. Veneto Sci. Atti, ser. 6, 5: 881-86. 1886;
Hemsley in J. Linn. Soc. 23: 90. 1886; in Kew Bull. 1891: 248; Pierre, Fl. for
Cochinch., fasc. 3, t. 198. 1889; Mouilefert, Traite Arbres et Arbr., Atlas t. 73. 1892-98;
Usef. PL Jap. 2: t. 599. 1895; Ramaley in Minnesota Bot. Studies 2: 79-80, pi. 3.
1899; Kawakami, List PL Formosa 13. 1910 (ao-gori); Gagnepain in Lecomte, Fl.
gen. Indoch. 1: 464. 1911; Tardieu-Blot. Suppl. 1: 403. 1945; Schneider, 111. Handb.
Laubh. kunde 2: 392. 1909; Wilson in Publ. Arnold Arb. 2: t. 480. 1911; Arnold
Arb. Exped. China .1910-11, t. 0263. 1912; Dunn & Tutcher in Kew Bull., Add. Ser.
10: 49. 1912; Bull. Soc. dendrol. France 79: 218. 1913; Shaw, Chin. For. Trees &
timb. Supply 256. 1914; Leveille, FL Kouy-Tcheou 406. 1915; Sargent, PL Wilson.
2 (2): 377. 1915; Ridley in Kew Bull. 1934: 215; Petelot, PL medic. Cambodge, Laos
& Vietnam 1: 125. 1952 (Arch. Recherch. Agron. 14) ; L.H. Bailey, Stand. Cyclop.
Hort. 3: 3239, fig. 3691. 1947 (cum var. variegata Bailey); Little, Check list Trees
U.S.A. 185. 1953. — Caucanthus platanifolia (L.) Rafinesque, Sylva tellur. 72. 1838;
Merrill, Index Rafin. 166. 1949. — Sterculia fiorniana J. F. Gmelin, Syst., ed. 13,
2 (2) : 1034. 1791. — Sterculia firmiana J. F. Gmelin, I.e. 1633. — Firmiana chinensis
Medic, ex Steudel, Nomencl. 814. 1821; ed. 2, 1: 642. 1840; ed. 2, 2: 640. 1841 (as
a syn. of Sterculia platanifolia L.f.) ; Sargent, I.e. (as a syn. of Firmiana simplex
Wight). — Sterculia tomentosa Thunb., Mus. Uppsal. App. 19: 168. 1796 (nomen) ;
Jap. Icon. Dec. IV: 38, t. 8, 17. 1802; G. Don, I.e. 517; R. Brown, I.e. (as a syn.
of Sterculia platanifolia L.f.); Juel, PL Thunb. 294. 1918; Petelot, I.e. (as a syn.
of Sterculia platanifolia L.f.). — Sterculia pyriformis de Bunge in Mem. Sav. etrang.
Ac. Sci. St. Petersbourg 2: 83. 1835 (Enum. PL Chin. 9. 1833); Walpers, Rep. 1:
335. 1842; R. Brown, I.e. (as a syn. of Firmiana platanifolia R. Br.) ; Hemsley, I.e.
90 (as a syn. of Sterculia platanifolia L.f.); Sargent, I.e. (as a syn. of Firmiana

1957] A. J. G. H. KOSTERMANS : The genus Firmiana 303

PIG. 9. Firmiana simplex (L.) W. F. Wight. — Leafy branch and inflorescence (x 0 4) •
male and female flowers (x 2) ; follicles (X 0.4).



306 R E I N W A R D T I A [VOL. 4 1957] A. J. G. H. ICOSTERMANS : The genus Firmiana 307

surface. Pinned to Sheet 875/12 in the Linnean Herbarium is another
Sheet (875/13), which bears a single leaf mounted with the under surface
showing; this leaf, also larger and less markedly lobed than the one on
Sheet 875/12, is obviously of the same species and presumably came from
the same plant. Neither of these sheets bears a determination by the elder
or younger Linnaeus, but Smith has named Sheet 875/12 Hibiscus simplex
and has further suggested that it is StercuUa platanifolia. This sheet also
bears an inscription by Linnaeus indicating that the specimen was from
Hortus Upsaliensis".

As Mr. Dandy points out, that Hibiscus simplex was described from
a living plant that had not flowered, and that Sterculia platanifolia was
described from a flowering specimen, it is possible that the plants on
Sheets 875/12, 13 could be the type of Hibiscus simplex and even of Ster-
culia platanifolia also, if the flowering plant described by the younger
Linnaeus was the mature stage of the young plant earlier described by
the elder Linnaeus.

Mr. Dandy suggested to find out, whether in the living plant the
domatia contain moisture, "in which case they could represent Linnaeus'
poris melliferis."

As this is really the case, we now may accept the plants on the two
sheets mentioned above as the type of Hibiscus simplex L., which is con-
specific with Firmiana platanifolia. Thanks to the tenacity of Mr. Dandy,
this problem now has been solved.

CHINA. North China, Peking: de Bunge s.n., fl. (male) (P) (type of S. pyriformis
Bunge) ; ibid., fl. (female), Fortune s.n. (P) ; Tsingtao, fl., (female). Zimmermunn 463
(P) ; Szechuan, young fr., Fang 2275 (P) ; Yunnan, May, fl. (female), Ducloux 3486
(P); Shensi, ster., Maximowiez s.n. (P) ; Hongkong, fl. (female), Hanee 5,12 (P);
TONKIN, near Xinh Binh, fl. (female), Bon 3893 (P) ; cult, near Ben Hue, fl., Pierre
3205 (P) ; JAPAN. Nagasaki, cult., fr., Maximowiez s.n. (BO, Petrop.);" Kurachibishi,
Uno s.n. (A). S. DOMINGO, cult, fl., Herb. Piret s.n. (P) ; MAURITIUS, cult., herb.
Sieber Fl. Maurit. exciscc. part. II no 298, female (P) ; N. AMERICA, locality not
indicated, cult., fl. (female), Bon s.n. (P).

USE. — From the bark of young trees a fibre is obtained.

7. FIRMIANA PAPUANA Mildbread—Fig. 10
Firmiana' papuana Mildbread in Engl. Bot. Jahrb. 62: 365. 1929; Ridley in Kew

Bull. 1934: 214 (nomen) ; Kosterm., Comm. For. Res. Inst. Indon. 54: 28, /. 8. 1956. —
Schlechter 17614 (Isotype in L).

Tree about 30—35 m tall; bole 20 m, buttressed up to 1 m, out 2 m.
Bark greyish, mottled with green; when the outer dead corky layer of

I

FIG. 10. Firmiana papuana Mildbread. — Leafy branch and inflorescence (x 0.6),
after Gray 4093 (BO) ; male flower (X 3), after Gray 4093 (BO), female flower (X 3),

after Cavenaugh N.G.F. 4053 (BO).

3—5 mm is scraped off a green layer is revealed; living bark 15—20 mm,
very pale straw coloured or orange with white streaks, yielding a little
sap. Sapwood and heartwood not different, very pale straw yellow to
nghtbrown. Branchlets rather slender, densely pilose (ramifications of
stellate hairs rather long, somewhat appressed). Leaves palmately veined,
chartaceous, dark-green, slightly squamose above, grey-green below
(Floyd), ovate, not lobed, 12—20 x 10—16 cm, base as a rule truncate, top
acute or subacuminate; above glossy green, soon glabrous (pilosity on
nerves tardily disappearing), pitted or densely reticulate (when dried),
nerves flat; lower surface covered with a dense, pale felt of woolly stellate
hairs; the 5 main nerves prominent; the 3 central ones with 2—3 pairs
of rather straight secondary nerves (at margin arcuate and running out) ;





310 R E I N W A R D T I A [VOL. 4

I have not seen the specimens H. Fung 20056 and Liang 61719, enum-
erated by Merrill under F. colorata. They may eventually belong to
F. hainanensis.

1. F.

2.
3.
4.

5.
6.

7.
8.

9.
10.
11.

12.
13.
14.
15.

16.
17.

F.
F
F

F
F

F
F

F
F
F

F
F
F
F

F
F

E X C L U D E D S P E C I E S

acerifolia Voigt, Hort. suburb. Calc. in Index XXII. 1845 (sphalm.)
= Sterculia acerifolia Wall.

affinis (Mast.) Terr. = Scaphium macropodum (Miq.) Beumee.
barteri (Mast.) K. Schum. = Hildegardia barteri (Mast.) Kosterm.
beccariama (Pierre) K. Schum. = Scaphium macropodum (Miq.)

Beumee.
borneensis Merrill = Seaphiwm macropodum (Miq.) Beumee.
campanulata Voigt, Hort. suburb. Calc. in Index p. XXII (sphalm.)

1845 = Pterocymbium javanicum R. Br.
heterophylla Pasq. = Cola heterophylla Schott & Endl.
guttata Voigt, Hort, suburb. Calc, Index XXII. 1845 (sphalm.) =

Sterculia guttata Roxb.
Umceaefolia Voigt, I.e. (sphalm.) = Sterculia lanceaefolia Roxb.
linearicarpa Terr. = Scaphium linearicarpum (Mast.) Pierre.
lychnophora (Hance) K. Schum. = Scaphium macropodum (Miq.)

Beumee.
merrittii Merrill = Hildegardia merrittii (Merr.) Kosterm.
migeodii Exell = Hildegardia migeodii (Exell) Kosterm.
ornata Voigt, I.e. XXII = Sterculia- ornata Wall, ex Voigt.
populifolia (Roxb.) Terr. = Hildegardia populifolia, (Roxb.)
Kosterm.

villosa Voigt, I.e. XXII = Sterculia villosa Roxb.
wallichii Terr. = Scaphium scasphigemm (G. Don) Guibourt.


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