Bangladesh J. Plant Taxon. 29(2): 373-401, 2022 (December)  DOI: https://doi.org/10.3329/bjpt.v29i2.63535  
© 2022 Bangladesh Association of Plant Taxonomists  

 

TAXONOMIC REVISION OF THE SUBFAMILY STERCULIOIDEAE 
BEILSCHM. IN BANGLADESH 

 
SHEIKH SUNZID AHMED AND M. OLIUR RAHMAN1 

 
Department of Botany, University of Dhaka, Dhaka-1000, Bangladesh 

 
Keywords: Sterculioideae; Systematics; Taxonomic treatment; Malvaceae; Bangladesh. 

 
Abstract 

 The subfamily Sterculioideae Beilschm. of Malvaceae has been revised for 
Bangladesh which revealed 16 taxa belonging to five genera, such as Firmiana Marsili, 
Heritiera Ait., Pterygota Schott & Endl., Scaphium Schott & Endl. and Sterculia L. The 
genus Sterculia is unveiled with the highest number of taxa, whereas both Pterygota and 
Scaphium are represented with single taxon. Dichotomous bracketed keys to genera and 
species are constructed for easy identification. Each taxon is presented with an updated 
nomenclature, description of the taxa along with illustrations, specimens examined, 
phenology, notes on distribution and economical importance. In most cases, chromosome 
number has been provided. 

 

Introduction 
 Understanding on the diversity and inter-relationships of plant life is based primarily upon the 
comparative data of revisionary studies (Stuessy, 1975). The process of undertaking a taxonomic 
revision is central to taxonomy. A revision involves restudy of a group to correct or improve its 
diagnosis, description or phylogeny. Taxonomic revision helps to update the system of 
classification as taxa are often subjected to phenotypic plasticity (Baur et al., 2014). Sterculioideae 
Beilschm. is one of the nine subfamilies of Malvaceae representing 13 genera and around 430 
species worldwide (Wilkie et al., 2006; APG IV, 2016). These 13 genera have been distributed in 
four major clades (APG IV, 2016), such as (a) Cola clade incorporating Cola Schott & Endl., 
Octolobus Welw., Pterygota Schott & Endl., Hildegardia Schott & Endl., Firmiana Marsili, 
Scaphium Schott & Endl. and Pterocymbium R.Br.; (b) Heritiera clade incorporating the only 
genus Heritiera Ait.; (c) Sterculia clade including the single genus Sterculia L.; (d) Brachychiton 
clade incorporating Brachychiton Schott & Endl., Acropogon Schltr., Argyrodendron Klotzsch 
and Franciscodendron B. Hyland & Steenis (Wilkie, 2006). According to Bentham and Hooker 
(1883), all these 13 genera belong to the order Malvales of the series Thalamiflorae, subclass 
Polypetalae under the class Dicotyledonae of the Kingdom Phanerogamia. Schumann (1890) 
classified all these genera under the order Malvales of the subclass Archichlamydeae. 
Archichlamydeae was grouped along with another subclass Metachlamydeae under the class 
Dicotyledonae of the subdivision Angiospermae which was incorporated into the division 
Embryophyta. Later, Hutchinson (1959) placed them under the order Tiliales of the division 
Lignosae of the sub-phylum Dicotyledones and phylum Angiospermae. Subsequently, Cronquist 
(1981) grouped them under the order Malvales of the superorder Malvanae, and the subclass 
Dilleniidae of the class Magnoliopsida. The presence of sheath cells in Sterculioideae are unique 
within Malvaceae sensu lato (Chattaway, 1932, 1938).    
 Members of Sterculioideae has a pan-tropical distribution, being found in New Caledonia and 
other Pacific islands, Australasia, South, South-east and East Asia, Madagascar, tropical Africa, 
Central America, the Caribbean and tropical South America with most diversely disbursed in 
                                                   
1Corresponding author. Email: oliur.bot@du.ac.bd  

https://doi.org/10.3329/bjpt.v29i2.63535
mailto:oliur.bot@du.ac.bd


374 AHMED AND RAHMAN 

 

South-east Asia (Wilkie, 2006). This subfamily harbors a good number of economically and 
medicinally important species. Heritiera fomes Buch.-Ham is a dominant species of Sundarban 
Mangrove Forest that shows special ecologically adaptive features and medicinal properties 
(Mahmud et al., 2014). Timbers obtained from Heritiera fomes, Sterculia hamiltonii (Kuntze) 
Adleb. are used for electric poles, posts and ridge plates (Alam, 2018). Firmiana colorata (Roxb.) 
R. Br., Sterculia villosa Roxb. and some other members are economically important for their use 
as avenue tree (Uddin et al., 2021). Pterygota alata (Roxb.) R.Br., Sterculia villosa, and Firmiana 
colorata are profusely used in traditional medicine for their significant medicinal values (Al 
Muqarrabun et al., 2015). Hooker (1874) reported seven species of Sterculioideae from the present 
territory of Bangladesh, and Prain (1903) documented five species from the current boundary of 
the country. Though several sporadic studies were carried out by different workers (Heinig, 1925; 
Raizada, 1941; Datta and Mitra, 1953; Sinclair, 1956; Ahmed et al., 2009; Rahman et al., 2012; 
Alam, 2018), however, taxonomic revision of the subfamily Sterculioideae has not been carried 
out in Bangladesh. Therefore, the current investigation aimed to revise the subfamily 
Sterculioideae in Bangladesh for the first time. 
 
Materials and Methods 
 A total of 25 field trips were conducted for collecting plant specimens from various parts of 
Bangladesh, covering all the seasons. The collected plant specimens were critically studied and 
identified. Identifications were confirmed by consulting standard literatures (Hooker, 1874; Prain, 
1903; Ahmed et al., 2009; Wilkie and Ahmad, 2011; Rahman et al., 2012; Alam, 2018). 
Specimens of Sterculioideae housed at Dhaka University Salar Khan Herbarium (DUSH), 
Bangladesh National Herbarium (DACB), Herbarium of Chittagong University (HCU) and 
Bangladesh Forest Research Institute Herbarium (BFRIH) were also studied and examined 
critically. Dichotomous bracketed keys to genera and species are constructed for ease of use. 
Nomenclature has been updated using standard recent Floras and authentic web-based data 
(https://theplantlist.org; https://powo.science.kew.org; https://tropicos.org; https://gbif.org; 
https://www.ipni.org). Each taxon is presented with detailed taxonomic description along with 
updated nomenclature, important synonyms, English and vernacular names, Flowering and 
fruiting period, Specimens examined, Chromosome number, Habitat, Distribution, Economic 
values and Illustrations (whenever possible). The taxa are arranged in an alphabetical order. 
Voucher specimens have been prepared following standard herbarium techniques (Hyland, 1972) 
and deposited at Dhaka University Salar Khan Herbarium (DUSH).  
 

Results  
Taxonomic Treatment 
 Sterculioideae is characterized by apetalous, exinvolucellate flowers with a fleshy, usually 
petaloid, gamosepalous calyx, absence of staminodes (estaminodes), monadelphous staminal 
column, androgynophore and apocarpous ovaries.  
Sterculioideae is represented by 5 genera and 16 species in Bangladesh. 
 

Key to genera of the subfamily Sterculioideae 
1 Leaves compound. Sterculia 
- Leaves simple. 2 
2 Fruits samara, 1-seeded, indehiscent; keels prominent. Heritiera 
- Fruits follicles, 1 to many seeded, dehiscent; keels absent. 3 

https://theplantlist.org;
https://powo.science.kew.org;
https://tropicos.org;
https://gbif.org;
https://www.ipni.org).


TAXONOMIC REVISION OF STERCULIOIDEAE 375 

 

3 Fruits woody; seeds long winged. Pterygota 
- Fruits membranous; seeds wingless. 4 
4 Seeds 2-4, borne on margin of carpels; leaves palmately lobed. Firmiana 
- Seeds solitary, not borne on margin of carpels; leaves not palmately lobed. Scaphium 

 
FIRMIANA Marsili 

 In Saggi, Sci. Acad. Padova 1: 114, 116 (1786); Miquel, Fl. Ind. Bat. 1: 178 (1859); Benth. & 
Hook. f., Gen. Pl. 1: 218 (1862); K. Schumann in Engler & Prantl, Nat. Pflanzenfam. 3: 97 (1895); 
Hutch., Gen. Fl. Pl. 2: 519 (1967); Erythropsis Lind., Melet. Bot.: 33 (1832); Ridley, FMP 1: 277 
(1922); Sterculia sect. Firmiana (Marsili) Mast. in Hook. f., Fl. Brit. Ind. 1: 359 (1874); King. J. 
As. Soc. Beng. 60: 60 (1891). 
 Trees or shrubs, deciduous. Leaves simple, palmately 3–5-lobed or entire. Inflorescence 
paniculate or rarely racemose, axillary or terminal. Flowers unisexual or polygamous, sometimes 
appearing before leaves. Calyx orange-red or golden, funnel-shaped or cylindrical, (4 or) 5-lobed 
or -partite, lobes short or divided nearly to base, reflexed. Petals absent. Male flowers with 
stamens 10–20, in capitate cluster at apex of androgynophore; anther 2-celled, anther cells curved; 
undeveloped pistil present. Female flowers with ovary 5- locular, ovoid or globose; styles basally 
connate; stigmas as many as carpels; ovules 2 to many per locule. Follicles stipitate, endocarp 
membranous, dehiscent long before maturity, foliaceous. Seeds 1 to many per follicle, globose, 
endosperm flat or plicate. 
 
Key to species of Firmiana  

1 Flowering when leafless; calyx lobes much shorter than tube. F. colorata 
- Flowering with leaves present; calyx lobes much longer than tube. F. simplex 

 
Firmiana colorata (Roxb.) R. Br. in Bennet & R. Br., Pl. Jav. Rar.: 235 (1844); Abedin and 
Ghafoor in Nasir & Ali (eds.), Fl. W. Pak. 99: 22 (1976); Malick in Sharma & Sanjappa, Fl. India 
3: 420 (1993); Verdcourt in Dassanayake et al., Rev. Handb. Fl. Ceyl. 9: 437 (1995); Ahmed et al. 
(ed.), Encycl. Fl. Fauna Bangladesh 10: 333 (2009). Sterculia rubicunda Wall., Numer. List 
[Wallich]: 1119 (1829); Erythropsis roxburghiana Schott & Endl., Melet. Bot.: 33 (1832); 
Clompanus colorata (Roxb.) Kuntze, Revis. Gen. Pl. 1: 78 (1891); Stercuila colorata Roxb., Pl. 
Corom. 1: 26, t. 25 (1795); Mast. in Hook. f., Fl. Brit. Ind. 1: 359 (1874); Prain, Beng. Pl. 1: 187 
(1903); Firmiana rubriflora Kosterm., Reinwardtia 6: 293 (1962).              (Fig. 1). 
 Vernacular names: Faisaudal, Budula, Ujal, Udal, Patagota, Deo Udal, Tey Udal (Chakma), 
Krasan-Baing (Marma), Silkamlambar, Sikalemak Gach (Tripura). 
 English name: The Coloured Sterculia. 
 A medium-sized, spreading, deciduous tree with a height of up to 15 m. Stem fluted, juvenile 
shoots covered with grayish stellate hairs. Branchlets sometimes grey-black at the time of 
desiccation. Bark smooth, very fibrous. Leaves 12-25 cm long and nearly as broad, crowded at the 
end of branchlets, palmately 3-5 lobed, lobes triangular, acuminate, leaves sometimes unlobed on 
adult trees, palmately 5-7 veined, glabrescent on adaxial and abaxial surfaces, base deeply cordate, 
petiolate, petioles 7-15 cm long. Flowers scarlet or orange-red, solitary or in clusters of 2-3 short 
panicles from the axils of fallen leaves, polygamous, the whole inflorescence densely covered with 
orange-red or dark brown stellate hairs. Sepals 5, calyx broadly tubular or funnel-shaped, slightly 
curved, toothed, abaxially densely stellate puberulent with orange hairs, adaxially densely stellate 



376 AHMED AND RAHMAN 

 

puberulent, lobes ovate-triangular, ca. 4 mm, apex acute. Petals absent. Androgynophore present, 
5-10 mm long, exserted, filamentous short, anthers 10, locules curved. Ovary superior, 5-locular, 
flask-shaped, glabrous; styles 5, free, short; stigma curved outward. Fruit a follicle, 8-11 x 3-5 cm, 
oblong with acute apex, follicle stalked, red or purple when mature, foliaceous, tongue-shaped, 
with apparent venation. Seeds 2, yellow, wrinkled or smooth, globose or ovoid, ca. 1 cm long, ca. 
6 mm in diameter. 
 Flowering and fruiting: March-June. 
 Chromosome number: 2n = 40 (Kumar and Subramaniam, 1986). 
 Ecology: The species occurs usually in mixed forests at high elevations and hilltops.  
 Representative specimens: Bandarban: Keokradong, Darjilingpara, 27 Mar 2019, Khandakar 
Kamrul Islam KKI 3582 (DACB). Chittagong: Raimalong hill, 6 Apr 1947, S.M. et Harley s.n. 
(DUSH); Dhurung, 23 Apr 1980, M.K. Alam et D.K. Das 3901 (BFRIH); Russian Hill, 17 Mar 
1982, Nashu et al. 4243 (BFRIH); Chunati, 27 Mar 1998, M.A. Rahman et al. 2566 (HCU); 
Hathazari, Mithachara, 19 Mar 1999, M.A. Rahman 4717 (HCU); Chittagong University 
Botanical Garden, 9 Oct 2010, M.A. Rahman 7129 (HCU); Dudhpukuria-Dhopachori Wildlife 
Sanctuary, 13 Jun 2022, Sunzid 65 (DUSH). Cox’s Bazar: Chakaria, Dulahazra, Bangabandhu 
Safari Park, 23 Jan 2017, Niyamul Kabir et al. NK 3207 (DACB). Dhaka: Sreepur, 20 Mar 1964, 
D.K. Das FRL348 (BFRIH). Jamalpur: Gandhigaon, Gajni forest, 5 May 1982, Mia et al. M. 695 
(DACB). Khagrachari: Mong-Ro-Para, 10 Jan 1989, M.K. Alam EB 111 (BFRIH). 
Mymensingh: Barta, 12 Feb 1959, M.S. Khan K. 342 (DUSH). Patuakhali: Andhermanik River, 
21 Apr 1960, Davidson et al. 129 (BFRIH). Rangamati: Pablakhali, 27 Apr 1965, D.K. Das s.n. 
(BFRIH). Sylhet: Satgaon forest, 8 Feb 1967, P. Bhattacharjee 155 (DUSH). Tangail: Madhupur, 
29 Mar 1988, A.M. Huq 78 (DACB). 
 Global distribution: Bhutan, Bangladesh, Brazil, Cambodia, China, India, Indonesia, Lao 
PDR, Malaysia, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand and Viet Nam. 
 Economic aspects: Fresh young seeds with an almond flavor are edible. In the Chittagong 
district of Bangladesh, the bark generates cordage that is used to tie animals and goats. The 
Chittagong Hill Tracts tribal people use the bark to manufacture bamboo busket straps called 
"Turung" which they wear on their foreheads. Ethnic people use this plant as a remedy to cholera 
for its promising anti-diarrheal activities. Besides that, some other diseases and disorders like 
hysteria, jaundice, urine infection, stomach ache, seminal emission, and spermaturia can be treated 
using this plant (Ashrafuzzaman and Sarwar, 2021). 
 
Firmiana simplex (L.) W. Wight, Bull. Bur. Pl. Industr. U.S.D.A. 142: 67 (1909); Meyer, Chin. 
Pl. Names: 20 (1911); Merrill, Enum. Philip. Fl. Pl. 3: 56 (1922); Cheo, Crypt. China. 1: 141 
(1931); Crossley, An Eva. Rec. Gen. Rec.: 23 (1977). Hibiscus simplex L., Sp. Pl. ed. 2: 977 
(1763). Sterculia urens Roxb., Pl. Coromandel 1: 25, t. 24 (1795). Cavallium urens Schott & 
Endl., Melet.: 33 (1832); Kavalama urens (Roxb.) Raf., Sylva Tellur.: 72 (1838); Clompanus 
urens (Roxb.) Kuntze, Revis. Gen. Pl. 1: 78 (1891).  
 Vernacular name: Not known. 
 English name: Not known. 
 A soft wooded deciduous tree, up to 15 m tall. Bark white papery or greenish, smooth on the 
outer surface. Twigs glabrescent with distinct raised leaf scars and lenticels. Leaves crowded at the 
end of branchlets, palmately 3-5 lobed, coriaceous, hairy on lower surface, 12-20 x 10-20 cm, 
acuminate to cuspidate at the apex, deeply cordate at the base, usually 5-nerved from the base; 
petiolate; petioles very long, up to 20 cm; stipules narrowly lanceolate, caducous. Inflorescence 



TAXONOMIC REVISION OF STERCULIOIDEAE 377 

 

terminal, many flowered, 10-18 cm long, glandular, pubescent. Flowers small, yellow, pedicellate, 
pedicels ca. 4 mm long, sometimes equal to the length of calyx. Calyx 5-lobed, campanulate, lobes 
oblong or narrowly triangular, 4-8 x 3-5 mm, twisted outward, abaxially puberulent or yellowish, 
adaxially villous only basally, calyx tube as long as lobes. Male flowers with androgynophore as 
long as calyx, thicker in lower part, glabrous. Stamens 10; anthers sessile; staminodes present, 10. 
Female flowers with 5-6 ovary, ovoid in shape and hairy; style hairy; stigma 5-6 lobed, recurved. 
Fruit a follicle, 2-6, oblong, ellipsoid or kidney-shaped, 4-6 cm long and 1-2 cm broad when 
young, densely rusty, pubescent. Seeds 3-6, oblong to ellipsoid, sometimes globose, 7 mm in 
diameter, wrinkled, black. 
 Flowering and fruiting: October-February. 
 Chromosome number: 2n = 40 (Raghavan and Arora, 1958). 
 Ecology: The species usually occurs in mixed deciduous forests. 
 Representative specimens: No specimen was available in any herbarium of Bangladesh. 
 Global distribution: Bangladesh, China, Cambodia, India, Japan, Sri Lanka, Taiwan, 
Thailand, United States of America and Viet Nam. 

 
Fig. 1. Firmiana colorata (Roxb.) R. Br.: A. Habit (x 0.2); B. Fruits (x 0.2). 



378 AHMED AND RAHMAN 

 

 Economic aspects: The species has been widely employed in the pharmaceutical, health-care, 
food, cosmetics, waste management, paper-textile, composite fiber, and leather sectors. Because 
the gum exudate, this tree has a high market value, it is popularly referred to as the 'Karaya Gum' 
tree. Because of its natural availability, economic effectiveness, non-hazardous nature, quick 
recovery, and physicochemical qualities, karaya gum polymers are an excellent alternative to 
synthetic ones. Furthermore, tribal communities use the tree for its indigenous cures for 
oligospermia, leucorrhoea, constipation, body swelling, throat infection, wound healing, and other 
diseases. Also, the presence of diverse bioactive compounds has made this plant an excellent 
source to conduct research on Computer Aided Drug Designing (CADD) (Dhiman et al., 2019). 

 
HERITIERA Ait. 

Hort. Kew. 3: 456 (1789); Miquel, Fl. Ind. Bat. 1: 179 (1859); Benth. & Hook. f., Gen. Pl. 1: 219 
(1862); Mast. in Hook. f., Fl. Brit. Ind. 1: 362 (1874); King, J. As. Soc. Beng. 60: 79 (1891); 
Kostermans, Publ. Counc. Sci. Ind. 1: 3 (1959); Backer & Bakhuizen f., FJ 1: 415 (1964); 
Smythies, CST: 127 (1965); Cockburn, TS 1: 228 (1976); Turner, Gard. Bull. Sing. 47: 475 
(1996); Phengklai, Fl. Thailand 7 (3): 573 (2001); Tarrietia Blume, Bijdr. Fl. Ned. Ind.: 227 
(1825). 
 Trees, usually with buttress. Leaves alternate, simple or palmately compound, abaxially 
usually scaly. Inflorescence cymose, paniculate, axillary, many-flowered, hairy or scaly. Flowers 
unisexual. Calyx campanulate or urceolate, 4–6-lobed. Petals absent. Male flowers with short 
androgynophore. Anthers 4–15, in 1 or 2 rings at apex of androgynophore, with pistillode. Female 
flowers with ovary with 3–5 coherent carpels, base with sterile anthers; ovules 1 per carpel. Style 
very short; stigma very small. Fruit woody or leathery, with keel-like prominence or wing, 
indehiscent. Seeds without endosperm. 
 
Key to species of Heritiera  

1 Samara with a longitudinal ridge; leaves with faint secondary nerves 
and veins. 

 
2 

- Samara winged or wing-like; leaves with prominent secondary nerves 
and veins. 

 
3 

2 Fruit body up to 2 x 1 cm; epicarp dull; pistil and stamens on a slender 
column with parallel sides. 

H. fomes 

- Fruit body not less than 5 x 3 cm; epicarp shining; pistil and stamens on 
a based column, narrowly in the distal half. 

H. littoralis 

3 Leaves with 8-11 pairs of lateral nerves; anthers irregularly clumped on 
androgynophores. 

H. macrophylla 

- Leaves with 4-5 pairs of lateral nerves; anthers in regular rings on 
androgynophores. 

H. papilio 

 
Heritiera fomes Buch.-Ham. in Symes, Embassy Ava ed. 2, 3: 319, t. 28 (1800); Mast. in Hook. 
f., Fl. Brit. Ind. 1: 363 (1874); Balfour, Cycl. Ind. East. South. Asia 3: 885 (1885); Winkler, 
Botanis. Hilfs. Pflanz. : 139 (1912); Heinig, List Pl. Chitt. Coll. & HT.: 7 (1925); Sinclair, Bull. 
Bot. Soc. Beng. 9(2): 88 (1955); Malick in Sharma & Sanjappa (eds.), Fl. India 3: 428 (1993); 
Ahmed et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 336 (2009). Heritiera minor (Gaertn.) 
Roxb., Hort. Bengal.: 50 (1814); Fometica punctata Raf., Syl. Tellur. :75 (1838); Amygdalus 
minor (Roxb.) Kuntze, Revis. Gen. Pl. 1: 75 (1891).             (Fig. 2).  



TAXONOMIC REVISION OF STERCULIOIDEAE 379 

 

 Vernacular names: Sundari, Sunder. 
 English name: Sundri.  
 A medium-sized evergreen tree, 15-18 m tall, branchlets lepidote, bark grey. Leaves simple, 
alternate, 10-17 x 3-6 cm, elliptic-lanceolate, tapering to rounded at the base, acute or rounded and 
mucronate at the apex, glabrous above, adpressed scaly beneath, petioles up to 2 cm long. Flowers 
small, unisexual, in axillary panicles. Sepals 5 or rarely 4-6 toothed, stellate-hairy inside. Petals 
absent. Male flowers with 5-10 stamens, united in a column with a ring of 2-celled anthers at the 
apex. Female flowers with 4-6 carpels, almost free, styles short, stigmas 5 thick, ovule solitary in 
each carpel. Fruit a cluster of woody, indehiscent, keeled or winged ripe carpel. Fruit a cluster of 
woody, indehiscent, keeled or winged ripe carpel. Seed solitary without vivipary but can float on 
the tidal water. 
 

 
 

Fig. 2. Heritiera fomes Buch.-Ham.: A. Habit (x 0.4); B. Flower (x 1.5). 
  

 



380 AHMED AND RAHMAN 

 

 Flowering and fruiting: September-December. 
 Chromosome number: 2n = 38 (Das et al., 1995). 
 Ecology: Moderately saline zone, well-drained soil inundated by tidal water of low salinity. 
Although it is a mangrove species, it can tolerate the conditions of higher altitudes (Ahmed et al., 
2009). 
 Representative specimens: Bagerhat: Mongla, Hiron-point, 3 Feb 1987, A.M. Huq et M.K. 
Mia H. 8128 (DACB); Barguna: Patharhat, 20 Mar 1989, Huq et al. 9187 (DACB); Taltoli, 
Tangragiri, 15 Oct 2019, Dr. M Sultana DMS 3056 (DACB). Cox’s Bazar: Chakaria, Palakata 
beat, 24 Apr 1966, D.K. Das FRL89 (BFRIH); Chokoria Sundarban, 27 Apr 2014, Shahriar 
Ahmed 50 (HCU); Chakaria, 27 Apr 2014, Md. Dulal Uddin 1 (HCU). Dhaka: Dhaka University 
Botanical Garden, 17 Aug 1953, Md. Shahjahan s.n. (DUSH); Dhaka University Botanical 
Garden, 30 Nov 2021, Sunzid 39 (DUSH). Khulna: Sarankhola range, 9 Mar 1963, D.K. Das 
FRL305 (BFRIH); Sundarbans, tidal forest, 20 Jun 1966 (DUSH), M. Ismail & M. Bhowmik 180 
(DUSH); Bhadra, Sundarbans, 23 Mar 1970, M.S. Khan K. 1919 (DUSH); Sundarbans, Jongra, 
Chota basta beel area, 5 Nov 2001, S.N. Uddin et Dr. Floris Deodatus N. 1151 (DACB). 
Satkhira: Sundarban, Burigualini, 11 Dec 1989, Huq et al. H. 9368 (DACB); Sundarbans, 
Noxabaki, 20 Aug 2002, S.N. Uddin N. 1360 (DACB).  
 Global distribution: Bangladesh, India, Myanmar and Thailand.  
 Economic aspects: The plant is used by traditional health practitioners to treat diabetes, 
hepatic disorders, gastrointestinal disorders, goiter, and skin diseases. It has potent antioxidant, 
antinociceptive, antihyperglycemic, antibacterial, and anticancer properties. Important chemical 
ingredients such as saponins, alkaloids, glycosides, tannins, steroids, flavonoids, gums, 
phytosterols, and reducing sugars have been discovered through phytochemical investigations. 
Twigs are used in the treatment of oral infections and toothache. Hot decoction of bark is used to 
get relief from goiter and diabetes. Decoction of leaves and seeds have efficacy to treat various 
gastrointestinal disorders (Mahmud et al., 2014). 
 
Heritiera littoralis Dryand. in Ait., Hort. Kew. ed. 1, 3: 546 (1789); Mast. in Hook. f., Fl. Brit. 
Ind. 1: 363 (1874); Ridl., Fl. Mal. Pen. 1: 279 (1922); Craib, Fl. Siam. Enum. 1: 170 (1925); 
Kosterm., Reinwardtia 4: 490 (1959); Ahmed et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 337 
(2009). Balanopteris minor Gaertn., Fruct. Sem. Pl. 2: 94 (1790); Balanopteris tothila Gaertn., 
Fruct. Sem. Pl. 2: 94 (1790); Sutherlandia littoralis (Aiton) J.F. Gmel., Syst. Nat. ed. 13: 1027 
(1792); Samadera littoralis (Aiton) Oken, Allg. Naturgesch. 3(2): 1205 (1841); Systemon fischeri 
Regel, Ind. Semin. 1856: 38 (1856); Heritiera fischeri Regel & Rach, Ind. Semin. 1858: 45 
(1859); Amygdalus litoralis (Aiton) Kuntze, Revis. Gen. Pl. 1: 75 (1891).                           (Fig. 3). 
 Vernacular names: Sundri, Aina. 
 English names: Looking Glass Tree, Looking Glass Mangrove. 
 Evergreen trees, up to 10-15 m tall; bark grayish-brown, sometimes pinkish-grey, vertically 
and superficially fissured, rough or smooth, inner bark brown-pink. Sapwood pale yellow. Young 
branchlets white scurfy scaly, mature branchlets lepidote. Stipules lanceolate, caducous; petiole 1-
2 cm; leaf blade simple, alternate, oblong-lanceolate, elliptic, or ovate, 10-20 x 5-10 cm, leathery, 
abaxially densely silver-white scurfy scaly, adaxially glabrous or nearly so, base obtuse, apex 
acute or obtuse. Inflorescence paniculate, axillary, ca. 8 cm, densely stellate hairy or with scales. 
Flowers pale green to pinkish red. Calyx red-brown, campanulate, 4-6 mm, both surfaces stellate 
hairy, lobes triangular, ca. 2 mm. Male flowers with thin disc, papillate. Androgynophore short 
and glabrous. Anthers 4 or 5, in 1 ring. Female flowers with carpels 4 or 5; stigmas as many as 



TAXONOMIC REVISION OF STERCULIOIDEAE 381 

 

carpels, short and curved downward. Fruit nutlike, woody, drying yellow-brown, nearly ellipsoid, 
ca. 6 x 3.5 cm, keeled on back, glabrous. Seeds ovoid, ca. 2 cm. 
 Flowering and fruiting: May-December. 
 

 
Fig. 3. Heritiera littoralis Dryand. - A habit sketch (x 0.4). 

 
 Chromosome number: 2n = 38 (Das et al., 1995). 
 Ecology: Inland zone of mangrove swamps. 
 Representative specimens: Khulna: s. loc., 13 Sep 1949, P. Maheshwari s.n. (DUSH); 
Dhaka: Dhaka University Botanical Garden, 6 Jun 2022, Sunzid 83 (DUSH). 
 Global distribution: Australia, Bangladesh, Cambodia, China, East Africa, India, Indonesia, 
Madagascar, Malaysia, Myanmar, Philippines, Sri Lanka, Taiwan, Tanzania, Thailand, Tonga, 
Vanuatu and Viet Nam. 
 Economic aspects: The timber is used as a good source of fuel for its high heat-yielding 
efficiency. Wood is suitable for the production of wrapping, writing and printing papers. Tannins 
are present in the bark which can be used for toughening fishing nets. Seed extract is used as 
medicine in the treatment of dysentery and diarrhea (Ahmed et al., 2009; Mitra et al., 2021). The 
crude methanol extract of leaves possesses free radical scavenging, anti-hemolytic, cytotoxic and 
anti-bacterial activities (Karim et al., 2020). 



382 AHMED AND RAHMAN 

 

Heritiera macrophylla Wall. ex Kurz in J. Asiat. Soc. Beng. 42(2): 61 (1873); Kurz, Fl. Burm. 1: 
141 (1877); Kanjilal et al., Fl. Assam 1: 155 (1934); Kosterm., Reinwardtia 4: 502 (1959); Malick 
in Sharma & Sanjappa (eds.), Fl. India 3: 430 (1993). Trochetia contracta Wall., Numer List.: 
1162 (1829); Amygdalus macrophylla (Wall.) Kuntze, Revis. Gen. Pl. 1: 75 (1891).   (Fig. 4). 
 Vernacular name: Not known. 
 English name: Large leaf looking glass tree. 
 A medium-sized evergreen tree, with umbrella shaped crown, arborescent, up to 30 m in 
height. Leaves simple, petiolate, petioles cylindrical, pubescent, rusty brown when mature; lamina 
ovate-oblong or elliptic-oblong, apex acute, base obtuse to slightly cordate sometimes, margin 
entire mostly, sometimes undulate in dry condition, veined with pinnate parallel venation, veins 
prominent adaxially, texture subcoriaceous, adaxial surface sometimes rusty brown while abaxial 
surface silvery. Flowers are unisexual, 3-4 mm across, white-yellowish to pinkish-rose in colour, 
present in foot-long panicle like cymes, in leaf axils. Male flowers with 6-8 stamens; anthers 
irregularly clumped on androgynophore. Female flowers with 1 carpel. Fruit a samara, ellipsoid, 
with one apical fish-tail wing. 
 

 

Fig. 4. Heritiera macrophylla Wall. ex Kurz.- A habit sketch (x 0.4). 



TAXONOMIC REVISION OF STERCULIOIDEAE 383 

 

 Flowering and fruiting: November-February. 
 Chromosome number: 2n = 38 (Das et al., 1995). 
 Ecology: Inland zone of mangrove swamps. 
 Representative specimens: Khulna: s. loc, 13 Sep 1949, P. Maheshwari s.n. (DUSH). 
 Global distribution: Bangladesh, Cambodia, China, India, Lao PDR, Myanmar, Thailand and 
Viet Nam. 
 Economic aspects: Mainly valued for its timber. The wood is hard with distinct annual rings 
and uniform medullary rays, joined by fine transverse lines. It takes a fine polish. It is used for 
posts and poles (Duncan, 2005). 
 
Heritiera papilio Bedd., Fl. Sylhet: t. 218 (1872); Mast. in Hook. f., Fl. Brit. Ind. 1: 363 (1874); 
Balfour, Cycl. Ind. East. South. Asia 2: 39 (1885); Gamble, Fl. Pres. Mad. 1: 104 (1915); Malick 
in Sharma & Sanjappa (eds.), Fl. India 3: 430 (1993); Ahmed et al. (ed.), Encycl. Fl. Fauna 
Bangladesh 10: 338 (2009); Uddin & Hassan, Vas. Fl. Chit. & Chit. Hill Tr. 2: 414 (2018). 
Amygdalus papilio (Bedd.) Kuntze, Revis. Gen. Pl. 1: 75 (1891); Cattimarus hospitus (L.) Kuntze, 
Revis. Gen. Pl. 1: 77 (1891); Heritiera acuminata All. ex Kurz in J. Bot. 12: 65, t. 141, f. 1, 1-3 
(1874); Kanjilal et al., Fl. Assam 1: 155 (1934).   
 Vernacular name: Sundari. 
 English name: Not known. 
 A small to medium-sized tree, 10-15 m in height, sometimes up to 30 m tall, arborescent. 
Young branches stellate-tomentose. Leaves simple, alternate, unifoliolate, 4-16 x 3-17 cm, 
lanceolate, oblanceolate or oblong to elliptic, base obtuse, obscurely acuminate at the apex, entire, 
texture coriaceous, petioles ca. 2 mm long, scaly. Flowers in axillary panicles, densely covered 
with stellate tomentose scales. Sepals 5, calyx campanulate. Petals absent. Male flowers with 
slender staminal column, anther thecae in a regular ring on an androgynophore. Female flowers 
with 5-6 sessile ovary, stellate scaly, sterile anther thecae present at the base of the ovary. Fruit a 
samara, 5-7 cm long, glabrous, 1-seeded. 
 Flowering and fruiting: April-December. 
 Ecology: Usually occurs in the hilly forests (Alam, 2018). 
 Representative specimen: No specimen was available in any herbarium of Bangladesh. 
 Global distribution: Bangladesh, India and Myanmar. 
 Economic aspects: Timber is hard which takes a fine polish and is used for temporary 
construction works for the manufacturing of poles, posts as well as agricultural implements 
(Ahmed et al., 2009). 
 

PTERYGOTA Schott & Endl. 
 Melet. Bot.: 32 (1832); Schumann in Engler & Prantl, Nat. Pflanzenfam. 3: 97 (1895); 
Kostermans, Reinwardtia 5: 415 (1961); Phengklai, Fl. Thailand 7 (3): 615 (2001); Bayer & 
Kubitzki in Kubitzki & Bayer (eds.), Fam. Gen. Vasc. Pl. 5: 264 (2003); Tetradia R.Br. in 
Bennett, Pl. Jav. Rar.: 233 (1844); Sterculia L. sect. Pterygota, Benth. & Hook. f., op. cit. 218; 
Mast. in Hook. f., Fl. Brit. Ind. 1: 360 (1874); King, J. As. Soc. Beng. 60: 60 (1891). 
 Trees. Leaves cordate, usually entire, but lobed when very young. Inflorescence axillary, 
racemose or paniculate. Flowers unisexual. Calyx campanulate, 5-lobed to base, lobes reflexed at 
apex. Petals absent. Male flowers with cylindrical androgynophore, enclosed by calyx. Filaments 
clustered into 5 groups, staminodes usually present; anthers sessile. Female flowers with very 



384 AHMED AND RAHMAN 

 

short androgynophore and 5 undeveloped staminodes. Carpels almost free; ovules many per 
carpel; stigma swollen, radiate. Follicle woody, subglobose, with long stipe and many seeds. 
Seeds with long and wide apical wing. 
 
Pterygota alata (Roxb.) R. Br. in Benn., Pl. Jav. Rar. : 234 (1834); Heinig, List Pl. Chitt. Coll. & 
HT.: 7 (1925); Kochumenn in Whitmore, Tr. Fl. Malaya 2: 371 (1973); Abedin & Ghafoor in 
Nasir & Ali (eds.), Fl. W. Pak. 99: 23 (1976); Malick in Sharma & Sanjappa (eds.), Fl. India 3: 
455 (1993); Ahmed et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 343 (2009). Erythropsis 
roxburghii Schott & Endl., Melet. Bot.: 33 (1832); Sterculia teynii Bedd., Fl. Sylv.: t. 230 (1872); 
Clompanus alata (Roxb.) Kuntze, Revis. Gen. Pl. 1: 78 (1891); Sterculia alata Roxb., Pl. Corom. 
3: 84, t. 287 (1819); Fl. Ind. ed. Carey 3: 152 (1832); Mast. in Hook. f., Fl. Brit. Ind. 1: 360 
(1874); Kurz, Fl. Burm. 1: 134 (1877); Prain, Beng. Pl. 1: 274 (1903).   (Fig. 5).  
 Vernacular names: Buddha Narikel, Kashmiri Badam, Narikeli, Pagla Gach, Tula, Kufala 
(Chakma), Lakha (Marma). 
 English name: Buddha Coconut Tree.  
 Large tree, arborescent, deciduous, up to 50 m in height, crown narrow, conical, branches 
horizontal, branchlets pubescent at first with golden yellow hairs. Bark greyish-brown to grey with 
horizontal wrinkles and shallow vertical fissures. Buttress root present. Leaves simple, alternate, 
usually clustered at the end of branchlets, stipulate, stipules subulate, caducous, petiolate, petiole 
cylindrical, glabrous, 50-150 x 2-2.5 mm, leaf blade cordate or broadly ovate, 10-23 x 8.5-19 cm, 
sometimes deeply cordate at the base, lobes absent, coriaceous, glabrous both adaxially and 
abaxially, margin entire to nearly entire, apex acute or obtuse, palmately reticulate veins present, 
veins 5-7, veinlets numerous, prominent on both surfaces. Inflorescence axillary, paniculate, 
shorter than petiole. Flowers brownish yellow, ebracteate, unisexual, male flowers or bisexual but 
functionally female flowers in rusty-tomentose racemes. Calyx campanulate, densely stellate, 
adaxially subglabrous red abaxially, 5-6 lobed, linear-lanceolate, free upwards, connected at the 
base, 16 x 4 mm each with 1.5 mm thickness. Petals absent. Male flowers with androgynophore, 
cone shaped, inserted, half as long as calyx, 6 mm long, bearing 4-6 groups of 4 anthers each. 
Female or bisexual flowers with 5 carpels, inserted ovary 2-3 mm long, globose and puberulent; 
style short, extrorse, recurved; stigmas rounded. Fruit is woody follicle, large, compressed 
globose, ca. 12 cm in diameter, puberulent outside, adaxially cork-like. Seeds many, ca. 40 per 
follicle, oblong, compressed in 2 rows with long and wide wing.  
 Flowering and fruiting: December-May. 
 Chromosome number: 2n = 40 (Kumar and Subramaniam, 1986). 
 Ecology: Highland and open sunny dryland. The species is also found to be planted along the 
roadsides as avenue tree (Alam, 2018). 
 Representative specimens: Bandarban: Ruma, 28 Feb 1988, M.K. Alam et M. Mohiuddin 
6007 (BFRIH). Bogra: Dosmail, RDA campus, 18 Nov 2020, Zinia Nasrin ZN 03 (DACB). 
Dhaka: Ramna, 1 Feb 1947, S.K. Sen s.n. (DUSH); Azimpur, Army Recruiting Office, 1 Nov 
1963, A.F. Muhammad 34 (DUSH); Dhaka University Campus, TSC, 10 Sep 1982, A.M. Huq 
5688 (DACB); Dhaka University Campus, TSC, 17 Oct 2021, Sunzid 33 (DUSH); Bangladesh 
National Botanical Garden, 6 Jun 2022, Sunzid 80 (DUSH).  
 Global distribution: Bangladesh, Bhutan, China, Cuba, India, Malaysia, Myanmar, Pakistan, 
Philippines, Thailand, United States of America and Viet Nam.  
 Economic aspects: Timber is white and has great potential for pulping because of long fibers. 
Extract of leaves have been used to demonstrate antioxidant activities. Therefore, the 



TAXONOMIC REVISION OF STERCULIOIDEAE 385 

 

phytochemicals can be used to design potential drugs for biogerontological research. Seeds are 
medicinally significant, edible and are often eaten after roasting. Seeds possess narcotic properties 
and often used as a substitute for opium (Agarwal and Jain, 2017). Crude ethanol extract of leaves 
is reported to have excellent in-vivo and in-vitro antioxidant effects (Jahan et al., 2014). The leaf 
extracts contain active compounds showing in-vitro antibacterial activity against multidrug 
resistant and biofilm forming strains of Staphylococcus spp. (Panda et al., 2020). 
 

 
Fig. 5. Pterygota alata (Roxb.) R. Br.: A. Habit (x 0.3); B. Sepal (x 0.5); C. Stamen (x 1.5). 

 
SCAPHIUM Schott & Endl. 

 Melet. Bot.: 33 (1832); Kostermans, J. Sci. Res. Indonesia 2: 3 (1953); Phengklai, Fl. 
Thailand 7 (3): 624 (2001); Bayer & Kubitzki in Kubitzki & Bayer (eds.) Fam. Gen. Vasc. Pl. 5: 
265 (2003); Cheek in Heywood et al., Flow. Pl. Fam. World: 311 (2007); Wilkie, Edin. J. Bot. 66 
(2): 283 (2009); Caryophyllum Miq., Fl. Ind. Bat. Suppl. 1: 401 (1861); Sterculia L. sect. 
Scaphium Benth. & Hook. f., Gen. Pl. 1: 218 (1862); Mast. in Hook. f., Fl. Brit. Ind. 1: 361 
(1874); King, J. As. Soc. Beng. 60: 60 (1891).  



386 AHMED AND RAHMAN 

 

 Trees. Leaves simple, spirally arranged, entire, stipules caducous. Flowers unisexual, small 
pale green, axillary or terminal panicles. Calyx 4-6 lobed, united at the base, glabrous inside. 
Corolla absent. Stamens 8 to 10 in male flowers, on a globose head at the terminal part of a thin 
staminal column. Female flowers 5 carpels, free, styles free, stigmas simple, pointed. Fruit a large 
follicle, boat-shaped, membranous. Seeds wingless, solitary.  
 
Scaphium scaphigerum (Wall. ex G. Don) G. Planch, Hist. Nat. Drogues Simples ed. 6, 3: 646 
(1869); Baillon, Hist. des Pl. 4: 110 (1867); Kurz, Fl. Burm. 1: 140 (1877); Balfour et al., Ann. 
Bot. 19: 356 (1905); Kosterm., J. Sci. Res. Indo. 2(1): 3 (1953); Ahmed et al. (ed.), Encycl. Fl. 
Fauna Bangladesh 10: 344 (2009). Scaphium wallichii Schott & Endl., Melet. Bot.: 33 (1832); 
Sterculia scaphigera Wall. Cat. No. 1130 (1828); Mast. in Hook. f., Fl. Brit. Ind. 1: 361 (1874); 
Clompanus scaphigera (Wall. ex G. Don.) Kuntze, Revis. Gen. Pl. 1: 78 (1891).    (Fig. 6). 
 Vernacular names: Shaogan, Shampan, Pogan, Sugan, Shagan. 
 English name: Water Malva Nut. 
 A deciduous tree, up to 35 m tall. Buttresses often present, 1–2 m tall. Inner bark reddish, 
fibrous, 10–15 mm thick. Outer bark grey-green to brown, cracked to flaking. Sapwood white. 
Terminal branches pale brown to brown, striate to smooth, lenticels absent, glabrous. Petiole pale 
brown, 2–12 cm long, 2–4 mm wide, smooth to striate, glabrous or occasionally stellate hairy, 
both ends darker. Stipules caducous or persistent only at apex of stalk, rounded to triangular, 2–4 
mm long, glabrous or with very minute simple hairs on both surfaces. Leaf lamina sub-leathery to 
leathery, occasionally papery, ovate, elliptic or elliptic-oblong, symmetric to asymmetric, base 
rounded to acute (occasionally truncate or slightly cordate), apex acute, 1.3–2.3 times as long as 
wide, 8–34 cm long, 6–17 cm at widest point, upper surface glabrous, occasionally with gland 
dots, drying pale brown-tan, shiny, lower surface glabrous, occasionally with gland dots, drying 
pale brown-tan, dull; midrib raised on both surfaces; basal veins 1–2 pairs. Tertiary veins flat on 
both surfaces, slightly paler than lamina below, inconspicuous above. Inflorescences 5–12 cm 
long, densely stellate hairy. Bracts caducous. Flower buds drying brown, ovoid, slightly elongated 
towards apex, sparsely to densely stellate hairy, lobe margin densely stellate hairy, more or less 
distinct. Pedicel absent to 2 mm long, 0.5–1 mm wide, densely stellate hairy. Flowers 5–8 mm 
long, yellow with purple base, when fresh, drying brown, 5-lobed, lobes 2/3 the length of the 
calyx, calyx outer surface sparsely stellate hairy, inner surface glabrous (including base), 
androgynophore 5–8 mm long, protruding, recurved at apex, sparsely simple hairy, sometimes 
with stellate hairs with long branches on upper half, glabrous on lower half, anthers ca. 10–12, 
glabrous or sparsely simple hairy, carpels rudimentary and sparsely hairy in male flower, fully 
developed and densely stellate hairy in female flower, 3–5, free or nearly so; style absent to 0.2 
mm long in male flower, 1–1.5 mm long in female flower; stigmas glabrous, as many as carpels, 
superficially fused. Fruit with 2–5 follicles per flower, follicles papery, 9–26 cm long, inner 
surface sparsely stellate hairy, shiny, outer surface sparsely to densely stellate hairy (especially 
dense along venation), dull, follicle stalk to 7–25 mm long. Seeds 1 per follicle, spherical to 
elliptic, glabrous, seed oriented away from follicle. 
 Flowering and fruiting: February-May. 
 Chromosome number: Not known.  
 Ecology: Near limestone deposits, there are evergreen forests. 
 Representative specimens: Bandarban: Kapru Para VCF, 25 May 2017, Syedul Alam et 
Rahman 27893 (BFRIH). 
 Global distribution: Bangladesh, Cambodia, China, Lao PDR, Malaysia, Myanmar, Thailand 
and Viet Nam. 



TAXONOMIC REVISION OF STERCULIOIDEAE 387 

 

 
 

Fig. 6. Scaphium scaphigerum (Wall. ex G. Don) G. Planch. - A habit sketch (x 0.3). 
 

 Economic aspects: In China, the fruit is used to treat dysentery. The fruit is macerated in 
water in Cambodia, Thailand, and Malaysia, resulting in a massive increase in the volume of the 
outer shell or pericarp, generating a big gelatinous mass. Sweetened jelly is served as a delicacy. It 
also aids in the treatment of diarrhea and dysentery (Ahmed et al., 2009). Studies on biological 
activities reveals that fruit gel powder of Scaphium scaphigerum exhibit glucose absorption 
inhibitory effect and antioxidant activity. The quality-controlled fruit gel powder controls the body 
weight (Phlicharoenphon et al., 2017).  
 

STERCULIA L. 
 Sp. Pl.: 1007 (1753); Miquel, Fl. Ind. Bat. 1: 172 (1859); Benth. & Hook. f., Gen. Pl. 1: 217 
(1862); Mast. in Hook. f., Fl. Brit. Ind. 1: 354 (1874); King, J. As. Soc. Beng. 60: 59 (1891); 
Schumann in Engler & Prantl, Nat. Pflanzenfam. 3: 96 (1895); Ridley, FMP 1: 267 (1922); Bull. 
Misc. Inform. Kew 6: 221 (1938); Prain, Beng. Pl. 1 (Reprint): 186 (1963); Backer & Bakhuizen 
f., FJ. 1: 411 (1964); Phengklai, Fl. Thailand 7 (3): 623 (2001); Bayer & Kubitzki in Kubitzki & 
Bayer (eds.), Fam. Gen. Vasc. Pl. 5: 263 (2003); Cheek in Heywood et al., Flow. Pl. Fam. World: 
311 (2007). 



388 AHMED AND RAHMAN 

 

 Trees or shrubs. Leaves simple, entire or palmately lobed, rarely palmately compound, margin 
entire or dentate. Inflorescence usually axillary, paniculate, rarely racemose. Flowers unisexual. 
Calyx 5-lobed or 5-partite. Petals absent. Male flowers with anthers clustered at top of 
androgynophore, enclosing undeveloped carpels. Female flowers with very short androgynophore, 
staminodes at top of androgynophore in whorl around base of carpels. Carpels 5; ovules 2 to many 
per carpel; styles connate at base; stigmas as many as carpels, free. Fruit a group of follicles, 
usually leathery, less often woody, dehiscent when mature. Seeds 1 to many per follicle, usually 
with endosperm. 
 
Key to species of Sterculia L. 

1 Leaves palmately lobed or digitate. 2 
- Leaves not lobed. 4 
2 Leaves digitate. 3 
- Leaves palmately lobed. S. villosa 
3 Leaflets whitish pubescent beneath; calyx lobes inflexed; staminal 

column 2 mm long. 
S. versicolor 

- Leaflets glabrous beneath when mature; calyx lobes inflexed; staminal 
column 10 mm long. 

S. foetida 

4 Calyx lobes broadly ovate, spreading. 5 
- Calyx lobes linear or linear-lanceolate, connivent at the top. 6 
5 Leaves glabrous above, rusty-tomentose beneath, cordate or 

subcordate at base. 
S. guttata 

- Leaves glabrous on both surfaces, more or less rounded at base. S. lanceifolia 
6 Fruits lanceolate; staminal column 4-5 mm long. S. lanceolata var. 

coccinea 
- Fruits ovate; staminal column less than 4 mm. 7 
7 Leaves glabrous or glabrescent on both surfaces; calyx urceolate; style 

glabrous. 
S. parviflora 

- Leaves densely stellate hairy on both surface; calyx campanulate; style 
hairy. 

S. balanghas. 

 
Sterculia balanghas L., Sp. Pl.: 1007 (1753); Wight, Illus. Ind. Bot. 1: 30 (1840); Mast. in Hook. 
f., Fl. Brit. Ind. 1: 358 (1874); Pickering, Chron. Hist. Pl. : 333 (1879); Gamble, Fl. Pres. Mad. 1: 
106 (1915); Merrill, Enum. Phill. Pl. 3: 53 (1922); Craib, Fl. Siam. Enum. 1: 169 (1925); Ahmed 
et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 345 (2009). Sterculia citrifolia Salisb., Prodr. Stirp. 
Chap. Allerton: 387 (1796); Balanghas telabo Raf., Sylva Tellur.: 72 (1838); Sterculia mollis 
Wall. ex R. Br., Pterocym.: 231 (1844); Sterculia ensifolia Mast. in Hook. f., Fl. Brit. Ind. 1: 358 
(1874); Sterculia balanghas L. var. angustifolia (Roxb.) Mast. in Hook. f., Fl. Brit. Ind. 1: 358 
(1874); Sterculia angustifolia Roxb., Fl. Ind. 3: 148 (1832); Kurz, Fl. Burm. 1: 138 (1877); Ridl., 
Fl. Mal. Pen. 1: 274 (1922); Craib, Fl. Siam. Enum. 1: 165 (1925); Raizada, Ind. For. 67: 245 
(1941).   
 Vernacular name: Balan Udal. 
 English name: Panama tree. 
 A medium-sized tree, young parts rusty tomentose, bark greyish-brown, smooth. Leaves 
simple, 10-30 x 5-13 cm, oblong-ovate, rounded at the base, apex blunt or acute, densely stellate 



TAXONOMIC REVISION OF STERCULIOIDEAE 389 

 

hairy on both surfaces, petioles 3-5 cm long, swollen at both ends, pubescent. Flowers small, 
yellow or greenish-purple, fragrant, on stellate hairy panicles, slightly drooping from ends of 
branches. Calyx campanulate, 5-lobed, 7-10 mm long, pubescent outside, hairy inside. Petals 
absent. Male flowers with 10 stamens, staminal column ca. 2 mm long, with a group of 2-loculed 
anthers at the tip, anthers sessile, staminodes 10. Female flowers with 5 carpels, free, hairy with 4-
8 ovules; styles cohering, densely hairy; stigmas recurved, 5-lobed. Fruit a follicle, 4-5, oblong, 
woody, almost sessile, curved with dark brown tomentose surfaces. Seeds 3-6, oblong-ovoid, 
black, shining.  
 Flowering and fruiting: January-May. 
 Chromosome number: Not known. 
 Ecology: Swampy areas and hilly forests. Also occurs in the deciduous forests. 
 Representative specimen: No specimen was available in any herbarium of Bangladesh. 
 Global distribution: Bangladesh, Cambodia, India, Lao PDR, Malay Peninsula, Myanmar, 
Nepal, Sri Lanka and Thailand. 
 Economic aspects: The plant has ornamental value. Seeds are edible and often consumed after 
roasting. The seeds are nearly palatable as chestnuts. Fiber is obtained from the bark which is used 
in Sri Lanka to build cottages. The wood is soft. Fruit of the species is considered to have cooling 
and laxative effects (Ahmed et al., 2009). 
 

Sterculia foetida L., Sp. Pl.: 1008 (1753); Roxb., Fl. Ind. ed. Carey 3: 154 (1832); Mast. in Hook. 
f., Fl. Brit. Ind. 1: 354 (1874); Kurz, Fl. Burm. 1: 135 (1877); Prain, Beng. Pl. 1: 187 (1903); 
Ridl., Fl. Mal. Pen. 1: 268 (1922); Craib, Fl. Siam. Enum. 1: 166 (1925); Abedin and Ghafoor in 
Nasir & Ali (eds..), Fl. W. Pak. 99: 16 (1976); Malick in Sharma & Sanjappa (eds.), Fl. India 3: 
459 (1993); Verdcourt in Dassanayake et al., Rev. Handb. Fl. Ceyl. 9: 431 (1995); Ahmed et al. 
(ed.), Encycl. Fl. Fauna Bangladesh 10: 346 (2009). Sterculia polyphylla R. Br., Pterocymb.: 227 
(1844); Sterculia mexicana var. guianensis Sagot, Ann. Sci. Nat. Bot. Ser. 6, 11: 153 (1881); 
Clompanus foetida (L.) Kuntze, Revis. Gen. Pl. 1: 77 (1891).  (Fig. 7). 
 Vernacular names: Jangli Badam, Jukigli Badam, Jangal Badam, Udal Badam. 
 English names: Wild Almond, Poon Tree. 
 Medium to large tree, arborescent, deciduous, upto 40 m tall. Branches verticillate and 
spreading. Bark grey to brown, lenticellate; inner bark brown to reddish brown fibrous. Sapwood 
white to cream. Twigs glabrous, swollen towards apex, 2.5 cm in diameter, leaf scars prominent, 
young shoots hairy. Leaves digitately compound, palmately 5-10 foliate, crowded at the end of 
thick branchlets. Stipules present, caducous, subulate, margin pubescent, glabrous adaxially, 
puberulent abaxially, 8 x 3 mm. Petiole 6 cm long, finely pubescent or puberulent. Leaflets 
elliptic-lanceolate, coriaceous, entire, veins pinnately parallel, 7.5-14.0 x 2.0-4.5 cm, apex acute to 
acuminate, base acute to cuneate, glabrous beneath when mature. Inflorescence terminal, 
subterminal or axillary panicles. Flowers unisexual and bisexual, greenish yellow becoming red, 
smell unpleasant; pedunculate, peduncle 5.5 to 8.5 cm long, pedicellate, pedicel 6 mm long; calyx 
campanulate, deeply divided into 5-6 lobes, adnate at the base, linear-oblong to lanceolate, 
subacute, stellate-hairy, 10 x 4 mm. Petals absent. Male flowers with 14-15 stamens, staminal 
column ca. 10 mm long, capitate, hairy at the base, 10-15 anthers grouped into a head, staminodes 
14. Female flowers inserted, ovary globose, densely villous, 5 loculed, each locule with 12-20 
ovules; styles cohering; stigmas very small, 5-lobed or flattened, glabrous. Fruit is follicle, follicle 
curved, arranged in a cluster, boat-shaped, woody, short beaked, smooth outside, fibrous inside, 
10-15 seeded, red when mature. Seeds ca. 20 in each follicle, ellipsoid, 2.5 cm long, 1.3 cm 
diameter, purple-black. 



390 AHMED AND RAHMAN 

 

 Flowering and fruiting: November-April. 
 Chromosome number: 2n = 40 (Kumar and Subramaniam, 1986). 
 Ecology: Usually grows in the hot, tropical lowlands and moderate highlands. Also found in 
the areas with or without a clear dry season. The species prefers a deep, fertile, moist but well-
drained soil in a sunny, sheltered position. 
 

 
  Fig. 7. Sterculia foetida L.: A. Habit (x 0.2); B. Flower (x 1.2); C. Fruit (x 0.2). 
 
 Representative specimens: Bandarban: Betchara Para, 13 Jun 1983, M.K. Alam 4606 
(BFRIH). Chittagong: Hamzarbagh, 13 Mar 1985, Mujib et al. 5345 (BFRIH); Hathazari, 
Fatehpur, Mithachara, 1 Mar 2017, Iqbal Mahmud IQ 606 (DACB). Cox’s Bazar: Kutubida, 27 
Dec 1983, Huq et al. H. 6653 (DACB); Bhomarighona, 24 Apr 2017, Ehsanul Huq EH 91 
(DACB); Maheshkhali, Dineshpur, 2 May 2017, Niyamul Kabir et al. NK 3644 (DACB); 
Chakaria, Fasiakhali, 10 May 2017, Niyamul Kabir et al. NK 3912 (DACB); Himchari, s. dies., 
M.A. Rahman 390 (HCU). Dhaka: s. loc., 15 Sep 1949, S.K. Sen s.n. (DUSH); Azimpur 
Maternity Hospital, 12 Aug 1964, A.F. Muhammad 180 (DUSH); Suhrawardi Udyan, 30 Oct 
2021, Sunzid 29 (DUSH). Munshiganj: Bikrompur, 20 Sep 1996, Nazrul Huq s.n. (DACB). 



TAXONOMIC REVISION OF STERCULIOIDEAE 391 

 

 Global distribution: Bangladesh, Cambodia, China, India, Indonesia, Malaysia, Myanmar, 
Pakistan, Philippines, Sri Lanka, Thailand, Viet Nam, Eastern Africa and North Australia.  
 Economic aspects: The leaves and seeds of the species possess anti-inflammatory and anti-
nociceptive activities. The seeds are fit for human consumption, representing a good source of fats 
(30%–36%) and proteins (11.4%). The leaves and bark are often used as diaphoretic, diuretic, and 
aperient agents and have been applied to the treatment of rheumatism, obesity, gonorrhea, edema, 
and skin disease. The plant serves as a repository of phytoconstituents, including alkaloids, 
flavonoids, terpenoids, phenols, and steroids. Therefore, these phytochemicals can serve as useful 
compounds in the searching of new small drug like potential molecules in the Computer Aided 
Drug Designing endevors (CADD). The ethanolic seed extracts have been reported to show 
antioxidant and anticancer activities (Alam et al., 2021). The sweet yellowish cotyledons are eaten 
after the black seed coat is removed. They are effective for rheumatism relief and are used as a 
laxative, diuretic, anti-epileptic, purgative and insect repellant. Seed oil is useful for lighting and 
painting, as well as treating itches and other skin disorders (Ashrafuzzaman and Sarwar, 2021). 
 
Sterculia guttata Roxb., Fl. Ind. ed. Carey 3: 148 (1832); Mast. in Hook. f., Fl. Brit. Ind. 1: 355 
(1874); Balfour, Cycl. Ind. East. South. Asia 3: 737 (1885); Nairne, Fl. Pl. W. Ind.: 34 (1894); 
Brandis, Ind. Trees : 82 (1921); Craib, Fl. Siam. Enum. 1: 166 (1925); Malick in Sharma & 
Sanjappa (eds.), Fl. India 3: 462 (1993); Ahmed et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 347 
(2009). Astrodendrum malabaricum Dennst., Schlussel Hort. Malab.: 30 (1818); Clompanus 
malabarica Kuntze, Revis. Gen. Pl. 1: 77 (1891).                                             
 Vernacular names: Raiphal, Bansal. 
 English name: Not known. 
 A large evergreen tree, young parts rusty stellate tomentose, bark dark green, warty, cracked 
outside, blaze white, mottled with brown. Leaves simple, ovate, rounded, or slightly cordate at the 
base, acute or abruptly shortly acuminate at the apex, entire, under surface and petioles tomentose, 
petioles 2-7 cm long, stipules ensiform, caducous. Flowers white and pale yellow, in terminal, 
rusty, tomentose, racemiform panicles, pedicels 2-3 mm long, hairy, with an offensive smell. 
Calyx campanulate, deeply 5-partite, lobes narrowly lanceolate, rusty tomentose outside, glandular 
and with scattered long hairs inside. Petals absent. Male flowers with 10-12 stamens, staminal 
column 6-8 mm long, slender, curved, anthers 10-15, sessile, very small, staminodes 10. Female 
flowers with 5 carpels, free, ovary surmounting the ring of anthers, globose, woolly; styles curved; 
stigmas 5-lobed. Fruit a follicle, red when ripe, coriaceous, compressed, tomentose outside, 
smooth and reddish inside, with 3-5 seeds in each follicle. Seeds black and large. 
 Flowering and fruiting: April-August. 
 Ecology: The species occurs along the margins of evergreen forests, and in semi-evergreen 
forests. Very rare in moist, low-lying areas. 
 Representative specimens: Bandarban: Keokradong, Darjiling Para, 26 Mar 2019, 
Khandakar Kamrul Islam KKI 3584 (DACB). Chittagong: Hazarikhil, 30 Mar 1976, D.K. Das 
s.n. (BFRIH). Sylhet: Lawachara, 17 Mar 1984, M.K. Alam 4975 (BFRIH). 
 Global distribution: Bangladesh, India, Myanmar, Sri Lanka, Thailand and Viet Nam. 
 Economic aspects: One kind of coarse rug is manufactured by beating and washing the inner 
bark in the Malabar Coast. Seeds are eaten after roasting. Seeds contain important bioactive 
compounds that might be used in the development of effective insecticides as shown in-vitro tests 
to control larvae of mosquitos (Katade et al., 2006). 
 



392 AHMED AND RAHMAN 

 

Sterculia lanceolata var. coccinea (Jack) Phengklai, Thai. Forest. Bull. Bot. 23: 99 (1995); 
Rehder et Sargent, Bard. Bibl. 2: 538 (1911); Craib, Contrib. Fl. Siam. 1: 24 (1912); Merrill, Bibl. 
Enum. Born. Pl.: 379 (1921). Sterculia coccinea Roxb. [Hort. Beng. : 50 (1814) nom. nud.], Fl. 
Ind. ed. Carey 3: 151 (1832) non Jack (1822); Mast. in Hook. f., Fl. Brit. Ind. 1: 359 (1874). 
Clompanus hamiltonii O. Kuntze, Rev. Gen. Pl. 1: 77 (1891). Sterculia hamiltonii (Kuntze) 
Adelb., C.A. Backer, Bekn. Fl. Java 4b (107): 23 (1944). Sterculia indica Men., J. Ann. Arb. 33: 
245 (1952).              (Fig. 8). 
 Vernacular name: Toni Udal. 
 English name: Not known.  
 Small tree, up to 15 m tall, ca. 30 cm in diameter. Arborescent, buttress absent. Bark grey, 
smooth; inner bark fibrous. Sapwood white. Twigs grey to pale brown, glabrous but apical parts 
sometimes with sparse stellate hairs, slender, 0.2-07 cm diameter. Stipules caducous, linear, up to 
0.7 cm long, creamy. Leaves alternate or spirally arranged, simple, lamina papery, broadly elliptic, 
ovate to slightly obovate-oblong, 7-17 x 3-7 cm, base acute to rounded, symmetric to slightly 
asymmetric, with 3-5 basal veins, apex often distinctly caudate with slender acumen to 2 cm long, 
occasionally acuminate; upper surface glabrous, drying brown, lower surface with sparse stellate 
hairs or glabrous; lateral veins 5-7 pairs, raised on both surfaces, arching to form intramarginal 
vein; petioles slender, very pale brown to yellow, glabrous, 1-7 cm long, swollen and darkened at 
both ends. Inflorescences sub-terminal or axillary racemes, often branched, up to 12 cm long, with 
sparse stellate hairs, each branch 3-flowered. Flowers unisexual, pale white to green-yellow, calyx 
tubular to urceolate, 2-15 mm long, 2-4 mm in diameter, outer surface with sparse stellate hairs, 
inner surface glabrous, 5-lobed, lobes narrowly oblong, 2-3 mm long, with dense simple hairs 
along margin, joined or not at apex, androgynophore 1-2 mm long, erect, glabrous. Male flowers 
with 10 anthers present at the globose head. Female flowers with ca. 1 mm long androgynophore; 
styles coherent, with sparse stellate hairs; stigmas obovate with sparse stellate hairs. Fruit a cluster 
of 1-5 orange-red leathery follicles; follicles up to 10 cm long, 1.5-3 cm wide, outer surface with 
dense minute stellate hairs, inner surface glabrous. Seeds 3-5 in each follicle, ellipsoid, 0.9-1.6 cm 
long, 0.6-1.2 cm in diameter, black in color.  
 Flowering and fruiting: April-September. 
 Ecology: Hill slopes along streams. 
 Representative specimens: Bandarban: Ruma, Thanapara, 10 May 2018, Khandakar Kamrul 
Islam KKI 2791 (DACB); Ruma, Bogalake, 26 Mar 2019, Khandakar Kamrul Islam KKI 3510 
(DACB). Chittagong: Badolchari, 25 Jan 1997, M.A. Rahmand et Aditi Khisa 665B (HCU); 
Himchari, Barachara, 29 Jun 1997, M.A. Rahman et al. 1422 (HCU); Dhopachori, 2 Sep 1999, 
M.A. Rahman et al. 5530 (HCU); Hazarikhil Wildlife Sanctuary, 19 Aug 2014, S.N. Uddin N. 
5469 (DACB). Cox’s Bazar: Ruma, Eid Ghor, 20 Mar 2018, Niyamul Kabir et Mehedi Hassan 
NK 7218 (DACB). Khagrachari: Dighinala, Kowser et al. KH 6689 (DACB). Maulvibazar: 
Madhabkunda, 25 Jun 2001, S.N. Uddin N. 1086 (DACB); Lawachara National Park, 17 Aug 
2009, S.N. Uddin N. 3896 (DACB); Kamalganj, Adampur, 20 Sep 2011, S.N. Uddin N. 4745 
(DACB). Rangamati: Kutukchari, Chegaiyachari, 16 Jun 1998, S.B. Uddin et al. 3125 (HCU); 
Bilaichari, Farua, 23 Jul 2009, S.N. Uddin N. 3802 (DACB). Sylhet: Golapganj, 3 Mar 1976, 
Atiqur Rahman 60 (DACB); Jainta-Sarighat, 3 Oct 1983, Huq et al. H. 5337 (BFRIH); Jaintapur, 
19 Oct 1986, A.M. Huq et M.K. Mia H. 7839 (DACB). 
 Global distribution: Bangladesh, Bhutan, India, Lao PDR, Malaysia, Myanmar, Nepal, 
Thailand and Viet Nam. 
 Economic aspects: Wood is hard, strong. Usually used for construction works especially for 
the production of poles, posts and ridge plates. Fruits are edible and enriched with nutrients, 



TAXONOMIC REVISION OF STERCULIOIDEAE 393 

 

usually consumed after cooking. Seeds are also edible and are eaten fried or roasted (Ahmed et al., 
2009; Alam, 2018). 
 

 
Fig. 8. Sterculia lanceolata var. coccinea (Jack) Phengklai - A habit sketch (x 0.3). 

 
Sterculia parviflora Roxb. ex G. Don., Gen. Hist. 1: 516 (1831); Roxb., Fl. Ind. ed. Carey 3: 147 
(1832); Mast. in Hook. f., Fl. Brit. Ind. 1: 356 (1874); Ridl., Fl. Mal. Pen. 1: 271 (1922); Malick in 
Sharma & Sanjappa (eds.), Fl. India 3: 468 (1993); Ahmed et al. (ed.), Encycl. Fl. Fauna 
Bangladesh 10: 348 (2009). Sterculia maingayi Mast. in Hook. f., Fl. Brit. Ind. 1: 359 (1874); 
Sterculia holttumii Ridl., Kew Bull. 1926: 471 (1926).   
 Vernacular name: Parvi Udal. 
 English name: Not known.  
 Arborescent, up to 35 m in height, ca. 100 cm in diameter. Buttress plank-like up to 1 m tall. 
Bark grey-orange to brown, smooth with occasional small scales and lenticels; inner bark red-
brown, 1 cm thick, fibrous. Sapwood white. Twigs 0.3-0.6 cm in diameter, slightly swollen at the 
apex, glabrous but sometimes stellate hairs on the very young parts, leaf scars conspicuous. 
Stipules caducous. Leaves alternate, simple, lamina papery to leathery, ovate, elliptic to elliptic-
oblong, 8-20 x 4-10 cm, base slightly cordate to truncate, basal veins 1-2 pairs, apex acute or very 
shortly acuminate; upper surface glabrous, lower surface with sparse, very minute, round, peltate 
scales; lateral veins excluding basal ones, 5-7 or 12 pairs, straight or arching, sunken above, 



394 AHMED AND RAHMAN 

 

prominent beneath; intercostal veins ladder-like, sunken above, often prominent beneath, petioles 
1.5-8 cm long, glabrous. Inflorescences sub-terminal or axillary panicles, much branched, erect, 9-
20 cm long, with dense stellate hairs or scurfy; bracts caducous. Flowers unisexual, pale yellow 
becoming reddish pink; calyx tube urceolate, 3-5 mm long, outer surface with dense stellate hairs, 
inner surface glabrous, lobes 5, lanceolate, 3-4 mm long, inner surface with sparse hairs, 
converging and joined at apex. Male flowers with slender androgynophore, 0.5-0.25 mm long, 
glabrous; anthers 10, forming a globose head at the apex of androgynophore. Female flowers with 
0.5-1 mm long androgynophore, base of carpels surrounded by 10 sterile anthers, carpels 5, 
glabrous or with sparse hairs; styles coherent, slender, ca. 0.5 mm long, glabrous or with a few 
hairs; stigmas 5, ovate, curved. Fruit a cluster of up to 5 orange-red leathery follicles; follicles 
oblong, 6-9 cm long, 2-3 cm wide, with dense minute stellate hairs. Seeds 2 or more in each 
follicle, ellipsoid, 1.2-2 cm long, 0.8-1.5 cm in diameter, black. 
 Flowering and fruiting: February-July. 
 Ecology: Lowland and mixed deciduous forests. Found in a variety of substrates including 
basalt and calcareous shale. 
 Representative specimen: No specimen was available in any herbarium of Bangladesh. 
Global distribution: Bangladesh, Cambodia, India, Malaysia, Myanmar, Thailand and Viet Nam. 
 Economic aspects: The white colored wood is used for indoor construction and plywood. 
Good quality fibers can be obtained from the bark of the species. The tree also has ornamental 
value as it is sometimes planted in the garden as well as in the roadside avenue (Alam 2018). 
  
Sterculia lanceifolia Roxb., Fl. Ind. (eds) 3: 150-151 (1832); Laudon et Spottiswoode, Gard. Mag. 
Reg. 4: 450 (1826); Kurz, Asiatic Soc. Beng., Journ. Asiat. Soc. 45(2): 120 (1876); Western 
Australia Museum, Rec. Wes. Aust. Mus. 23: 115 (2006). Sterculia ovalifolia Wall., Numer. List: 
1132 (1829); Sterculia lanceifolia G. Don., Sylhet 1: 517 (1831); Southwellia roxburghiana 
Spach, Hist. Nat. Veg. 14: 402 (1847); Clompanus roxburghii (Wall.) Kuntze, Revis. Gen. Pl. 1: 
78 (1891).   (Fig. 9). 
 Vernacular name: Ushli. 
 English name: Not known. 
 Trees or shrubs. Bark grayish. Branchlets puberulent. Stipules subulate, minutely hairy, 
caducous. Leaves petiolate. Petioles 2.5-3.5 cm, both ends pulvinate. Lamina lanceolate, linear-
lanceolate or oblong-lanceolate, 10-23 x 2.5-7.5 cm, minutely pilose, becoming glabrous, lateral 
veins 9-10 on each side of midrib, slightly connected near margin, base rounded or obtuse, apex 
obtusely acuminate. Inflorescence racemose, or rarely paniculate, axillary, 5-7 cm, much shorter 
than leaves, few flowered, stellate pilose. Pedicel slender, 5-8 mm. Calyx red, campanulate, 
divided nearly to base, ca. 7 mm, abaxially sparsely puberulent, lobes oblong-lanceolate, ca. 5mm, 
much longer than calyx tube, spreading outward. Male flowers with curved androgynophore, 
surface glabrous. Female flowers with globose ovary, densely puberulent. Follicle oblong or 
oblong-lanceolate, ca. 7 cm in length, seeds 4-8, abaxially densely red hirsute, apex beaked. Seeds 
black, ovoid.  
 Flowering and fruiting: February-August. 
 Ecology: The species is commonly found in the hilly forests.  
 Representative specimens: Maulvibazar: Kamalganj, 7 Mar 2011, S.N. Uddin N4472 
(DACB). 
 Global distribution: Bangladesh, China, India, Lao PDR, Malaysia, Myanmar, Nepal, 
Thailand and Viet Nam. 



TAXONOMIC REVISION OF STERCULIOIDEAE 395 

 

 
    Fig. 9. Sterculia lanceifolia Roxb. - A habit sketch (x 0.3) 
 
 Economic aspects: Timber serves as the source of fuel. It can also be used for different 
construction purpose and infrastructure development. Seeds are edible, often used in the form of 
condiments for seasoning of foods (Uphof, 1959). 
 
Sterculia versicolor Wall., Pl. Asiat. Rar. 1: 48, t. 59 (1830); Mast. in Hook. f., Fl. Brit. Ind. 1: 
355 (1874); Kurz, Fl. Burm. 1: 135 (1877); Just, Botanis. Jahres. 6(2): 966 (1878); Ahmed et al. 
(ed.), Encycl. Fl. Fauna Bangladesh 10: 349 (2009); Uddin & Hassan, Vas. Fl. Chit. & Chit. Hill 
Tr. 2: 427 (2018). Southwellia versicolor (Wall.) Wight, Ill. Ind. Bot. 1: 77 (1838); Clompanus 
versicolor Kuntze, Revis. Gen. Pl. 1: 78 (1891).    (Fig. 10). 
 Vernacular name: Ranga Udal. 
 English name: Not known. 
 Large tree with spreading crown, 24-27 m tall, trunk straight, stout, bark grey, branchlets with 
prominent scars of fallen leaves. Leaves peltate, digitately 5-7 foliate, elliptic-lanceolate, 18.5-
28.9 x 4-16 cm, tapering at the base, acute or shortly acuminate at the apex, subcoriaceous, 
glabrous above, puberulous or glabrescent beneath, petioles 8-15 cm long. Flowers fragrant, 
orange-yellow or pale-yellow, many, in erect panicles crowded at the ends of branchlets, pedicels 
short. Calyx campanulate, hairy, ca. 1 cm long, lobes oblong, inflexed. Male flowers with ca. 2 
mm long staminal column, curved, filaments short, anthers 2-loculed. Female flowers with 5-



396 AHMED AND RAHMAN 

 

lobed ovary, hairy with sterile anthers at the base; styles ca. 2 mm long, villous, curved with 
radiating sub-peltate stigmas. Fruit a follicle, slightly compressed, coriaceous. Seeds 7-15 mm 
long, oblong. 
 

 
  Fig. 10. Sterculia versicolor Wall. - A habit sketch (x 0.5). 
 
 Flowering and fruiting: March-July. 
 Ecology: The species occurs in the hill forests. 
 Representative specimens: Chittagong: Maheshkhali, 7 Mar 1978, M.S. Khan et al. K. 4895 
(DACB). 
 Global distribution: Bangladesh, India and Myanmar. 
 Economic aspects: Mainly valued for timber. Timber can serve as a useful source of fuel 
wood for its high heat yielding capacity. Seeds are edible, often eaten after roasting. Fiber 
obtained from bark is used to make ropes and cordage (Ahmed et al., 2009; Alam, 2018). 
 
Sterculia villosa Roxb., Fl. Ind. ed. Carey 3: 153 (1832); Mast. in Hook. f., Fl. Brit. Ind. 1: 355 
(1874); Prain, Beng. Pl. 1: 187 (1903); Heinig, List Pl. Chitt. Coll. & HT.: 7 (1925); Sinclair, Bull. 
Bot. Soc. Beng. 9(2): 88 (1955); Malick in Sharma & Sanjappa (eds.), Fl. India 3: 472 (1993); 
Ahmed et al. (ed.), Encycl. Fl. Fauna Bangladesh 10: 350 (2009). Sterculia armata Mast. in Hook. 
f., Fl. Brit. Ind. 1: 357 (1874); Clompanus armata (Mast.) Kuntze, Revis. Gen. Pl. 1: 78 (1891); 



TAXONOMIC REVISION OF STERCULIOIDEAE 397 

 

Clompanus villosa (Roxb. ex Sm.) Kuntze, Revis. Gen. Pl. 1: 78 (1891); Sterculia ornata Wall. ex 
Kurz in J. Asiat. Soc. Beng. 42 (2): 228 (1873); Kurz, Fl. Burm. 1: 136 (1877); Brandis, Ind. 
Trees: 81 (1921); Craib, Fl. Siam. Enum. 1: 168 (1925); Sterculia lantsangensis Hu, Bull. Fan 
Mem. Inst. Biol. Bot. 8 (1): 42 (1937).               (Fig. 11). 
 Vernacular names: Udal, Chandul, Sambeing (Marma), Frit (Marma), Crukhaoya Bang 
(Marma), Lambuk (Tripura), Sibo toigo kalai (Tripura), Ya Sing (Murong). 
 English name: Elephant Rope Tree. 
 Small to medium-sized tree, arborescent, deciduous, 10-15 m tall, bark grey-white, ca. 2.5 cm 
thick. Branchlets robust with heart shaped leaf scars of fallen leaves, brown stellate pubescent to 
tomentose when young. Leaves simple, crowded at the end of branchlets when young; stipulate, 
stipules lanceolate, ca. 10 mm long, petiolate, petiole cylindrical, 15-40 cm long, minutely 
tomentose, leaf blade deeply cordate at the base sometimes, entire, coriaceous, minutely 
pubescent, apex caudate, abaxially densely yellow-brown stellate tomentose, adaxially sparsely 
pubescent, 30-40 x 30-45 cm, palmately 3-7 lobed, each lobe again 3 lobed, central lobe broadly 
ovate, veins 5, palmately reticulate. Inflorescence subterminal on branchlets, paniculate, densely 
ferruginous stellate tomentose. Flowers bisexual, incomplete, irregular, ebracteate, pinkish-yellow 
in crowded drooping panicles from the end of the thick shoots, male and female flowers 
intermixed, pedunculate, peduncle 3-18 cm long, pedicellate, pedicel 5 mm long. Calyx broadly 
campanulate, pinkish inside, 5 lobed, lobes ovate, abaxially pubescent, adaxially glabrous, 7 x 4 
mm. Petals absent. Male flowers with 10 stamens, staminal column 2-3 mm long, recurved, 
glabrous, anthers 10, sessile, staminodes 10. Female flowers with 5 carpels, ovary globose, 
strigose, 5 loculed, many-ovuled; style inserted, recurved, hairy; stigmas flattened or with 5 lobes, 
extrorse. Fruit is follicle, 3-5 cm in diameter, oblong to ellipsoid, apex shortly beaked, sessile, 
leathery, rusty pubescent, many seeded, red when ripe. Seeds 3-5, oblong, smooth, black.  
 Flowering and fruiting: February-May. 
 Chromosome number: 2n = 40 (Kumar and Subramaniam, 1986). 
 Ecology: Hill slopes in mixed deciduous forests. 
 Representative specimens: Bandarban: Roangchari, 16 Jun 2001, M. Rahman 8703 
(BFRIH). Chittagong: Pablakhali-Saratali, 30 Apr 1977, Huq et Rahman H. 3275 (DACB); s. 
loc., 3 May 1977, A.M. Huq 3354 (DACB); Chunati Wildlife Sanctuary, 8 May 1994, A.M. Huq 
et M.K. Mia 10372 (DACB); Mirsori, Mohamaya lake, 21 Feb 2018, Moniruzzaman et al. MAK 
7737 (DACB); Fatikchari, Hazarikhil, 18 Apr 2018, Moniruzzaman et Kowser MK 8275 (DACB). 
Cumilla: Cumilla University Campus, 29 Aug 2016, Tajul et al. TOK 197 (DACB). Dhaka: 25 
Mar 1964, D.K. Das FKH 365 (BFRIH); Savar, 5 Jun 1978, Soejarto et M. Rahman 4974 
(DACB); Dhamrai, 29 Jan 1982, A.M. Huq 5519 (DACB); Dhaka University Botanical Garden, 
19 Aug 2021, Sunzid 3 (DUSH). Khagrachari: Panchuri, 5 Sep 2016, Kowser et al. KH 392 
(DACB). Mymensingh: Jamalpur, Gajni forest, 7 May 1982, Mia et al. M. 817 (DACB); 
Bangladesh Agricultural University Botanical Garden, 2 Jan 2022, Sunzid 50 (DUSH). Rangpur: 
Rangpur Sadar, 15 Apr 1994, M. Mohiuddin 1107 (BFRIH). Sherpur: Jhenaighati, Gazni, 9 Feb 
1985, Khan et al. K. 7041 (DACB). Tangail: Pirgacha, 14 Jun 1989, M.K. Mia 211 (DACB). 
 Global distribution: Bangladesh, Cambodia, China, India, Lao PDR, Myanmar, Nepal, 
Pakistan and Thailand.  
 Economic aspects: Ropes, cordage, and bags are made from a coarse fiber produced from the 
inner bark. The tree produces a transparent gum that is utilized in medicinal preparation. Baked or 
roasted seeds are popular. The Chittagong Hill Tracts tribal people use the bark to build straps for 
their long bamboo basket, known as "Turung" which they wear on their heads (Ahmed et al., 
2009). Traditionally the plant is used as an agent in diuretic, cooling and aphrodisiac purposes. 



398 AHMED AND RAHMAN 

 

The plant is also utilized by Indians as traditional remedy for inflammation. Sherbet, prepared 
from the petiole of the plant along with water and sugar is given in urinary problems and 
rheumatism. The bark and the petiole are used as a remedy in seminal weakness. White exudates 
of the tree are used for throat infection. Root infusion is taken as food adjunct while the whole 
plant extract is useful for skin diseases. The plant also has anthelmintic, antidiabetic, 
antimicrobial, membrane stabilization and antithrombotic activity. Some chemical constituents 
like flavonoids, chrysoeriol, diosmetin-7-O-β-D-glucoside and hrysoeriol-7-O-β-D-glucoside have 
been isolated from the species. These important phytochemicals can be used for computer aided 
drug discovery against suitable target using molecular docking, ADMET (absorption, distribution, 
metabolism, excretion and toxicity) and molecular dynamics simulation approaches (Hossain et 
al., 2013). 
 

 
Fig. 11. Sterculia villosa Roxb.: A. Habit (x 0.3); B. Flower (x 1.7); C. Calyx (x 1.3); D. L.S. of 

flower (x 1.0). 
 
Discussion 
 Taxonomic revisions are required to keep track of potential changes and implications, as 
every new species discovered has an impact on the currently accepted system of classification 
(Stuessy, 1975). Phenotypic plasticity further necessitates the need of taxonomic revision as it is 
responsible for the expression of different types of phenotypes from the same genotype depending 
on the environmental effect (DeWitt et al., 1998). The present revisionary studies in the subfamily 
Sterculioideae with 16 taxa is the first on its nature in Bangladesh. Sterculioideae in Bangladesh 



TAXONOMIC REVISION OF STERCULIOIDEAE 399 

 

incorporates a good number of economically and medicinally important taxa (Alam, 2018). 
Several attempts have been made worldwide to shed light on the current state of the member taxa 
of Sterculioideae. Taylor (1989) revised neotropical taxa of Sterculia L. which solved the 
controversy regarding the sexuality of the two floral types. The genus Brachychiton was 
previously included under Sterculia and its generic status was ambiguous. Later, Guymer (1988) 
resolved it into a distinct genus using characteristic features of follicles, seed coats and embryo. 
The problem of species delimitation in the genus Scaphium was solved by Wilkie (2009) with 
recognition of eight species which was four in the previous revision conducted by Kostermans 
(1953). 
 Several floristic studies have reported the occurrence of the member taxa of Sterculioideae 
throughout the country but no detailed revisionary studies have been carried out so far. The 
present investigation revealed five genera of Sterculioideae in Bangladesh such as Firmiana, 
Heritiera, Pterygota, Scaphium and Sterculia with two, four, one, one and eight taxa, respectively. 
Among the taxa studied, four taxa belonging to three genera could not be examined because of 
inavailability of these species and no specimens belonging to these species were deposited in any 
herbaria of the country. These four taxa were Firmiana simplex, Heritiera papilio, Sterculia 
balanghas and Sterculia parviflora. Firmiana simplex was reported as Sterculia urens Roxb. by 
Mia et al. (2011) based on the specimen examined by Hooker & Thomson in 1850, where no 
specific location was mentioned. Ahmed et al. (2009) stated the occurrence of Heritiera papilio 
from the forests of Sylhet district. Sterculia balanghas was reported by Raizada (1941) from the 
Chittagong Hill Tracts. Hooker (1874) reported Sterculia parviflora from the territory which at 
present falls under Sylhet region.  
 Based on the field observations, herbarium specimens and relevant literature (Ahmed et al., 
2009; IUCN, 2012; Mia et al, 2011; Rahman et al., 2012; Alam, 2018; Ashrafuzzaman and 
Sarwar, 2021) the status of the taxa of the subfamily Sterculioideae has been determined. Among 
the studied taxa, Sterculia villosa and S. lanceifolia var. coccinea (= Sterculia hamiltonii) were 
found as common and considered as Least Concern (LC). Firmiana colorata, Sterculia foetida, 
Pterygota alata, Heritiera fomes, H. littoralis and H. macrophylla are rarely found in nature and 
going to be Near Threatened (NT). Scaphium scaphigerum was found as Vulnerable (VU), while 
Firmiana simplex, Sterculia guttata and Sterculia versicolor were found to be Endangered (EN).  
In the recent past, Ara et al. (2013) reported Sterculia versicolor as Endangered in Bangladesh that 
was supported by the present study. As a continuation of the current revision, we aim to apply the 
techniques of molecular systematics including DNA barcoding and phylogenetic analyses which 
would further strengthen the systematic position of the taxa of Sterculioideae in Bangladesh. 
 
Acknowledgements 
 The authors gratefully acknowledge the financial support from the Ministry of Science and 
Technology for conducting this study. Thanks are due to the authorities of the DACB, DUSH, 
HCU and BFRIH for providing facilities to examine specimens and to consult their libraries.  
 
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(Manuscript received on 10 July, 2022; revised on 16 November, 2022)