TILIACEAE 1. TILIA Linnaeus, Sp. Pl. 1: 514. 1753.

TILIACEAE 

椴树科 duan shu ke 

Tang Ya (唐亚) 1 ; Michael G. Gilbert 2 , Laurence J. Dorr 3 

Trees, shrubs, or herbs. Leaves simple, alternate or rarely opposite, basally veined, entire or serrate, sometimes lobed; stipule, 

when present, caducous or persistent. Inflorescences cymose or cymose-paniculate. Flowers bisexual or unisexual (plants dioecious), 

actinomorphic. Bracts caducous or sometimes large and persistent. Sepals (4 or)5, free or sometimes basally connate, valvate. Petals 

as many as sepals, sometimes absent, free, usually glandular on adaxial surface. Androgynophore present or absent. Stamens 

numerous, rarely 5, free or connate into fascicles at base; anthers 2-loculed, dehiscence longitudinal or apical; petaloid staminodes 

alternating with petals or absent. Ovary superior, 2–6-loculed, sometimes more; ovules 1 to many per locule; placentation axile; style 

simple, sometimes free; stigma acute or peltate, usually lobed. Fruit usually a drupe, capsule, or schizocarp, sometimes a berry or 

samara, 2–10-loculed. Seeds without aril; endosperm copious; embryo erect; cotyledons flat. 

About 52 genera and ca. 500 species: primarily in tropical and subtropical areas; 11 genera and 70 species (32 endemic) in China. 

Molecular data have shown that the members of the Tiliaceae as here defined fall clearly into four clades that can either be treated as subfamilies 

within an enlarged Malvaceae or as families in their own right. In this view the Tiliaceae/Tilioideae is restricted to Tilia and Craigia, along with the 

Central American Mortoniodendron Standley & Steyermark; Colona, Corchorus, Grewia, Microcos, and Triumfetta are placed in the Sparrmanniaceae/Grewioideae; 

Berrya and Diplodiscus are placed in the Brownlowiaceae/Brownlowioideae; and Burretiodendron and Excentrodendron form a 

basal group of uncertain placement, possibly most closely allied to the Pentapetaceae/Dombeyoideae which includes mostly genera here placed in the 

Sterculiaceae. 

Plagiopteron Griffith, included in the Tiliaceae in FRPS (49(1): 48–50. 1989), is now regarded as either a member of the Celastraceae or a 

separate family, Plagiopteraceae (see Fl. China 11). 

Chang Hung-ta & Miau Ru-huai. 1989. Tiliaceae. In: Chang Hung-ta, ed., Fl. Reipubl. Popularis Sin. 49(1): 47–123. 

1a. Herbs or subshrubs. 

2a. Androgynophore absent; petals eglandular; fruit a smooth capsule ........................................................................ 3. Corchorus 

2b. Androgynophore present; petals glandular; fruit a spiny or bristly capsule ............................................................ 7. Triumfetta 

1b. Trees or shrubs. 

3a. Each inflorescence with a single large, persistent, band-shaped bract partially connate with peduncle .......................... 1. Tilia 

3b. Inflorescence without such a bract. 

4a. Fruit without wings, at the most ridged. 

5a. Calyx campanulate; fruit a capsule; androgynophore absent ................................................................... 8. Diplodiscus 

5b. Calyx with free sepals; fruit a drupe, often 2- or 4-lobed; short androgynophore present. 

6a. Inflorescences paniculate, terminal; stigma subulate, not lobed ............................................................ 5. Microcos 

6b. Inflorescences cymose, usually axillary; stigma swollen, lobed .............................................................. 6. Grewia 

4b. Fruit with well-defined wings. 

7a. Calyx campanulate; ovary 3-loculed; fruit with 3 pairs of long wings ............................................................ 9. Berrya 

7b. Calyx divided to base or near to base; ovary 3–5-loculed; fruit with 3–5 wings. 

8a. Flowers bisexual. 

9a. Basal veins 3; inflorescences axillary, cymose; petals absent ............................................................ 2. Craigia 

9b. Basal veins 5–7; inflorescences usually terminal, paniculate; petals 5 .............................................. 4. Colona 

8b. Flowers unisexual or polygamous. 

10a. Leaf blade leathery, glabrous, basal veins 3 .................................................................. 10. Excentrodendron 

10b. Leaf blade papery, stellate scaly or stellate pilose, basal veins 5–9 ............................... 11. Burretiodendron 

1. TILIA Linnaeus, Sp. Pl. 1: 514. 1753. 

椴树属 duan shu shu 

Trees deciduous. Leaves alternate; stipule caducous; leaf blade base usually slightly oblique, cordate, truncate, or rounded, margin 

usually serrate or sometimes entire, often with hairy domatia in axils of veins. Inflorescences axillary, cymose, 3- to many-flowered. 

Bracts adnate to inflorescence peduncle, band-shaped, large, persistent. Flowers bisexual. Sepals 5, with adaxial nectary at 

base. Petals 5, white or yellow, imbricate. Stamens many, free or connate into 5 fascicles; anthers 2-locular, dorsifixed; staminodes 

when present petaloid, tightly enclosing style and stigma, alternating with petals. Ovary 5-loculed, ovoid, usually hairy after anthesis; 

1 

Department of Environmental Sciences, Sichuan University, Chengdu, Sichuan 610065, People’s Republic of China. 

2 

Missouri Botanical Garden c/o Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom. 

3 

United States National Herbarium, Department of Botany, National Museum of Natural History, MRC-166, Smithsonian Institution, P.O. Box 37012, Washington, DC 

20013-7012, U.S.A. 

240


ovules 2 per locule; style glabrous; stigma apparent, 5-lobed. Fruit a nut or capsule, globose, obovoid, ellipsoid, or ovoid, pericarp 

usually hairy, mostly woody or fragile and indehiscent, rarely leathery and dehiscent when dry. Seeds 1 or 2. 

Between 23 and 40 species: primarily in temperate and subtropical regions; 19 species (15 endemic) in China. 

Tilia is very distinct by its unique bracts adnate to the peduncle of the inflorescence. It is in the process of being monographed by C. D. Pigot, 

who has studied many populations in China, but this work is not yet available. An initial publication (Pigott, Edinburgh J. Bot. 59: 239–246. 2002) 

indicated that he intends to accept only 23 species in total, 13 of these occurring in China, and he intends to include many of the more recently 

described Chinese species within the very variable Tilia tuan. Chromosome numbers were given for all those taxa that he accepts. Tang and Zhuge 

(Acta Phytotax. Sin. 34: 254–264. 1996) recognized 14 “good” species in China, 17 in E Asia, and 25 in total, but Tang noted that more study for a 

taxonomic revision is needed. 

Modern molecular analyses indicate that Tilia and Craigia form a very distinct lineage that could be treated as a distinct family or subfamily 

restricted to just these two genera plus the poorly known Central American genus Mortoniodendron Standley & Steyermark. 

1a. Fruit exocarp leathery, dehiscent when dry ............................................................................................................. 1. T. endochrysea 

1b. Fruit exocarp woody or thickly leathery, indehiscent. 

2a. Exocarp thickly leathery, fragile; bract with stalk. 

3a. Leaf blade 4–6 cm; fruit 5-angled. 

4a. Cymes 5–8 cm; leaf blade apex usually 3-lobed; petaloid staminodes present; fruit obovoid ............ 15. T. mongolica 

4b. Cymes 3–5 cm; leaf blade apex usually not lobed; staminodes absent; fruit ovoid ............................ 16. T. amurensis 

3b. Leaf blade 5–10 cm; fruit not angled or only weakly so. 

5a. Ovary glabrous; cymes (20–)40–100-flowered, with bracteoles .................................................... 19. T. jiaodongensis 

5b. Ovary hairy; cymes 3–16-flowered, without bracteoles. 

6a. Leaf blade orbicular, brown when dry; fruit ovoid; sepals abaxially sparsely stellate hairy; 

pedicel 5–8 mm; bracts 3.5–6 cm ...................................................................................................... 17. T. japonica 

6b. Leaf blade ovate or ovate-triangular, greenish when dry; fruit obovoid; sepals abaxially 

glabrous; pedicel 10–15 mm; bracts 5–8.5 cm ........................................................................... 18. T. paucicostata 

2b. Exocarp woody, hard; bract sessile or with short stalk, basal 1/3–1/2 adnate to peduncle. 

7a. Fruit not angled, apex rounded. 

8a. Leaf blade margin entire or with a few teeth near apex .................................................................................. 10. T. tuan 

8b. Leaf blade prominently serrate. 

9a. Bracts with stalk 5–8 cm. 

10a. Leaf blade broadly ovate, abaxially hairy in vein axils only ................................................. 13. T. chingiana 

10b. Leaf blade ovate, suborbicular, or ovate-oblong, abaxially uniformly tomentose or 

puberulent. 

11a. Branchlets hairy; bracts 12–16 cm ......................................................................................... 10. T. tuan 

11b. Branchlets glabrous; bracts 6–9 cm ............................................................................. 7. T. likiangensis 

9b. Bracts usually sessile. 

12a. Leaf blade submembranous, abaxially glabrous or hairy in axils of veins only ............ 11. T. membranacea 

12b. Leaf blade thicker, not membranous, abaxially uniformly hairy. 

13a. Leaf blade marginal teeth awnlike, 1.5–5 mm; bracts 3.5–9 cm .............................. 2. T. mandshurica 

13b. Leaf blade marginal teeth smaller, not awnlike; bracts 8–12 cm. 

14a. Cymes 3–22-flowered; leaf blade base usually oblique ............................................... 10. T. tuan 

14b. Cymes 3–12-flowered; leaf blade base symmetrically cordate ........................ 12. T. miqueliana 

7b. Fruit prominently or weakly 5-angled, apex acute or obtuse. 

15a. Old leaves abaxially glabrous or hairy in axils of veins only; branchlets glabrous or rarely hairy 

when very young. 

16a. Cymes 50–200-flowered ........................................................................................................ 14. T. taishanensis 

16b. Cymes 1–15-flowered. 

17a. Bract with stalk 0.4–2 cm. 

18a. Leaf blade suborbicular, marginal teeth 3–5 mm; bracts 7–10 × 1–1.3 cm .............. 3. T. henryana 

18b. Leaf blade broadly ovate, marginal teeth small and dense; bracts 

6–6.5 × 1.5–1.8 cm .................................................................................................... 8. T. callidonta 

17b. Bract sessile. 

19a. Leaf blade (5.5–)10–14(–17) cm; fruit weakly 5-angled ............................................... 9. T. nobilis 

19b. Leaf blade 5–10 cm; fruit prominently 5-angled ....................................................... 4. T. chinensis 

15b. Old leaves abaxially uniformly densely hairy; branchlets hairy or not. 

20a. Branchlets hairy. 

21a. Leaf blade marginal teeth less than 1.5 mm, lateral veins 7–9 pairs; fruit 

ovoid-globose ....................................................................................................................... 4. T. chinensis 

241

242 


21b. Leaf blade marginal teeth 1.5–5 mm, lateral veins 5–7 pairs; fruit globose or obovoid. 

22a. Branchlets and leaves yellow stellate tomentose; leaf blade orbicular, marginal 

teeth with long mucros 3–5 mm ................................................................................. 3. T. henryana 

22b. Branchlets and leaves gray stellate tomentose; leaf blade ovate-orbicular, 

marginal teeth triangular ....................................................................................... 2. T. mandshurica 

20b. Branchlets glabrous. 

23a. Leaf blade marginal teeth 3–5 mm; bract stalk 0.7–2 cm ................................................... 3. T. henryana 

23b. Leaf blade marginal teeth less than 2 mm; bract stalk usually less than 5 mm. 

24a. Leaf blade broadly elliptic, abaxially tomentose; bracts sessile or subsessile. 

25a. Cymes 1–3-flowered; petals 8–9 mm ............................................................... 4. T. chinensis 

25b. Cymes 7–20-flowered; petals 6–7 mm .................................................................. 5. T. oliveri 

24b. Leaf blade ovate-oblong or obliquely ovate, abaxially puberulent or puberulent; 

bracts stalked. 

26a. Petiole glabrous, leaf margin deeply serrate, teeth extending into awns 

..................................................................................................................... 6. T. kueichouensis 

26b. Petiole stellate pubescent, leaf margin serrulate ........................................... 7. T. likiangensis 

1. Tilia endochrysea Handel-Mazzetti, Anz. Akad. Wiss. 

Wien, Math.-Naturwiss. Kl. 63: 9. 1926. 

白毛椴 bai mao duan 

Tilia begoniifolia Chun & H. D. Wong (1935), not Steven 

(1846); T. croizatii Chun & H. D. Wong; T. hypoglauca Rehder; 

T. lepidota Rehder; T. leptocarya Rehder; T. leptocarya 

var. triloba Rehder; T. scalenophylla Ling; T. vitifolia Hu & F. 

H. Chen (1951), not Host (1831). 

Trees to 20 m tall. Bark gray with longitudinal dark gray 

fissures; branchlets bright green when young, glabrous or minutely 

hairy; buds 3–3.5 mm, glabrous. Stipules oblong, 7–10 

mm; petiole 3–7 cm, nearly glabrous; leaf blade dark brown 

when dry, broadly to narrowly ovate or almost triangular, sometimes 

apically 3–5-lobed, 7–16 × 5–13 cm, abaxially gray-white 

stellate tomentose, sometimes glabrescent, adaxially glabrous, 

base obliquely cordate or truncate, margin sparsely denticulate, 

teeth 5–12 mm apart, apex acuminate or acute. Cymes 9–16 

cm, 8–32-flowered; peduncle 3.5–9 cm, nearly glabrous. Bracts 

narrowly oblong, 6–13 × 1.5–3 cm, adnate to peduncle for 1– 

1.5 cm, abaxially gray-white stellate puberulent, adaxially glabrous 

or sparsely hairy, base cuneate to cordate, apex rounded; 

stalk 1–3 cm. Pedicel 2–15 mm, stellate pilose. Sepals narrowly 

ovate, 6–8 mm, gray-brown puberulent. Petals 0.9–1.2 cm. Stamens 

ca. 40, as long as sepals; staminodes slightly shorter than 

petals. Ovary hairy; style 4–5 mm, glabrous; stigma 5-lobed. 

Fruit globose, leathery, dehiscent into 5-valves. Fl. Jul–Aug. 2n 

= 82*. 

● Species-rich mixed evergreen and deciduous woodlands, often 

on steep sides of narrow valleys and along rivers on well-drained brown 

soils; 600–1200 m. S Anhui, Fujian, Guangdong, Guangxi, Hunan, 

Jiangxi, Zhejiang. 

Pigott (Bot. Mag. 23: 56–61. 2006) gave a discussion of the 

taxonomy and ecology of Tilia endochrysea. 

2. Tilia mandshurica Ruprecht & Maximowicz, Bull. Cl. 

Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 15: 124. 1856. 

糠椴 kang duan 

Trees to 20 m tall, to 0.5(–0.7) m d.b.h. Bark dark gray, 

longitudinally fissured when old; branchlets gray-white stellate 

tomentose when young; buds tomentose. Petiole robust, 2–5 

cm, stellate tomentose, glabrescent; leaf blade ovate-orbicular, 

8–10 × 7–9 cm, abaxially densely gray stellate tomentose, adaxially 

glabrous, lateral veins 5–7 pairs, base obliquely cordate or 

truncate, margin dentate, teeth triangular, 1.5–5 mm, with or 

without awns, 4–7 mm apart, apex acute. Cymes 6–12(–20)flowered, 

6–9 cm; peduncle hairy. Bracts narrowly oblong or 

narrowly oblanceolate, 5–9 × 1–2.4 cm, adnate to peduncle for 

1/3–1/2 of length, abaxially stellate puberulent, adaxially glabrous, 

base obtuse, apex rounded; stalk 4–5 mm. Pedicel 4–6 

mm, hairy. Sepals ca. 5 mm, abaxially stellate puberulent, silky, 

adaxially villous. Petals 7–8 mm. Stamens as long as sepals; 

staminodes slightly smaller than petals. Ovary stellate tomentose; 

style 4–5 mm, glabrous. Fruit globose, ovoid or obovoid, 

weakly to prominently 5-angled, 7–9 mm, sometimes tuberculate; 

exocarp woody, indehiscent. Fr. Jul, fr. Sep. 2n = 82. 

Hebei, Heilongjiang, N Jiangsu, Jilin, Liaoning, Nei Mongol, 

Shandong [Japan, Korea, Russia (Siberia)]. 

1a. Fruit obovoid or obovoid-cylindrical, 

prominently 5-angled ......................... 2d. var. megaphylla 

1b. Fruit globose or ovoid, rarely obscurely 

angled. 

2a. Fruit prominently tuberculate ..... 2b. var. tuberculata 

2b. Fruit not tuberculate. 

3a. Fruit globose, weakly 

5-angled ........................... 2a. var. mandshurica 

3b. Fruit ovoid, rarely obscurely 

angled ........................................... 2c. var. ovalis 

2a. Tilia mandshurica var. mandshurica 

糠椴(原变种) kang duan (yuan bian zhong) 

Tilia pekingensis Ruprecht ex Maximowicz. 

Leaf blade marginal teeth with awns. Bracts 5–9 × 1–2.4 

cm. Fruit globose, weakly 5-angled, not tuberculate. 

Hebei, Heilongjiang, N Jiangsu, Jilin, Liaoning, Nei Mongol, 

Shandong [Japan, Korea, Russia (Siberia)]. 

2b. Tilia mandshurica var. tuberculata Liou & Li, Ill. Man. 

Woody Pl. N.-E. Prov. 565. 1955. 

瘤果糠椴 liu guo kang duan

Leaf blade marginal teeth with awns. Bracts 3.5–5.5 cm. 

Fruit globose, prominently tuberculate. 

● Liaoning. 

2c. Tilia mandshurica var. ovalis (Nakai) Liou & Li, Ill. Man. 

Woody Pl. N.-E. Prov. 565. 1955. 

卵果糠椴 luan guo kang duan 

Tilia ovalis Nakai, Bot. Mag. (Tokyo) 35: 15. 1921. 

Leaf blade smaller, marginal teeth without awns. Bract 5– 

9 cm. Fruit ovoid, rarely obscurely 5-angled, not tuberculate. 

Jilin [Japan]. 

2d. Tilia mandshurica var. megaphylla (Nakai) Liou & Li, Ill. 

Man. Woody Pl. N.-E. Prov. 418. 1955. 

棱果辽椴 leng guo liao duan 

Tilia megaphylla Nakai, Bot. Mag. (Tokyo) 27: 130. 1913. 

Leaf blade larger, marginal teeth with awns. Bract 5–9 cm. 

Fruit obovoid or obovoid-cylindrical, prominently 5-angled, 

densely stellate hairy, not tuberculate. 

Heilongjiang [Korea]. 

3. Tilia henryana Szyszyłowicz, Hooker’s Icon. Pl. 20: t. 1927. 

1890. 

毛糯米椴 mao nuo mi duan 

Trees to 25 m. Bark pale gray, fissured; branchlets and 

buds yellow stellate tomentose or glabrous. Petiole 3–5 cm, 

yellow tomentose; leaf blade orbicular, 6–10 × 6–10 cm, abaxially 

densely yellow stellate tomentose or hairy only in vein 

axils, adaxially glabrous, lateral veins 5–6 pairs, extending into 

3–5 mm awns, base cordate, sometimes oblique, margin serrate, 

apex broadly rounded, shortly caudate. Cymes 30–100-flowered, 

10–12 cm; peduncle stellate puberulent. Bracts narrowly 

oblanceolate, 7–10 × 1–1.3 cm, adnate to peduncle for 3–5 cm, 

abaxially yellow stellate puberulent, adaxially sometimes glabrous, 

base narrow, apex obtuse; stalk 0.7–2 cm. Pedicel 7–9 

mm, hairy. Sepals narrowly ovate, 4–5 mm, abaxially hairy. 

Petals 6–7 mm. Stamens as long as sepals; staminodes shorter 

than petals. Ovary hairy; style ca. 4 mm. Fruit obovoid, 5angled, 

7–9 mm, stellate hairy; exocarp woody, hard, indehiscent. 

Fl. Jun. 2n = 164*. 

● Anhui, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Zhejiang. 

1a. Branchlets yellow stellate tomentose; 

leaf blade abaxially densely yellow 

stellate tomentose .................................... 3a. var. henryana 

1b. Branchlets glabrous or nearly so; leaf 

blade abaxially hairy in vein axils, 

otherwise glabrous ................................ 3b. var. subglabra 

3a. Tilia henryana var. henryana 

毛糯米椴(原变种) mao nuo mi duan (yuan bian zhong) 

Branchlets yellow stellate tomentose. Leaf blade abaxially 

densely yellow stellate tomentose. Bracts adaxially yellow stellate 

puberulent. 


243 

● Anhui, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Zhejiang. 

3b. Tilia henryana var. subglabra V. Engler, Monogr. Tilia, 

125. 1909. 

糯米椴 nuo mi duan 

Branchlets and bud glabrous or nearly so. Leaf blade abaxially 

hairy in vein axils only, marginal teeth 3–5 mm. Bracts 

adaxially glabrous. 

● Anhui, Jiangsu, Jiangxi, Zhejiang. 

4. Tilia chinensis Maximowicz, Trudy Imp. S.-Peterburgsk. 

Bot. Sada 11: 83. 1890. 

华椴 hua duan 

Trees to 30 m tall. Branchlets usually brown when dry, 

glabrous, rarely sparsely pilose or densely tomentose; winter 

buds big, bud scale glabrous. Petiole 3–8 cm, usually sparsely 

pilose; leaf blade broadly elliptic or orbicular to ovate-orbicular, 

(5–)7–13 × (4.5–)6–9 cm, papery, abaxially densely stellate 

tomentose, adaxially glabrous, basal veins 5–7, lateral veins 7– 

9 pairs, base usually oblique, cordate, truncate, or rounded, 

margin serrate, apex acuminate or shortly acuminate. Cymes 1– 

3-flowered, usually not exceeding bract, shorter than leaves; 

peduncle hairy. Bracts long band-shaped, 4–9(–12) × 1–2 cm, 

adnate to peduncle for 2/5–1/2 of length, abaxially hairy, adaxially 

glabrous, sessile or with stalk to 0.5 cm. Sepals narrowly 

ovate, 6–7 mm, abaxially stellate hairy, adaxially stellate tomentose 

or glabrous at central part, margin densely hairy. Petals 

8–9 mm, glabrous. Stamens 30–45, in 5 fascicles; staminodes 5, 

smaller than petals. Ovary 5-angled, gray-yellow tomentose; 

style glabrous; stigma peltate or capitate. Fruit ellipsoid or globose, 

prominently 5-angled, 10–14 × 8–9 mm, gray-yellow 

tomentose; exocarp woody, hard, indehiscent. Fl. Jun–Jul, fr. 

Aug–Oct. 2n = 164*. 

● Forests; 1800–3100(–3900) m. Gansu, Henan, Hubei, Shaanxi, 

Sichuan, Xizang, Yunnan. 

1a. Old leaves abaxially subglabrous and hairy 

in axils of veins only ................................ 4b. var. investita 

1b. Old leaves abaxially densely hairy. 

2a. Branchlets glabrous ......................... 4a. var. chinensis 

2b. Branchlets hairy ................................. 4c. var. intonsa 

4a. Tilia chinensis var. chinensis 

华椴(原变种) hua duan (yuan bian zhong) 

Tilia baroniana Diels; T. laetevirens Rehder & E. H. Wilson; 

T. yunnanensis Hu. 

Branchlets glabrous or rarely hairy when very young. Leaf 

blade 7–13 × 6–9 cm, abaxially densely stellate tomentose 

when old, marginal awns 3–5 mm. Fruit ellipsoid or globose. 

● Forests; 1800–3100(–3900) m. Gansu, Henan, Hubei, Shaanxi, 

Sichuan, Xizang, Yunnan. 

4b. Tilia chinensis var. investita (V. Engler) Rehder, J. Arnold 

Arbor. 12: 75. 1931. 

秃华椴 tu hua duan

244 

Tilia baroniana var. investita V. Engler, Monogr. Tilia, 

132. 1909. 

Branchlets glabrous or very nearly so. Leaf blade 5–10 × 

4.5–9 cm, abaxially subglabrous and hairy in vein axils only 

when old. Fruit ellipsoid or globose. 

● Hubei, Shaanxi, Sichuan, ?Xizang, Yunnan. 

4c. Tilia chinensis var. intonsa (E. H. Wilson) Y. C. Hsu & R. 

Zhuge, J. SouthW. Forest. Coll. 11(1): 3. 1991. 

多毛椴 duo mao duan 

Tilia intonsa E. H. Wilson in Sargent, Pl. Wilson. 2: 365. 

1915; T. fulvosa Hung T. Chang. 

Branchlets densely tomentose. Leaf blade marginal teeth 

shorter than 1.5 mm. Fruit ovoid-globose. 

● Sichuan. 

5. Tilia oliveri Szyszyłowicz, Hooker’s Icon. Pl. 20: sub t. 

1927. 1890. 

鄂椴 e duan 

Trees 6–26 m tall. Bark dark gray, smooth with shallow 

furrows and rounded ridges; branchlets glabrous; winter buds 

big, glabrous or hairy. Petiole 1.5–5 cm, glabrous or rarely 

pilose; leaf blade broadly elliptic, ovate-orbicular, or triangularovate, 

6–14 × 4.5–10 cm, abaxially densely gray-white stellate 

tomentose, adaxially glabrous, lateral veins 5–6 pairs, base 

shallowly cordate to obliquely truncate, margin serrate to serrulate, 

teeth often awned, apex shortly acuminate. Cymes 7–20flowered, 

as long as or shorter than bracts. Bracts band-shaped, 

sometimes broader distally, 5–8 × 1–2.5 cm, adnate to peduncle 

for 2/5 of length, sessile. Sepals triangular-ovate, 5–6 mm, 

abaxially densely stellate puberulent, adaxially densely tomentose. 

Petals 5, 6–7 mm, glabrous. Stamens 45, in 5 fascicles, 

glabrous; staminodes smaller than petals, glabrous, stalked. 

Ovary densely stellate tomentose; style glabrous. Fruit obovoid, 

globose, or ellipsoid, 7–10 mm, gray-white tomentose and verrucose; 

exocarp woody, hard, indehiscent. Fl. Jun–Aug, fr. 

Aug–Sep. 2n = 82*. 

● Evergreen or mixed evergreen and deciduous forests; 1300– 

2300 m. Gansu, Hubei, Hunan, Shaanxi, Sichuan. 

1a. Leaf blade abaxially gray-white tomentose, 

marginal teeth often extending into distinct 

awn ............................................................... 5a. var. oliveri 

1b. Leaf blade abaxially loosely yellow-brown 

or gray stellate tomentose to pubescent, 

marginal teeth short and broad ........... 5b. var. cinerascens 

5a. Tilia oliveri var. oliveri 

鄂椴(原变种) e duan (yuan bian zhong) 

Tilia pendula V. Engler ex C. K. Schneider. 

Petiole 1.5–5 cm; leaf blade 6–10 cm, broadly elliptic, 

ovate-orbicular, or triangular-ovate, abaxially gray-white tomentose, 

adaxially glabrous, marginal teeth often prominently 

awned. 


● Evergreen or mixed evergreen and deciduous forests; 1300– 

2300 m. Gansu, Hubei, Hunan, Shaanxi, Sichuan. 

5b. Tilia oliveri var. cinerascens Rehder & E. H. Wilson in 

Sargent, Pl. Wilson. 2: 367. 1915. 

灰背椴 hui bei duan 

Tilia populifolia Hung T. Chang (1982), not Lesquereux 

[fossil]. 

Petiole 3–5 cm; leaf blade to 14 cm, abaxially loosely 

yellow-brown or gray stellate tomentose to pubescent, marginal 

teeth short and broad. 

● Forests; 1600–2300 m. Hubei. 

6. Tilia kueichouensis Hu, Acta Phytotax. Sin. 8: 198. 1963. 

黔椴 qian duan 

Tilia nanchuanensis Hung T. Chang. 

Trees to 10 m tall. Branchlets and terminal bud glabrous. 

Petiole 2–3 cm, glabrous; leaf blade obliquely ovate, 9.5–13.5 × 

5–7.5 cm, papery, abaxially sparsely stellate puberulent, adaxially 

glabrous, lateral veins 6 pairs, base cordate or truncate, 

margin deeply serrate, teeth extended into awns, apex acute. 

Bracts narrowly oblong, shorter than infructescence stalk, 6.5–8 

× 1–1.2 cm, adnate to peduncle for 1/2 of length, base narrowed, 

apex obtuse; stalk short. Flowers unknown. Infructescence 

stalk and stipe terete, slender, glabrous. Fruit globose, 

obscurely ridged, ca. 8 mm in diam., densely appressed hairy, 

verruculose, apex acute; exocarp woody, hard, indehiscent. Fr. 

Aug–Sep. 

● Chongqing, N Guizhou, ?Yunnan. 

7. Tilia likiangensis Hung T. Chang, Acta Phytotax. Sin. 20: 

171. 1982. 

丽江椴 li jiang duan 

Trees. Branchlets and terminal bud gray stellate or glabrescent. 

Petiole slender, 2–3.5 cm, stellate pubescent; leaf 

blade ovate-oblong, 9–12 × 4–6 cm, papery, abaxially gray 

appressed-pubescent, adaxially glabrous, lateral veins 6–7 pairs, 

base oblique, cordate or truncate, margin serrulate, apex acuminate. 

Cymes 6–16-flowered, 7–11 cm; peduncle glabrous. 

Bracts narrowly oblanceolate, 6–9 × 1–1.5 cm, abaxially gray 

stellate pilose, adaxially glabrous, basal 1/2 tapering, decurrent, 

apex obtuse; stalk 0.5–0.7 cm. Pedicel grooved, 7–9 mm, gray 

hairy. Sepals narrowly ovate, 4–5 mm, abaxially gray hairy. 

Petals ovate, 6–7 mm. Stamens 35, 3–4 mm; staminodes ca. 5 

mm; stalk ca. 1.5 mm. Ovary tomentose; style 3–4 mm. Fruit 

with exocarp woody, hard, indehiscent. 

● Forests; ca. 2300 m. NW Yunnan. 

8. Tilia callidonta Hung T. Chang, Acta Phytotax. Sin. 20: 


美齿椴 mei chi duan 

Trees to 10 m tall. Branchlets and terminal bud pilose. 

Petiole 3–4.5 cm, nearly glabrous; leaf blade broadly ovate, 6–9 

× 5–7 cm, abaxially yellowish green, glabrescent, hairy only in

vein axils, adaxially glabrous, lateral veins 5–6 pairs, base symmetrical 

or oblique, cordate, sometimes truncate, margin regularly 

serrulate, apex acute. Cymes 9–12-flowered, 5–8 cm; peduncle 

slender, 5–7 cm, hairy. Bracts narrowly oblong, 6–6.5 × 

1.5–1.8 cm, proximal 1.5–2.5 cm adnate to peduncle, luminous 

adaxially, stellate pilose abaxially, base cuneate, apex obtuse; 

stalk 1–1.5 cm. Flowers not seen. Fruiting pedicel 7–15 mm, 

hairy. Fruit narrowly obovoid, prominently 5-angled, 10–12 × 

ca. 5 mm, tomentose, apex pointed; exocarp woody, hard, indehiscent. 

● NW Yunnan. 

9. Tilia nobilis Rehder & E. H. Wilson in Sargent, Pl. Wilson. 

2: 363. 1915. 

大叶椴 da ye duan 

Trees 4–12 m tall. Branchlets glabrous; terminal bud 

glabrous or sparsely pilose. Petiole 4–10 cm, glabrous; leaf 

blade nearly orbicular, (5.5–)10–14(–17) × (5–)7–8(–13) cm, 

papery, abaxially glabrous or fascicled hairy in vein axils only, 

adaxially glabrous, basal veins 5, lateral veins 6–8 pairs, minor 

veinlets prominently raised abaxially, base oblique, cordate, 

truncate, or rounded, margin serrate, apex acuminate or shortly 

caudate. Cymes 3–16-flowered, usually longer than or equaling 

bracts; peduncle adnate to bract for 2/5–1/2 length. Bracts 

broadly oblanceolate or narrowly oblong, 8–16 × 1.5–2.5 cm, 

abaxially nearly glabrous or sparsely pilose, adaxially glabrous; 

stalk to 0.5 cm. Sepals abaxially subglabrous or sparsely pilose, 

adaxially villous at base, margin densely hairy. Petals oblanceolate, 

glabrous, shortly stalked. Stamens 30–45; staminodes spatulate, 

long stalked. Ovary tomentose or nearly glabrous in bud; 

style glabrous; stigma prominently swollen, 2-lobed. Fruit 

nearly globose, sometimes prominently 5-angled, ca. 10 mm in 

diam., yellowish tomentose; exocarp woody, hard, indehiscent. 

Fl. Jun–Jul, fr. Aug–Oct. 2n = ca. 328*. 

● Forests; 1800–2500 m. ?Henan, Sichuan, Yunnan. 

10. Tilia tuan Szyszyłowicz, Hooker’s Icon. Pl. 20: t. 1926. 

1890. 

椴树 duan shu 

Trees 10–20 m tall. Bark gray, longitudinally exfoliate; 

branchlets glabrous or tomentose; terminal bud glabrous or 

hairy. Petiole 1–6 cm, glabrous or tomentose; leaf blade narrowly 

ovate or ovate-oblong to ovate-orbicular, 6.5–17 × 3.5– 

11 cm, abaxially glabrous to densely shortly gray-white or 

brown tomentose, adaxially glabrous, lateral veins 3–11 pairs, 

base oblique, rounded, truncate, or cordate, margin entire or 

with a few minute teeth near apex or prominently dentate, apex 

acuminate or acute. Cymes 3–22-flowered, 5–14 cm, glabrous 

or hairy. Bracts band-shaped to oblanceolate, 6–16 × 1–3 cm, 

adnate to peduncle for ca. 1/2 of length, stellate pilose or tomentose 

to glabrous, base cuneate to rounded, apex obtuse to 

acuminate, sessile or stalk 0.5–0.8 cm. Pedicel 4–9 mm, glabrous 

or tomentose. Sepals ovate to ovate-lanceolate, 4–6 mm, 

abaxially puberulent, adaxially villous at base, margin densely 

hairy. Petals 6–8 mm, glabrous, shortly clawed. Stamens 35– 

50, in 5 fascicles, glabrous; staminodes 5, oblanceolate, prominently 

keeled. Ovary ovoid, densely gray-white stellate tomen- 


245 

tose; style 3–4 mm, glabrous. Fruit globose or obovoid-globose, 

not ridged, 7–11 × 7–9 mm, brown or gray hairy, verrucose; 

exocarp woody, hard, indehiscent. Fl. Jun–Jul, fr. Jul–Nov. n = 

82*. 

● Forests; 1200–2400 m. Guangxi, Guizhou, Hubei, Hunan, 

Jiangsu, Jiangxi, Sichuan, Yunnan, Zhejiang. 

Tilia tuan has long been recognized as the most variable Tilia 

within China (e.g., by Rehder and Wilson in Sargent, Pl. Wilson. 2: 368. 

1915). Pigott (Edinburgh J. Bot. 59: 239–246. 2002) indicated that he 

believes that a number of the more local species in China should be 

included within T. tuan, a view that has been at least partially followed 

in this account. 

1a. Leaf blade entire or with a few teeth near 

apex ............................................................... 10a. var. tuan 

1b. Leaf blade prominently dentate. 

2a. Cymes 16–22-flowered; bracts 

8–12 cm, usually sessile ............... 10b. var. chinensis 

2b. Cymes 3–6-flowered; bracts 

12–16 cm, stalk 5–8 mm .............. 10c. var. chenmoui 

10a. Tilia tuan var. tuan 

椴树(原变种) duan shu (yuan bian zhong) 

Tilia angustibracteata Hung T. Chang; T. gracilis Hung T. 

Chang; T. hupehensis W. C. Cheng ex Hung T. Chang; T. integerrima 

Hung T. Chang; T. mesembrinos Merrill; T. mofungensis 

Chun & H. D. Wong; T. oblongifolia Rehder; T. obscura 

Handel-Mazzetti; T. omeiensis Fang; T. tristis Chun ex Hung T. 

Chang; T. tuan var. cavaleriei Engler & H. Léveillé; T. tuan f. 

divaricata V. Engler; T. tuan var. pruinosa V. Engler. 

Branchlets glabrous or tomentose. Leaf blade narrowly 

ovate or ovate-oblong to ovate-orbicular, 6.5–17 × 3.5–11 cm, 

abaxially glabrous to densely shortly gray-white or brown 

tomentose, adaxially glabrous, lateral veins 3–11 pairs, base 

oblique, rounded, truncate, or cordate, margin entire or with a 

few minute teeth near apex. Cymes 3–22-flowered. Bracts 6–16 

× 1–3 cm, sessile or stalk 0.5–0.8 cm. 

● Forests; 1200–2000 m. Guangxi, Guizhou, Hubei, Hunan, 

Jiangsu, Jiangxi, Sichuan, Yunnan, Zhejiang. 

10b. Tilia tuan var. chinensis (Szyszyłowicz) Rehder & E. H. 

Wilson in Sargent, Pl. Wilson. 2: 369. 1915. 

毛芽椴 mao ya duan 

Tilia miqueliana Maximowicz var. chinensis Szyszyłowicz, 

Hooker’s Icon. Pl. 20: sub t. 1927. 1890; T. oblongifolia 

var. sangzhiensis B. R. Liao & W. X. Wang. 

Branchlets and terminal buds tomentose. Leaf blade 

broadly ovate, 10–12 × 7–10 cm, abaxially gray stellate tomentose, 

base usually oblique, margin prominently dentate. Cymes 

16–22-flowered. Bracts 8–12 cm, sessile. 

● Guizhou, Hubei, Hunan, Jiangsu, Sichuan, Zhejiang. 

10c. Tilia tuan var. chenmoui (W. C. Cheng) Y. Tang, comb. 

et stat. nov. 

长苞椴 chang bao duan 

Basionym: Tilia chenmoui W. C. Cheng, Contr. Biol. 

Lab. Sci. Soc. China, Bot. Ser. 10: 170. 1936.

246 

Branchlets densely yellowish stellate tomentose. Leaf 

blade abaxially tomentose, margin prominently dentate. Cymes 

3–6-flowered. Bracts 12–16 cm; stalk 5–8 mm. 

● Forests on streamsides; 2100–2400 m. W Yunnan (Binchuan). 

11. Tilia membranacea Hung T. Chang, Acta Phytotax. Sin. 

20: 173. 1982. 

膜叶椴 mo ye duan 

Trees to 10 m tall. Bark smooth; branchlets glabrous. Petiole 

slender, 2–3.5 cm, glabrous; leaf blade ovate, 6–10 × 4–6 

cm, nearly membranous, both surfaces glabrous or abaxially 

hairy on veins only, lateral veins 6–8 pairs, prominent on both 

surfaces, base cordate, margin serrulate, teeth 1–1.5 mm, apex 

acute. Bracts subsessile, 6–8 × 1.5–1.8 cm, adnate to peduncle 

for 2–2.5 cm, abaxially sparsely pilose, adaxially glabrous, base 

rounded, apex obtuse. Flowers unknown. Infructescences 6–7 

cm, with 2–3 capsules, slightly shorter than bracts; peduncle 4– 

5 cm; stipe 0.8–1 cm. Fruit subglobose, not ridged, 6–7 mm, 

hairy; exocarp woody, hard, indehiscent. 

● Hunan, Jiangxi. 

12. Tilia miqueliana Maximowicz, Bull. Acad. Imp. Sci. Saint- 

Pétersbourg 26: 434. 1880. 

南京椴 nan jing duan 

Tilia franchetiana C. K. Schneider; T. kinashii H. Léveillé 

& Vaniot; T. kwangtungensis Chun & H. D. Wong; T. miqueliana 

var. longipes P. C. Chiu. 

Trees to 10(–20) m tall. Bark gray-white; branchlets yellow-brown 

tomentose; terminal bud ovoid, yellow-brown tomentose. 

Petiole terete, 3–4 cm, stellate tomentose; leaf blade 

ovate-orbicular, (5–)9–12 × (4–)7–9.5 cm, abaxially gray or 

gray-yellow stellate tomentose, adaxially glabrous, lateral veins 

6–8 pairs, base cordate, rarely oblique, margin serrate, apex 

acute. Cymes 3–12-flowered, 6–8 cm; peduncle gray tomentose. 

Bracts on 2–3 mm stalk, sometimes sessile, narrowly oblanceolate, 

8–12 × 1.5–2.5 cm, adnate to peduncle for 4–6 cm, 

both surfaces stellate puberulent, base narrowly cuneate, apex 

obtuse. Pedicel 8–12 mm. Sepals 5–6 mm, gray hairy. Petals 

slightly longer than sepals. Stamens slightly shorter than sepals; 

staminodes 5, smaller. Ovary hairy; style as long as petals. Fruit 

globose, not angled, stellate puberulent, verrucose; exocarp 

woody, hard, indehiscent. Fl. Jul. 2n = 164. 

Anhui, Guangdong, Jiangsu, Jiangxi, Zhejiang [Japan]. 

13. Tilia chingiana Hu & W. C. Cheng, Contr. Biol. Lab. Sci. 

Soc. China, Bot. Ser. 10: 79. 1935. 

短毛椴 duan mao duan 

Tilia breviradiata (Rehder) Hu & W. C. Cheng, nom. superfl.; 

T. orocryptica Croizat; T. tuan Szyszyłowicz var. breviradiata 

Rehder. 

Trees to 15 m tall. Bark gray, smooth; branchlets glabrous 

or at first minutely hairy; terminal bud sparsely puberulent. 

Petiole 2.5–4 cm, hairy, glabrescent; leaf blade broadly ovate, 

5–10 × 4–9 cm, abaxially usually glabrescent, hairy in vein 

axils, adaxially glabrous, lateral veins 6–7 pairs, base oblique, 

truncate or cordate, margin serrate, apex acuminate or acute. 


Cymes 4–10-flowered, 5–8 cm; peduncle stellate puberulent. 

Bracts on 5–8 mm stalk, narrowly oblanceolate, 7–9 cm, adnate 

to peduncle for 1/2 of length, hairy, more densely so abaxially, 

base cuneate, apex obtuse. Pedicel 7–9 mm. Sepals lanceolate, 

ca. 5 mm, abaxially stellate puberulent. Petals ca. 7 × 3 mm. 

Stamens ca. 4 mm; staminodes 5, shorter than petals. Ovary 

hairy; style glabrous. Fruit globose, stellate puberulent, verrucose; 

exocarp woody, hard, indehiscent. Fl. Jun–Jul. 2n = 82*. 

● Anhui, Jiangsu, Jiangxi, Zhejiang. 

14. Tilia taishanensis S. B. Liang, Bull. Bot. Res., Harbin 5(1): 

146. 1985. 

泰山椴 tai shan duan 

Trees. Branchlets and bud glabrous. Petiole 3–7 cm, glabrous; 

leaf blade suborbicular or broadly ovate, 5–8 × 5–7 cm, 

abaxially hairy only in axils of veins, glabrous adaxially, lateral 

veins 7–8 pairs, base shallowly cordate or obliquely truncate, 

margin serrulate, apex acute. Cymes 50–200-flowered, 8–13 

cm. Bracts sessile, narrowly oblong, 5–8 × 1–1.2 cm, glabrous, 

base rounded, apex obtuse. Sepals narrowly ovate, 4–5 mm, 

both surfaces puberulent. Petals oblong, 7–8 mm. Staminodes 

present. Ovary ovoid, densely gray tomentose. Fruit obovoid, 

prominently 5-angled, 5–8 × 3–5 mm, densely brown puberulent; 

exocarp woody, hard, indehiscent. 

● About 600 m. Shandong. 

The prominently 5-angled fruits and serrulate leaf margins of Tilia 

taishanensis suggest that this might be a form of T. chinensis with abnormally 

large inflorescences. 

15. Tilia mongolica Maximowicz, Bull. Acad. Imp. Sci. Saint- 

Pétersbourg 26: 433. 1880. 

蒙椴 meng duan 

Trees to 10 m. Bark grayish, exfoliating irregularly; 

branchlets glabrous; winter buds ovoid, glabrous. Petiole slender, 

2–3.5 cm, glabrous; leaf blade broadly ovate or orbicular, 

4–6 × 3.5–5.5 cm, abaxially hairy only in axils of veins, adaxially 

glabrous, lateral veins 4–5 pairs, base slightly cordate or 

obliquely truncate, margin coarsely serrate, apex acuminate, usually 

3-lobed. Cymes 6–12-flowered, 5–8 cm; peduncle glabrous. 

Pedicel slender, 5–8 mm. Bracts on ca. 10 mm stalk, narrowly 

oblong, 3.5–6 × 0.6–1 cm, adnate to peduncle for 1/2 of 

length, both surfaces glabrous, base obtuse, apex obtuse. Sepals 

lanceolate, 4–5 mm, abaxially subglabrous. Petals 6–7 mm. Stamens 

as long as sepals; staminodes slightly smaller. Ovary 

hairy; style glabrous. Fruit obovoid, 5-angled or obscurely 

angled, 6–8 mm; exocarp thickly leathery, fragile, hairy, indehiscent. 

Fl. Jul. 2n = 164*. 

● Hebei, Henan, W Liaoning, Nei Mongol, Shanxi. 

16. Tilia amurensis Ruprecht, Mém. Acad. Imp. Sci. Saint 

Pétersbourg, Sér. 7, 15(2) [Fl. Caucasi]: 253. 1869–1870. 

紫椴 zi duan 

Trees to 25 m tall, 1 m d.b.h. Bark dark gray, exfoliating; 

branchlets white or reddish stellate pubescent, glabrescent; 

terminal bud glabrous, bud scales 3. Petiole slender, 2–3.5 cm, 

glabrous; leaf blade broadly ovate or ovate-orbicular, 4.5–6 × 

4–5.5 cm, abaxially greenish, hairy only in vein axils, adaxially

glabrous, lateral veins 4–5 pairs, base cordate or rarely obliquely 

truncate, margin serrate, teeth ca. 1 mm, pointed, apex acute, 

acuminate, truncate or slightly emarginate or 3-lobed. Cymes 

3–20-flowered, slender, 3–5 cm, glabrous. Bracts on 1–3.5 cm 

stalk, narrowly band-shaped, 3–7 × 0.5–0.8 cm, adnate to peduncle 

for 1/3–1/2 of length, both surfaces glabrous. Pedicel 7– 

10 mm. Sepals broadly lanceolate, 5–6 mm, abaxially stellate 

puberulent. Petals 6–7 mm. Stamens 20, 5–6 mm; staminodes 

absent. Ovary canescent-tomentose; style ca. 5 mm, glabrous. 

Fruit ovoid-globose, 5-angled or obscurely angled, 5–8 mm, 

stellate tomentose; exocarp thickly leathery, fragile, indehiscent. 

Fl. Jul. 2n = 82. 

Mixed Abies and Picea forests; 1300–1400 m. Heilongjiang, Jilin, 

Liaoning [Korea, Russia]. 

Tilia amurensis is similar to T. japonica but differs in having 

smaller leaves and bracts and a shorter cyme. 

1a. Leaf base truncate or slightly cordate; 

branchlets reddish stellate pilose ............. 16c. var. taquetii 

1b. Leaf base cordate; branchlets white 

stellate pubescent. 

2a. Leaf blade 4.5–6 × 4–4.5 cm, hairy 

only in vein axils on abaxial 

surface ......................................... 16a. var. amurensis 

2b. Leaf blade 3–4.5 × 2.5–4.5 cm, 

densely hairy on abaxial surface .. 16b. var. araneosa 

16a. Tilia amurensis var. amurensis 

紫椴(原变种) zi duan (yuan bian zhong) 

Tilia amurensis var. tricuspidata Liou & Li. 

Branchlets white stellate pubescent, glabrescent. Leaf 

blade 4.5–6 × 4–4.5 cm, hairy only in vein axils on abaxial 

surface, base cordate, apex acute or acuminate, sometimes 3lobed. 

Bract stalk 1–3.5 cm. 2n = 82. 

Heilongjiang, Jilin, Liaoning [Korea, Russia]. 

Plants with 3-lobed leaf tips have been placed in var. tricuspidata. 

16b. Tilia amurensis var. araneosa C. Wang & S. D. Zhao, 

Bull. Bot. Res., Harbin 1(4): 135. 1981. 

毛紫椴 mao zi duan 

Branchlets white stellate pubescent. Petiole ca. 1 cm, hairy; 

leaf blade 3–4.5 × 2.5–4.5 cm, abaxially greenish, densely hairy, 

base cordate, apex caudate. 

● Mixed Abies and Picea forests; 1300–1400 m. Jilin. 

16c. Tilia amurensis var. taquetii (C. K. Schneider) Liou & 

Li, Woody Fl. N.E. China, 420. 1955. 

小叶紫椴 xiao ye zi duan 

Tilia taquetii C. K. Schneider, Repert. Spec. Nov. Regni 

Veg. 7: 200. 1909; T. amurensis subsp. taquetii (C. K. Schneider) 

Pigott; T. koreana Nakai. 

Branchlets and cymes reddish stellate pilose. Leaf blade 

base usually truncate or slightly cordate. Bract stalk 1–1.5 cm. 

2n = 82. 

Heilongjiang, Jilin, Liaoning [Korea, Russia]. 


247 

17. Tilia japonica (Miquel) Simonkai, Math. Term. Közlem. 

22: 326. 1888. 

华东椴 hua dong duan 

Tilia cordata Miller var. japonica Miquel, Ann. Mus. Bot. 

Lugduno-Batavi 3: 18. 1867; T. eurosinica Croizat; T. ulmifolia 

Scopoli var. japonica (Miquel) O. H. Sargent ex Mayr. 

Trees. Branchlets villous, soon glabrescent; terminal bud 

ovoid, glabrous. Petiole slender, 3–4.5 cm, glabrous; leaf blade 

brown when dry, orbicular or nearly so, 5–10 × 4–9 cm, 

leathery, abaxially hairy only in vein axils, adaxially glabrous, 

lateral veins 6–7 pairs, base cordate or rarely truncate, margin 

sharply serrulate, apex acute. Cymes 6–16-flowered, 5–7 cm; 

peduncle with proximal bract. Bracts on 1–1.5 cm stalk, narrowly 

oblanceolate or narrowly oblong, 3.5–6 × 1–1.5 cm, 

adnate to peduncle for 1/2 of length, glabrous. Pedicel 5–8 mm. 

Sepals narrowly oblong, 4–4.5 mm, sparsely stellate puberulent. 

Petals 6–7 mm. Stamens ca. 5 mm; staminodes slightly shorter 

than petals. Ovary hairy; style 3–4 mm. Fruit ovoid, not angled, 

stellate hairy; exocarp thickly leathery, fragile, indehiscent. 2n 

= 164. 

Anhui, Jiangsu, Shandong, Zhejiang [Japan]. 

18. Tilia paucicostata Maximowicz, Trudy Imp. S.-Peterburgsk. 

Bot. Sada 11: 82. 1890. 

少脉椴 shao mai duan 

Trees 10–15 m tall. Branchlets usually yellow-brown 

when dry, slender, glabrous; terminal buds minute, glabrous or 

slightly hairy at tip. Petiole slender, 2–5 cm, glabrous; leaf 

blade ovate-orbicular or ovate-triangular, (3–)6–10 × 2.5–6 cm, 

thinly leathery, both surfaces glabrous or abaxially sparsely tomentose 

in vein axils, lateral veins 5–6 pairs, basal pair usually 

1/2 or more as long as blade, veinlets prominent, base truncate 

or obliquely cordate, rarely rounded, margin serrate or sparsely 

dentate, apex caudate-acuminate or acuminate. Cymes 3–10flowered, 

usually shorter or rarely longer than bract; peduncle 

adnate with proximal 1/2 bract. Bracts on slender 0.5–2 cm 

stalk, oblanceolate, 4–6 cm, glabrous. Sepals 4–5 mm, abaxially 

glabrous, adaxially villous on proximal half. Petals ca. 4 mm, 

glabrous. Stamens 25–35, in 5 fascicles, glabrous; staminodes 

5, linear-oblanceolate, distal 1/2 slightly wider, 1/2 as long as 

petals. Ovary ovoid, densely stellate puberulent; style glabrous. 

Fruit obovoid, 6–7 mm, apex usually beaked; exocarp thickly 

leathery, fragile, indehiscent. Fl. Jun–Aug, fr. Sep–Oct. 2n = 

164*. 

● Forests; 1300–2400 m. Gansu, Hebei, Henan, Hubei, Hunan, 

Shaanxi, Sichuan, Yunnan. 

1a. Branchlets puberulent; leaf blade 

abaxially gray stellate puberulent ... 18b. var. yunnanensis 

1b. Branchlets and leaf blade abaxially 

glabrous. 

2a. Leaf blade 6–10 × 3–6 cm, 

margin serrate .......................... 18a. var. paucicostata 

2b. Leaf blade 3.5–5.5 × 2.5–4 cm, 

margin sparsely dentate ........... 18c. var. dictyoneura

248 

18a. Tilia paucicostata var. paucicostata 

少脉椴(原变种) shao mai duan (yuan bian zhong) 

Tilia paucicostata var. firma V. Engler; T. paucicostata 

var. tenuis V. Engler. 

Branchlets glabrous. Leaf blade 6–10 × 3–6 cm, abaxially 

glabrous, margin serrate. Fruit obovoid, 6–7 mm. 

● Forests; 1300–2400 m. Gansu, Henan, Hubei, Hunan, Shaanxi, 

Sichuan, Yunnan. 

18b. Tilia paucicostata var. yunnanensis Diels, Notes Roy. 

Bot. Gard. Edinburgh 5: 285. 1912. 

毛少脉椴 mao shao mai duan 

Tilia paucicostata var. ningshanensis P. H. Yang. 

Branchlets and terminal buds puberulent. Leaf blade abaxially 

gray stellate puberulent, adaxially glabrous. Fruit unknown. 

● Forests; 2000–2400 m. Gansu, Sichuan, Yunnan. 

18c. Tilia paucicostata var. dictyoneura (V. Engler ex C. K. 

Schneider) Hung T. Chang & E. W. Miau, Fl. Reipubl. Popularis 

Sin. 49(1): 72. 1989. 

红皮椴 hong pi duan 

Tilia dictyoneura V. Engler ex C. K. Schneider, Ill. 

Handb. Laubholzk. 2: 369. 1909. 

Branchlets glabrous. Leaf blade triangular-ovate, 3.5–5.5 


× 2.5–4 cm, abaxially glabrous, margin sparsely denticulate. 

Bracts with stalk. Fruit 5–6 mm, without ridges. 

● Forests. Gansu, Hebei, Henan, Hunan, Shaanxi. 

19. Tilia jiaodongensis S. B. Liang, Bull. Bot. Res., Harbin 

5(1): 145. 1985. 

胶东椴 jiao dong duan 

Trees. Branchlets and buds glabrous. Petiole 3–5 cm, glabrous; 

leaf blade ovate-orbicular, 5–8 × 5–7 cm, abaxially hairy 

only in axils of veins, adaxially glabrous, lateral veins 6–7 

pairs, base cordate or shallowly cordate, margin serrulate, teeth 

2–3 mm, apex acute. Cymes (20–)40–100-flowered, 6–13 cm, 

glabrous. Bracts on 1–2 cm stalk, oblanceolate, 5–9 × 0.8–1.5 

cm, glabrous, base obliquely cuneate, apex obtuse; bracteoles 

3–4-verticillate at bases of pedicels, present at each branching 

of inflorescence, ovate-lanceolate or elliptic-lanceolate, 0.3–3 × 

0.2–1 cm, densely brown stellate tomentose. Pedicel 3–7 mm. 

Sepals narrowly ovate, 4–5 mm, abaxially stellate hairy, adaxially 

sparsely stellate at apex and densely villous at base. Petals 

obovate, 4–5 mm, glabrous. Staminodes shorter than petals. 

Ovary ovoid, prominently 5-angled; style short. Fruit subglobose, 

minutely 5-angled, ca. 5 mm in diam., thickly leathery, 

fragile, densely brown puberulent, indehiscent. 

● Sunny slopes; ca. 600 m. Shandong. 

The very large bracteolate inflorescence of Tilia jiaodongensis is 

unusual and is probably aberrant. The species was not accepted by 

Pigott. It was originally compared to T. mongolica from which it was 

distinguished by the acute leaf apex and glabrous ovary. The nature of 

the ovary requires clarification as the fruit is described as hairy. 

2. CRAIGIA W. W. Smith & W. E. Evans, Trans. & Proc. Bot. Soc. Edinburgh 28: 69. 1921. 

滇桐属 dian tong shu 

Trees deciduous; buds with scales. Leaves long petiolate; leaf blade elliptic or oblong, leathery or papery, basal veins 3, margin 

denticulate. Inflorescences axillary, cymose. Flowers bisexual. Pedicel articulate. Sepals 5, fleshy, valvate. Petals absent. Androgynophore 

absent. Stamens and staminodes many, in 2 or 3 series, outer series of 10 staminodes, in pairs, inner stamens 20, in 5 

fascicles; filaments slightly connate; anthers 2-loculed. Ovary superior, 5-loculed, sessile; ovules 6 per locule; styles 5. Fruit a 5winged 

capsule, ellipsoid, loculicidally dehiscent, wings veined, membranous. Seeds 1, 2, or 4 per locule, oblong. 

Two species: China and adjacent Vietnam; two species (one endemic) in China. 

The 5-winged capsules of Craigia superficially resemble the fruits of Burretiodendron, which has led to confusion. Craigia can be distinguished 

from Burretiodendron by its petaloid staminodes and the lack of petals. Both species of Craigia are included in the IUCN Red List; C. kwangsiensis is 

possibly extinct. 

1a. Leaf blade elliptic, base rounded; branchlets and leaves glabrous abaxially .......................................................... 1. C. yunnanensis 

1b. Leaf blade oblong, base cuneate; branchlets and leaves hairy abaxially .............................................................. 2. C. kwangsiensis 

1. Craigia yunnanensis W. W. Smith & W. E. Evans, Trans. & 

Proc. Bot. Soc. Edinburgh 28: 69. 1921. 

滇桐 dian tong 

Burretiodendron combretoides Chun & F. C. How; B. 

yunnanense Kostermans. 

Trees 6–20 m tall. Branchlets glabrous; buds gray-white 

hairy. Petiole 1.5–5 cm; leaf blade dark greenish, elliptic, 10–20 

× 5–11 cm, abaxially glabrous, 2 lateral veins 8–10 mm from 

margin, ascending less than 1/2 length of blade, lateral veins 5– 

7 pairs, base rounded, margin denticulate, apex acute. Cymes 

2–5-flowered, ca. 3 cm. Sepals purplish pink, oblong, ca. 1 cm, 

hairy abaxially. Stamens of inner series shorter than sepals. 

Ovary glabrous. Capsule ca. 3.5 × 2.5–3 cm, wings 5-angled. 

Seeds ca. 1 cm. 

Forests, open forests; 500–1600 m. W Guangxi (Jingxi, Napo), S 

Guizhou (Dushan), SE Xizang (Mêdog), SE and W Yunnan (Malipo, 

Ruili, Xichou) [N Vietnam]. 

Craigia yunnanensis is listed as “Endangered” in the IUCN Red 

List. The synonym Burretiodendron yunnanense is an independent 

name based on a different type.

2. Craigia kwangsiensis H. H. Hsue, Acta Phytotax. Sin. 13(1): 

107. 1975. 

桂滇桐 gui dian tong 

Branchlets, buds, stalk, and stipe stellate pubescent. Petiole 

1.8–2.5 cm; leaf blade oblong, 7–9 × 2.5–4 cm, dark greenish 

when dry, abaxially yellow-brown pubescent, 2 lateral veins 

close to margin, ascending to 1/3 length of blade, lateral veins 

5–6 pairs, veinlets prominent, base cuneate, margin serrate, 


3. CORCHORUS Linnaeus, Sp. Pl. 1: 529. 1753. 

黄麻属 huang ma shu 

249 

apex acuminate. Infructescence cymelike; peduncle 1–1.5 cm. 

Stipe 1–1.2 cm. Capsule 2.5–3 × 2–2.4 cm, wings transversely 

veined. Seeds 4 per locule, ca. 8 mm. Fr. Nov. 

● Evergreen broad-leaved forests on limestone; ca. 1400 m. NW 

Guangxi (Tianlin). 

Craigia kwangsiensis is known only from the type. It has not been 

relocated and it is possibly extinct due to deforestation. It is listed as 

“Critically endangered” in the IUCN Red List. 

Herbs or subshrubs. Leaves petiolate; stipules filiform; leaf blade papery, basal veins usually 3, usually with linear appendages 

at or near base, margin serrate, serrulate, or crenate. Flowers solitary or several arranged in cymes, axillary or extra-axillary, bisexual, 

yellow. Sepals 4 or 5. Petals 4 or 5; glands absent. Stamens 15 to many, on androgynophore, free; staminodes absent. Ovary 2–5loculed; 

ovules many per locule; style short; stigma peltate or disk-shaped, entire or lobed. Fruit a capsule, cylindrical or globose, 

sometimes angled, 2–5-valved, loculicidally dehiscent. Seeds many. 

Between 40 and 100 species: primarily in tropical regions; four species in China. 

Corchorus oenotheroides H. Léveillé is the basionym of Indigofera oenotheroides (H. Léveillé) Lauener in the Fabaceae (see Fl. China 10). 

Corchorus polygonatum H. Léveillé is a synonym of Tricyrtis pilosa Wallich in the Liliaceae (see Fl. China 24: 152. 2000). 

Several species have been grown as a fiber crop. Corchorus capsularis and C. olitorius are the source of the important fiber jute. 

1a. Capsule globose, apex truncate or slightly emarginate; leaf blade glabrous .............................................................. 1. C. capsularis 

1b. Capsule cylindrical, apex angled or beaked; leaf blade hairy at least abaxially on veins. 

2a. Capsule apex 3–5-angled; leaf blade abaxially sparsely pilose, glabrescent ......................................................... 2. C. aestuans 

2b. Capsule apex beaked; leaf blade abaxially sparsely hairy only on veins. 

3a. Ovary 5-loculed; leaf blade margin serrulate, petiole 0.8–3.5 cm ................................................................... 3. C. olitorius 

3b. Ovary 3-loculed; leaf blade margin crenate, petiole 0.5–1 cm ................................................................... 4. C. trilocularis 

1. Corchorus capsularis Linnaeus, Sp. Pl. 1: 529. 1753. 

黄麻 huang ma 

Herbs woody, erect, 1–2 m tall. Petiole ca. 2 cm, puberulent; 

leaf blade ovate-lanceolate or narrowly lanceolate, 5–12 × 

2–5 cm, glabrous, basal veins ascending to mid leaf, lateral 

veins 8–10 pairs, base rounded, margin coarsely serrate, apex 

acuminate. Flowers solitary or several arranged in cymes, axillary; 

peduncle and pedicel short. Sepals 4 or 5, 3–4 mm. Petals 

obovate, ± as long as sepals, glabrous. Stamens 18–22; androgynophore 

short, glabrous. Ovary 5-loculed, glabrous; stigma 

lobed. Capsule globose, obtusely angled, 5-valved, ca. 1 cm in 

diam., verrucose, apex truncate or slightly emarginate. Fl. summer, 

fr. late autumn. 

Widely cultivated. Anhui, Fujian, Guangdong, Guangxi, Guizhou, 

Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Taiwan, 

Yunnan, Zhejiang [Bangladesh, India, Indonesia, Japan, Malaysia, 

Myanmar, Pakistan, Philippines, Sri Lanka]. 

2. Corchorus aestuans Linnaeus, Syst. Nat., ed. 10, 2: 1079. 

1759. 

甜麻 tian ma 

Herbs annual, to 1 m tall. Stem red-brown, slightly yellowish 

puberulent; branches slender. Petiole 0.9–1.6 cm, with 

yellowish, long, rigid hairs; leaf blade ovate or broadly ovate, 

4.5–6.5 × 3–4 cm, sparsely pilose on both surfaces, glabrescent, 

basal veins 5–7, lateral veins 3–6 pairs, base rounded, margin 

serrate, basal pair of teeth usually elongating into filiform or 

caudate appendages, apex shortly acuminate or acute. Flowers 

solitary or several together in cymes, axillary or leaf-opposed; 

peduncle and pedicel very short or absent. Sepals 5, purple-red 

abaxially, narrowly oblong, ca. 5 mm, semi-foveolately boatshaped 

distally, apex awned. Petals 5, yellow, nearly as long as 

sepals, obovate. Stamens many, yellow, ca. 3 mm. Ovary 3–5loculed, 

long cylindrical, puberulent; style rodlike; stigma beaklike, 

5-toothed. Capsule cylindrical, angled, 3–5-valved, 1.2– 

2.5 × ca. 0.5 cm, apex 3–5-angled, awns bifurcate. Seeds separated 

by transverse septum. Fl. summer–autumn. 

Widely cultivated. Anhui, Fujian, Guangdong, Guangxi, Guizhou, 

Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang 

[Bangladesh, Bhutan, India, Indonesia, Malaysia, Myanmar, Nepal, 

Pakistan, Philippines, Sri Lanka, Thailand, Vietnam; tropical Africa, 

Australia, Central America, West Indies]. 

1a. Ovary puberulent, stigma 5-toothed; 

capsule 3–4-loculed ................................. 2a. var. aestuans 

1b. Ovary woolly, stigma entire; capsule 

usually 5-loculed ................................. 2b. var. brevicaulis 

2a. Corchorus aestuans var. aestuans 

甜麻(原变种) tian ma (yuan bian zhong) 

Stems usually erect, internodes as long as or longer than 

leaf blade. Stamens many. Ovary puberulent; stigma 5-toothed. 

Capsule 3–4-loculed, 6–8-angled, ca. 2.5 cm. Fl. summer– 

autumn.

250 

Widely cultivated. Anhui, Guangdong, Guangxi, Hainan, Hubei, 

Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Bangladesh, 

Bhutan, India, Laos, Malaysia, Myanmar, Nepal, Pakistan, Sri 

Lanka, Vietnam; tropical Africa, Australia, Central America, West 

Indies]. 

2b. Corchorus aestuans var. brevicaulis (Hosokawa) T. S. Liu 

& H. C. Lo, Fl. Taiwan 3: 695. 1977. 

短茎甜麻 duan jing tian ma 

Corchorus brevicaulis Hosokawa, Trans. Nat. Hist. Soc. 

Taiwan 22: 226. 1932. 

Stems almost procumbent, internode shorter than leaf 

blade. Leaf blade membranous, basal appendages subulate filiform. 

Stamens 16 to many. Ovary woolly; stigma entire. Capsule 

usually 5-loculed, 10-angled, ca. 1.2 cm (excluding awns). 

Fl. Aug. 

● Taiwan. 

3. Corchorus olitorius Linnaeus, Sp. Pl. 1: 529. 1753. 

长蒴黄麻 chang shuo huang ma 

Herbs woody, 1–3 m tall. Stipule ovate-lanceolate, ca. 1 

cm; petiole 0.8–3.5 cm, puberulent; leaf blade oblong-lanceolate, 

7–10 × 2–4.5 cm, glabrous, basal veins 5, lateral veins 7– 

10 pairs, base rounded, margin serrulate, apex acuminate. Flowers 

solitary or 1–3 arranged in cymes, axillary, opposite to 

leaves; peduncle and pedicel short. Sepals oblong, hairy at base, 

Columbia Persoon. 


4. COLONA Cavanilles, Icon. 4: 47. 1798. 

一担柴属 yi dan chai shu 

apex long awned. Petals oblong, as long as or slightly shorter 

than sepals, stalked at base; stalk ciliate. Stamens on very short 

androgynophore, glabrous. Ovary 5-loculed, hairy; style glabrous; 

stigma disk-shaped, lobed. Capsule cylindrical, slightly 

curved, 10-angled, robust, 5–6-valved, 3–8 cm, apex beaked, 

with single awn. Seeds obconic, slightly angled, separated by 

septum. Fl. summer–autumn. 

Cultivated, weed of disturbed areas. Anhui, Fujian, Guangdong, 

Guangxi, Hainan, Hunan, Jiangxi, Yunnan [widely distributed throughout 

tropics]. 

4. Corchorus trilocularis Linnaeus, Syst. Nat., ed. 12, 2: 369; 

Mant. Pl. 1: 77. 1767. 

三室黄麻 san shi huang ma 

Herbs annual or perennial, many branched, pilose. Stipule 

subulate; petiole 0.5–1 cm, pilose; leaf blade elliptic or ovateoblong, 

1.5–3 × 1–2 cm, hairy only on veins abaxially, glabrous 

adaxially, lateral basal veins 1/3 as long as leaf blade, base 

broadly rounded, margin crenate, apex obtuse. Cymes opposite 

to leaves, 1–3-flowered; peduncle very short. Sepals ca. 5 mm. 

Petals nearly spatulate, 5–6 mm. Stamens 15–20; filaments 5–6 

mm. Ovary 3-loculed. Capsule 1–3-fascicled, long cylindrical, 

5–7 × ca. 0.3 cm, apex beaked. Seeds separated by septum. Fl. 

Aug, fr. Sep. 

Moist shady places, weed of cultivated fields. NW Yunnan [Afghanistan, 

Bhutan, India, Indonesia (Java), Pakistan, Sri Lanka; tropical 

and N Africa, S Asia, Australia]. 

Trees or shrubs. Leaves long petiolate; leaf blade ovate, usually hairy abaxially, basal veins 5–7, margin entire or serrulate, apex 

acute, sometimes 3–5-lobed. Inflorescences usually terminal, cymose paniculate. Bracts small; bracteoles present. Flowers bisexual. 

Sepals 5, free. Petals 5, glandular at base. Stamens many, free or in 5 bundles, borne on short androgynophore. Ovary 3–5-loculed; 

ovules 2–4 per locule; stigma thin, lobed or entire. Fruit a capsule, subglobose, 3–5-winged, septicidally dehiscent. 

About 20 species: tropical Asia; two species in China. 

1a. Leaf blade broadly obovate-orbicular or nearly orbicular, 11–20 cm wide, abaxially and adaxially roughly hairy; 

petiole 1.5–5.5 cm ....................................................................................................................................................... 1. C. floribunda 

1b. Leaf blade oblong, 4–7 cm wide, abaxially stellate tomentose, adaxially glabrous; petiole 0.5–1 cm .......................... 2. C. thorelii 

1. Colona floribunda (Wallich ex Kurz) Craib, Bull. Misc. 

Inform. Kew 1925: 21. 1925. 

一担柴 yi dan chai 

Columbia floribunda Wallich ex Kurz, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 42: 63. 1873. 

Trees 8–10 m tall. Branchlets gray-brown stellate puberulent. 

Petiole 1.5–5.5 cm; leaf blade broadly obovate-orbicular or 

nearly orbicular, 14–21 × 11–20 cm, with rough, gray-brown 

stellate hairs on both surfaces, more dense abaxially, basal 

veins 5–7, veinlets prominently raised abaxially, base slightly 

cordate, margin serrulate, apex acute or acuminate, sometimes 

3–5-lobed. Inflorescences terminal, to 27 cm. Flowers ca. 8 mm 

in diam. Sepals lanceolate, ca. 4 mm, stellate tomentose abax- 

ially. Petals yellow, spatulate, nearly as long as sepals. Stamens 

ca. 40, nearly as long as petals, in 5 bundles. Ovary stellate tomentose, 

3–5-loculed; stigma hairy. Capsule 1–1.5 cm in diam., 

stellate hairy, 3–5-winged, wings ca. 5 mm. Fl. Jun, fr. Nov. 

Secondary montane forests; 300–2000 m. S Yunnan [India, Laos, 

Myanmar, Thailand, Vietnam]. 

The basionym is often given as “Grewia floribunda Wallich ex 

Voigt” (Hort. Suburb. Calcutt. 128. 1845) but that name, as a nomen 

nudum, was not validly published. 

2. Colona thorelii (Gagnepain) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 

9: 808. 1926. 

狭叶一担柴 xia ye yi dan chai

Columbia thorelii Gagnepain, Notul. Syst. (Paris) 1: 132. 

1910; Colona sinica Hu. 

Trees 5–15 m tall, many branched. Branchlets gray-brown 

stellate tomentose. Stipule hastate; petiole 0.5–1 cm; leaf blade 

oblong, 8–15 × 4–7 cm, thinly leathery, densely gray stellate 

tomentose abaxially, adaxially glabrous, smooth or slightly 

rough, basal veins 5, lateral veins 2–3 pairs, base rounded, margin 

entire or slightly serrulate near apex, apex acute or acuminate. 

Inflorescences terminal or axillary at twig tips, 4–9 cm. 


5. MICROCOS Linnaeus, Sp. Pl. 1: 514. 1753. 

破布叶属 po bu ye shu 

251 

Bracts triangular, ovate, or lanceolate, 5–6 mm; bracteoles 

obovate. Pedicel 3–5 mm. Sepals narrowly lanceolate, 6–7 mm, 

abaxially gray hairy, adaxially glabrous. Petals 5–6 mm. Stamens 

as long as petals. Ovary 3-loculed, hairy; style ca. 5 mm; 

stigma 2-lobed. Capsule 2–2.5 cm in diam., gray tomentose, 3winged, 

wing ca. 10 mm wide. Fl. Sep–Oct, fr. Nov–Dec. 

Mountains; 200–800 m. S and SE Yunnan [Laos, Malaysia, Myanmar, 

Thailand]. 

Shrubs or small trees. Leaves alternate, shortly petiolate; leaf blade ovate, oblong, or lanceolate, leathery, basal veins 3, margin 

entire or lobed in distal 1/2. Inflorescences terminal or axillary, cymose paniculate. Flowers bisexual. Sepals 5, free. Petals 5 or rarely 

absent, glandular at base adaxially. Stamens many, free, borne distally on androgynophore. Ovary superior, usually 3-loculed; ovules 

4–7 per locule; style simple; stigma subulate, usually not lobed. Fruit a drupe, globose, obovoid, or pyriform, not furrowed, without 

drupelets. 

About 60 species: primarily in Africa, Asia; three species in China. 

1a. Mature leaf blade tomentose abaxially, apex acute ............................................................................................... 3. M. stauntoniana 

1b. Mature leaf blade glabrescent or nearly so abaxially, apex acuminate. 

2a. Ovary glabrous; leaf blade ovate-oblong; sepals 5–7 mm .................................................................................. 1. M. paniculata 

2b. Ovary densely villous; leaf blade oblong or lanceolate; sepals 8–10 mm ............................................................... 2. M. chungii 

1. Microcos paniculata Linnaeus, Sp. Pl. 1: 514. 1753. 

破布叶 po bu ye 

Fallopia nervosa Loureiro; Grewia microcos Linnaeus, 

nom. illeg. superfl.; G. nervosa (Loureiro) Panigrahi; Microcos 

nervosa (Loureiro) S. Y. Hu. 

Shrubs or small trees 3–12 m tall. Bark rough; branchlets 

hairy. Stipule filiform, lanceolate, 5–7 mm; petiole 1–1.5 cm, 

hairy; leaf blade ovate or oblong, 8–18 × 4–8 cm, thinly leathery, 

very sparsely stellate at first and glabrescent both abaxially 

and adaxially, basal veins 3, laterals more than 1/2 as long as 

leaf blade, base rounded, margin finely crenate, apex acuminate. 

Panicles terminal, 4–10 cm, stellate. Bracts lanceolate. 

Pedicel short. Sepals oblong, 5–7 mm, hairy abaxially. Petals 

oblong, 3–4 mm, hairy in proximal 1/2; glands ca. 2 mm. Stamens 

shorter than sepals. Ovary globose, glabrous. Drupe 

nearly globose or obovoid, ca. 1 cm; stipe short. Fl. Jun–Jul. 

Guangdong, Guangxi, Hainan, Yunnan [Cambodia, India, Indonesia, 

Laos, Malaysia, Myanmar, Sri Lanka, Thailand, Vietnam]. 

2. Microcos chungii (Merrill) Chun, Sunyatsenia 4: 196. 1940. 

海南破布叶 hai nan po bu ye 

Grewia chungii Merrill, Philipp. J. Sci. 23: 252. 1923. 

Trees 5–15 m tall. Branchlets yellow-brown pubescent. 

Petiole 1–1.5 cm, stellate; leaf blade oblong or lanceolate, 11– 

20 × 3.5–6 cm, nearly leathery, very sparsely stellate and gla- 

brescent abaxially, glabrous adaxially, base rounded obtuse, 

margin entire or sparsely serrate distally, apex long acuminate. 

Panicles terminal or axillary. Bracts yellow-brown puberulent. 

Sepals 5, narrowly lanceolate, 8–10 mm, stellate puberulent on 

both surfaces, more densely abaxially. Petals yellow, narrowly 

oblong, 3–4 mm, sparsely pubescent abaxially, with hairy 

glands at base adaxially; glands ca. 1/3 as long as petals. Ovary 

broadly ovoid, densely villous. Drupe pyriform, 12–22 × 9–12 

mm, densely gray-yellow stellate pubescent; stipe robust, hairy. 

Fl. summer–autumn, fr. winter. 

Montane forests. Hainan, S Yunnan [Vietnam]. 

3. Microcos stauntoniana G. Don, Gen. Hist. 1: 551. 1831. 

毛破布叶 mao po bu ye 

6. GREWIA Linnaeus, Sp. Pl. 2: 964. 1753. 

扁担杆属 bian dan gan shu 

Trees 2–5 m tall. Branchlets stellate tomentose. Petiole ca. 

1 cm, tomentose; leaf blade ovate or long ovate, 9–16 × 4.5–9 

cm, densely brown tomentose abaxially, glabrous adaxially, lateral 

pair of basal veins ascending to mid leaf blade, base 

rounded or slightly cordate, apex acute. Panicles 4–7 cm. Pedicel 

2–4 mm. Sepals oblong, 5–6 mm, tomentose abaxially. Petals 

ca. 1/2 as long as sepals; glands 1–1.5 mm. Stamens as long 

as sepals. Ovary glabrous or hairy at base; style slightly longer 

than stamens, glabrous. Drupe obovoid. Fl. summer–autumn. 

Hainan [Cambodia, Indonesia, Laos, Malaysia, Myanmar, Thailand, 

Vietnam]. 

Trees or shrubs. Branchlets usually stellate. Leaves alternate; stipule minute, caducous; petiole short; leaf blade basal veins 3–5, 

margin serrate or rarely lobed. Flowers bisexual, polygamous, or unisexual (plants monoecious), usually arranged in 3- to many-

252 


flowered cymes, solitary or fascicled, axillary or opposite leaves, rarely terminal. Peduncle and pedicel usually hairy. Bracts caducous. 

Sepals 5, yellow, white, or lilac, rarely brown, free, valvate, abaxially hairy, adaxially usually glabrous, rarely hairy. Petals 5, 

mostly yellow or white in Flora area, shorter than sepals; glands usually scalelike, borne on base of petals, margin usually villous. 

Androgynophore short, usually glabrous. Stamens many, free, irregular in length; anthers globose, dorsifixed, dehiscence longitudinal. 

Ovary 2–4-loculed; ovules 2–8 per locule; style simple; stigma swollen, peltate, entire or 2–4-lobed. Drupe usually with 2 or 4 

drupelets. Endosperm copious; cotyledons flat. 

About 90 species: tropical regions of Old World; 27 species (13 endemic) in China. 

Grewia nitida Jussieu (Ann. Mus. Natl. Hist. Nat. 4: 90. 1804) could not be treated here because no material was seen by the authors. 

Grewia salviifolia Linnaeus f. is the basionym of Alangium salviifolium (Linnaeus f.) Wangerin in the Alangiaceae (see Fl. China 13: 307. 2007). 

1a. Ovary and drupe globose, not lobed; drupelets 1 or 2 per fruit. 

2a. Climbing lianas; leaf blade oblong or lanceolate, base cordate, abaxially subglabrous ........................... 5. G. kwangtungensis 

2b. Erect shrubs or trees; leaf blade broadly ovate-orbicular to ovate-oblong, base oblique or symmetrically 

rounded, abaxially tomentose, rarely glabrous. 

3a. Leaf blade base oblique. 

4a. Leaf blade abaxially densely softly stellate tomentose; cymes in clusters of 1–3 ................................. 1. G. eriocarpa 

4b. Leaf blade abaxially sparsely pilose or subglabrous; cymes in clusters of 2–6 ........................................ 4. G. tiliifolia 

3b. Leaf blade base symmetrical, rounded. 

5a. Petiole 5–10 mm, leaf blade broadly ovate, 4.5–6 cm wide; drupe 6–8 mm in diam. .......................... 2. G. celtidifolia 

5b. Petiole 3–4 mm, leaf blade nearly orbicular, 7.5–10 cm wide; drupe 4–5 mm in diam. ................. 3. G. yinkiangensis 

1b. Ovary and drupe obviously 2–4-lobed, lobes ± globose, each with 1 drupelet. 

6a. Leaf blade ovate, rhomboid, suborbicular, or obovate-elliptic, base oblique or symmetrical, lateral basal 

veins ascending beyond middle of blade. 

7a. Leaf blade abaxially glabrescent or sparsely pilose, never tomentose. 

8a. Leaf blade margin serrulate, leaf blade elliptic to obovate-elliptic or suborbicular ..................................... 7. G. biloba 

8b. Leaf blade margin serrate or irregularly serrate, leaf blade rhomboid, ovate, or elliptic. 

9a. Leaf blade 2–4.5 cm wide, rhomboid or ovate; branchlets rust-brown stellate tomentose .......... 6. G. rhombifolia 

9b. Leaf blade 1–2 cm wide, ovate or elliptic; branchlets sparsely stellate hairy ............................... 8. G. piscatorum 

7b. Leaf blade abaxially tomentose, coarsely hairy or pilose. 

10a. Leaf blade adaxially glabrous or softly hairy. 

11a. Leaf blade elliptic or ovate-elliptic, margin crenate, branchlets softly yellow tomentose .... 9. G. angustisepala 

11b. Leaf blade oblong-lanceolate, margin denticulate; branchlets glabrescent .............................. 27. G. multiflora 

10b. Leaf blade adaxially coarsely hairy. 

12a. Leaf blade longer than 10 cm, base regular. 

13a. Erect shrubs or small trees; leaf blade 21–28 cm ............................................................ 10. G. permagna 

13b. Climbing shrubs; leaf blade 8–13 cm ............................................................................. 11. G. acuminata 

12b. Leaf blade shorter than 10 cm, base regular or oblique. 

14a. Leaf blade ovate-orbicular or elliptic, 2–3 cm wide ................................................................. 7. G. biloba 

14b. Leaf orbicular or broadly ovate-orbicular, 4–9 cm wide. 

15a. Cymes many per axil; peduncle 3–6 mm; stigma 2-lobed ................................... 12. G. abutilifolia 

15b. Cymes 1 or 2 per axil; peduncle ca. 10 mm; stigma 5-lobed ................................. 13. G. urenifolia 

6b. Leaf blade lanceolate or oblong, base always symmetrical, lateral basal veins reaching mid leaf only. 

16a. Old leaves abaxially subglabrous or sparsely hairy on veins, adaxially hairy only in vein axils at most. 

17a. Leaf blade leathery; peduncle robust, with long hairs; petals 2.5–3 mm .......................................... 14. G. concolor 

17b. Leaf blade thinly leathery; peduncle slender, with short hairs; petals 4–5 mm .......................... 15. G. macropetala 

16b. Old leaves abaxially tomentose or rough-hairy, adaxially hairy. 

18a. Leaf blade broadly oblong, base broadly cuneate or obtuse on one side; petiole 7–9 mm .................. 25. G. henryi 

18b. Leaf blade lanceolate, oblong-lanceolate, triangular-lanceolate, or band-shaped, base rounded or 

slightly cordate, rarely obtuse; petiole 1–8 mm. 

19a. Leaf blade triangular-lanceolate, base wide .......................................................................... 16. G. sessiliflora 

19b. Leaf blade lanceolate, oblong-lanceolate, or band-shaped, base rounded and narrow. 

20a. Teeth on leaf blade margin irregular in size, leaf blade leathery, densely 

tomentose ............................................................................................................ 24. G. cuspidatoserrata 

20b. Teeth on leaf blade margin regular and similar in size, leaf blade papery or 

leathery, glabrous or sparsely to densely hairy. 

21a. Leaf blade band-shaped, 13–20 × 1–3 cm. 

22a. Cymes 1–3 per axil; stamens 20–30; leaf veinlets adaxially obscure ............. 23. G. falcata


22b. Cymes 2 to several per axil; stamens 10–20; leaf veinlets adaxially 

prominent ............................................................................................................. 26. G. lacei 

21b. Leaf blade lanceolate or oblong-lanceolate, 4–14 × 2–4 cm. 

23a. Leaf blade abaxially gray-white appressed stellate pilose ......................... 22. G. retusifolia 

23b. Leaf blade abaxially yellow-brown or gray-brown tomentose or coarsely 

hairy. 

24a. Leaf blade abaxially stellate pilose. 

25a. Leaf blade abaxially sparsely yellow stellate pilose ............. 21. G. brachypoda 

25b. Leaf blade abaxially densely white stellate pilose ............ 20. G. latiglandulosa 

24b. Leaf blade abaxially thickly and softly or coarsely tomentose. 

26a. Leaf blade coarsely yellow-brown tomentose; petiole 2–3 mm .. 19. G. hirsuta 

26b. Leaf blade softly gray-brown tomentose; petiole 3–8 mm. 

27a. Leaf blade leathery; petiole 3–5 mm ............................... 17. G. chuniana 

27b. Leaf blade papery; petiole 5–8 mm ................................ 18. G. oligandra 

1. Grewia eriocarpa Jussieu, Ann. Mus. Natl. Hist. Nat. 4: 93. 

1804. 

毛果扁担杆 mao guo bian dan gan 

Grewia boehmeriifolia Kanehira & Sasaki; G. lantsangensis 

Hu. 

Shrubs or small trees to 8 m tall. Branchlets softly gray 

stellate tomentose. Stipule filiform, lanceolate, 5–10 mm; petiole 

5–10 mm; leaf blade ovate or ovate-oblong, 6–13 × 3–6 cm, 

papery, black-brown when dry, softly gray stellate tomentose 

abaxially, sparsely stellate adaxially, lateral basal veins to 3/4 as 

long as leaf blade, lateral veins 3–4 pairs, base obliquely 

rounded or truncate, margin serrulate, apex acuminate or acute. 

Cymes 1–3, axillary, 1.5–3 cm; peduncle 3–8 mm. Pedicel 3–5 

mm. Bracts lanceolate. Flowers bisexual. Sepals narrowly oblong, 

6–8 mm, hairy on both surfaces. Petals ca. 3 mm, eglandular. 

Androgynophore absent. Stamens irregular in length, 

shorter than sepals. Ovary hairy; style puberulent. Drupe subglobose, 

furrowed, 6–8 mm in diam., stellate hairy; drupelets 1 

or 2. 

Guangdong, Guangxi, Guizhou, Taiwan, Yunnan [Bhutan, Cambodia, 

India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Philippines, 

Sri Lanka, Thailand, Vietnam]. 

2. Grewia celtidifolia Jussieu, Ann. Mus. Natl. Hist. Nat. 4: 93. 

1804. 

朴叶扁担杆 pu ye bian dan gan 

Grewia asiatica Linnaeus var. celtidifolia (Jussieu) L. F. 

Gagnepain; G. simaoensis Y. Y. Qian; G. yunnanensis Hung T. 

Chang. 

Shrubs. Branchlets softly gray-brown tomentose. Petiole 

5–6 mm, softly tomentose; leaf blade broadly ovate, 8–10 × 

4.5–6 cm, shortly yellow-brown stellate tomentose abaxially, 

glabrescent adaxially, lateral basal veins 3/4 as long as leaf 

blade, ca. 1 cm from margin, each with secondary veins 5–6, 

lateral veins 2–3 pairs distally, base rounded, margin serrate, 

apex shortly acute. Cymes 1–1.5 cm, yellow-brown stellate 

tomentose; peduncle to 1 cm. Pedicel 2–3 mm. Sepals lanceolate, 

6–7 mm, tomentose abaxially, pubescent adaxially. Petals 

3–4 mm, glandular at base. Stamens 3–4 mm. Androgynophore 

hairy. Ovary 2-loculed, tomentose; style ca. 2 mm. Drupe 

subglobose, shallowly furrowed, 6–8 mm in diam., stellate 

hairy; exocarp not separating from endocarp; drupelets 1 or 2. 

Fl. Apr–Jun, fr. Jul–Aug. 

253 

Open forests, shrublands; 100–1800 m. Guangdong, Guangxi, 

Guizhou, Taiwan, Yunnan [Cambodia, Indonesia, Laos, Malaysia, 


3. Grewia yinkiangensis Y. C. Hsu & R. Zhuge, Fl. Yunnan. 

6: 412. 1995. 

盈江扁担杆 ying jiang bian dan gan 

Shrubs. Branchlets striate, gray-yellow stellate pilose. Petiole 

very short, 3–4 mm, gray-yellow pilose; leaf blade nearly 

orbicular, 8–12 × 7.5–10 cm, papery, densely stellate tomentose 

abaxially, sparsely minutely hairy adaxially, basal veins 5, lateral 

veins 2–3 pairs, ascending, base rounded, margin entire 

basally, glandular dentate distally, apex obtuse or acute. Cymes 

2–4 per leaf axil, slender, 3-flowered. Infructescence stipe 1.8– 

2.5 cm; stipe 8–12 mm, densely pilose. Drupe subglobose, 4–5 

mm in diam., sparsely stellate pilose; drupelets 1 or 2. Fl. Apr– 

May, fr. Jun–Jul. 

● Open woodlands, scrub; ca. 1000 m. SW Yunnan. 

4. Grewia tiliifolia Vahl, Symb. Bot. 1: 35. 1790 [“tiliaefolia”]. 

椴叶扁担杆 duan ye bian dan gan 

Grewia rotunda C. Y. Wu ex Hung T. Chang. 

Trees 5–8 m tall. Branchlets gray stellate tomentose. 

Petiole ca. 1 cm, softly tomentose; leaf blade nearly orbicular or 

broadly ovate-orbicular, (1.7–)8–13(–36) × (1–)6.5–9.5(–24) 

cm, papery, black-brown when dry, with sparse solitary hairs or 

glabrescent and hairy only on veins adaxially, lateral basal 

veins 2/3 as long as leaf blade, 1.5–2.7 cm from margin, each 

with 5 or 6 secondary veins, dark brown, abaxially sparsely 

hairy or glabrous, base obliquely cordate, margin serrulate, 

apex shortly acute. Cymes 2–6 per leaf axil, 3-flowered; peduncle 

1–1.5 cm. Pedicel 6–7 mm, gray-brown tomentose. Sepals 

oblong lanceolate, 7–8 mm, softly gray-yellow tomentose 

on both surfaces. Petals yellow, shorter and narrower than sepals. 

Stamens many, in 5 fascicles, slightly connate at base, 

slightly shorter than sepals. Ovary 2-loculed, hairy; style 

slightly longer than stamens. Drupe subglobose, not divided; 

exocarp not separating from endocarp; drupelets 1 or 2. Fl. 

Apr–Jun, fr. Jul–Aug. 

Open woodlands, shrublands, grasslands; 800–1600 m. Guangxi,

254 

SW Yunnan [Cambodia, India, Laos, Malaysia, Myanmar, Thailand, 

Vietnam; E Africa]. 

5. Grewia kwangtungensis Hung T. Chang, Acta Phytotax. 

Sin. 20: 175. 1982. 

广东扁担杆 guang dong bian dan gan 

Shrubs, climbing. Branchlets brown, shortly tomentose. 

Petiole 5–6 mm; leaf blade oblong or lanceolate, 6–10 × 2–3 

cm, membranous, abaxially puberulent only on veins, adaxially 

hairy, glabrescent, lateral basal veins 1/3 as long as leaf blade, 

2–3 mm from margin, lateral veins 3–5 pairs, minor veins 

prominently raised abaxially, base cordate or slightly cordate, 

margin serrulate, apex acuminate or caudate-acuminate. Flowers 

unknown. Infructescence in subterminal axils; peduncle 3–6 

mm, shortly brown tomentose; fruiting pedicel 7–9 mm, shortly 

brown tomentose. Androgynophore short, glabrous, with hairs 

on apex. Drupe red, globose, 6–7 mm in diam., hairy, becoming 

glabrescent, glossy; drupelets 1 or 2. 

● Evergreen forest; 800–900 m. Guangdong. 

6. Grewia rhombifolia Kanehira & Sasaki, Trans. Nat. Hist. 

Soc. Taiwan 18: 335. 1928. 

菱叶扁担杆 ling ye bian dan gan 

Shrubs. Branchlets rust-brown stellate tomentose. Stipule 

subulate, 5–7 mm; petiole 5–7 mm; leaf blade rhomboid or 

ovate, rarely ovate-orbicular, 2.5–5.5 × 2–4.5 cm, sparsely stellate, 

abaxially hairy only on veins, adaxially glabrescent, lateral 

basal veins to 1/2 as long as leaf blade, lateral veins 4–5 pairs, 

base broadly cuneate to nearly rounded, margin irregularly 

serrate, apex slightly acute or obtuse. Cymes axillary; peduncle 

5–8 mm. Pedicel 5–7 mm. Bracts subulate, ca. 4 mm. Sepals ca. 

6 × 1.5–2 mm, glabrous abaxially, adaxially stellate tomentose. 

Petals oblong, ca. 1.5 mm; glands nearly round, hairy. Stamens 

many, with long silk hairs. Ovary hairy; style glabrous. Drupe 

2–4-lobed; drupelets 1 per lobe. Fl. Jun. 

● Taiwan. 

7. Grewia biloba G. Don, Gen. Hist. 1: 549. 1831. 

扁担杆 bian dan gan 

Shrubs or small trees 0.5–1(–4) m tall, many branched. 

Branchlets pubescent or nearly glabrous. Stipule subulate, 3–4 

mm; petiole 4–8 mm, densely puberulent; leaf blade ovateorbicular 

or obovate-elliptic, sometimes shallowly 3-lobed, 4–9 

× 2.5–4 cm, thinly leathery, abaxially sparsely stellate hairy to 

stellate tomentose, adaxially more sparsely stellate hairy, lateral 

basal veins more than 1/2 as long as leaf blade, lateral veins 

3–5 pairs, base cuneate or obtuse, margin serrulate, apex acute. 

Cymes axillary, many flowered; peduncle less than 1 cm. Pedicel 

3–6 mm. Bracts subulate, 3–5 mm. Sepals narrowly oblong, 

4–7 mm, abaxially hairy, adaxially white and glabrous. Petals 

1–1.5 mm. Androgynophore ca. 0.5 mm, hairy. Stamens ca. 2 

mm. Ovary hairy; style as long as sepals. Drupe red, 2–4-lobed; 

drupelets 1 per lobe. Fl. May–Jul. 

Anhui, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hubei, 

Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, 

Yunnan, Zhejiang [Korea]. 


1a. Leaf blade suborbicular, 1–1.5 × ca. 

1 cm .................................................... 7c. var. microphylla 

1b. Leaf blade elliptic or obovate-elliptic, 

4–9 × 2.5–4 cm. 

2a. Leaf blade sparsely stellate hairy 

abaxially .............................................. 7a. var. biloba 

2b. Leaf blade densely softly stellate 

tomentose abaxially ...................... 7b. var. parviflora 

7a. Grewia biloba var. biloba 

扁担杆(原变种) bian dan gan (yuan bian zhong) 

Grewia biloba var. glabrescens (Bentham) Rehder; G. glabrescens 

Bentham; G. parviflora Bunge var. glabrescens (Bentham) 

Rehder & E. H. Wilson; G. tenuifolia Kanehira & Sasaki. 

Leaf blade elliptic or obovate-elliptic, 4–9 × 2.5–4 cm, 

abaxially sparsely stellate hairy. 

Anhui, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hunan, 

Jiangsu, Jiangxi, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Yunnan, 

Zhejiang [Korea]. 

7b. Grewia biloba var. parviflora (Bunge) Handel-Mazzetti, 

Symb. Sin. 7: 612. 1933. 

小花扁担杆 xiao hua bian dan gan 

Grewia parviflora Bunge, Enum. Pl. China Bor. 9. 1833; 

G. chanetii H. Léveillé; G. parviflora var. velutina Pampanini. 

Leaf blade elliptic or obovate-elliptic, 4–9 × 2.5–4 cm, 

abaxially densely softly stellate tomentose. Flowers smaller. 

● Anhui, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hubei, 

Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Shanxi, Sichuan, Yunnan, 

Zhejiang. 

7c. Grewia biloba var. microphylla (Maximowicz) Handel- 

Mazzetti, Symb. Sin. 7: 612. 1933. 

小叶扁担杆 xiao ye bian dan gan 

Grewia parviflora var. microphylla Maximowicz, Trudy 

Imp. S.-Peterburgsk. Bot. Sada 11: 81. 1890. 

Leaf blade suborbicular, 1–1.5 cm, sparsely pilose abaxially. 

Flowers larger. 

● Sichuan, Yunnan. 

8. Grewia piscatorum Hance, Ann. Sci. Nat., Bot., sér. 5, 5: 

208. 1866. 

海岸扁担杆 hai an bian dan gan 

Shrubs small, many branched, extending, slender. Branchlets 

sparsely stellate. Stipule filiform, ca. 2 mm; petiole 2–3 

mm; leaf blade ovate or elliptic, 1–2.3 × 1–2 cm, nearly membranous, 

glabrous adaxially, greenish when dry, luminous, glabrous 

or sparsely pilose abaxially, lateral basal veins ca. 1/2 

as long as leaf blade, lateral veins 2–3 pairs, base rounded or 

slightly cordate, margin serrate, apex obtuse or shortly acute. 

Cymes 1 or 2 in subterminal axils, usually 3-flowered; peduncle 

ca. 5 mm. Pedicel as long as peduncle or slightly shorter. Bracts 

subulate, 2–4 mm. Sepals narrowly oblong, 5–7 mm, abaxially

hairy, adaxially glabrous. Petals 1.5–2 mm. Androgynophore 

very short. Stamens many, 4–5 mm. Ovary hairy; style glabrous. 

Drupe 2- or 4-lobed, furrowed; drupelets 1 per lobe. 

● Coastal areas. Fujian, Hainan, Taiwan. 

This species is very similar to Grewia biloba var. microphylla. It 

differs in having leaves that are nearly membranous, not thinly leathery, 

and abaxially nearly glabrous, not stellate hairy. 

9. Grewia angustisepala Hung T. Chang, Acta Phytotax. Sin. 

20: 175. 1982. 

狭萼扁担杆 xia e bian dan gan 

Shrubs 1–2.5 m tall. Branchlets softly yellow-brown tomentose. 

Petiole 3–5 mm, softly yellow-brown tomentose; leaf 

blade ovate-elliptic, thinly leathery, 13–16 × 6–9 cm, densely 

softly yellow-brown tomentose abaxially, stellate but glabrescent 

and hairy only on veins adaxially, lateral basal veins to 2/3 

as long as blade, 8–15 mm from margin, each with secondary 

veins 6–8, lateral veins 4–5 pairs in distal 1/2, base rounded or 

slightly cordate, slightly oblique, margin dentate (teeth alternately 

small and large, triangular, 1–2.5 mm), apex acuminate. 

Cymes 1 or 2 in axils of leaves, 2–3 cm; peduncle 1–1.7 cm. 

Pedicel 6–8 mm. Bracts lanceolate, 6–8 mm, hairy. Sepals narrowly 

lanceolate, 7–8 × 1–1.5 mm, abaxially tomentose, adaxially 

white and glabrous. Petals 3–4 mm. Stamens 5–6 mm. 

Ovary tomentose; style glabrous. Drupe 2-lobed, lobes globose. 

● Open forests; 800–1200 m. Yunnan. 

10. Grewia permagna C. Y. Wu ex Hung T. Chang, Acta Phytotax. 

Sin. 20: 176. 1982. 

大叶扁担杆 da ye bian dan gan 

Grewia rugulosa C. Y. Wu ex Hung T. Chang. 

Shrubs or small trees. Bark gray-brown; branchlets 

coarsely yellow-brown tomentose. Petiole 1.7–2.5 cm, coarsely 

stellate; leaf blade nearly orbicular, 21–28 × 8–23 cm, leathery, 

softly yellow-brown tomentose abaxially, coarsely hairy adaxially, 

lateral basal veins ascending to near apex of leaf blade, 

3.5–5 cm from margin, each with secondary veins 8–11, lateral 

veins 5–6 pairs, base rounded or slightly obliquely cordate, 

margin dentate, apex acute. Flowers not seen. Cymes axillary, 

ca. 1.5 cm; peduncle 7–11 mm, coarsely hairy. Drupe globose, 

7–8 mm in diam., slightly hairy. Fr. winter. 

● Scrub; ca. 1200 m. Yunnan. 

11. Grewia acuminata Jussieu, Ann. Mus. Natl. Hist. Nat. 4: 


密齿扁担杆 mi chi bian dan gan 

Grewia densiserrulata Hung T. Chang. 

Shrubs climbing. Branchlets densely brown stellate, 

shortly coarsely hairy. Petiole 3–4 mm, densely yellow stellate; 

leaf blade ovate to ovate-oblong, 8–13 × 4–6 cm, leathery, 

stellate, shortly coarsely hairy adaxially, densely brown stellate 

abaxially, lateral basal veins to 1/2 as long as leaf blade, 6–8 

mm from margin, each with 10 secondary veins, lateral veins 

5–6 pairs distally, base rounded, margin densely serrulate, teeth 


255 

less than 0.4 mm, ca. 1 mm apart, apex acute. Cymes solitary in 

leaf axils, 2- or 3-flowered; peduncle 1–1.5 cm. Pedicel 12–18 

mm. Sepals to 2 cm. Petals ca. 7 mm. Androgynophore 4–8 

mm. Stamens many; filaments 8–12 mm. Ovary 2-loculed; 

style ca. 1 cm. Fruiting peduncle 1–2 cm, pedicels 1.5–2 cm, 

both coarsely stellate hairy. Drupe 4-lobed, ca. 2 cm in diam.; 

drupelets coarsely hairy; androgynophore 5–6 mm, stellate tomentose. 

Scrub; 800–900 m. S Yunnan [Cambodia, India, Indonesia (Java), 

Laos, Malaysia, Myanmar, Thailand, Vietnam]. 

12. Grewia abutilifolia W. Vent ex Jussieu, Ann. Mus. Natl. 

Hist. Nat. 4: 92. 1804. 

苘麻叶扁担杆 qing ma ye bian dan gan 

Grewia hirsutovelutina Burret; G. kainantensis Masamune; 

Sterculia tiliacea H. Léveillé. 

Shrubs or small trees 1–5 m tall; branchlets yellow-brown 

stellate, coarsely hairy. Petiole 1–2 cm, coarsely stellate tomentose; 

leaf blade broadly ovate to nearly orbicular, usually lobed 

distally, 7–11 × 5–9 cm, papery, densely yellow and brown, 

coarsely stellate tomentose abaxially, adaxially sparsely 

coarsely stellate hairy, lateral basal veins more than 1/2 as long 

as leaf blade, each with 7–9 secondary veins, lateral veins distal, 

3–4 pairs, base rounded or slightly cordate, margin serrulate, 

apex acute. Cymes 3–7 per leaf axil; peduncle 3–6 mm. 

Pedicel 4–8 mm. Bracts filiform, caducous. Sepals white, narrowly 

oblong, 6–8 × ca. 2 mm, abaxially hairy, adaxially glabrous. 

Petals 2–3 mm. Androgynophore glabrous. Stamens 4–5 

mm. Ovary villous; style as long as sepals; stigma 2-lobed. 

Drupe obscurely 2- or 4-lobed, tomentose; drupelets 1 per lobe. 

Scrub, slopes on grasslands. Guangdong, Guangxi, Guizhou, Hainan, 

Taiwan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, 


13. Grewia urenifolia (Pierre) Gagnepain, Notul. Syst. (Paris) 

1: 126. 1909 [“urenaefolia”]. 

稔叶扁担杆 ren ye bian dan gan 

Grewia abutilifolia W. Vent ex Jussieu var. urenifolia 

Pierre, Fl. Forest. Cochinch. t. 164. 1888 [“urenaefolia”]. 

Shrubs 1–2 m tall. Branchlets coarsely stellate. Petiole 1– 

1.5 cm, coarsely hairy; leaf blade broadly ovate, 6–8 × 4–6.5 

cm, coarsely stellate adaxially, densely coarsely stellate abaxially, 

base cordate. Cymes 1–2 in axils of leaves; peduncle ca. 1 

cm. Pedicel 4–6 mm. Flower buds 6–7 mm. Sepals ca. 6 mm, 

hairy abaxially, adaxially glabrous. Petals minute, gland base 

hairy. Ovary hairy; stigma 5-lobed. Drupe 2–4-lobed; drupelets 

1 per lobe. 

Secondary scrub. Guangxi, Hainan, Yunnan [Cambodia, Laos, 

Malaysia, Myanmar, Thailand, Vietnam]. 

This species is similar to Grewia abutilifolia but differs in having 

only 1 or 2 cymes per leaf axil, longer peduncles and pedicels, and a 5lobed 

stigma. 

14. Grewia concolor Merrill, Lingnan Sci. J. 14: 35. 1935. 

同色扁担杆 tong se bian dan gan

256 

Shrubs climbing. Branchlets rust-brown stellate tomentose. 

Stipules lanceolate, ca. 5 × 2 mm; petiole 5–7 mm, hairy; leaf 

blade oblong, 7–12 × 2.5–4 cm, leathery, sparsely stellate and 

villous with long solitary hairs at first on both surfaces, glabrescent 

or with sparsely solitary hairs only on veins on both surfaces, 

lateral basal veins less than 1/2 as long as leaf blade, 

lateral veins 5–6 pairs, base slightly oblique, nearly rounded to 

shallowly cordate, margin serrulate, apex shortly acute or obtuse. 

Cymes 1 or 2 per leaf axil, 2- or 3-flowered; peduncle ca. 

1 cm, brown hairy. Flowers unisexual. Pedicel 5–9 mm, brown 

hairy. Bracts subulate, ca. 5 mm, hairy abaxially. Sepals narrowly 

lanceolate, 7–8 mm, adaxially hairy. Petals 2.5–3 mm; 

glands obovoid, hairy. Stamens many. Ovary villous; style glabrous; 

stigma 5-lobed. Drupe 2–4-lobed, lobes globose; drupelets 

1 per lobe. Fl. Jul–Aug. 

● Forests, open scrub. Fujian, Hainan. 

15. Grewia macropetala Burret, Repert. Spec. Nov. Regni Veg. 

33: 74. 1933. 

长瓣扁担杆 chang ban bian dan gan 

Shrubs 1–3 m tall. Branchlets coarsely brown stellate. 

Petiole 4–5 mm, stellate pubescent; leaf blade oblong-lanceolate, 

4–7 × 1.5–2 cm, thinly leathery, sparsely coarsely stellate 

pubescent, glabrescent, lateral basal veins to 1/2 as long as leaf 

blade, lateral veins 3–4 pairs, veinlets prominent on both surfaces, 

base narrowly rounded, margin irregularly serrulate, apex 

acuminate. Cymes 1 or 2 per leaf axil, 3-flowered; peduncle 

slender, 4–14 mm. Pedicel 4–12 mm, stellate pubescent. Bracts 

filiform, 3–4 mm. Sepals ca. 6 mm, abaxially hairy, adaxially 

glabrous. Petals 4–5 mm. Androgynophore short, glabrous. Stamens 

16–18, as long as or slightly longer than petals. Ovary 

hairy; style as long as stamens; stigma 2–4-lobed. Drupe 2–4lobed, 

hairy, glabrescent; drupelets 1 per lobe. Fl. May–Jun. 

● Secondary forests. Guangdong, Guangxi, Yunnan. 

16. Grewia sessiliflora Gagnepain, Notul. Syst. (Paris) 1: 167. 

1910. 

无柄扁担杆 wu bing bian dan gan 

Shrubs. Branches and branchlets softly yellow-brown tomentose. 

Petiole 1–3 mm; leaf blade triangular-lanceolate, 6–10 

× 3–5 cm, abaxially densely softly yellow-brown tomentose, 

adaxially softly stellate tomentose, lateral basal veins to 1/2 as 

long as leaf blade, 7–13 mm from margin, lateral veins 3–4 

pairs, base oblique, rounded or shallowly cordate, margin serrate, 

apex narrowly acuminate. Cymes 1 or 2 per leaf axil, 4–5 

cm; peduncle 3–4 cm, tomentose. Bracts filiform, 7–8 mm, 

tomentose. Sepals lanceolate, 8–9 × ca. 1.5 mm, abaxially tomentose, 

adaxially glabrous. Petals 4–5 mm. Stamens slightly 

longer than petals. Ovary tomentose; style longer than stamens. 

Drupe 2-lobed, lobes globose, glossy, sparsely stellate hairy. Fl. 

Jun–Jul. 

Secondary forests at low elevations. S Guangdong, Guangxi 

[Laos, N Thailand, N Vietnam]. 

The diagnostic features of this species include the triangular-lanceolate 

leaves covered with yellow-brown soft indumentum and the long 

cymes; at 4–5 cm, these are the longest cymes in the genus in China. 


17. Grewia chuniana Burret, Notizbl. Bot. Gart. Berlin- 

Dahlem 13: 488. 1936. 

崖县扁担杆 ya xian bian dan gan 

Shrubs. Old branches dark brown, glabrous; branchlets 

densely rust-brown tomentose. Stipules subulate, 5–6 mm; petiole 

3–5 mm, densely tomentose; leaf blade oblong-lanceolate, 

7–11 × 3–4 cm, leathery, abaxially densely gray-brown stellate 

tomentose, adaxially sparsely hairy, base rounded or obliquely 

shallowly cordate, margin serrulate, apex acuminate. Cymes 3flowered; 

peduncle 1–1.2 cm, tomentose. Pedicel 5–7 mm, tomentose. 

Bracts subulate, ca. 5 mm. Sepals 5–6 mm, abaxially 

densely tomentose, adaxially glabrous. Petals ca. 2.5 mm; 

glands scalelike, enclosed by ring of long hairs. Stamens many, 

shorter than sepals, slightly longer than petals. Ovary hairy; 

style as long as sepals; stigma many-lobed. Drupe 2–4-lobed; 

drupelets 1 per lobe. Fl. Aug–Sep. 

● Coastal scrub. Hainan. 

This species is similar to Grewia hirsuta but differs in having leaf 

blades that are gray-brown, not yellowish brown, stellate tomentose, 

and not blackened when dried, and longer cymes: peduncle 10–12 mm, 

not 3–7 mm. 

18. Grewia oligandra Pierre, Fl. Forest. Cochinch. t. 163. 

1888. 

寡蕊扁担杆 gua rui bian dan gan 

Shrubs. Old branches dark brown; branchlets stellate tomentose. 

Stipules subulate, ca. 4 mm, caducous; petiole 5–8 

mm; leaf blade lanceolate or oblong-lanceolate, 9–10.5 × 2–3.5 

cm, papery, dark brown when dried, abaxially densely softly 

gray-brown tomentose, adaxially sparsely coarsely shortly stellate 

hairy, base rounded, margin serrulate, teeth alternately large 

and small. Cymes 3–5-flowered; peduncle 4–7 mm, tomentose. 

Pedicel 3–4 mm, tomentose. Bracts 3–4 mm. Sepals 5–6 mm, 

abaxially tomentose, adaxially glabrous. Petals oblong, 2–3 mm; 

glands scalelike, enclosed by ring of hairs. Stamens shorter than 

sepals. Ovary hairy; stigma many-lobed. Drupe 2–4-lobed, lobes 

globose, ca. 1 cm in diam., glossy. Fl. Aug. 

Guangdong, Guangxi, Hainan [Cambodia, Laos, Malaysia, Myanmar, 

Thailand, Vietnam]. 

This species is similar to Grewia cuspidatoserrata but differs by 

its soft gray-brown, not rust-brown, indumentum on the abaxial leaf 

blade surface and longer cymes: peduncle 4–7 mm, not less than 3 mm. 

19. Grewia hirsuta Vahl, Symb. Bot. 1: 34. 1790. 

粗毛扁担杆 cu mao bian dan gan 

Shrubs or small trees. Branchlets coarsely gray-brown 

stellate hairy. Petiole 2–3 mm, tomentose; leaf blade lanceolate, 

6–14 × 2–3.5 cm, leathery, black-brown when dried, abaxially 

yellow-brown stellate tomentose, adaxially sparsely coarsely 

stellate, lateral basal veins to 1/2 as long as leaf blade, lateral 

veins 4–5 pairs, base narrow, shallowly cordate, margin serrulate, 

apex acuminate or rarely obtuse. Cymes 1–5 per leaf axil, 

3- or 4-flowered; peduncle 3–7 mm, tomentose. Pedicel 3–5 

mm, tomentose. Bracts lanceolate, 3–4 mm. Sepals narrowly 

lanceolate, 6–7 × ca. 1.5 mm, abaxially tomentose, adaxially

glabrous. Petals narrowly ovate, ca. 3 × 1.5 mm. Stamens 4–5 

mm. Ovary villous; style longer than stamens, glabrous; stigma 

4-lobed. Drupe globose or 2-lobed, sparsely coarsely hairy; 

drupelets 2 per lobe. Fl. Jun–Jul. 

Guangdong, S Guangxi [Bangladesh, Cambodia, India, Laos, Malaysia, 

Myanmar, Nepal, Sri Lanka, Thailand, Vietnam]. 

20. Grewia latiglandulosa Z. Y. Huang & S. Y. Liu, J. Trop. 

Subtrop. Bot. 13: 367. 2005. 

阔腺扁担杆 kuo xian bian dan gan 

Shrubs 1–1.8 m tall. Stem and branches grayish brown; 

branchlets pale brown, densely white stellate pilose. Stipules 

linear-lanceolate, 3–5 mm; petiole 2–3 mm, stellate pilose; leaf 

blade elliptic or oblong, 2.2–10 × 1.2–3.4 cm, papery, abaxially 

densely stellate pilose, adaxially drying green, minutely stellate 

pilose, lateral basal veins to 1/2 as long as leaf blade, lateral 

veins 3–6 pairs, base rounded or nearly rounded, slightly 

oblique, margin serrulate, apex acuminate or long acuminate. 

Cymes 1 or 2 per leaf axil, 1–3-flowered; peduncle 8–15 mm, 

densely white stellate pilose. Pedicel 5–10 mm. Sepals 10–12 × 

2–3 mm, abaxially densely stellate pilose, adaxially white, glabrous, 

apex obtuse. Petals yellow, ovate-rotund, 2.5–3 × 1.5–2 

mm; gland scalelike, ca. 2/3 as large as petal, margin pilose. 

Androgynophore ca. 1 mm, pilose. Stamens many; filaments 

white, unequal; anthers yellow. Ovary depressed-globose, 

densely white stellate pilose; style ca. 7 mm, stellate pilose 

below middle; stigma 4- or 5-lobed. Drupe yellow-green drying 

blackish, 2–4-lobed, sparsely white stellate pilose, glossy; 

drupelets ca. 5 × 5–6 mm. 

● Guangxi (Tianyang). 

21. Grewia brachypoda C. Y. Wu, J. W. China Border Res. 

Soc. 16: 162. 1946. 

短炳扁担杆 duan bing bian dan gan 

Shrubs 0.5–1.5 m tall. Branchlets densely yellow-brown 

tomentose, two-year-old branches slightly coarsely tomentose. 

Petiole 1–4 mm, densely tomentose; leaf blade oblong-lanceolate 

or oblong, 3–9 × 2–3.5 cm, leathery, abaxially densely yellow-brown 

to gray-brown stellate tomentose, adaxially coarsely 

stellate hairy, fallen hairs leaving verrucose prominence, lateral 

basal veins 1/2 as long as blade or longer, lateral veins 3–4 

pairs, base rounded, margin serrate, apex acute. Cymes solitary 

in axils, 3-flowered; peduncle 0.7–1 cm, tomentose. Pedicel 4– 

6 mm, tomentose. Bracts subulate, hairy, caducous. Sepals 5–7 

× 2–2.5 mm, abaxially stellate hairy, adaxially glabrous. Petals 

2–3 mm. Stamens 3–5 mm. Ovary villous; style 3–4 mm, glabrous; 

stigma swollen. Drupe 2-lobed, 1–1.2 cm, glossy, glabrous. 

● Scrub, dry grassy slopes. SW Sichuan, N and SW Yunnan. 

22. Grewia retusifolia Pierre, Fl. Forest. Cochinch. t. 168. 

1888. 

钝叶扁担杆 dun ye bian dan gan 

Shrubs or small trees. Branchlets slender, appressed yellow 

stellate tomentose. Petiole 3–5 mm; leaf blade oblong-lanceolate, 

6–8.5 × 2.2–3.4 cm, abaxially gray-white adnate stel- 


257 

late puberulent, adaxially coarsely shortly stellate hairy, glabrescent, 

lateral basal veins to 1/2 as long as leaf blade, 4–7 mm 

from margin, lateral veins 3–4 pairs, base rounded, margin irregularly 

denticulate, apex obtuse or slightly acute. Cymes 1 or 

2 per leaf axil, 3–5-flowered; peduncle ca. 1 cm, slender, hairy. 

Pedicel ca. 3 mm. Bracts filiform, lanceolate, 3–5 mm. Sepals 

narrowly lanceolate, 5–6 mm, abaxially stellate tomentose, 

adaxially glabrous. Petals ca. 2.5 mm. Androgynophore slightly 

longer than petals. Ovary hairy; style longer than stamens; stigma 

4-lobed. Drupe 4-lobed, lobes globose, glossy dark reddish 

brown. 

Secondary forests. S Guangxi (Nanning) [Indonesia, Vietnam; N 

Australia]. 

23. Grewia falcata C. Y. Wu, J. W. China Border Res. Soc. 16: 


镰叶扁担杆 lian ye bian dan gan 

Shrubs or small trees. Branchlets softly yellow-brown tomentose. 

Petiole 3–5 mm, tomentose; leaf blade band-shaped, 

slightly curved or straight, or falcate, 6–18 × 1–3 cm, leathery 

or papery, abaxially softly yellow-brown or rust-brown tomentose, 

adaxially sparsely stellate pilose and more densely on 

veins, lateral basal veins obscure, 1/4–1/3 as long as leaf blade, 

1–3 mm from margin, lateral veins 4–8 pairs, veinlets prominent, 

base obtuse, margin serrulate, apex acuminate. Cymes 1–3 

per leaf axil; peduncle 3–5 mm, yellow-brown tomentose. Pedicel 

3–4 mm, yellow-brown tomentose. Sepals lanceolate, 8–9 × 

2–2.5 mm, abaxially tomentose, adaxially glabrous. Petals oblong, 

3–4 mm, glands on basal 1/2. Stamens 20–30, 5–6 mm. 

Ovary densely yellow-brown villous; style longer than stamens; 

stigma 4-lobed. Drupe 4-lobed, lobes globose, hairy, glossy. 

Open forests; 800–1700 m. SW Guangxi, Yunnan [Cambodia, 

Laos, Malaysia, Myanmar, Thailand, Vietnam]. 

This species is similar to Grewia cuspidatoserrata but differs in 

having the leaf margin with uniform teeth, not alternately small and 

large, the indumentum yellow-brown, not gray-brown, and the leaf 

blade narrowly band-shaped, not oblong-lanceolate or lanceolate. 

24. Grewia cuspidatoserrata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 

9: 718. 1926 [“cuspidato-serrata”]. 

尖齿扁担杆 jian chi bian dan gan 

Shrubs. Old branches brown; branchlets rust-brown tomentose. 

Stipules lanceolate, ca. 7 mm, caducous; petiole 4–6 

mm, tomentose; leaf blade oblong-lanceolate or lanceolate, 7– 

16 × 2.5–4.5 cm, papery or leathery, abaxially densely softly 

rust-brown tomentose, adaxially coarsely stellate, lateral basal 

veins 1/3 to more than 1/2 as long as leaf blade, lateral veins 

distal, 4–5 pairs, base nearly rounded, slightly oblique, margin 

double-serrate, teeth 1–2.5 mm, alternately small and large, narrow 

and acute, apex shortly acute or slightly obtuse. Cymes 1 to 

several per leaf axil; peduncle less than 3 mm. Pedicel 5–8 mm. 

Bracts lanceolate, ca. 3 mm. Sepals oblong, 6–7 × ca. 2 mm, 

abaxially hairy, adaxially glabrous. Petals ca. 3 mm, abaxially 

hairy at base; glands obovoid, enclosed by ring of hairs. Stamens 

many, with long hairs in between. Ovary hairy. Fruit 2–4lobed. 

Seeds 1 or 2 per drupelet. Fl. Apr–May.

258 

● Open forests; 1200–1500 m. S Yunnan. 

25. Grewia henryi Burret, Notizbl. Bot. Gart. Berlin-Dahlem 

9: 674. 1926. 

黄麻叶扁担杆 huang ma ye bian dan gan 

Shrubs or small trees 1–6 m tall. Branchlets yellow-brown 

coarsely stellate. Petiole 7–9 mm, coarsely stellate; leaf blade 

broadly oblong, 11–19 × 3–4.5 cm, thinly leathery, abaxially 

greenish or coarsely yellow-green stellate, glabrescent, adaxially 

yellow-green when dry, sparsely shortly stellate hairy, lateral 

basal veins 1/3–1/2 as long as blade, 3–8 mm from margin, 

lateral veins 4–6 pairs, base broadly cuneate, rarely unequally 

obtuse, margin serrulate, apex acuminate. Cymes 1 or 2 per leaf 

axil, 3–4-flowered; peduncle 1–2.5 cm. Pedicel 5–11 mm. Sepals 

lanceolate, 1–1.3 cm, abaxially tomentose, adaxially glabrous. 

Petals long ovate, 4–5 mm. Stamens 5–7 mm. Ovary 4loculed, 

hairy; style 6–7 mm; stigma 4-lobed. Drupe 4-lobed. 

● Fujian, Guangdong, Guangxi, Guizhou, Jiangxi, Yunnan. 

26. Grewia lacei Drummond & Craib, Bull. Misc. Inform. Kew 

1911: 21. 1911. 

细齿扁担杆 xi chi bian dan gan 

Trees to 5 m tall. Branchlets densely tomentose. Petiole ca. 

5 mm, hairy; leaf blade narrowly lanceolate, 15–20 × 2–2.5 cm, 

papery, densely rust-brown tomentose abaxially, glabrous or 

sparsely hairy adaxially, lateral basal veins 1/4–1/3 as long as 

leaf blade, lateral veins 7–8 pairs, base slightly oblique, cuneate, 

margin obscurely serrulate, apex obtuse or acute. Cymes 2 

to several per leaf axil, 1–3-flowered; peduncle ca. 2 mm. 

Flowers polygamo-monoecious. Pedicel 3–5 mm. Sepals linear, 

ca. 6 mm, margin involute, apex obtuse, densely coarsely yel- 

Bartramia Linnaeus. 


low-brown hairy. Petals ca. 1.5 × 1 mm, abaxially minutely 

hairy, apex erose. Stamens 10–20; filaments ca. 2 mm. Ovary 

densely yellow-brown tomentose; style ca. 2 mm; stigma 

swollen, lobed. Drupe nearly square, 4-lobed, yellow villous. 

Fl. Aug–Sep, fr. Oct–Nov. 

Open lowland forests; 500–700 m. S Yunnan [Laos, Myanmar, 

Thailand]. 

27. Grewia multiflora Jussieu, Ann. Mus. Natl. Hist. Nat. 4: 

89. 1804. 

光叶扁担杆 guang ye bian dan gan 

7. TRIUMFETTA Linnaeus, Sp. Pl. 1: 444. 1753. 

刺蒴麻属 ci shuo ma shu 

Grewia didyma Roxburgh ex G. Don; G. disperma Rottler 

ex Sprengel; G. glabra Blume; G. jinghongensis Y. Y. Qian; G. 

serrulata Candolle. 

Shrubs and small trees. Branchlets sparsely stellate, glabrescent. 

Stipule filiform, 1–1.5 × ca. 0.1 cm, caducous; petiole 

6–8 mm, tomentose; leaf blade oblong-lanceolate, 7–17 × 2.5– 

4.5 cm, papery, glabrous or very sparsely hairy adaxially, hairs 

simple and stellate, in vein axils, lateral basal veins to 1/2 as 

long as leaf blade, lateral veins 5–6 pairs, base cuneate or 

broadly cuneate, margin serrulate, apex long acuminate. Cymes 

2–3 in axils, 3-flowered; peduncle slender, 1.2–2.5 cm. Pedicel 

1–1.2 cm, sparsely stellate. Bracts subulate, ca. 4 mm. Sepals 

ca. 1.3 × 0.3–0.5 cm, densely yellow-brown pubescent abaxially. 

Petals ca. 3 mm. Androgynophore 2–4 mm. Drupe 4lobed, 

glabrous; exocarp not separating. Fl. Jul–Sep, fr. Oct. 

S Yunnan [India, Indonesia, Malaysia, Myanmar, Nepal, Pakistan; 

Australia]. 

This species has been confused with Grewia laevigata Vahl (see 

Chung, Edinburgh J. Bot. 62: 1–27. 2005). 

Herbs, rarely subshrubs, annual or perennial, erect or procumbent, hairs with swollen base, stellate or with 1 ray and apparently 

simple. Leaves alternate, simple or palmately 3–5-lobed, palmately veined, margin serrate. Flowers solitary or arranged in small 

cymes or fascicles, axillary, opposite to leaves, bisexual; peduncle very short. Sepals 5, free, valvate, usually with hooked appendages 

at tip. Petals as many as sepals, free, thickened glands on base of adaxial surface. Stamens 5 to numerous; filaments free; 

anthers dorsifixed, subglobose, dehiscence longitudinal, borne on androgynophore; androgynophore fleshy, segmented, short, with 5 

glands opposite to petals. Ovary 2–5-loculed; ovules 2 per locule; style simple; stigma 2–5-lobed. Fruit a capsule nearly globose, 3– 

6-valved, spiny or strigose, loculicidally dehiscent or indehiscent, spine tips pointed, straight or hooked. Seeds with endosperm; 

cotyledons fleshy, epigeous. 

Between 100 to 160 species: primarily in tropical and subtropical areas, several species are widespread weeds; seven species in China. 

1a. Capsule dehiscent, spines 3–10 mm; leaf blade not lobed. 

2a. Capsule spines glabrous or hairy at base only; leaf blade hairs mostly unbranched, occasionally few-rayed on 

main veins; flower bud nearly glabrous ...................................................................................................................... 3. T. annua 

2b. Capsule spines uniformly hairy; leaf blade hairs mostly obviously stellate, abaxially tomentose; flower bud 

obviously hairy. 

3a. Capsule spines with straight or gently curved apex; pedicel ca. 1.5 mm .............................................................. 1. T. cana 

3b. Capsule spines with strongly hooked apex; pedicel 3–5 mm .............................................................................. 2. T. pilosa 

1b. Capsule indehiscent, spines 1.5–4 mm; blade of lower leaves often 3–5-lobed or coarsely dentate. 

4a. Blade of upper leaves 0.7–1.5 cm wide, abaxial hairs apparently simple .......................................................... 4. T. grandidens 

4b. Blade of upper leaves 1.5–6 cm wide, abaxial hairs stellate.


5a. Leaf blade base cordate, abaxial surface densely yellow-brown tomentose, apex obtuse; stamens 

20–35 ........................................................................................................................................................... 7. T. procumbens 

5b. Leaf blade base broadly cuneate or rounded, abaxial surface pilose or pubescent, apex acute or 

acuminate; stamens 10 or 15–20. 

6a. Pedicels less than 1 mm; stamens 10 ................................................................................................... 5. T. rhomboidea 

6b. Pedicels 2–3 mm; stamens 15 or more .................................................................................................. 6. T. semitriloba 

1. Triumfetta cana Blume, Bijdr. 126. 1825. 

毛刺蒴麻 mao ci shuo ma 

Triumfetta tomentosa Bojer; T. tomentosa var. calvescens 

Franchet. 

Herbs woody or subshrubs to 2 m tall. Branchlets yellowbrown 

tomentose, hairs stellate. Petiole 1–3 cm; leaf blade 

ovate or ovate-lanceolate, 4–8 × ca. 2.4 cm, abaxially densely 

stellate tomentose, adaxially sparsely stellate hairy, basal veins 

3–5, 2 lateral ones ascending beyond 1/2 blade, base rounded, 

margin irregularly serrate, apex acuminate. Cymes 1 to several 

per axil; peduncle ca. 3 mm. Flower buds densely appressed 

gray-white puberulent. Pedicel ca. 1.5 mm. Sepals narrowly 

oblong, ca. 7 mm, tomentose. Petals oblong, slightly shorter 

than sepals, base shortly stalked, stalk ciliate. Stamens 8–10 or 

rarely more. Ovary 4-loculed, spiny; stigma 3–5-lobed. Capsule 

globose, ca. 4 mm in diam., spiny, dehiscent into 4 valves; 

spines slender, 5–7 mm, puberulent, curved, tip straight. Seeds 

2 per cell. Fl. summer–autumn. 

Secondary forests, scrub. Fujian, Guangdong, Guangxi, Guizhou, 

Hainan, Xizang, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, 

Myanmar, Nepal, Thailand, Vietnam]. 

2. Triumfetta pilosa Roth, Nov. Pl. Sp. 223. 1821. 

长勾刺蒴麻 chang gou ci shuo ma 

Herbs woody at base, or subshrubs to 1(–1.5) m tall. 

Branchlets yellow-brown stellate tomentose, simple hairs also 

present. Petiole 1–5 cm; leaf blade ovate or narrowly ovate, 3– 

14 × 1.3–4.5 cm, thickly papery, abaxially densely yellowbrown 

stellate tomentose, adaxially sparsely stellate, basal veins 

3, lateral 2 ascending beyond 1/2 blade, base rounded or subcordate, 

margin irregularly serrate, apex acuminate or acute. 

Cymes 1 to several per node; peduncle 5–8 mm. Flower buds 

pilose. Pedicel 3–5 mm. Bracts lanceolate, ca. 1 mm. Sepals 

narrowly lanceolate, 6–10(–12) × 1–1.5 mm, hairy, apex cucullate, 

apical appendage 0.5–1 mm. Petals as long as sepals. Stamens 

10(–30). Ovary hairy. Capsule globose, body 3–5 mm in 

diam., spiny, overall 1.1–1.3 × 1.2–1.4 cm, 4-loculed, dehiscent; 

spines 6–8 mm, hairy, tip hooked. Fl. summer–autumn. 

Dry scrub. Guangdong, Guangxi, Guizhou, Sichuan, Yunnan 

[Bhutan, ?Cambodia, India, ?Laos, Malaysia, Nepal, New Guinea, Sri 

Lanka, Thailand, ?Vietnam; tropical Africa, Australia]. 

3. Triumfetta annua Linnaeus, Syst. Nat., ed. 12, 2: 327; 

Mant. Pl. 1: 73. 1767. 

单毛刺蒴麻 dan mao ci shuo ma 

Herbs annual, or subshrubs. Branchlets yellow-brown tomentose. 

Stipule narrowly triangular; petiole 1–7.5 cm, sparsely 

simple-pilose; leaf blade ovate or ovate-lanceolate, 5–13 × 3–7 

cm, papery, sparsely pilose, both surfaces simple-pilose, basal 

259 

veins 3–5, 2 lateral ones ascending beyond 1/2 blade, base 

rounded or subcordate, margin serrate, apex caudate or acuminate. 

Peduncle very short. Pedicel 3–6 mm, villous. Bracts 2– 

3 mm, villous. Flower bud nearly glabrous. Sepals ca. 5 mm, 

tips appendaged. Petals oblanceolate, slightly shorter than sepals. 

Stamens 10. Ovary 3- or 4-loculed, spiny; style very short; 

stigma 2–3-lobed. Capsule impressed globose, ca. 6 mm in 

diam., spiny, dehiscent, 3- or 4-valved; spines 3–7 mm, glabrous 

or hairy at base, tip hooked. Fl. autumn. 

Open areas, roadsides. Guangdong, Guangxi, Guizhou, Hubei, 

Hunan, Jiangxi, Sichuan, Yunnan, Zhejiang [Bhutan, India, Malaysia, 

Nepal, Pakistan; Africa]. 

Triumfetta annua was incorrectly named as T. suffruticosa Blume 

by some authors, including Merrill; it has also been confused with T. japonica 

Makino, which is restricted to Japan, Korea, and the Philippines. 

4. Triumfetta grandidens Hance, J. Bot. 15: 329. 1877. 

粗齿刺蒴麻 cu chi ci shuo ma 

Herbs woody, creeping or not, many branched; stems pubescent, 

hairs simple, or glabrous. Petiole 5–10 mm, hairy; 

lower leaf blades rhomboid, 3–5-lobed; upper ones oblong, 1– 

2.5 × 0.7–1.5 cm, both surfaces glabrous or abaxially hairy on 

veins, basal veins not strongly ascending, base cuneate, margin 

serrate, apex obtuse. Cymes 10–20 mm; peduncle 5–7 mm. 

Pedicel 2–3 mm. Sepals filiform, ca. 6 mm, abaxially puberulent. 

Petals broadly ovate, slightly shorter than sepals, shortly 

stalked. Stamens 8–10. Ovary 2- or 3-loculed. Capsule globose, 

spiny, indehiscent; spines 2–4 mm, puberulent or glabrous, tip 

shortly hooked. Fl. winter–spring. 

Sandy coasts, coastal wastelands. Guangdong, Hainan [Cambodia, 

Malaysia, Thailand, Vietnam]. 

Triumfetta grandidens has been included within T. repens (Blume) 

Merrill & Rolfe, but the combination of small leaves with relatively 

very large triangular marginal teeth is very distinctive. 

1a. Stems and capsule pubescent or 

puberulent ............................................ 4a. var. grandidens 

1b. Stems and capsule glabrous ........................ 4b. var. glabra 

4a. Triumfetta grandidens var. grandidens 

粗齿刺蒴麻(原变种) cu chi ci shuo ma (yuan bian zhong) 

Triumfetta dunalis O. Kuntze. 

Branchlets and leaves hairy. Stems and capsule pubescent 

or puberulent. 

Guangdong, Hainan [Malaysia, Vietnam]. 

4b. Triumfetta grandidens var. glabra R. H. Miao ex Hung T. 

Chang, Acta Phytotax. Sin. 20: 178. 1982. 

秃刺蒴麻 tu ci shuo ma 

Stems, branchlets, leaves, and capsule glabrous.

260 

● Coastal wastelands. Hainan. 

5. Triumfetta rhomboidea Jacquin, Enum. Syst. Pl. 22. 1760. 

刺蒴麻 ci shuo ma 

Bartramia indica Linnaeus; Triumfetta angulata Lamarck; 

T. bartramii Linnaeus, nom. illeg. superfl.; T. indica Lamarck; 

T. trilocularis Roxburgh; T. velutina Vahl. 

Subshrubs or herbs. Branchlets gray-brown tomentose. 

Petiole 1–5 cm; lower leaf blades broadly ovate-orbicular, 

rhomboid, or broadly ovate, 3-lobed, 3–9.5 × 2–8.5 cm, abaxially 

stellate pilose, adaxially sparsely hairy, basal veins 3–5, 2 

lateral ones reaching tip of lobes, base broadly cuneate or 

rounded, margin irregularly bluntly serrate, apex acute; upper 

leaf blades oblong-lanceolate, not lobed. Cymes 3–5 per axil; 

peduncle to 2 mm. Flower buds cylindrical, apex slightly wider, 

ca. 4 mm. Pedicel less than 1 mm. Sepals narrowly oblong, 4.5– 

5.5 mm, villous, appendaged at tip. Petals yellow, slightly 

shorter than sepals, hairy along margins. Stamens 10. Ovary 

spiny. Capsule globose, ca. 3 mm in diam., spiny, indehiscent; 

spines ca. 2 mm, gray-yellow puberulent, tip hooked. Seeds 2– 

6. Fl. summer–autumn. 2n = 32. 

Forest margins, open areas, wastelands; 100–1500 m. Fujian, 

Guangdong, Guangxi, Taiwan, Yunnan [throughout tropics; type from 

West Indies]. 

Triumfetta indica Lamarck (Encycl. 3: 420. 1792) is not based on 

Bartramia indica Linnaeus (Sp. Pl. 1: 389. 1753) and therefore blocks 

the transfer of B. indica to Triumfetta; hence the correct name for this 

species is T. rhomboidea. 

The whole plant is used medicinally. 

6. Triumfetta semitriloba Jacquin, Enum. Syst. Pl. 22. 1760. 

菲岛刺蒴麻 fei dao ci shuo ma 

Shrubs or perennial herbs, to 1 m or more tall. Branchlets 

pubescent, hairs simple and stellate. Petiole 0.5–6 cm; leaf 

blade broadly ovate, sometimes obscurely 3-lobed, 4–8 × 3–6 

cm, membranous, abaxially grayish green, stellate pubescent, 


adaxially pubescent, hairs simple, basal veins 3, base rounded, 

rarely shallowly cordate, margin irregularly serrate-dentate, 

apex acuminate; upper leaves usually oblong. Cymes 2 or 3 per 

axil; peduncle 1–2(–3) mm. Flower buds cylindrical, ca. 4 mm, 

pubescent. Pedicel 2–3 mm. Sepals linear, ca. 6 mm, abaxially 

stellate pubescent, appendage subapical, ca. 0.5 mm, pubescent. 

Petals narrowly elliptic-obovate, ca. as long as sepals, Stamens 

15 or more, filaments glabrous. Ovary 3-loculed, spiny; style 3– 

5 mm; stigma ± 3-lobed. Capsule globose, body 3–5 mm in 

diam., 3-loculed, indehiscent, glabrescent; spines 1.5–2 mm, 

retrorsely pilosulose at base, conical at base, tip hooked. Seeds 

1 or 2 per cell. Fl. Oct–Jan. 

Fields, wastelands. S Taiwan [Philippines; tropical America]. 

Triumfetta semitriloba is very easily confused with T. rhomboidea. 

7. Triumfetta procumbens G. Forster, Fl. Ins. Austr. 35. 1786. 

铺地刺蒴麻 pu di ci shuo ma 

Shrubs, stems procumbent, to 3 m. Branchlets yellowbrown, 

shortly stellate hairy. Petiole 2–3 mm; leaf blade ovateorbicular, 

sometimes 3-lobed, 2–4.5 × 1.5–4.5 cm, thickly papery 

to slightly fleshy, abaxially densely yellow-brown tomentose, 

adaxially more sparsely shortly stellate tomentose, 

basal veins 5–7, base cordate, margin crenate, apex obtuse. 

Cymes solitary, 2–5-flowered; peduncle 5–10(–35) mm. Pedicel 

2–3(–10) mm. Sepals 9–13 × 2–2.5 mm, abaxially moderately 

densely stellate hairy, adaxially densely stellate tomentose 

near base, tips with subapical appendage terete, 1–2 mm, 

glabrous. Petals linear-obovate, 7.5–9 × ca. 4 mm, claw tomentose. 

Androgynophore ca. 0.5 mm. Stamens 20–35; filaments 

5–8 mm, glabrous. Capsule globose, 4-loculed, 1.5–2 cm in 

diam., spiny, indehiscent, glabrous or with sparse simple and 

stellate hairs; spines robust, 3–5 mm, puberulent, curved at tip. 

Seeds 1 or 2 per cell. Fr. May–Sep. 

Coastal strands near sea level. South China Sea [Japan, Malaysia; 

Australia, Indian Ocean islands (Diego Garcia, Seychelles), SW Pacific 

islands]. 

8. DIPLODISCUS Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 31: 235. Jun–Jul 1858. 

Hainania Merrill; Pityranthe Thwaites (Nov–Dec 1858). 

海南椴属 hai nan duan shu 

Trees or shrubs. Leaves simple, entire, long or shortly petiolate; stipules minute, caducous; leaf blade ovate-orbicular, base 

asymmetrical, usually cordate, basal veins 5–7, margin entire, slightly sinuate, or minutely denticulate distally; foliar nectaries 

absent. Inflorescences terminal, paniculate. Flowers bisexual. Bracts minute, caducous. Involucral bracts absent. Calyx campanulate, 

2–5-lobed, lobes unequal in length. Petals 5, oblanceolate. Stamens 20–30, free or slightly connate in 5 fascicles; filaments long; 

anthers minute, 2-thecate, anther cells not connate; staminodes 5, opposite to petals, lanceolate. Ovary syncarpous, superior, 5loculed; 

ovules up to 5 per locule; style slender; stigma pointed. Fruit a capsule, obovoid, 4–5-angled, loculicidally dehiscent 

[?sometimes indehiscent]. Seeds 1–3 per cell, sometimes reduced to 1 per fruit, glabrous, scaly, or hairy. 

Nine or ten species: China, Borneo, Malaysia, Philippines, Sri Lanka; one species (endemic) in China. 

A close morphological relationship between Diplodiscus, Hainania, and Pityranthe has long been recognized, and various pairs of these three 

genera have been synonymized. Merrill distinguished Hainania from Pityranthe, represented by a single species endemic to Sri Lanka, on the basis of 

Hainania having more numerous ovules and hairy seeds. Diplodiscus was separated from Hainania and Pityranthe by the fruit, which contains only 

one (rarely two) glabrous seeds. Kostermans (Reinwardtia 5: 372. 1961), followed by Meijer and Robyns (Rev. Handbook Fl. Ceylon 7: 428–430.


1991), reduced Pityranthe to synonymy under Diplodiscus, which has priority by about five months. Bayer and Kubitzki (Fam. Gen. Vasc. Pl. 5: 258. 

2003) made Hainania a synonym of Pityranthe and thus also of Diplodiscus. 

1. Diplodiscus trichospermus (Merrill) Y. Tang, M. G. Gilbert 

& Dorr, comb. nov. 

海南椴 hai nan duan 

Basionym: Hainania trichosperma Merrill, Lingnan Sci. J. 

14: 36. 1935; Pityranthe trichosperma (Merrill) Kubitzki. 

Trees to 15 m tall, to 45 cm d.b.h. Bark gray-white, old 

branches dark brown, glabrous; branchlets densely gray-brown 

tomentose. Petiole 2.5–5.5 cm, hairy; leaf blade 6–14 × 4–10 

cm, abaxially densely appressed gray-yellow stellate puberulent, 

adaxially glabrous or nearly so, base subcordate or trun- 

261 

cate, apex acuminate or acute. Panicles to 26 cm; peduncle 

densely gray-yellow stellate puberulent. Pedicel 5–7 mm, hairy. 

Calyx 3–4 mm, abaxially densely yellowish stellate puberulent. 

Petals yellow or white, 6–7 mm, oblanceolate, glabrous. Stamens 

glabrous; staminodes ca. 2.5 mm, apex pointed. Ovary 

ovoid, densely stellate puberulent; style simple. Capsule 2–2.5 

cm, valves densely yellowish stellate puberulent abaxially, 

deeply furrowed and glabrous adaxially. Seeds ellipsoid, ca. 4 

mm, densely yellow-brown villous. Fl. autumn, fr. winter. 

● Open forests; 200–300 m. Guangxi, Hainan. 

9. BERRYA Roxburgh, Pl. Coromandel 3: 60. 1820 [“Berria,” “1819”], nom. cons. 

Espera Willdenow. 

六翅木属 liu chi mu shu 

Trees. Leaves alternate; leaf blade basal veins 5–7, base cordate, margin entire. Inflorescences terminal or axillary, paniculate. 

Flowers many, minute. Calyx campanulate, 3–5-divided. Petals 5, white or pink, spatulate or filiform. Stamens many, free; staminodes 

absent. Ovary 3(or 4)-loculed; stigma 3-lobed. Fruit a capsule, globose, 3(or 4)-valved, loculicidally dehiscent, each valve with 

2 erect wings. Seeds 1 or 2(–4) per cell, villous; endosperm fleshy; cotyledons fallacious. 

Six species: primarily in tropical Asia and Polynesia; one species in China. 

1. Berrya cordifolia (Willdenow) Burret, Notizbl. Bot. Gart. 

Berlin-Dahlem 9: 606. 1926. 

六翅木 liu chi mu 

Espera cordifolia Willdenow, Ges. Naturf. Freunde Berlin 

Neue Schriften 3: 450. 1801; Berrya ammonilla Roxburgh. 

Trees 4–6(–35) m tall, 10–20 cm d.b.h. Stipules filiform, 

1–2 cm, glabrous; petiole 3.5–10 cm; leaf blade ovate to ovateoblong, 

10–20(–25) × 6–8(–14) cm, thinly leathery, very 

sparsely stellate pubescent, soon glabrous, lateral veins 4 or 5 

pairs, base cordate, apex acuminate. Panicles to ca. 20 cm, 

hairy. Pedicel ca. 10 mm. Calyx 3–5-lobed, 3–5 mm, lobes triangular-ovate, 

hairy. Petals white or pink, filiform, 6–8 mm, 

becoming reflexed. Stamens 5–6 mm. Ovary hairy; style short. 

Capsule with persistent calyx, globose, 1–1.3 cm wide, pubescent, 

with 6(or 8) horizontally spreading thin wings, wings 2.5– 

3 × 0.7–1 cm. Seeds ca. 6 × 3 mm, with brown or yellow, caducous 

bristles. 

Taiwan [Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, 

Philippines, Sri Lanka, Thailand, Vietnam]. 

Berrya cordifolia is a valued timber tree in some parts of its range. 

10. EXCENTRODENDRON Hung T. Chang & R. H. Miao, Acta Sci. Nat. Univ. Sunyatseni 

17(3): 21. 1978. 

蚬木属 xian mu shu 

Trees, evergreen. Branchlets and buds glabrous. Leaves alternate, petiolate; leaf blade ovate or oblong, leathery, glabrous, basal 

veins 3, with glands in axils of veins, base cuneate or rounded, margin entire. Inflorescences paniculate. Flowers unisexual (plants 

monoecious or dioecious). Bracts caducous. Male flowers (4 or)5-merous, rarely more. Sepals (4 or)5, oblong, valvate, free or 

slightly connate at base, abaxially pilose, adaxially glabrous, glandular at base. Petals (4 or)5, obovate or oblanceolate, slightly 

shorter than sepals, shortly clawed. Stamens 25–35; filaments filiform, slightly connate at base, in 5 fascicles; anthers 2-loculed, 

basifixed. Ovary 5-loculed; ovules 2 per locule; style 5, very short. Fruit a capsule, ellipsoid or obconic, 5-loculed, loculicidally 

dehiscent, wings 5, thin. Seeds 1 per cell. 

Two species: China, Vietnam; two species (one endemic) in China. 

Many authors follow Zhuge (J. Arnold Arbor. 71: 371. 1990) and do not accept the segregation of Excentrodendron from Burretiodendron (but 

see Gao et al., Acta Phytotax. Sin. 44: 538–550. 2006). 

1a. Capsule 2–3 cm, ellipsoid; leaf blade orbicular-ovate or elliptic-ovate ..................................................................... 1. E. tonkinense 

1b. Capsule 5–5.5 cm, obconic; leaf blade oblong .......................................................................................................... 2. E. obconicum

262 

1. Excentrodendron tonkinense (A. Chevalier) Hung T. 

Chang & R. H. Miao, Acta Sci. Nat. Univ. Sunyatseni 17(3): 

23. 1978. 

蚬木 xian mu 

Pentace tonkinensis A. Chevalier, Bull. Écon, Indochine, 

n.s., 20: 803. 1918; Burretiodendron hsienmu Chun & F. C. 

How; B. tonkinense (A. Chevalier) Kostermans; Excentrodendron 

hsienmu (Chun & F. C. How) Hung T. Chang & R. H. 

Miao; E. rhombifolium Hung T. Chang & R. H. Miao. 

Trees to 40 m. Petiole 3.5–6.5(–10) cm; leaf blade green, 

orbicular-ovate or elliptic-ovate, 8–14(–18) × 5–8(–12) cm, 

leathery, glossy, abaxially yellow-brown fascicled hairy in vein 

axils, lateral basal veins to 1/2 as long as blade, 10–15 mm from 

margin, each with 4 or 5 secondary veins, base rounded, apex 

acuminate or caudate-acuminate. Male inflorescences paniculate, 

7–13-flowered, 5–9 cm; female inflorescences racemelike, 

1–3-flowered. Flowers unisexual. Pedicel not articulate. Sepals 

oblong-lanceolate, 1–1.5 cm, abaxially brown stellate puberulent, 

adaxially glabrous, without glands or a few inner petals 

with 2 globose glands. Petals broadly obovate to spatulate, 8–9 

× 5–6 mm, base obviously clawed. Stamens 25–35; filaments 

4–6 mm; anthers ca. 3 mm. Female flowers unknown. Capsule 


ellipsoid, 2–3 cm. 

Evergreen forests on limestone. Guangxi, SE Yunnan [Vietnam]. 

Excentrodendron tonkinense (as Burretiodendron hsienmu) is included 

in the IUCN Red List as “Vulnerable (B1+2c).” The wood is 

very hard and is prized for making chopping boards. 

2. Excentrodendron obconicum (Chun & F. C. How) Hung T. 

Chang & R. H. Miao, Acta Sci. Nat. Univ. Sunyatseni 17(3): 

24. 1978. 

长蒴蚬木 chang shuo xian mu 

Burretiodendron obconicum Chun & F. C. How, Acta 

Phytotax. Sin. 5: 11. 1956. 

Trees, to 20 m. Petiole 5–6(–8) cm; leaf blade oblong, 

(7–)11–15 × (4–)5–6(–10) cm, leathery, abaxially hairy in vein 

axils, lateral basal veins 1/2 as long as leaf blade, 7–10 mm 

from margin, each with 4 or 5 secondary veins, without obvious 

lateral veins at base, lateral veins 3–4 pairs, base cuneate, apex 

acuminate. Flowers unknown. Infructescences with 1–4 capsules, 

10–11 cm; fruiting pedicel articulate. Capsule narrowly 

obconic, 5–5.5 cm, proximal 1/2 narrow. Seed ca. 1 cm. 

● Evergreen forests on limestone. Guangxi. 

11. BURRETIODENDRON Rehder, J. Arnold Arbor. 17: 47. 1936. 

柄翅果属 bing chi guo shu 

Trees deciduous or semi-evergreen. Buds with scales. Leaves petiolate; stipules caducous; leaf blade cordate, papery or leathery, 

basal veins 5–9, margin minutely denticulate or nearly entire. Flowers unisexual or bisexual (plants polygamo-dioecious or dioecious). 

Male flowers in cymes. Bracts 2–3, caducous. Pedicel articulate. Sepals 5, valvate, free or shortly connate at base, abaxially 

stellate puberulent, adaxially sometimes glandular at base. Petals 5, free, imbricate, slightly longer than sepals, clawed. Androgynophore 

absent. Stamens 25–30; filaments connate at base, in 5 fascicles; anthers oblong, 2-loculed, basifixed, dehiscence longitudinal. 

Rudimentary ovary enclosed in stamens; style short. Female flowers solitary. Fruit a winged capsule, oblong, loculicidally 

dehiscent, base shortly stalked, wings 5, thin. Seed 1 per cell, narrowly obovoid. 

Four species: China, Myanmar, Thailand; two species (one endemic) in China. 

1a. Mature leaves densely stellate puberulent abaxially; sepals glandular at base; staminodes absent ........................... 1. B. esquirolii 

1b. Mature leaves glabrous; sepals not glandular; staminodes 5 ..................................................................................... 2. B. kydiifolium 

1. Burretiodendron esquirolii (H. Léveillé) Rehder, J. Arnold 

Arbor. 17: 48. 1936. 

柄翅果 bing chi guo 

Pentace esquirolii H. Léveillé, Repert. Spec. Nov. Regni 

Veg. 10: 147. 1911; Burretiodendron longistipitatum R. H. 

Miao; Eriolaena esquirolii H. Léveillé. 

Trees deciduous, to 20 m tall. Branchlets gray-brown, 

stellate pubescent. Petiole 2–4 cm; leaf blade elliptic, broadly 

elliptic, or broadly obovate, 9–14 × 6–9 cm, papery, stellate 

puberulent abaxially, stellate puberulent adaxially, densely 

gray-brown, basal veins 5–7, 4 lateral veins with 4–5 secondary 

veins, base obliquely cordate, margin denticulate, apex acute. 

Cymes 3-flowered, as long as petiole. Bracts 2, ovate, ca. 7 mm, 

hairy. Male flowers pedicellate, ca. 2 cm in diam. Sepals oblong, 

ca. 1 × 0.4 cm, stellate tomentose abaxially, glandular at 

base adaxially, glands as long as 1/3 sepals. Petals broadly obovate, 

ca. 11 × 7 mm, base with 3–4 mm claw, apex nearly trun- 

cate. Stamens ca. 30, ca. 7 mm; anthers ca. 2 mm; staminodes 

absent. Infructescence with 1–4 fruits, infructescence stalk ca. 1 

cm, stellate. Stipe slightly shorter than infructescence stalk, not 

articulate, stellate. Capsule ellipsoid, 3.5–4 cm, base rounded; 

stalk 3–4 mm. Seeds obovate, ca. 1 cm. 

Evergreen forests; 100–700 m. Guangxi, SW Guizhou, S and SE 

Yunnan [Myanmar, W Thailand]. 

Burretiodendron esquirolii is included in the IUCN Red List as 

“Vulnerable (B1+2c).” It is relatively common in Guizhou and Yunnan. 

2. Burretiodendron kydiifolium Y. C. Hsu & R. Zhuge, J. 

Arnold Arbor. 71: 378. 1990. 

元江柄翅果 yuan jiang bing chi guo 

Trees deciduous or semi-evergreen, to 15 m tall. Branchlets 

densely brown stellate scaly. Petiole 3.5–10 cm; leaf blade 

suborbicular, sometimes 3-lobed, 7–15 × 7–13 cm, papery, 

glabrous, basal veins 7–9, lateral veins 3 or 4 pairs, base cor-

date, margin entire, apex acute. Flowers unisexual. Male flowers 

in 3–7-flowered cymes. Female flowers solitary or rarely in 

2- or 3-flowered cymes. Bracteoles 3, large, enveloping flower 

bud. Sepals eglandular. Petals flabelliform, apex praemorse, not 

glandular adaxially. Stamens 25–30, without pollen in ap- 


263 

parently bisexual flowers; staminodes 5, longer than stamens. 

Ovary 5-loculed, 5-angled; styles 5, clavate. Capsule ellipsoid, 

3–4 cm. Fl. Apr–May, fr. Jun–Jul. 

● Sparse forests in dry valleys; 400–900 m. S Yunnan.

TILIACEAE 1. TILIA Linnaeus, Sp. Pl. 1: 514. 1753.

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