RUTACEAE

RUTACEAE 

芸香科 yun xiang ke 

Zhang Dianxiang (张奠湘) 1 ; Thomas G. Hartley 2 , David J. Mabberley 3 

Shrubs, trees, or sometimes herbs, sometimes scrambling or scandent, sometimes armed, with aromatic volatile oils contained in 

glands visible at surface of at least leaves, young branchlets, inflorescences, flower parts, fruit, or cotyledons in seed. Stipules absent 

[or stipular excrescences rarely present]. Leaves alternate, opposite [or whorled], simple (petiole neither apically swollen nor articulate 

with leaf blade), 1-foliolate (in individual specimens at least some 1-foliolate leaves with petiole apically swollen and/or articulate with 

leaf blade), or variously compound. Flowers bisexual or unisexual, usually 3–5-merous, actinomorphic or rarely zygomorphic, hypogynous 

[or rarely perigynous]. Perianth in 2 series, with clearly differentiated calyx and corolla or sometimes in 2 irregular series or 1 

series, with ± undifferentiated tepals. Sepals distinct or connate to their full length. Petals distinct [or rarely coherent or connate for part 

of their length]. Stamens usually as many as or 2 × as many as petals or sometimes more numerous; filaments distinct or sometimes 

coherent or connate for at least part of their length; anthers introrse or sometimes latrorse, longitudinally dehiscent. Disk [rarely lacking] 

within androecium, nectariferous, flattened, annular, cup-shaped, pulvinate, or sometimes columnar, bell-shaped, conic, or hourglass-shaped. 

Gynoecium of 1–5 distinct 1-loculed carpels or 2 to many partially to completely connate carpels; placentation axile 

[very rarely becoming parietal]; ovules 1 to many per locule. Fruit of 2–5 follicles [drupes or samaras] or a single follicle, capsule, or 

berry [or samara]. Seeds with relatively large embryo; endosperm present and fleshy or lacking. 

About 155 genera and ca. 1600 species: nearly cosmopolitan but mainly tropical and subtropical; 22 genera (one endemic, one introduced) and 

126 species and hybrid species (49 endemic, at least two introduced) in China. 

Oil glands of Rutaceae, when viewed from the surface of plant parts they occupy, are usually pellucid. They also appear to be ± isodiametric and to 

have ± definite patterns of distribution. In blades of leaves, for example, where they are most commonly observed, they are usually ± evenly scattered 

throughout, or sometimes they are restricted to the margins. Rarely they are alleged to occur only along the secondary veins of the blades. 

In a cladistic analysis of selected genera of Rutaceae and related families based on rbcL and atpB molecular data (but only on rbcL data for 

Harrisonia), M. W. Chase, C. M. Morton, and J. A. Kallunki (Amer. J. Bot. 86: 1191–1199. 1999) recommended the placement of Harrisonia 

(traditionally Simaroubaceae) in Rutaceae. We do not agree with this classification, particularly because Harrisonia appears to lack oil glands, and 

suggest that the genus is most correctly placed in Cneoraceae, in which it is treated herein. 

Several taxa of cultivated Rutaceae treated by C. C. Huang (Fl. Reipubl. Popularis Sin. 43(3). 1997) are not treated here: Casimiroa edulis La 

Llave, which is native to Mexico, has been introduced as a cultivated plant to the Xishuangbanna Botanical Garden in Yunnan; Limonia acidissima 

Linnaeus (Feronia limonia (Linnaeus) Swingle), which is native to India and Sri Lanka, has been reported as cultivated in Taiwan by C. C. Huang (Fl. 

Reipubl. Popularis Sin. 44(3): 212. 1997) but was not reported as a cultivated plant by T. C. Huang in A Checklist of the Vascular Plants of Taiwan (Fl. 

Taiwan 6: 81–83. 1979); Flindersia amboinensis Poiret, which is native to Indonesia and New Guinea, is cultivated in S China; Ptelea trifoliata 

Linnaeus, which is native to North America, is cultivated in Beijing and Liaoning; and Ruta graveolens Linnaeus, which is native to the Mediterranean 

region, is widely cultivated in China. 

Huang Chengchiu. 1997. Rutaceae. In: Huang Chengchiu, ed., Fl. Reipubl. Popularis Sin. 43(2): ii–viii, 1–250. 

1a. Leaves opposite. 

2a. Leaves mostly odd-pinnate; inflorescences terminal or terminal and axillary; plants dioecious. 

3a. Axillary buds exposed; ovary in female flowers with carpels connate at base, otherwise contiguous; 

fruit of 1–5 follicles ............................................................................................................................................ 3. Tetradium 

3b. Axillary buds concealed in excavated base of petioles; ovary in female flowers syncarpous; fruit a 

drupaceous berry ....................................................................................................................................... 10. Phellodendron 

2b. Leaves digitately 3-foliolate or 1-foliolate; inflorescences axillary or basal to leaves; plants usually 

monoclinous or dioecious. 

4a. Leaves digitately 3-foliolate or 1-foliolate; ovary in bisexual and female flowers with carpels connate 

at base, otherwise contiguous; fruit of 1–4 basally connate follicles .................................................................. 4. Melicope 

4b. Leaves 1-foliolate; ovary in bisexual and female flowers syncarpous or rarely (Acronychia) separated 

apically by septicidal fissures; fruit a drupaceous berry. 

5a. Flowers male or female; stamens glabrous, similar in male and female flowers but lacking pollen 

in latter; gynoecium rudimentary and minute in male flowers, ± as long as petals in female 

flowers; seed coat with thick inner layer of dense black sclerenchyma and spongy outer 

layer bounded externally by a shiny black pellicle ......................................................................... 12. Maclurodendron 

5b. Flowers bisexual; staminal filaments with an indumentum; gynoecium ± half as long as petals; 

seed coat with thick inner layer of dense black sclerenchyma and an outer layer of compact 

dull to ± shiny parenchymatous tissue ..................................................................................................... 11. Acronychia 

1 South China Botanical Garden, Chinese Academy of Sciences, Longdong, Shahe, Guangzhou, Guangdong 510650, People’s Republic of China. 

2 Centre for Plant Biodiversity Research, CSIRO, G.P.O. Box 1600, Canberra, A.C.T. 2601, Australia. 

3 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom. 

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1b. Leaves alternate. 

6a. Plants herbaceous perennials. 

7a. Leaves mostly simple ................................................................................................................................... 6. Haplophyllum 

7b. Leaves mostly compound. 

8a. Flowers zygomorphic, 5-merous; stamens 10; fruit of 5 basally connate follicles, 1–2 cm ...................... 8. Dictamnus 

8b. Flowers actinomorphic, 4-merous; stamens usually 8; fruit follicular or capsular, to 0.5 cm. 

9a. Leaves pinnately to ternately decompound; fruit of 4 distinct follicles .................................. 5. Boenninghausenia 

9b. Leaves mostly digitately 3-foliolate; fruit a capsule ...................................................................... 7. Psilopeganum 

6b. Plants woody. 

10a. Fruit follicular or drupaceous; endocarp cartilaginous or leathery; seeds with endosperm; flowers 

male, female, or sometimes bisexual; functional stamens as many as petals except in plants of 

Zanthoxylum with undifferentiated perianth which have 5–9 tepals and 3–8(–10) functional stamens; 

functional gynoecium 1–7-loculed, syncarpous or with distinct or only basally connate carpels. 

11a. Leaves mostly compound; plants usually armed. 

12a. Leaves odd-pinnately 3- to many foliolate or sometimes digitately 3-foliolate (occasional 

leaves even-pinnate, 2-foliolate, or 1-foliolate); functional gynoecium 1–5-loculed, 

with distinct carpels or carpels basally connate; fruit of 1–5 distinct or basally connate 

follicles ...................................................................................................................................... 1. Zanthoxylum 

12b. Leaves digitately 3-foliolate (occasional leaves 1- or 2-foliolate); functional gynoecium 

4–7-loculed, syncarpous; fruit a 4–7-loculed drupaceous berry ..................................................... 9. Toddalia 

11b. Leaves simple; plants unarmed. 

13a. Inflorescences axillary, between leaves, or basal to leaves; flowers male or female; functional 

gynoecium 4-loculed; ovaries basally connate; fruit of 1–4 basally connate follicles ........................ 2. Orixa 

13b. Inflorescences terminal; flowers male, female, or sometimes bisexual; functional gynoecium 

2–5-loculed, syncarpous; fruit drupaceous berry, with 1–5 1-seeded leathery pyrenes ............... 13. Skimmia 

10b. Fruit baccate; endocarp membranous or fleshy; seeds without endosperm; flowers bisexual or bisexual 

and male; stamens at least 2 × as many as petals or rarely fewer; functional gynoecium 2- to many 

loculed, syncarpous. 

14a. Flowers bisexual or bisexual and male; stamens more than 2 × as many as petals; gynoecium 2- to 

many loculed; ovules 2 to many per locule; fruit with stalked or rarely sessile pulp vesicles; 

leaves odd-pinnately 3(or 5)-foliolate, digitately 3-foliolate, 1-foliolate, or simple. 

15a. Plants evergreen, rarely deciduous (Citrus trifoliata); fruit with leathery (or rarely soft) 

exocarp and spongy mesocarp; seeds embedded in pulp vesicles; leaves 1-foliolate, 

simple, or rarely digitately 3-foliolate ............................................................................................... 21. Citrus 

15b. Plants deciduous; fruit with thin, parenchymatous exocarp and woody mesocarp; seeds 

embedded in a clear glutinous substance; leaves odd-pinnately 3(or 5)-foliolate ............................. 22. Aegle 

14b. Flowers bisexual; stamens 2 × as many as petals or rarely fewer; gynoecium 2–5-loculed; ovules 1 

or 2 per locule; fruit either without pulp vesicles or with sessile pulp vesicles; leaves odd-pinnately 

3- to many foliolate, digitately 3-foliolate, 1-foliolate, or simple (occasional leaves 2-foliolate). 

16a. Radial walls of locules in gynoecium becoming curved after anthesis; cotyledons in seed 

thin and flat, convolute, and/or folded ..................................................................................... 14. Micromelum 

16b. Radial walls of locules in gynoecium straight; cotyledons in seed plano-convex, neither 

convolute nor folded. 

17a. Terminal and axillary buds and usually young inflorescences with a rust-colored 

villosulous indumentum (sometimes partly bleached); style persistent in fruit ............... 15. Glycosmis 

17b. Terminal and axillary buds and young inflorescences without a rust-colored villosulous 

indumentum; style or at least its distal portion usually deciduous in fruit. 

18a. Leaves 1-foliolate or simple. 

19a. Fruit without pulp vesicles; stamens distinct; flowers axillary, solitary or 

in few-flowered fascicles; petiole 0.4–2.5 cm, usually bent and/or twisted 

and ± apically swollen; erect or scrambling shrubs or woody climbers, 

usually armed ............................................................................................... 19. Paramignya 

19b. Fruit usually with pulp vesicles; stamens distinct or with filaments variously 

connate or coherent; flowers in terminal and/or axillary inflorescences, in 

fascicles, racemes, or panicles; petiole 0.2–1.3 cm, usually not bent, twisted, 

or swollen; shrubs or trees, armed or unarmed ................................................ 20. Atalantia 

18b. Leaves, or most of them, compound. 

20a. Leaves mostly digitately 3-foliolate; woody climbers, armed .......................... 18. Luvunga 

20b. Leaves odd-pinnate; shrubs or trees, unarmed.


21a. Flower buds globose, pyriform, or rarely ovoid; staminal filaments ± 

abruptly dilated toward base, ± straight or geniculate; style 0.5–2.5 × as 

long as ovary; disk columnar, conic, bell-shaped, or ± hourglass-shaped; 

seeds glabrous .......................................................................................... 16. Clausena 

21b. Flower buds ellipsoid to obovoid or narrowly so or subcylindric; staminal 

filaments linear or sublinear, ± straight; style 3–7 × as long as ovary; disk 

annular, pulvinate, or columnar; seeds glabrous or villous ..................... 17. Murraya 

1. ZANTHOXYLUM Linnaeus, Sp. Pl. 1: 270. 1753. 

花椒属 hua jiao shu 

Zhang Dianxiang (张奠湘); Thomas G. Hartley 

Shrubs sometimes scrambling, trees, or woody climbers, evergreen or deciduous, dioecious, rarely monoecious or polygamo- 

monoecious [or monoclinous], usually armed [or exclusively unarmed]. Leaves alternate, odd-pinnately 3- to many foliolate or 

sometimes digitately 3-foliolate (occasional leaves even-pinnate, 2-foliolate, or 1-foliolate). Inflorescences terminal, axillary, or basal 

to leaves, paniculate, thyrsiform, corymbiform, racemose, or umbelliform. Perianth in 2 series and differentiated with 4 or 5 sepals and 

4 or 5 petals or grading to 1 series and undifferentiated with 5–9 tepals. Sepals distinct or basally connate. Petals valvate or imbricate in 

bud. Stamens distinct, 4 or 5 in plants with sepals and petals, 3–8(–10) in plants with tepals, rudimentary or lacking in female flowers. 

Disk flattened, pulvinate, or columnar. Gynoecium 1–5-carpelled, rudimentary or lacking in male flowers; ovaries basally connate, 

otherwise ± contiguous or distinct, 1-loculed; ovules 2 per locule. Styles in compound gynoecium apical or subapical, coherent or 

contiguous to spreading-ascending or recurved; stigmas capitate, coherent or distinct. Style in simple gynoecium off-centered, erect or 

variously curved or bent; stigma capitate to peltate. Fruit follicles 1–5, distinct or basally connate, apex often with a stylar beak; 

abortive carpels, if any, often persistent. Seeds globose to ovoid, persistent in dehisced fruit; seed coat with a thick or rarely ± thin inner 

layer of dense black sclerenchyma and spongy-fleshy outer layer bounded externally by a shiny black or reddish pellicle; endosperm 

copious or rarely scant; embryo straight or ± curved; cotyledons ± orbicular to broadly elliptic, flattened or rarely plano-convex; 

hypocotyl superior. 

Two hundred or more species: pantropical and extending to temperate latitudes in E Asia and E North America; 41 species (25 endemic) in China. 

1a. Perianth in 2 irregular series or 1 series, with 5–9 ± undifferentiated tepals; stamens in male flowers 4–8(–10). 

2a. Gynoecium in female flowers 1(or 2)-carpelled and leaf rachises not winged but canaliculate with sides of 

channel rarely slightly spreading at apex of rachis ................................................................................................. 32. Z. wutaiense 

2b. Gynoecium in female flowers 2–5-carpelled or if 1-carpelled then leaf rachises with wing to 6 mm wide on 

each side. 

3a. Fruit follicles stipitate. 

4a. Leaflet blades 0.7–1.3 cm wide ........................................................................................................................ 41. Z. stipitatum 

4b. Leaflet blades usually at least 1.5 cm wide. 

5a. Leaves 3–5(–11)-foliolate; follicles apically shortly beaked .......................................................... 29. Z. dimorphophyllum 

5b. Leaves 5–15-foliolate; follicles not apically beaked ...................................................................................... 40. Z. simulans 

3b. Fruit follicles not stipitate. 

6a. Leaf rachises usually conspicuously winged. 

7a. Wings of leaf rachis extending to as much as 3 mm on each side; secondary veins of leaflet blades 

evident, 10–28 on each side of midvein; anthers in male flowers reddish purple prior to anthesis; 

gynoecium in female flowers 2–5-carpelled ....................................................................................... 30. Z. acanthopodium 

7b. Wings of leaf rachis extending to as much as 6 mm on each side; secondary veins of leaflet blades 

generally faint especially adaxially, 7–15 on each side of midvein; anthers in male flowers yellow 

prior to anthesis; gynoecium in female flowers 1–3-carpelled ..................................................................... 31. Z. armatum 

6b. Leaf rachises not conspicuously winged but sometimes with a distinct margin. 

8a. Fruit follicles pubescent; branchlets covered with setiform prickles and compressed pseudostipular 

prickles decurrent along branchlet and forming a wing joining them between nodes ......................... 39. Z. pteracanthum 

8b. Fruit follicles glabrous; branchlets without compressed pseudostipular prickles joined by a wing. 

9a. Leaflet blades to 3 × 1.5 cm. 

10a. Leaflet blades reddish brown to blackish brown when dry .................................................................. 37. Z. piasezkii 

10b. Leaflet blades grayish green to yellowish green when dry ................................................................. 38. Z. pilosulum 

9b. Leaflet blades usually larger than 3 × 1.5 cm. 

11a. Leaf near inflorescence on fertile branchlet 3-foliolate, with terminal leaflet 4–6 cm wide ............ 36. Z. motuoense 

11b. Leaf near inflorescence on fertile branchlet more than 5-foliolate, with terminal leaflet less 

than 3.5 cm wide. 

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12a. Leaf rachises terete, glabrous ..................................................................................................... 35. Z. austrosinense 

12b. Leaf rachises with a distinct margin or pubescent. 

13a. Leaflet blades abaxially flocculent along midvein and adaxially glabrous or both surfaces 

pubescent .................................................................................................................................... 33. Z. bungeanum 

13b. Leaflet blades abaxially glabrous and adaxially hirsutulous to sparsely puberulent ........... 34. Z. undulatifolium 

1b. Perianth in 2 series, with 4 or 5 sepals and 4 or 5 petals; stamens in male flowers 4 or 5. 

14a. Flowers 5-merous; trees or rarely shrubs. 

15a. Shrubs; leaflet blades rarely wider than 2 cm, adaxially with trichomes and abaxially glabrous ............... 28. Z. schinifolium 

15b. Trees; leaflet blades wider than 2 cm, adaxially glabrous or both surfaces with trichomes. 

16a. Fertile branchlets unarmed, with small pith; leaf rachises winged. 

17a. Gynoecium in female flowers 2(or 3)-carpelled; leaflet blades asymmetric, oblique at base .................. 23. Z. avicennae 

17b. Gynoecium in female flowers 3(or 4)-carpelled; leaflet blades at base rounded to broadly cuneate, 

symmetric or asymmetric ....................................................................................................................... 24. Z. micranthum 

16b. Fertile branchlets armed, with large pith; leaf rachises not winged. 

18a. Leaflet blades glabrous. 

19a. Leaflet blades dark green to light yellowish green when dry, abaxially glaucous ............................ 25. Z. ailanthoides 

19b. Leaflet blades reddish brown to blackish brown, abaxially not glaucous ...................................... 26. Z. myriacanthum 

18b. Leaflet blades abaxially and/or adaxially with trichomes. 

20a. Leaflet blades abaxially and adaxially with trichomes .................................................................... 26. Z. myriacanthum 

20b. Leaflet blades abaxially with trichomes, otherwise glabrous. 

21a. Leaflet blades abaxially with a soft woolly villous indumentum .............................................................. 27. Z. molle 

21b. Leaflet blades abaxially pubescent .................................................................................................. 25. Z. ailanthoides 

14b. Flowers 4-merous; woody climbers, shrubs, or rarely trees. 

22a. Gynoecium in female flowers 2- or 3-carpelled and styles usually recurved; follicles stipitate ........ 29. Z. dimorphophyllum 

22b. Gynoecium in female flowers 1-carpelled or if 2–4-carpelled then styles usually erect to 

spreading-ascending; follicles not stipitate. 

23a. Inflorescences terminal, cymose-corymbiform; pedicel at least 1 cm in fruit and purplish red like follicles. 

24a. Fruit pedicel 1–1.5 cm, ca. 1 mm wide; leaflet blades with numerous oil glands, midvein adaxially 

impressed and puberulent, margin of blade serrulate ............................................................................. 20. Z. oxyphyllum 

24b. Fruit pedicel 1.5–4.5 cm, less than 1 mm wide; leaflet blades with inconspicuous oil glands, midvein 

adaxially ridged, plane, or impressed, margin of blade crenulate or entire toward base. 

25a. Midvein on adaxial surface of leaflet blades ridged or plane toward apex, petiolules pubescent 

adaxially ............................................................................................................................................ 21. Z. stenophyllum 

25b. Midvein on adaxial surface of leaflet blades impressed, petiolules glabrous ......................................... 22. Z. esquirolii 

23b. Inflorescences axillary and terminal, thyrsiform; pedicel rarely to 1 cm in fruit and not purplish red. 

26a. Fruit follicles with prickles and/or trichomes. 

27a. Fruit follicles with prickles and trichomes ...................................................................................... 19. Z. echinocarpum 

27b. Fruit follicles with trichomes, without prickles. 

28a. Leaflets opposite; leaflet blades 7–19 × 5–8 cm ................................................................................. 17. Z. collinsiae 

28b. Leaflets alternate to opposite; leaflet blades 6–8 × 2.5–3.5 cm ............................................................. 18. Z. liboense 

26b. Fruit follicles with neither prickles nor trichomes. 

29a. Fruit follicles 1–1.5 cm; outer part of pericarp (exocarp and mesocarp) wider than endocarp ................ 16. Z. dissitum 

29b. Fruit follicles to 0.9 cm; outer part of pericarp not wider than endocarp. 

30a. Leaflets opposite. 

31a. Trees; leaflet blades asymmetric ................................................................................................... 3. Z. integrifolium 

31b. Woody climbers or shrubs; leaflet blades symmetric. 

32a. Leaflet blades abaxially without oil glands along secondary veins, apex retuse at tip; petiolules 

obsolete or to 5 mm ............................................................................................................................. 1. Z. nitidum 

32b. Leaflet blades abaxially with oil glands along secondary veins, apex not retuse at tip; 

petiolules ca. 1 mm ....................................................................................................................... 2. Z. xichouense 

30b. Leaflets alternate or partly opposite. 

33a. Leaflet blades abaxially, leaf rachises, and inflorescences tomentulose ....................................... 4. Z. tomentellum 

33b. Leaflet blades abaxially, leaf rachises, and inflorescences glabrous or puberulent. 

34a. Leaves with numerous oil glands which project outward when dry. 

35a. Fruit follicles not apically beaked; leaflet blade oil glands light yellow to brown when dry, 

pellucid ..................................................................................................................................... 5. Z. glomeratum 

35b. Fruit follicles usually apically beaked; leaflet blade oil glands brown or blades blackish 

brown when dry. 

36a. Leaves 3- or 7(or 9)-foliolate; leaflet blades at least 5 cm, lustrous when dry ................... 6. Z. macranthum


36b. Leaves 7–51-foliolate; leaflet blades rarely to 5 cm, dull when dry. 

37a. Leaves 7–15-foliolate; leaflet blades symmetric or subsymmetric, midvein 

adaxially impressed ................................................................................................... 7. Z. rhombifoliolatum 

37b. Leaves 17–51-foliolate; leaflet blades asymmetric, midvein adaxially plane 

to ridged ............................................................................................................................. 13. Z. multijugum 

34b. Leaves with few oil glands or oil glands inconspicuous. 

38a. Leaf rachises with straight prickles 5–10 mm ............................................................................. 8. Z. leiboicum 

38b. Leaf rachises unarmed or with curved prickles rarely to 5 mm. 

39a. Fruit follicles 7–8 mm; margin of leaflet blades entire ................................................................ 9. Z. laetum 

39b. Fruit follicles 5–6 mm; margin of leaflet blades crenate at least near apex or rarely entire. 

40a. Pedicel of fruit and rachis of infructescences glabrous or sparsely puberulent .................. 10. Z. scandens 

40b. Pedicel of fruit and rachis of infructescences puberulent or pubescent. 

41a. Leaflet blades abaxially pubescent at least on midvein toward base or secondary 

veins ............................................................................................................................. 11. Z. kwangsiense 

41b. Leaflet blades abaxially glabrous. 

42a. Fruit pedicel 7–10 mm; petiolules 4–10 mm, leaflet blade apices caudate ............... 12. Z. khasianum 

42b. Fruit pedicel rarely more than 5 mm; petiolules 1–4 mm, leaflet blade apices acute, 

shortly acuminate, or shortly caudate. 

43a. Leaflet blades 2–5 × 0.7–2.5 cm, apex acute or shortly acuminate; inflorescences 

to 3 cm; innovations glaucous .................................................................................... 14. Z. calcicola 

43b. Leaflet blades 6–10 × 2.5–4 cm, apex shortly caudate; inflorescences to 

26 cm; innovations not glaucous ....................................................................... 15. Z. yuanjiangense 

1. Zanthoxylum nitidum (Roxburgh) Candolle, Prodr. 1: 727. 

1824. 

两面针 liang mian zhen 

Shrubs, erect or scrambling, or sometimes woody climbers. 

Trunk winged. Stems, branchlets, and leaf rachises usually with 

prickles. Rachis of inflorescences and abaxial surface of leaflet 

blades glabrous or hirsutulous. Leaves (3 or)5–11-foliolate; 

petiolules obsolete or to 5 mm; leaflet blades opposite, broadly 

ovate, subcordate, elliptic, narrowly elliptic, or rarely ovate, 

3–12 × 1.5–6(–8) cm, leathery, secondary and tertiary veins 

abaxially ridged when dry, margin crenate at least toward apex 

or entire, apex acuminate to caudate with a retuse tip. Inflorescences 

axillary. Flowers 4-merous. Perianth in 2 series. Sepals 

ca. 1 mm wide, apically purplish green. Petals pale yellowish 

green, ovate-elliptic to oblong, ca. 3 mm. Stamens in 

male flowers 5–6 mm; mature anthers ellipsoid to globose. 

Petals in female flowers broader than those in male flowers. 

Gynoecium in female flowers 4-carpelled; carpels globose. 

Styles stout, coherent at anthesis; stigma capitate. Fruit pedicel 

2–5 mm; follicles reddish brown, (5–)5.5–7 mm in diam., apex 

beaked. Fl. Mar–May, fr. Sep–Nov or May. 

Below 800 m. Fujian, Guangdong, Guangxi, S and SE Guizhou, 

Hainan, S Hunan, Taiwan, Yunnan, S Zhejiang [India, Indonesia, Japan 

(Ryukyu Islands), Malaysia, Myanmar, Nepal, New Guinea, Philippines, 

Thailand, Vietnam; Australia, SW Pacific islands]. 

1a. Rachis of inflorescences and abaxial surface 

of leaflet blades glabrous ........................... 1a. var. nitidum 

1b. Rachis of inflorescences and abaxial 

surface of leaflet blades hirsutulous .. 1b. var. tomentosum 

1a. Zanthoxylum nitidum var. nitidum 

两面针(原变种) liang mian zhen (yuan bian zhong) 

Fagara nitida Roxburgh, Fl. Ind. 1: 439. 1820; F. hamiltoniana 

(Wallich ex J. D. Hooker) Engler; F. hirtella (Ridley) 

Engler; F. oblongifolia Bakhuizen f.; F. pendjaluensis Bakhuizen 

f.; F. torva (F. Mueller) Engler; F. warburgii Perkins; Zanthoxylum 

asperum C. C. Huang var. glabrum C. C. Huang; Z. 

hamiltonianum Wallich ex J. D. Hooker; Z. hirtellum Ridley; Z. 

torvum F. Mueller. 

Rachis of inflorescences and abaxial surface of leaflet 

blades glabrous. Petiolules 2–5 mm or leaflets subsessile; leaflet 

blades broadly ovate, subcordate, or narrowly elliptic, 1.5–6 cm 

wide, margin crenate or entire, apex caudate with a retuse tip. 

Fruit follicles 5.5–7 mm in diam. Fr. Sep–Nov. 

Below 800 m. Fujian, Guangdong, Guangxi, S and SE Guizhou, 

Hainan, S Hunan, Taiwan, Yunnan, S Zhejiang [India, Indonesia, Japan 

(Ryukyu Islands), Malaysia, Myanmar, Nepal, New Guinea, Philippines, 

Thailand, Vietnam; Australia, SW Pacific islands]. 

1b. Zanthoxylum nitidum var. tomentosum C. C. Huang, 

Guihaia 7: 5. 1987. 

毛叶两面针 mao ye liang mian zhen 

Rachis of inflorescences and abaxial surface of leaflet 

blades hirsutulous. Petiolules 1–3 mm; leaflet blades elliptic or 

rarely ovate, 3–5(–8) cm wide, margin revolute and entire or 

apically crenate, apex acuminate. Fruit follicles ca. 5 mm in 

diam. Fr. May. 

● Hillside thickets. E Guangxi (Pingnan). 

2. Zanthoxylum xichouense C. C. Huang, Acta Phytotax. Sin. 

16(2): 83. 1978. 

西畴花椒 xi chou hua jiao 

Woody climbers, glabrous except for inflorescence 

rachises. Branchlets dark reddish purple when young, unarmed 

or spinose. Leaves 3–7-foliolate; petiolules ca. 1 mm; leaflet 

blades opposite, elliptic to obovate, 5–10 × 3–5 cm, thinly 

papery, oil glands inconspicuous, reticulate veinlets ridged when 

dry, base broadly cuneate, margin entire, apex acuminate. In- 

55

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fructescences axillary. Pedicel 4–7 mm in fruit. Fruit follicles 

crimson, globose, ca. 5.5 mm in diam., apex beaked; exocarp, 

mesocarp, and endocarp thin. Seeds 4–4.5 mm in diam. Fr. Oct. 

● Forests; 1400–1500 m. SE Yunnan (Xichou). 

3. Zanthoxylum integrifolium (Merrill) Merrill, Enum. Philipp. 

Fl. Pl. 2: 327. 1923. 

兰屿花椒 lan yu hua jiao 

Fagara integrifolia Merrill, Philipp. J. Sci. 1(Suppl. 1): 68. 

1906. 

Trees 13–20 m tall. Branchlets prickly. Leaves (13–)17– 

25-foliolate; rachis unarmed; petiolules 5–7 mm; leaflet blades 

opposite, oblong to obovate, 1.5–2 × 0.6–0.8 cm, papery, glabrous, 

adaxially lustrous when dry, oil glands inconspicuous, 

midvein adaxially impressed, secondary veins adaxially ridged, 

base oblique, margin entire, apex cuspidate with a blunt tip. 

Inflorescences terminal or axillary, to 25 cm. Flowers 4-merous. 

Perianth in 2 series. Petals white, ca. 3 mm. Male flowers: stamens 

4; rudimentary gynoecium shortly conic. Female flowers 

1-carpelled. Fruit pedicel 3–5 mm; follicle single, ca. 7 mm in 

diam. Seeds ca. 6 mm in diam. 

Taiwan (Lan Yu) [Philippines]. 

4. Zanthoxylum tomentellum J. D. Hooker, Fl. Brit. India 1: 

493. 1875. 

毡毛花椒 zhan mao hua jiao 

Fagara tomentella (J. D. Hooker) Handel-Mazzetti; F. 

tomentella var. mekongensis Handel-Mazzetti. 

Woody climbers. Young branchlets, leaf rachises, petiolules, 

abaxial surface of leaflet blades, and rachis of infructescences 

grayish yellow to grayish brown tomentulose. Leaves 

7–15-foliolate; petiolules to 5 mm, thick; leaflet blades alternate, 

ovate to elliptic, 4–8 × 2–4 cm, leathery, midvein adaxially plane 

or ridged, base obliquely rounded, margin entire or apically 

crenate, apex mucronate to cuspidate. Inflorescences axillary. 

Flowers 4-merous. Perianth in 2 series. Petals pale yellowish 

white. Male flowers: stamens 4, longer than petals; connective 

with an oil gland at apex; rudimentary gynoecium shorter than 

petals. Female flowers: carpels globose. Infructescences 3–12 

cm. Fruit pedicel 2–5 mm, thick; follicles reddish brown, 

becoming brownish black when dry, 5–6 mm in diam., oil glands 

slightly protruding, apex beaked. Seeds ca. 5 mm in diam. Fl. 

Apr–May, fr. Sep–Oct. 2n = 72. 

Moist valleys; 2000–3000 m. NW and W Yunnan [Bhutan, NE 

India, Myanmar, Nepal]. 

5. Zanthoxylum glomeratum C. C. Huang, Acta Phytotax. Sin. 

16(2): 82. 1978. 

密果花椒 mi guo hua jiao 

Shrubs. Branchlets and leaves glabrous, unarmed or with 

prickles. Leaves 5–9-foliolate; petiolules 4–8 mm; leaflet blades 

alternate, lanceolate, long elliptic, or rarely ovate, 6–12 × 2.5–5 

cm, thickly papery, oil glands numerous, reticulate veinlets 

abaxially inconspicuous and adaxially ridged, margin entire, 


apex acuminate to caudate with a rounded to retuse tip. Inflorescences 

axillary, thyrsiform. Female inflorescences 2–6 cm. 

Flowers 4-merous. Pedicel 1–2 mm. Perianth in 2 series. Petals 

pale yellowish green, ovate-elliptic, ca. 3 mm. Infructescences 

globose to conic. Fruit pedicel 5–8 mm; follicles pale yellow to 

brownish yellow when dry, ca. 7 mm, oil glands impressed, apex 

not beaked. Seeds dark brown, ca. 6 mm in diam. Fl. Apr, fr. 

Sep. 

● Upland forests and thickets; ca. 1500 m. Guangxi (Damiao 

Shan), SE Guizhou (Rongjiang). 

6. Zanthoxylum macranthum (Handel-Mazzetti) C. C. Huang, 

Acta Phytotax. Sin. 6: 70. 1957. 

大花花椒 da hua hua jiao 

Fagara macrantha Handel-Mazzetti, Sinensia 5: 17. 1934. 

Woody climbers, unarmed or echinate on abaxial surface of 

leaf rachis. Branchlets dark gray, rugose. Leaves 3- or 7(or 

9)-foliolate; leaflet blades opposite, ovate, elliptic, or oblanceolate, 

5–10 × 1.5–4 cm, thickly leathery, abaxially lustrous when 

dry, oil glands visible and slightly protruding when dry, midvein 

impressed and puberulent, base symmetric or rarely oblique, 

margin entire or crenate and becoming revolute. Inflorescences 

axillary, thyrsiform. Female flowers subsessile or with a pedicel 

to 1 mm, male flowers with a longer pedicel. Flowers 4-merous. 

Perianth in 2 series. Sepals purplish green. Petals pale yellowish 

green, broadly elliptic, ca. 3 mm. Male flowers: filaments ca. 4 

mm; rudimentary gynoecium shortly club-shaped, apically 

2-lobed. Female flowers: carpels 4. Infructescences 3–5 cm. 

Fruit pedicel 2–3 mm; follicles reddish brown, 5.6–6 mm in 

diam., with a flange, oil glands impressed, apex beaked. Seeds 

ca. 5 mm in diam. Fl. Apr–May, fr. Aug–Sep. 

● Upland open forests and thickets; 500–3100 m. Chongqing 

(Nanchuan), Guizhou, SW Henan, W Hubei, Hunan, Sichuan (Emei 

Shan), S Yunnan (Xishuangbanna), SE Xizang. 

7. Zanthoxylum rhombifoliolatum C. C. Huang, Acta Phytotax. 

Sin. 6: 67. 1957. 

菱叶花椒 ling ye hua jiao 

Shrubs 1–2 m tall, erect, glabrous. Branchlets and leaf 

rachises unarmed or prickly. Young branchlets purplish red. 

Leaves 7–15-foliolate; petiolules 1–5 mm; leaflet blades subopposite 

or alternate, rhombic, elliptic, or broadly ovate, 1.5–5 × 

1–2.5 cm, leathery, oil glands numerous and slightly protruding 

when dry, midvein adaxially impressed, secondary veins 4–7 on 

each side of midvein, base oblique or symmetric and narrowly to 

broadly cuneate, margin crenate, apex mucronate to caudate. 

Inflorescences terminal or axillary, 4–10 cm. Flowers 4-merous. 

Perianth in 2 series. Sepals purplish green, ovate to broadly 

deltoid, ca. 0.3 mm. Petals 2–3 mm. Male flowers subsessile; 

rudimentary gynoecium shortly club-shaped, 2-cleft to entire. 

Fruit pedicel 1–3 mm; follicles red when mature, ca. 5 mm in 

diam., oil glands numerous and slightly protruding. Seeds ca. 4 

mm in diam. Fl. May, fr. Sep. 

● Upland open forests; 500–1000 m. Chongqing (Nanchuan, 

Zhongxian), Guizhou (Zheng’an).

8. Zanthoxylum leiboicum C. C. Huang, Acta Phytotax. Sin. 

16(2): 82. 1978. 

雷波花椒 lei bo hua jiao 

Zanthoxylum calcicola C. C. Huang var. macrocarpum C. 

C. Huang. 

Shrubs to 2 m tall, erect. Branchlets and leaf rachises with 

prickles. Leaves 17–31-foliolate; petiolules 1–3 mm, puberulent; 

leaflet blades subopposite to alternate, broadly ovate to 

obovate with rounded or cuspidate apex, or ovate-obdeltoid 

with abrupt apex, 2–4 × 1.5–2.5 cm, thickly papery, oil glands 

inconspicuous, midvein impressed or apically plane, basally 

puberulent, margin apically crenulate. Inflorescences axillary, 

subracemosely cymose, 4–10 cm. Fruit pedicel 1–3 mm; follicles 

dark blackish brown when dry, 0.5–1.5 cm in diam., 

rugulose, oil glands inconspicuous, apex beaked. Seeds ca. 5 

mm in diam. Fr. Jul. 

● Thickets on dry hill slopes, river valleys; 400–1500 m. SW Sichuan. 

9. Zanthoxylum laetum Drake, J. Bot. (Morot) 6: 274. 1892. 

拟蚬壳花椒 ni xian ke hua jiao 

Zanthoxylum dissitoides C. C. Huang. 

Woody climbers to 4 m tall. Trunk, branches, branchlets, 

and leaf rachises with prickles. Rachis of young leaves, 

petiolules, and midvein of leaflet blades puberulent. Leaves 

5–13-foliolate; petiolules 2–6 mm; leaflet blades alternate, 

ovate, ovate-elliptic, or rarely oblong, 8–15 × 4–7 cm, adaxially 

lustrous, oil glands sparse, inconspicuous, and pellucid, midvein 

plane to adaxially impressed, secondary veins 9–14 on each side 

of midvein, base symmetric or rarely oblique, margin entire, 

apex acuminate. Inflorescences axillary. Flowers 4-merous. 

Pedicel ca. 4 mm. Perianth in 2 series. Sepals pale purplish 

green, narrowly ovate, to 1 mm. Petals yellowish green, broadly 

ovate, ca. 4 mm. Male flowers: filaments pale yellowish green, 

linear, 6–8 mm; rudimentary gynoecium terete, 4-lobed. Female 

flowers: staminodes shortly linear. Fruit pedicel 2–5 mm; 

follicles reddish brown with purplish red margin, 7–9 mm in 

diam., with a flange, oil glands impressed when dry, apex 

beaked. Seeds brownish black, subglobose, 6–7 mm in diam. Fl. 

Mar–May, fr. Sep–Dec. 

Moist thickets and forests; 500–1300 m. Guangdong (Zhanjiang), 

SW Guangxi, Hainan, S Yunnan [N Vietnam]. 

10. Zanthoxylum scandens Blume, Bijdr. 249. 1825. 

花椒簕 hua jiao le 

Fagara chinensis Merrill; F. cuspidata (Champion ex 

Bentham) Engler; F. cyrtorhachia Hayata; F. laxifoliolata Hayata; 

F. leiorhachia Hayata; F. scandens (Blume) Engler; Zanthoxylum 

chinense (Merrill) Chung; Z. cuspidatum Champion ex 

Bentham; Z. cyrtorhachium (Hayata) C. C. Huang; Z. laxifoliolatum 

(Hayata) C. C. Huang; Z. leiorhachium (Hayata) C. C. 

Huang. 

Shrubs or woody climbers. Trunks, branches, branchlets, 

and leaf rachises with prickles. Leaves 5–25-foliolate; leaflet 


blades alternate or opposite on distal part of leaf rachis, ovate, 

ovate-elliptic, or obliquely oblong, 4–10 × 1.5–4 cm, both 

surfaces black or blackish brown when dry, abaxially lustrous, 

oil glands inconspicuous, midvein impressed and glabrous or 

plane and puberulent, base oblique to subsymmetric and obtuse 

to broadly cuneate, margin entire or apically crenulate, apex 

mucronate, caudate, cuspidate, or long acuminate and with an 

obtuse to retuse tip. Inflorescences terminal or axillary. Flowers 

4-merous. Perianth in 2 series. Sepals pale purplish green, 

broadly ovate, ca. 0.5 mm. Petals pale yellowish green, 2–3 mm. 

Male flowers: stamens 4, 3–4 mm; connective with an oil gland 

at apex; rudimentary gynoecium subtended by pulvinate disk, 

2–4-lobed. Female flowers: carpels 3 or 4; staminodes ligulate. 

Fruit follicles purplish red, grayish brown to black when dry, 

4.5–5.5 mm in diam., oil glands slightly protruding, plane, or 

impressed, apex beaked. Seeds subglobose, 4–5 mm in diam. Fl. 

Mar–May, fr. Jul–Aug. 2n = 68. 

Lowland forests, open forests, thickets; near sea level to 1500 m. S 

Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, 

Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India, Indonesia, 

Japan (Ryukyu Islands), Malaysia, Myanmar]. 

11. Zanthoxylum kwangsiense (Handel-Mazzetti) Chun ex C. 

C. Huang, Acta Phytotax. Sin. 6: 71. 1957 [“kwangsiensis”]. 

广西花椒 guang xi hua jiao 

Fagara kwangsiensis Handel-Mazzetti, Sinensia 3. 186. 

1933. 

Woody climbers. Young branchlets, leaf rachises, and inflorescences 

pubescent. Branchlets unarmed. Leaves 5–9-foliolate; 

leaflet blades subopposite or rarely alternate, lanceolate, 

ovate, or oblanceolate, 4–10 × 2–3 cm, papery, midvein abaxially 

pubescent and adaxially impressed and puberulent, base 

broadly cuneate, margin subentire or apically crenate, apex 

mucronate to caudate. Inflorescences axillary or terminal, 

thyrsiform, 2–15 cm. Fruit pedicel 5–10 mm; follicles ca. 5 mm, 

apex beaked. Seeds subglobose, ca. 4 mm in diam. 

● Hillside thickets and open forests; 600–700 m. Chongqing 

(Fengjie, Wushan), N Guangxi, S Guizhou (Libo). 

Hartley (J. Arnold Arbor. 47: 177. 1966) placed Fagara kwangsiensis 

in synonymy of Zanthoxylum scandens. Zanthoxylum kwangsiense 

was inadvertently omitted. 

12. Zanthoxylum khasianum J. D. Hooker, Fl. Brit. India 1: 

494. 1875. 

云南花椒 yun nan hua jiao 

Zanthoxylum yunnanense C. C. Huang. 

Shrubs or trees, to 3 m tall. Branchlets gray, with prickles. 

Young branchlets, inflorescences, leaf rachises, and adaxial surface 

of leaflet blades with long grayish yellow trichomes. Leaves 

5–13-foliolate; petiolules 4–10 mm, winged; leaflet blades alternate 

or rarely subopposite, ovate to elliptic, asymmetric, 3.5–9 

× 2–4 cm, papery, midvein adaxially impressed, secondary veins 

10–16 on each side of midvein, base cuneate, margin crenulate 

or apically crenulate-serrulate, apex caudate. Inflorescences terminal 

or axillary, thyrsiform; bracts minute. Flowers 4-merous. 

Perianth in 2 series. Sepals ovate, ca. 0.5 mm, margin ciliolate. 

57

58 

Petals long elliptic, 2–3 mm. Female flowers: staminodes ca. 1 

mm. Fruit pedicel 7–10 mm, villous; follicles oblique, 5–6 mm, 

apex beaked. Fl. May, fr. Jul–Aug. 

Hillside thickets and open forests; 1500–2500 m. W Yunnan 

[India]. 

Hartley (J. Arnold Arbor. 47: 177. 1966) placed Zanthoxylum 

khasianum and Z. yunnanense in synonymy of Z. scandens. 

13. Zanthoxylum multijugum Franchet, Pl. Delavay. 124. 

1889. 

多叶花椒 duo ye hua jiao 

Fagara mengtzeana Hu; F. multijuga (Franchet) Hu; 

Zanthoxylum multifoliolatum Hemsley. 

Woody climbers. Young branchlets pale reddish brown, 

glabrous. Stems, branches, and leaf rachises with prickles. 

Leaves 19–51-foliolate; leaflet blades subsessile, subopposite to 

alternate, lanceolate, elliptic, or ovate, asymmetric, 1.5–5 × 

0.5–2 cm, oil glands numerous and pellucid, midvein puberulent 

when young and adaxially slightly ridged to plane, margin 

crenate, apex blunt, rounded, or mucronate with a blunt tip. 

Inflorescences axillary. Flowers 4(or 5)-merous. Perianth in 2 

series. Sepals ovate, to 1 mm, apex blunt to rounded. Petals pale 

yellowish green, oblong, ca. 2 mm. Male flowers: stamens 4; 

anthers broadly ovate; connective with an oil gland at apex; 

rudimentary gynoecium long conic. Female flowers: pedicel 2–5 

mm, to 1 cm in fruit. Fruit follicles brownish red, ca. 5 mm in 

diam., apex beaked. Seeds 4–4.5 mm in diam. Fl. May–Jun, fr. 

Oct–Nov. 

● Hillside thickets and open forests; 1500–2200 m. Guizhou, C and 

N Yunnan. 

14. Zanthoxylum calcicola C. C. Huang, Acta Phytotax. Sin. 6: 

65. 1957. 

石山花椒 shi shan hua jiao 

Shrubs or woody climbers, 2–3 m tall. Branchlets and leaf 

rachises with prickles. Branchlets lenticellate, puberulent. 

Leaves 9–31-foliolate; petiolules, leaf rachises adaxially, and 

leaflet blades adaxially on midvein puberulent to pubescent; 

petiolules 1–3 mm; leaflet blades lanceolate, obliquely oblong, 

or rarely ovate, 2–5 × 0.7–2.5 cm, oil glands inconspicuous, 

midvein adaxially plane or basally impressed, secondary veins 

9–12 on each side of midvein, base oblique and subrounded to 

broadly cuneate, margin subapically crenulate, apex acute or 

shortly acuminate, glandular retuse at tip. Inflorescences axillary. 

Flowers 4(or 5)-merous. Perianth in 2 series. Sepals 0.5–1 

mm. Petals 2–3 mm. Infructescences paniculate, 3–6 cm. Fruit 

pedicel ca. 5 mm; follicles 5–6 mm, oil glands impressed when 

dry. Seeds 3.5–4.5 mm in diam. Fl. Mar–Apr, fr. Sep–Nov. 

● Upland open forests; 500–1600 m. W Guangxi, SW Guizhou, SE 

Yunnan. 

15. Zanthoxylum yuanjiangense C. C. Huang, Acta Phytotax. 

Sin. 16(2): 81. 1978. 

元江花椒 yuan jiang hua jiao 

Woody climbers. Branchlets with prickles. Leaf rachises, 


inflorescences, and adaxial surface of petiolules minutely 

puberulent but other parts glabrous. Leaves 7–15-foliolate; 

petiolules 2–4 mm; leaflet blades alternate to subopposite, 

elliptic, ovate, or rarely obovate, 6–10 × 2.5–4 cm, subleathery, 

oil glands inconspicuous on mature leaflets, midvein adaxially 

impressed, secondary and tertiary veins ridged on both surfaces, 

base rounded to broadly cuneate, margin entire, apex shortly 

caudate. Inflorescences terminal or axillary, thyrsiform, to 26 

cm. Flowers 4-merous. Perianth in 2 series. Sepals purplish 

green, broadly ovate, ca. 1 mm. Petals oblong, 3–4 mm. Stamens 

4; connective with an oil gland at apex. Infructescences ca. 12 

cm wide. Fruit follicles dark brownish black, ca. 6 mm in diam., 

oil glands inconspicuous, apex beaked. Seeds 5–5.5 mm in diam. 

Fl. May, fr. Nov. 

● Upland thickets and moist secondary forests; 400–600 m. S 

Yunnan. 

16. Zanthoxylum dissitum Hemsley, J. Linn. Soc., Bot. 23: 

106. 1886. 

蚬壳花椒 xian ke hua jiao 

Woody climbers. Stem grayish white. Branchlets, leaf 

rachises, and midvein of leaflet blades with brownish red 

prickles. Leaves (3 or)5–9-foliolate; petiolules 3–10 mm; leaflet 

blades alternate to subopposite, symmetric or rarely oblique, to 

20 × 1–8 cm, thickly papery to subleathery, glabrous, oil glands 

inconspicuous, midvein adaxially impressed, margin entire, 

repand, denticulate, or serrate, apex acuminate to caudate. 

Inflorescences axillary, to 10 cm; rachis pubescent. Flowers 

4-merous. Perianth in 2 series. Sepals purplish green, broadly 

ovate, less than 1 mm. Petals pale yellowish green, broadly 

ovate, 4–5 mm. Male flowers: pedicel 1–3 mm; stamens 4; 

filaments 5–6 mm; rudimentary gynoecium 4-lobed. Female 

flowers without staminodes. Fruit follicles brown, densely 

pressed together in infructescence, 1–1.5 cm; outer part of 

pericarp (exocarp and mesocarp) extended beyond endocarp on 

each side of suture in dehisced fruit, smooth. Seeds 8–10 mm in 

diam. Fl. Apr–May or Nov–Dec, fr. Sep–Nov. 

● Upland thickets and open forests, forests; 300–2600 m. S Gansu, 

Guangdong, Guangxi, Guizhou, Hainan, SW Henan, Hubei, Hunan, S 

Shaanxi, Sichuan, Yunnan. 

1a. Branchlets with congested straight 

spines ..................................................... 16c. var. hispidum 

1b. Branchlets without congested straight spines. 

2a. Margin of leaflet blades acutely 

serrate .................................... 16d. var. acutiserratum 

2b. Margin of leaflet blades entire, 

repand, or denticulate. 

3a. Leaflet blades ca. 6 × as long 

as wide, margin entire ............ 16a. var. dissitum 

3b. Leaflet blades 7–10 × as long 

as wide; margin repand or 

denticulate .......................... 16b. var. lanciforme 

16a. Zanthoxylum dissitum var. dissitum 

蚬壳花椒(原变种) xian ke hua jiao (yuan bian zhong) 

Fagara dissita (Hemsley) Engler; Zanthoxylum bodinieri 

H. Léveillé.

Branchlets without congested straight spines. Leaflet 

blades narrowly elliptic, broadly elliptic, or subrounded, margin 

entire. Fl. Apr–May, fr. Sep–Oct. 

● Upland thickets; 300–2600 m. S Gansu, Guangdong, Guangxi, 

Guizhou, Hainan, SW Henan, Hubei, Hunan, S Shaanxi, Sichuan, 

Yunnan. 

16b. Zanthoxylum dissitum var. lanciforme C. C. Huang, 

Acta Phytotax. Sin. 16(2): 82. 1978. 

长叶蚬壳花椒 chang ye xian ke hua jiao 

Leaflet blades narrowly linear-lanceolate, 10–20 × 1–2 cm, 

base attenuate, margin repand or denticulate, apex acuminate. Fl. 

Nov–Dec. 

● Forests; ca. 1000 m. Guangxi (Damiao Shan), Guizhou. 

16c. Zanthoxylum dissitum var. hispidum (Reeder & S. Y. 

Cheo) C. C. Huang, Acta Phytotax. Sin. 6: 78. 1957. 

刺蚬壳花椒 ci xian ke hua jiao 

Fagara dissita var. hispida Reeder & S. Y. Cheo, J. Arnold 

Arbor. 32: 69. 1951. 

Branchlets with congested straight spines. Inflorescence 

rachises with prickles. Fl. Apr–May, fr. Sep–Nov. 

● Upland open forests; 1500–1800 m. C and SW Sichuan, NE 

Yunnan (Suijiang). 

16d. Zanthoxylum dissitum var. acutiserratum C. C. Huang, 


针边蚬壳花椒 zhen bian xian ke hua jiao 

Leaflet blades oblong, 6–10 × 2–3 cm, margin acutely 

serrate. Fl. Apr–May, fr. Aug–Sep. 

● Upland open forests; ca. 2400 m. Sichuan (Ganluo). 

17. Zanthoxylum collinsiae Craib, Bull. Misc. Inform. Kew 

1926: 165. 1926 [“collinsae”]. 

糙叶花椒 cao ye hua jiao 

Zanthoxylum asperum C. C. Huang; Z. scabrum Guillaumin. 

Woody climbers. Branchlets with prickles. Younger 

branchlets, leaf rachises, petiolules, and abaxial surface of leaflet 

blades villous. Leaves 5–9-foliolate; petiolules 2–4 mm; leaflet 

blades opposite, broadly ovate to ovate-elliptic, 7–19 × 5–8 cm, 

papery, black to blackish brown when dry, slightly lustrous, 

adaxially hirsutulous, midvein with prickles, secondary veins 

8–12 on each side of midvein, base oblique and rounded to 

subcordate, margin entire to crenate, apex mucronate, caudate, 

or rarely acuminate and with a retuse tip and an oil gland. 

Inflorescences axillary, 3–5 cm. Flowers 4-merous. Pedicel 

puberulent. Perianth in 2 series. Sepals puberulent, ca. 1 mm. 

Petals ca. 3 mm. Male flowers: stamens 4; rudimentary 

gynoecium ca. 1.5 mm. Fruit pedicel 2–4 mm; follicles ca. 6 mm 

in diam., pubescent, oil glands impressed when dry. Seeds 4.5–5 

mm in diam. Fl. Apr–May, fr. Sep–Oct. 

Hillside open forests and thickets; 500–1000 m. NW Guangxi, SW 

Guizhou, S Yunnan [Laos, NE Thailand, N Vietnam]. 


Hartley (J. Arnold Arbor. 47: 181. 1966) placed Zanthoxylum collinsiae, 

Z. asperum, and Z. scabrum in the synonymy of Z. nitidum. 

18. Zanthoxylum liboense C. C. Huang, Guihaia 7: 6. 1987. 

荔波花椒 li bo hua jiao 

Shrubs or woody climbers, to 1.5 m tall or long. Branchlets 

and leaf rachises with prickles. Older branchlets brownish black, 

rugulose, pubescent. Leaf rachises, inflorescences, and petiolules 

pubescent. Leaves 5–9-foliolate; petiolules 2–5 mm; leaflet 

blades subopposite to alternate, ovate to elliptic, 6–8 × 2.5–3.5 

cm, subleathery, abaxially pubescent, midvein adaxially plane or 

apically impressed and puberulent, secondary veins 10–13 on 

each side of midvein, base rounded, margin entire, apex 

mucronate to acuminate. Infructescences axillary, 3–4 cm. Fruit 

pedicel 6–10 mm; fruit 4-carpelled, usually only 2 or 3 carpels 

developing; follicles dark brownish black when dry, 7–8 mm in 

diam., pubescent when young but perhaps only on undeveloped 

carpels, oil glands inconspicuous, apex beaked. Seeds 5–6 mm 

in diam. Fr. Aug–Sep. 

● Forests, thickets; 700–800 m. S Guizhou (Libo). 

19. Zanthoxylum echinocarpum Hemsley, Ann. Bot. (Oxford) 

9: 149. 1895. 

刺壳花椒 ci ke hua jiao 

Woody climbers. Young branchlets, leaf rachises, petiolules, 

and leaflet blades abaxially on midvein pubescent. Branchlets 

and leaves with prickles. Leaves (3 or)5–11-foliolate; 

petiolules 2–5 mm; leaflet blades alternate or opposite, ovate, 

ovate-elliptic, or long elliptic, 7–13 × 2.5–5 cm, thickly leathery, 

base rounded to cordate, margin entire or subentire. Inflorescences 

axillary or terminal. Flowers 4-merous. Perianth in 2 

series. Sepals pale purplish green. Petals 2–3 mm. Male flowers: 

stamens 4. Female flowers: carpels (3 or)4(or 5). Fruit pedicel 

obsolete or to 3 mm; follicles with prickles to 1 cm. Seeds 6–8 

mm in diam. Fl. Apr–May, fr. Oct–Dec. 

● Forests, hillside open forests and thickets; 200–1800 m. N 

Guangdong, Guangxi, Guizhou, W Hubei, Hunan, Sichuan, SE Yunnan. 

1a. Abaxial surface of leaflet blades 

glabrous except for pubescent 

midvein ......................................... 19a. var. echinocarpum 

1b. Abaxial surface of leaflet blades 

villous ............................................... 19b. var. tomentosum 

19a. Zanthoxylum echinocarpum var. echinocarpum 

刺壳花椒(原变种) ci ke hua jiao (yuan bian zhong) 

Fagara echinocarpa (Hemsley) Engler. 

Leaflet blades abaxially glabrous except for pubescent 

midvein. Prickles of follicles pubescent. 

● Forests; 200–1000 m. N Guangdong, Guangxi, Guizhou, W 

Hubei, Hunan, Sichuan, SE Yunnan. 

19b. Zanthoxylum echinocarpum var. tomentosum C. C. 

Huang, Acta Phytotax. Sin. 16(2): 82. 1978. 

毛刺壳花椒 mao ci ke hua jiao 

59

60 

Branchlets, leaf rachises, petioles, abaxial surface of leaflet 

blades, and inflorescence rachises villous. Prickles of follicles 

pilose when mature. 

● Hillside open forests and thickets; 300–1800 m. NW Guangxi 

(Tian’e), SW Guizhou (Anlong), SE Yunnan. 

20. Zanthoxylum oxyphyllum Edgeworth, Trans. Linn. Soc. 

London 20: 42. 1846. 

尖叶花椒 jian ye hua jiao 

Fagara oxyphylla (Edgeworth) Engler; Zanthoxylum 

alpinum C. C. Huang; Z. taliense C. C. Huang; Z. tibetanum C. 

C. Huang. 

Shrubs or small trees. Branchlets and leaf rachises with 

prickles. Leaf rachises adaxially and midvein of leaflet blades 

adaxially pubescent. Old leaves subglabrous. Leaves 11–19- 

foliolate; petiolules to 2 mm; leaflet blades alternate or opposite, 

lanceolate or rarely ovate, 5–12 × 1.5–2.5 cm, abaxially gray 

when dry, oil glands numerous, midvein impressed adaxially, 

secondary veins anastomosing near margin, reticulate veinlets 

ridged when dry, base cuneate, margin serrulate, apex acuminate. 

Inflorescences terminal, cymose-corymbose, to 30-flowered. 

Perianth in 2 series. Sepals 4, purplish green. Petals ca. 3 

mm. Male flowers: rudimentary gynoecium 2–4-parted, lobes 

linear. Fruit pedicel 1–1.5 cm, 1–1.5 mm in diam.; follicles 

purplish red, 6–7 mm, oil glands impressed when dry, apex 

beaked. Seeds ca. 5 mm in diam. Fl. May–Jun, fr. Sep–Oct. 2n = 

72. 

Open forests, forest margins; 1800–2900 m. S Xizang, W Yunnan 

[Bhutan, NE India, Myanmar, Nepal]. 

21. Zanthoxylum stenophyllum Hemsley, Ann. Bot. (Oxford) 

9: 147. 1895. 

狭叶花椒 xia ye hua jiao 

Fagara stenophylla (Hemsley) Engler; Zanthoxylum pashanense 

N. Chao. 

Shrubs or small trees. Stems and branches grayish white. 

Young branchlets pale purplish red. Branchlets and midvein of 

leaflet blades abaxially with prickles. Leaves 9–23-foliolate; 

petiolules 1–3 mm, adaxially pubescent; leaflet blades alternate, 

lanceolate and 2–11 × 1–4 cm, narrowly lanceolate and 2–3.5 × 

0.4–0.7 cm, or ovate and 8–16 × 6–8 mm, oil glands inconspicuous, 

midvein adaxially ridged to plane and puberulent but 

glabrous in fruit, reticulate veinlets ridged on both surfaces, base 

cuneate to subrounded, margin crenulate, apex acuminate to 

mucronate. Inflorescences terminal, cymose-corymbiform, to 

30-flowered. Flowers 4-merous. Perianth in 2 series. Sepals ca. 

0.5 mm. Petals 2.5–3 mm. Male flowers: pedicel 2–5 mm; 

stamens 4; connective without an oil gland at apex; rudimentary 

gynoecium disciform; styles coherent. Female flowers: pedicel 

0.6–1.5 cm; staminodes absent. Fruit pedicel purplish red, 1–3 

cm, glabrous; follicles pale purplish red to dark red, 4.5–5 cm in 

diam., oil glands impressed when dry, apex beaked. Seeds ca. 4 

mm in diam. Fl. May–Jun, fr. Aug–Sep. 

● Upland thickets; 700–2400 m. S Gansu (Chengxian, Huixian), W 

Henan, W Hubei, NE Hunan, SW Shaanxi, Sichuan. 


22. Zanthoxylum esquirolii H. Léveillé, Repert. Spec. Nov. 

Regni Veg. 13: 266. 1914. 

贵州花椒 gui zhou hua jiao 

Fagara chaffanjonii (H. Léveillé) Handel-Mazzetti; F. 

esquirolii (H. Léveillé) Handel-Mazzetti; Zanthoxylum chaffanjonii 

H. Léveillé. 

Shrubs or small trees. Branchlets and leaf rachises with 

prickles. Branchlets pale purplish red and glaucous when dry. 

Leaves 5–13-foliolate; petiolules 3–6 mm; leaflet blades alternate, 

ovate, lanceolate, or rarely broadly ovate, 3–10 × 1.5–4.5 

cm, oil glands inconspicuous, midvein adaxially impressed, base 

subrounded to broadly cuneate, margin crenulate or entire 

toward base, apex obliquely caudate with a retuse tip. Inflorescences 

terminal, cymose-corymbiform, to 30-flowered. 

Flowers 4-merous. Perianth in 2 series. Petals ca. 3 mm. Female 

flowers (3 or)4-carpelled. Fruit pedicel to 4.5 mm, 0.5–1 mm in 

diam.; follicles purplish red, ca. 5 mm in diam., oil glands 

impressed, apex beaked. Seeds ca. 4 mm in diam. Fl. May–Jun, 

fr. Sep–Nov. 

● Upland open forests and thickets; 700–3200 m. Guizhou, Sichuan, 

Yunnan. 

23. Zanthoxylum avicennae (Lamarck) Candolle, Prodr. 1: 

726. 1824. 

簕花椒 le dang hua jiao 

Fagara avicennae Lamarck, Encycl. 2: 445. 1788; Zanthoxylum 

avicennae var. tonkinense Pierre; Z. avicennae var. 

touranense Pierre; Z. lentiscifolium Champion ex Bentham. 

Trees to 15 m tall, deciduous. Branchlets and leaves 

glabrous, with prickles. Leaves 11–21-foliolate; rachis winged; 

leaflet blades opposite or rarely subopposite, obliquely ovate, 

rhomboidal, obovate, or falcate, 2.5–7 × 1–3 cm, oil glands 

visible on fresh leaves or inconspicuous, margin entire or apically 

crenate, apex mucronate to blunt. Inflorescences terminal, 

many flowered; rachis purplish red. Flowers 5-merous. Pedicel 

purplish red. Perianth in 2 series. Sepals green, broadly ovate. 

Petals yellowish white, ca. 2.5 mm. Male flowers: stamens 5; 

rudimentary gynoecium disciform, 2-lobed. Female flowers: 

carpels 2(or 3); staminodes small. Fruit pedicel 3–6 mm; follicles 

pale purplish red, 4–5 mm in diam., oil glands numerous, 

large, and slightly protruding, apex not beaked. Seeds 3.5–4.5 

mm in diam. Fl. Jun–Aug, fr. Oct–Dec. 

Secondary forests in lowland flat areas, hillsides, valleys; 400–700 

m. S Fujian, Guangdong, Guangxi, Hainan, S Yunnan [India, Indonesia, 

Malaysia, Philippines, Thailand, Vietnam]. 

24. Zanthoxylum micranthum Hemsley, Ann. Bot. (Oxford) 

9: 147. 1895. 

小花花椒 xiao hua hua jiao 

Fagara biondii Pampanini; F. micrantha (Hemsley) Engler. 

Trees to 15 m tall, deciduous. Branchlets glabrous, with 

prickles. Leaves 9–17-foliolate; petiolules 1.5–5 mm; leaflet 

blades opposite or on basal part of rachis subopposite, lanceo-

late, 5–8 × 1–3 cm, oil glands numerous, midvein impressed, 

secondary veins 8–12 on each side of midvein, base symmetrically 

or obliquely rounded to broadly cuneate, margin 

crenate, apex acuminate. Inflorescences terminal, many flowered. 

Flowers 5-merous. Perianth in 2 series. Sepals broadly 

ovate, ca. 0.3 mm wide. Petals pale yellowish white, 1.5–2 mm. 

Male flowers: stamens 5, ca. 3 mm; rudimentary gynoecium 

disciform, 3-lobed or not divided. Female flowers 3(or 4)-carpelled. 

Fruit follicles pale purplish red but pale yellow to grayish 

brown when dry, ca. 5 mm in diam., oil glands small, apex not 

beaked. Seeds to 4 mm. Fl. Jul–Aug, fr. Oct–Nov. 

● Hillside open forests; 300–1200 m. Guizhou, SW Henan, W 

Hubei, Hunan, Sichuan, Yunnan. 

25. Zanthoxylum ailanthoides Siebold & Zuccarini, Abh. 

Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 4(2): 138. 1845. 

椿叶花椒 chun ye hua jiao 

Trees to 15 m tall, deciduous. Branchlets and inflorescence 

rachises glabrous, with prickles. Leaves 11–27-foliolate; leaflet 

blades opposite, narrowly lanceolate but subovate basally on 

rachis, 7–18 × 2–6 cm, abaxially grayish green or glaucescent, 

oil glands numerous, midvein adaxially impressed, secondary 

veins 11–16 on each side of midvein, base symmetrically or 

subobliquely rounded, margin crenate, apex acuminate. Inflorescences 

terminal, many flowered. Flowers 5-merous, subsessile. 

Perianth in 2 series. Sepals broadly triangular, ca. 0.8 mm. 

Petals pale yellowish white, ca. 2.5 mm. Male flowers: stamens 

5; rudimentary gynoecium disciform, 2- or 3-lobed. Female 

flowers (3 or)4-carpelled. Fruit pedicel 1–3 mm; follicles pale 

reddish brown but pale gray to brownish gray when dry, ca. 4.5 

mm in diam., oil glands numerous, impressed when dry, apex 

not beaked. Seeds ca. 4 mm in diam. Fl. Aug–Sep, fr. Oct–Dec. 

Upland thickets; 300–1500 m. Fujian, Guangdong, Guangxi, 

Guizhou, S Jiangxi, SE Sichuan, Taiwan, SE Yunnan, Zhejiang [Japan 

(including Bonin and Ryukyu Islands), Korea, Philippines]. 

1a. Leaflet blades glabrous on both surfaces 

........................................................... 25a. var. ailanthoides 

1b. Leaflet blades abaxially pubescent ..... 25b. var. pubescens 

25a. Zanthoxylum ailanthoides var. ailanthoides 

椿叶花椒(原变种) chun ye hua jiao (yuan bian zhong) 

Fagara ailanthoides (Siebold & Zuccarini) Engler; F. 

boninshimae G. Koidzumi; F. emarginella (Miquel) Engler; F. 

hemsleyana (Makino) Makino; Zanthoxylum ailanthoides var. 

inerme Rehder & E. H. Wilson; Z. emarginellum Miquel; Z. 

hemsleyanum Makino. 

Leaflet blades glabrous on both surfaces, abaxially 

glaucous. 

Upland thickets; 300–1500 m. Fujian, Guangdong, Guangxi, 

Guizhou, S Jiangxi, SE Sichuan, Taiwan, SE Yunnan, Zhejiang [Japan 

(including Bonin and Ryukyu Islands), Korea, Philippines]. 

Zanthoxylum hemsleyanum Makino was based on Z. emarginellum 

sensu Hemsley (1895), not Miquel (1867). 

25b. Zanthoxylum ailanthoides var. pubescens Hatusima, 

Acta Phytotax. Geobot. 4: 210. 1935. 


毛椿叶花椒 mao chun ye hua jiao 

Leaf rachises and abaxial surface of leaflet blades pubescent. 

● Taiwan. 

This taxon is not recognized as a separate variety in Fl. Taiwan, ed. 

2, 3: 537. 1993. 

26. Zanthoxylum myriacanthum Wallich ex J. D. Hooker, Fl. 

Brit. India 1: 496. 1875. 

大叶臭花椒 da ye chou hua jiao 

Trees to 15 m tall, deciduous. Branchlets toward apex and 

rachis of inflorescences with prickles. Leaves without prickles, 

7–17-foliolate; leaflet blades opposite, broadly ovate, ovate- 

elliptic, or oblong, but suborbicular basally on rachis, 10–20 × 

4–10 cm, both surfaces glabrous, oil glands numerous, large, 

turning red or blackish brown and slightly protruding when dry, 

midvein adaxially impressed, base symmetrically or obliquely 

rounded to broadly cuneate, margin crenulate-serrulate. Inflorescences 

terminal, to 35 × 30 cm, many flowered. Flowers 

5-merous. Perianth in 2 series. Sepals broadly ovate, ca. 0.3 mm. 

Petals white, ca. 2.5 mm. Male flowers: stamens 5; rudimentary 

gynoecium disciform, 3-lobed. Female flowers: carpels (2 

or)3(or 4); staminodes small. Fruit follicles reddish brown, ca. 

4.5 mm in diam., oil glands numerous, apex not beaked. Seeds 

ca. 4 mm in diam. Fl. Jun–Aug, fr. Sep–Nov. 

Hillside forests, forests; 200–1500 m. Fujian, Guangdong, Guangxi, 

SE Guizhou, Hainan, S Hunan, Jiangxi, S Yunnan, S Zhejiang [Bhutan, 

India, Indonesia, Malaysia, Myanmar, Philippines, Vietnam]. 

1a. Petiolules, leaflet blades, leaf rachises, 

and inflorescences not villous ...... 26a. var. myriacanthum 

1b. Petiolules, leaflet blades, leaf rachises, 

and inflorescences villous ................... 26b. var. pubescens 

26a. Zanthoxylum myriacanthum var. myriacanthum 

大叶臭花椒(原变种) da ye chou hua jiao (yuan bian zhong) 

Euodia odorata H. Léveillé; Fagara diabolica (Elmer) 

Engler; F. gigantea Handel-Mazzetti; F. myriacantha (Wallich 

ex J. D. Hooker) Engler; F. odorata (H. Léveillé) Handel- 

Mazzetti; F. rhetsoides (Drake) Reeder & S. Y. Cheo; 

Zanthoxylum diabolicum Elmer; Z. giganteum (Handel-Mazzetti) 

Rehder; Z. odoratum (H. Léveillé) H. Léveillé; Z. rhetsoides 

Drake. 

Leaf rachises, petiolules, leaflet blades, and inflorescence 

rachises not villous. 

Hillside forests; 200–1500 m. Fujian, Guangdong, Guangxi, SE 

Guizhou, Hainan, S Hunan, Jiangxi, S Yunnan, S Zhejiang [Bhutan, 

India, Indonesia, Malaysia, Myanmar, Philippines, Vietnam]. 

26b. Zanthoxylum myriacanthum var. pubescens (C. C. 

Huang) C. C. Huang, Guihaia 11: 9. 1991. 

毛大叶臭花椒 mao da ye chou hua jiao 

Zanthoxylum rhetsoides var. pubescens C. C. Huang, Acta 

Phytotax. Sin. 6: 48. 1957; Z. utile C. C. Huang. 

61

62 

Leaf rachises, petiolules, leaflet blades, and inflorescence 

rachises villous. 

● Forests; ca. 1400 m. S Yunnan (Xishuangbanna). 

27. Zanthoxylum molle Rehder, J. Arnold Arbor. 8: 150. 1927. 

朵花椒 duo hua jiao 

Euodia mollicoma Hu & F. H. Chen; Fagara mollis 

(Rehder) Reeder & S. Y. Cheo. 

Trees to 10 m tall, deciduous. Bark brownish black. Young 

branches and rachis of inflorescences with prickles. Young 

branches dark purplish red. Leaves (5–)13–19-foliolate; rachis 

pubescent; leaflet blades opposite, subsessile, broadly ovate, 

elliptic, or rarely suborbicular, 8–15 × 4–9 cm, thickly leathery, 

abaxially with soft grayish white to grayish yellow woollyvillous 

indumentum, oil glands inconspicuous, midvein adaxially 

impressed, secondary veins 11–17 on each side of midvein, 

base symmetrically or rarely obliquely rounded to subcordate, 

margin entire or crenulate, apex acute. Inflorescences terminal, 

many flowered. Flowers 5-merous. Pedicel pale purplish red, 

pubescent. Perianth in 2 series. Petals white, 2–3 mm. Male 

flowers: rudimentary gynoecium 3-lobed. Female flowers: 

carpels 3; staminodes small. Fruit follicles pale purplish red but 

pale yellow to grayish brown when dry, 4–5 mm in diam., oil 

glands numerous and impressed when dry, apex not beaked. 

Seeds 3.5–4 mm in diam. Fl. Jun–Aug, fr. Oct–Nov. 

● Upland open forests and thickets; 100–900 m. Anhui, Guizhou, 

Henan, Hunan, Jiangxi, Yunnan, Zhejiang. 

28. Zanthoxylum schinifolium Siebold & Zuccarini, Abh. 

Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 4(2): 137. 1845. 

青花椒 qing hua jiao 

Fagara pteropoda (Hayata) Y. C. Liu; F. schinifolia 

(Siebold & Zuccarini) Engler; Zanthoxylum mantschuricum 

Bennett; Z. pteropodum Hayata. 

Shrubs 1–2 m tall. Stems and branchlets with prickles. 

Young branchlets dark purplish red. Leaves 7–19-foliolate; 

petiolules obsolete or to 3 mm; leaflet blades opposite, or 

alternate toward base of rachis, broadly ovate, broadly ovaterhombic, 

or lanceolate, 5–10(–70) × 4–6(–25) mm, papery, oil 

glands numerous or inconspicuous, midvein adaxially impressed, 

base symmetrically or sometimes obliquely rounded to 

broadly cuneate, margin serrate to subentire, apex mucronate to 

acuminate. Inflorescences terminal. Flowers 5-merous. Perianth 

in 2 series. Petals pale yellowish white, ca. 2 mm. Male flowers: 

rudimentary gynoecium 2- or 3-lobed. Female flowers: carpels 

3(–5). Fruit follicles reddish brown but dark green to brownish 

black when dry, 4–5 mm in diam., oil glands small, apex not 

beaked. Seeds 3–4 mm in diam. Fl. Jul–Sep, fr. Sep–Dec. 

Upland open forests and thickets; below 800 m. Anhui, Fujian, N 

Guangdong, N Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, 

Jiangsu, Jiangxi, Liaoning, Shandong, Taiwan, Zhejiang [Japan (including 

Ryukyu Islands), Korea]. 

29. Zanthoxylum dimorphophyllum Hemsley, Ann. Bot. (Oxford) 

9: 150. 1895. 

异叶花椒 yi ye hua jiao 


Trees to 10 m tall, deciduous. Branches grayish black, with 

few prickles or unarmed; young branchlets and shoots rust- 

colored pubescent or glabrous; axillary buds rust-colored 

villosulous. Leaves 3–5(–11)-foliolate; leaflet blades ovate, 

elliptic, or sometimes obovate, (2–)4–9(–20) × (1–)2–3.5(–7) 

cm, oil glands numerous, midvein adaxially plane or impressed 

and puberulent, reticulate veinlets slightly ridged when dry, base 

symmetric, margin crenate and with or without spines, apex 

blunt, rounded, mucronate, or acuminate, usually with a retuse 

tip. Inflorescences terminal; bracts rust-colored villosulous. 

Perianth in 2 series with 4 sepals and 4 petals or grading to 2 

irregular series or 1 series with 7 or 8 ± undifferentiated 2–3 mm 

tepals; when differentiated, sepals 0.2–0.5 mm and petals ca. 1.5 

mm. Male flowers: stamens 4–6; disk pulvinate; rudimentary 

gynoecium obsolete or 2- or 3-carpelled. Female flowers: 

staminodes 4 or 5; rudimentary anthers without pollen; gynoecium 

2- or 3-carpelled; styles recurved. Fruit follicles purplish 

red, 6–8 mm in diam., sparsely pubescent when young, with 

sparse oil glands, stipitate, apex shortly beaked. Seeds 5–7 mm 

in diam. Fl. Apr–Jun, fr. Sep–Nov. 2n = 36, 68. 

Moist areas in upland forests, hillside open forests and thickets; 

300–2400 m. S Gansu, N Guangdong, Guangxi, Guizhou, Hainan, SW 

Henan, W Hubei, Hunan, S Shaanxi, Sichuan, Taiwan, Yunnan [Thailand, 

Vietnam]. 

C. C. Huang (Fl. Reipubl. Popularis Sin. 43(2): 40. 1997) 

mistakenly placed Zanthoxylum dimorphophyllum in synonymy of Z. 

ovalifolium Wight, which ranges from India to NE Australia and is not 

known to occur in China. 

1a. Margin of leaflet blades with spines 

............................................................ 29c. var. spinifolium 

1b. Margin of leaflet blades without spines. 

2a. Leaves 3- or 5-foliolate 

.......................................... 29a. var. dimorphophyllum 

2b. Leaves (5 or)7–11-foliolate 

............................................. 29b. var. multifoliolatum 

29a. Zanthoxylum dimorphophyllum var. dimorphophyllum 

异叶花椒(原变种) yi ye hua jiao (yuan bian zhong) 

Fagara dimorphophylla (Hemsley) Engler; F. robiginosa 

Reeder & S. Y. Cheo; Zanthoxylum acanthopodium Candolle 

var. deminutum (Rehder) Reeder & S. Y. Cheo; Z. dimorphophyllum 

var. deminutum Rehder; Z. evoidiifolium Guillaumin; Z. 

pistaciiflorum Hayata; Z. robiginosum (Reeder & S. Y. Cheo) C. 

C. Huang. 

Leaves 3- or 5-foliolate; margin of leaflet blades without 

spines. 

Moist areas in upland forests; 300–2400 m. S Gansu, N Guangdong, 

Guangxi, Guizhou, Hainan, W Hubei, Hunan, S Shaanxi, Sichuan, 

Taiwan, Yunnan [Thailand, Vietnam]. 

29b. Zanthoxylum dimorphophyllum var. multifoliolatum C. 

C. Huang, Acta Phytotax. Sin. 16(2): 81. 1978. 

多异叶花椒 duo yi ye hua jiao 

Zanthoxylum ovalifolium Wight var. multifoliolatum (C. C. 

Huang) C. C. Huang.

Leaves (5 or)7–11-foliolate; margin of leaflet blades 

without spines. 

● Hillside thickets. Yunnan (Kunming). 

29c. Zanthoxylum dimorphophyllum var. spinifolium Rehder 

& E. H. Wilson in Sargent, Pl. Wilson. 2: 126. 1914. 

刺异叶花椒 ci yi ye hua jiao 

Zanthoxylum dissitum Hemsley var. spinulosum Z. M. Tan; 

Z. ovalifolium Wight var. spinifolium (Rehder & E. H. Wilson) 

C. C. Huang. 

Leaves 3- or 5-foliolate; margin of leaflet blades with 

spines. 

● Hillside open forests and thickets; 400–2100 m. Guizhou, SW 

Henan (Funiu Shan), W Hubei, Hunan, S Shaanxi (Qin Ling), Sichuan. 

30. Zanthoxylum acanthopodium Candolle, Prodr. 1: 727. 

1824. 

刺花椒 ci hua jiao 

Zanthoxylum acanthopodium var. oligotrichum Z. M. Tan; 

Z. acanthopodium var. timbor J. D. Hooker; Z. acanthopodium 

var. villosum C. C. Huang. 

Shrubs, woody climbers, or trees to 6 m tall. Bark grayish 

black. Branchlets rust-colored pubescent to nearly glabrous, 

usually with prickles. Leaves 3–9-foliolate; rachis wings to 3 

mm on each side; leaflet blades sessile, opposite, ovate-elliptic 

to lanceolate, 6–10 × 2–4 cm, papery, both surfaces glabrous to 

rust-colored pubescent, oil glands inconspicuous, secondary 

veins 10–28 on each side of midvein and evident, margin entire 

or crenate. Inflorescences axillary. Perianth in 2 irregular series 

or 1 series, with 6–8 ± undifferentiated tepals. Tepals pale 

yellowish green, narrowly lanceolate, ca. 1.5 mm. Male flowers: 

stamens 5 or 6; anthers reddish purple before anthesis; disk 

pulvinate; rudimentary carpels 2–5. Female flowers: rudimentary 

stamens lacking; carpels 2–5, sparsely hirsute to glabrous, 

abaxially often with a conspicuous oil gland; styles recurved. 

Fruit follicles usually purplish red, ca. 4 mm in diam., glabrous 

or sometimes with sparse trichomes, oil glands large and 

protruding. Seeds ca. 3 mm in diam. Fl. Apr–May, fr. Sep–Oct. 

2n = 64. 

Upland open forests and thickets; 1400–3200 m. W Guangxi, 

Guizhou, Sichuan, Xizang, Yunnan [Bangladesh, Bhutan, India, Indonesia, 

Laos, Malaysia, Myanmar, Nepal, Thailand, Vietnam]. 

31. Zanthoxylum armatum Candolle, Prodr. 1: 727. 1824. 

竹叶花椒 zhu ye hua jiao 

Shrubs, woody climbers, or trees to 5 m tall, deciduous. 

Branchlets and leaflet blades abaxially on midvein usually with 

prickles. Young branchlets and inflorescence rachises glabrous 

or rust-colored pubescent. Leaves 3–9(or 11)-foliolate; rachis 

glabrous or rust-colored pubescent, wings to 6 mm on each side; 

leaflet blades subsessile, opposite, lanceolate, ovate, or elliptic, 

3–12 × 1–3 cm, base attenuate to broadly cuneate, secondary 

veins 7–15 on each side of midvein and generally faint, margin 

crenate or entire and often revolute when dry, apex acute to 

acuminate. Inflorescences terminal on short lateral branchlets 


and sometimes axillary, 1–7 cm, with less than 30 flowers. 

Perianth in 2 irregular series or 1 series, with 6–8 ± undifferentiated 

0.3–1.5 mm tepals. Male flowers: stamens 4–6; 

anthers yellow prior to anthesis; connective apex with oil gland; 

disk pulvinate; rudimentary carpels lacking. Female flowers: 

carpels 2 or 3, abaxially often with a conspicuous oil gland; 

styles recurved; staminodes ligulate or lacking. Fruit follicles 

usually purplish red, 4–5 mm in diam., with a few protruding oil 

glands. Seeds blackish brown, 3–4 mm in diam. Fl. Apr–May, fr. 

Aug–Oct. 2n = 66. 

Found in many habitats; below 3100 m. Anhui, Fujian, S Gansu, 

Guangdong, Guangxi, Guizhou, S Henan, Hubei, Hunan, Jiangsu, 

Jiangxi, Shaanxi, Shandong, S Shanxi, Sichuan, N Taiwan, Xizang, Yunnan, 

Zhejiang [Bangladesh, Bhutan, India, Indonesia, Japan (including 

Ryukyu Islands), Kashmir, Korea, Laos, Myanmar, Nepal, Pakistan, 

Philippines, Thailand, Vietnam]. 

Ohba (Fl. Jap. 22c: 37. 2001) treats the plants from Japan and most 

of China as Zanthoxylum armatum var. subtrifoliolatum (Franchet) 

Kitamura. 

1a. Young branchlets and inflorescence rachises 

glabrous or young branches sparsely 

pubescent ............................................... 31a. var. armatum 

1b. Young branchlets and inflorescence 

rachises rust-colored pubescent ...... 31b. var. ferrugineum 

31a. Zanthoxylum armatum var. armatum 

竹叶花椒(原变种) zhu ye hua jiao (yuan bian zhong) 

Zanthoxylum alatum Roxburgh; Z. alatum var. planispinum 

(Siebold & Zuccarini) Rehder & E. H. Wilson; Z. alatum var. 

subtrifoliolatum Franchet; Z. arenosum Reeder & S. Y. Cheo; Z. 

armatum var. subtrifoliolatum (Franchet) Kitamura; Z. planispinum 

Siebold & Zuccarini. 

Young branchlet apices and inflorescence rachises glabrous, 

or young branches sparsely pubescent. Leaflet blades 

abaxially flocculent on midvein. 

Found in many habitats; below 3100 m. Anhui, Fujian, S Gansu, 

N Guangdong, Guangxi, Guizhou, S Henan, Hubei, Hunan, Jiangsu, 

Jiangxi, S Shaanxi, Shandong, S Shanxi, Sichuan, N Taiwan, Xizang, 

Yunnan, Zhejiang [Bangladesh, Bhutan, India, Indonesia, Japan (including 

Ryukyu Islands), Kashmir, Korea, Laos, Myanmar, Nepal, Pakistan, 


31b. Zanthoxylum armatum var. ferrugineum (Rehder & E. 

H. Wilson) C. C. Huang, Guihaia 7: 1. 1987. 

毛竹叶花椒 mao zhu ye hua jiao 

Zanthoxylum alatum f. ferrugineum Rehder & E. H. Wilson 

in Sargent, Pl. Wilson. 2: 215. 1914. 

Young branchlet apices, inflorescence rachises, and sometimes 

leaf rachises rust-colored pubescent. 

nan. 

● Guangdong, Guangxi, Guizhou, Hunan, Shaanxi, Sichuan, Yun- 

32. Zanthoxylum wutaiense I. S. Chen, Taiwan Sci. 26: 56. 

1972. 

屏东花椒 ping dong hua jiao 

63

64 

Shrubs. Branchlets with prickles or unarmed, glabrous. 

Leaves 5–13-foliolate; rachis canaliculate, sides of channel 

rarely slightly spreading apically; leaflet blades sessile, opposite, 

oblong to oblong-lanceolate, 2.5–10 × 0.6–2 cm, leathery, midvein 

narrowly ridged adaxially, secondary veins 7–16 on each 

side of midvein, base attenuate to cuneate, margin crenulate, 

apex acuminate at least in terminal blade. Inflorescences axillary, 

paniculate. Flowers yellow. Pedicel ca. 3 mm. Perianth in 2 

irregular series or 1 series, with 5–8 ± undifferentiated tepals. 

Male flowers: stamens 5–8. Female flowers: pistil subtended by 

columnar disk, 1(or 2)-carpelled; carpel ovoid to ellipsoid, with 

a conspicuous lateral oil gland; style short; stigma capitate. Fruit 

follicle ovoid, ca. 5 mm in diam. Fl. Apr. 

● Exposed slopes in secondary broad-leaved forests; 1300–1400 

m. Taiwan (Pingdong). 

33. Zanthoxylum bungeanum Maximowicz, Bull. Acad. Imp. 

Sci. Saint-Pétersbourg 16: 212. 1871. 

花椒 hua jiao 

Trees 3–7 m tall, deciduous. Stems and branchlets with 

prickles; stem prickles with a flat base. Young branchlets 

pubescent. Leaves 5–13-foliolate; rachis marginate; leaflet 

blades sessile, opposite, ovate, elliptic, or rarely lanceolate, 

sometimes suborbicular near leaf rachis base, 2–7 × 1–4.5 cm, 

both surfaces pubescent or abaxial surface flocculent along 

midvein, midvein adaxially impressed, margin crenate. Inflorescences 

axillary but terminal on lateral branchlets; rachis and 

pedicel pubescent or glabrous. Perianth in 2 irregular series or 1 

series, with 6–8 yellowish green ± undifferentiated tepals. Male 

flowers: stamens 5–8; rudimentary gynoecium 2-lobed. Female 

flowers 2–5-carpelled. Fruit follicles purplish red, 4–5 mm in 

diam., pustulose glandular, apex shortly beaked or beak lacking. 

Seeds 3.5–4.5 mm. Fl. Apr–May, fr. Aug–Oct. 

Found in many habitats; below 3200 m. Anhui, Fujian, Gansu, 

Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, 

Liaoning, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, SE 

Xinjiang, S and SE Xizang, Yunnan, Zhejiang [Bhutan]. 

The dried fruit follicles of Zanthoxylum bungeanum are used as a 

culinary spice and are particularly popular in Sichuan cuisine (Sichuan 

pepper). In Japan, Z. piperitum (Linnaeus) Candolle is sometimes similarly 

used. 

1a. Leaflet blades pubescent on both 

surfaces or sometimes adaxial surface 

glabrous ............................................... 33c. var. pubescens 

1b. Leaflet blades flocculent along midvein 

abaxially, otherwise glabrous. 

2a. Leaflet blades with oil glands only 

at margin ................................... 33a. var. bungeanum 

2b. Leaflet blades with scattered oil 

glands .......................................... 33b. var. punctatum 

33a. Zanthoxylum bungeanum var. bungeanum 

花椒(原变种) hua jiao (yuan bian zhong) 

Zanthoxylum bungei Hance; Z. bungei var. imperforatum 

Franchet; Z. fraxinoides Hemsley; Z. simulans Hance var. 

imperforatum (Franchet) Reeder & S. Y. Cheo. 


Leaflet blades flocculent along midvein abaxially, otherwise 

glabrous, oil glands only at margin. 

Found in many habitats; below 3200 m. Anhui, Fujian, Gansu, 

Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, 

Liaoning, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, SE 

Xinjiang, S and SE Xizang, Yunnan, Zhejiang [Bhutan]. 

33b. Zanthoxylum bungeanum var. punctatum C. C. Huang, 


油叶花椒 you ye hua jiao 

Leaf rachises, infructescences, and follicles reddish brown 

when dry. Leaflet blades with oil glands conspicuous, scattered. 

Fr. Jul–Aug. 

● Open forests; 2000–2500 m. W Sichuan. 

33c. Zanthoxylum bungeanum var. pubescens C. C. Huang, 

Acta Phytotax. Sin. 6: 24. 1957. 

毛叶花椒 mao ye hua jiao 

Young branchlets, rachis of leaves and inflorescences, and 

both surfaces of leaflet blades pubescent, or sometimes leaflet 

blades adaxially glabrous. Fl. May–Jun, fr. Oct–Nov. 

● 1700–3200 m. Gansu, Qinghai (Xunhua), S Shaanxi, NW and W 

Sichuan, Yunnan. 

34. Zanthoxylum undulatifolium Hemsley, Ann. Bot. (Oxford) 

9: 148. 1895. 

浪叶花椒 lang ye hua jiao 

Trees to 3 m tall. Young branchlets and leaf rachises 

rust-colored pubescent, with few prickles or unarmed. Leaves 

3–5(–7)-foliolate; terminal leaflet with petiolule 6–10 mm, 

lateral leaflets subsessile; leaflet blades opposite, ovate to 

ovate-lanceolate, 3–8 × 1.5–3.5 cm, abaxially glabrous, adaxially 

sparsely puberulent, midvein adaxially plane, secondary 

veins 6–10 on each side of midvein, base broadly cuneate to 

rounded, margin undulate and crenulate, apex mucronate to 

acuminate. Inflorescences terminal, corymbose. Perianth in 2 


Fruit pedicel reddish brown, 0.7–1.4 cm; follicles reddish brown, 

ca. 5 mm in diam., oil glands large and impressed, apex not 

beaked. Seeds ca. 4 mm in diam. Fl. Apr–May, fr. Aug–Oct. 

● Forests, thickets; 1600–3200 m. W Hubei, S Shaanxi, E Sichuan, 

NE Yunnan. 

35. Zanthoxylum austrosinense C. C. Huang, Acta Phytotax. 

Sin. 6: 53. 1957. 

岭南花椒 ling nan hua jiao 

Shrubs or trees, rarely to 3 m tall, dioecious or rarely 

polygamo-monoecious, all parts glabrous. Branchlets blackish 

brown, unarmed. Leaves 5–11-foliolate; rachis terete; lateral 

leaflets opposite, sessile or subsessile; terminal leaflet with 

petiolule 1–3 mm; leaflet blades ovate to lanceolate, 6–11 × 3–5 

cm, oil glands clear but dark reddish brown to blackish brown 

when dry, midvein adaxially impressed or plane, secondary 

veins 11–15 on each side of midvein, base rounded to subcordate 

or oblique, margin serrulate, apex acuminate. Inflorescences 

terminal, usually on lateral branchlets, to 30-flowered. Pedicel 

5–8 mm. Perianth in 2 irregular series or 1 series, with 7–9 ±

undifferentiated tepals. Tepals basally pale yellowish green and 

apically dark purplish red, lanceolate to oblanceolate, ca. 1.5 

mm. Bisexual flowers: stamens 3 or 4; carpels 4. Male flowers: 

stamens 6–8. Female flowers: carpels 3 or 4; styles recurved; 

stigmas capitate. Fruit pedicel dark purplish red, 1–2 cm; 

follicles dark purplish red, ca. 5 mm in diam., with sparse 

protruding oil glands, apex subrounded or shortly beaked. Seeds 

ca. 4 × 3–4 mm. Fl. Mar–Apr, fr. Aug–Sep. 

● Hillside open forests and thickets, upland thickets; 300–1700 m. 

S Anhui, Fujian, N Guangdong, NE Guangxi, SW Hubei, Hunan, 

Jiangxi, Zhejiang. 

1a. Leaflet blades dark reddish brown to 

blackish brown when dry, glabrous 

......................................................... 35a. var. austrosinense 

1b. Leaflet blades dark grayish green 

when dry, adaxially hirsutulous .......... 35b. var. pubescens 

35a. Zanthoxylum austrosinense var. austrosinense 

岭南花椒(原变种) ling nan hua jiao (yuan bian zhong) 

Zanthoxylum austrosinense var. stenophyllum C. C. Huang. 

Leaflet blades dark reddish brown to blackish brown when 

dry, glabrous. 

● Hillside open forests and thickets; 300–900 m. S Anhui, Fujian, 

N Guangdong, NE Guangxi, SW Hubei, Hunan, Jiangxi, Zhejiang. 

35b. Zanthoxylum austrosinense var. pubescens C. C. Huang, 


毛叶岭南花椒 mao ye ling nan hua jiao 

Leaflet blades dark grayish green when dry, adaxially hirsutulous. 

● Upland thickets; ca. 1700 m. Hunan (Sangzhi). 

36. Zanthoxylum motuoense C. C. Huang, Acta Phytotax. Sin. 

16(2): 83. 1978. 

墨脱花椒 mo tuo hua jiao 

Trees to 15 m tall, deciduous. Branchlets with prickles near 

leaf scars. Leaves 3- or 5-foliolate; rachis not winged, unarmed, 

with trichomes; leaflet blades broadly obovate to broadly 

elliptic, 3–6 × 2–4 cm but terminal one to 9 × 6 cm, both 

surfaces pubescent, oil glands inconspicuous, base subrounded 

to cuneate, margin crenulate, apex subrounded or rarely acuminate. 

Infructescences paniculate, 4–8 cm. Fruit pedicel with 

short trichomes; follicles elliptic, ca. 4.5 mm in diam., with 

large protruding oil glands. Seeds ca. 4 mm in diam. Fr. Sep– 

Oct. 

● Upland thickets; ca. 1100 m. SE Xizang (Mêdog). 

37. Zanthoxylum piasezkii Maximowicz, Trudy Imp. S.-Peterburgsk. 

Bot. Sada 11: 93. 1889. 

川陕花椒 chuan shan hua jiao 

Shrubs or trees, 1–3 m tall, all parts glabrous. Stems and 

branches with brownish red prickles. Leaves 7–17-foliolate; 

rachis marginate; leaflet blades sessile, orbicular, broadly elliptic, 

or obovate-rhombic, 3–25 × 3–8 mm, thickly leathery, pale 

brown to blackish brown when dry, midvein impressed, secondary 

veins inconspicuous, base symmetric or slightly oblique, 


margin apically crenate. Inflorescences terminal. Perianth in 2 


Tepals broadly deltoid, ca. 1.5 mm or longer. Male flowers: 

pedicel 5–8 mm; stamens 5 or 6; connective blackish brown 

when dry, with oil gland on tip; rudimentary gynoecium projecting 

outward and cushion-shaped. Female flowers: carpels 2 

or 3(or 4); styles recurved. Fruit follicles purplish red, 4–5 mm in 

diam., with a few protruding oil glands. Seeds 3–4 mm in diam. 

Fl. May, fr. Jun–Jul. 

● 1700–2500 m. S Gansu, W Henan, S Shaanxi, Sichuan. 

38. Zanthoxylum pilosulum Rehder & E. H. Wilson in Sargent, 

Pl. Wilson. 2: 123. 1914. 

微柔毛花椒 wei rou mao hua jiao 

Shrubs 1–2 m tall. Branchlets with prickles, puberulent 

when young. Leaves (5 or)7–11-foliolate; leaflet blades sessile, 

ovate to ovate-elliptic, 0.5–3 × 0.4–1.5 cm but terminal one to 5 

× 2.5 cm, thinly papery, grayish green when dry, oil glands 

inconspicuous, base symmetric, margin irregularly and remotely 

crenulate, apex mucronate to acuminate. Inflorescences terminal; 

rachis with short trichomes. Perianth in 2 irregular series or 

1 series, with 5–8 ± undifferentiated tepals. Male flowers: tepals 

lanceolate, 1.2–1.5 mm; stamens 5 or 6. Female flowers: carpels 

(2–)4. Fruit follicles purplish red, 4–5 mm in diam., with sparse 

protruding small oil glands. Fl. Apr–May, fr. Jul–Aug. 

● 2500–3100 m. S Gansu, S Shaanxi, NE and W Sichuan, Yunnan. 

39. Zanthoxylum pteracanthum Rehder & E. H. Wilson in 

Sargent, Pl. Wilson. 2: 123. 1914. 

翼刺花椒 yi ci hua jiao 

Shrubs or trees, 2–3 m tall. Branchlets grayish green, 

glabrous, densely covered with setiform prickles and strongly 

compressed pseudostipular decurrent prickles forming a wing 

which joins them between nodes. Leaves 7–13-foliolate; rachis 

marginate, glabrous; leaflet blades subsessile, opposite or subopposite, 

lanceolate to ovate-oblong, 2–4 × 0.8–1.5 cm, oil 

glands inconspicuous, base slightly oblique and subrounded to 

broadly cuneate, margin apically crenulate, apex acuminate with 

a mucronate, obtuse, or emarginate tip. Inflorescences terminal; 

rachis base with prickles. Fruit pedicel 5–8 mm; developed 

follicles 1–3 with carpels in female flowers at least 3, ca. 5 mm, 

pubescent. Seeds 3–4 mm. 

● Upland thickets; ca. 1000 m. W Hubei (Xingshan). 

40. Zanthoxylum simulans Hance, Ann. Sci. Nat., Bot., sér. 5, 

5: 208. 1866. 

野花椒 ye hua jiao 

Fagara podocarpa (Hemsley) Engler; F. setosa (Hemsley) 

Engler; Zanthoxylum acanthophyllum Hayata; Z. argyi H. 

Léveillé; Z. bungei Hance var. inermis Franchet; Z. podocarpum 

Hemsley; Z. setosum Hemsley; Z. simulans var. podocarpum 

(Hemsley) C. C. Huang. 

Shrubs or small trees. Stems and branchlets with prickles. 

Young branchlets and leaflet blades abaxially on midvein pubescent, 

sometimes also secondary veins pubescent, sometimes 

65

66 

all parts glabrous. Leaves 5–15-foliolate; rachis winged; leaflet 

blades sessile or at base of leaf rachis with a short petiolule, 

opposite, ovate, ovate-elliptic, or lanceolate, 2.5–7 × 1.5–4 cm, 

adaxially with spines, oil glands numerous, translucent, and 

slightly protruding when dry, midvein impressed, base slightly 

oblique, margin crenate, apex acute to mucronate or with a retuse 

tip. Inflorescences terminal, 1–5 cm. Perianth in 2 irregular 

series or 1 series, with 5–8 ± undifferentiated tepals. Tepals pale 

yellowish green, narrowly lanceolate, broadly ovate, or subdeltoid, 

ca. 2 mm. Male flowers: stamens 5–8(–10); filaments pale 

green; connective with an oil gland at apex; rudimentary gynoecium 

pale green. Female flowers: carpels 2 or 3; styles recurved. 

Fruit follicles reddish brown, ca. 5 mm in diam., base attenuate 

into a 1–2 mm stipe, oil glands numerous and slightly protruding. 

Seeds 4–4.5 mm. Fl. Mar–May, fr. Jul–Sep. 2n = ca. 132*. 

● Plains or upland forests. Anhui, Fujian, Gansu, N Guangdong, 

NE Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Qinghai, 

Shaanxi, Shandong, Taiwan, Zhejiang. 

41. Zanthoxylum stipitatum C. C. Huang, Guihaia 7: 2. 1987. 


2. ORIXA Thunberg, Nov. Gen. Pl. 3: 56. 1783. 

臭常山属 chou chang shan shu 


梗花椒 geng hua jiao 

Shrubs or trees, 1–3 m tall, glabrous except for leaflet 

blades. Prickles to 1.5 cm. Branchlets blackish brown when dry. 

Leaves 7–17-foliolate; leaflet blades subsessile or with petiolule 

to 2 mm, opposite, lanceolate to ovate but suborbicular at base of 

leaf rachis, 1–5 × 0.7–1.3 cm, papery, abaxially purplish red 

when young, turning grayish green when mature, and reddish 

brown to dark blackish red when dry, oil glands sparse and 

protruding on both surfaces when dry, midvein abaxially rustcolored 

flocculent and adaxially impressed, margin serrulate. 

Inflorescences terminal. Perianth in 2 irregular series or 1 series, 

with 6–8 ± undifferentiated tepals. Tepals usually lanceolate, 

2–3 mm. Male flowers: stamens 5–8; connective with oil gland 

at tip. Female flowers: carpels 3 or 4; styles recurved. Rachis of 

infructescences purplish red. Fruit pedicel purplish red, 5–8(–10) 

mm; follicles purplish red, ca. 5 × 4 mm, base attenuate into a 

1–3 mm stipe, oil glands slightly protruding when dry. Seeds ca. 

4 × 3.5 mm. Fl. Apr–May, fr. Jul–Aug. 

● Forests; 100–800 m. NW Fujian, N Guangdong, NE Guangxi, S 

Hunan. 

Shrubs or small trees, deciduous, dioecious, unarmed. Buds with imbricate scales. Leaves alternate, simple. Inflorescences 

axillary, between leaves or basal to leaves, racemose or in female plants often reduced to solitary flowers. Sepals 4, basally connate. 

Petals 4, imbricate in bud. Stamens 4 but rudimentary in female flowers, distinct. Disk 4-lobed, broadly flattened in male flowers. 

Gynoecium 4-carpelled but rudimentary or lacking in male flowers; ovaries basally connate, otherwise coherent, each with 1 ovule; 

style terminal, of 4 coherent stylar elements; stigma capitate, 4-lobed. Fruit of 1–4 basally connate follicles each with a stylar beak; 

endocarp cartilaginous, discharged elastically with seed at dehiscence. Seeds black, subglobose to broadly ellipsoid, glossy; seed coat 

sclerenchymatous, thin and brittle; endosperm ± scant; embryo straight; cotyledons suborbicular, flattened; hypocotyl partly included 

between cotyledons. 

One species: E Asia. 

1. Orixa japonica Thunberg, Nov. Gen. Pl. 3: 57. 1783. 

臭常山 chou chang shan 

Celastrus orixa (Lamarck) Siebold & Zuccarini, nom. superfl.; 

Euodia ramiflora A. Gray; Ilex orixa (Lamarck) Sprengel; 

Orixa racemosa Z. M. Tan; O. subcoriacea Z. M. Tan; 

Othera orixa Lamarck; Sabia cavaleriei H. Léveillé; S. feddei H. 

Léveillé. 

Shrubs or trees, to 3 m tall. Petiole 3–8 mm; leaf blade 

obovate to elliptic, 4–15 × 2–6 cm, papery, base obtuse, cuneate, 

or attenuate, apex obtuse, acute, or acuminate. Sepals 1–1.5 mm. 

Petals greenish, 3–4 mm. Fruit follicles 8–10 mm, ribbed, apex 

usually with a stylar beak. Seeds 4–4.5 mm in diam. Fl. Apr– 

May, fr. Sep–Nov. 

Open or dense forests on sunny slopes; 500–1300 m. Anhui, Fujian, 

Guizhou, S Henan, Hubei, NW Hunan, Jiangsu, N Jiangxi, SE Shaanxi, 

Sichuan, NW Yunnan, Zhejiang [Japan, S Korea]. 

3. TETRADIUM Loureiro, Fl. Cochinch. 1: 91. 1790. 

四数花属 si shu hua shu 


Boymia A. Jussieu; Megabotrya Hance ex Walpers; Philagonia Blume. 

Shrubs or trees, evergreen or deciduous, dioecious or rarely polygamo-dioecious. Axillary buds exposed. Leaves opposite, odd- 

pinnate (occasional leaves even-pinnate); lateral leaflet blades often ± inequilateral, especially at base. Inflorescences terminal or 

terminal and axillary, thyrsiform. Sepals 4 or 5, basally connate. Petals 4 or 5, narrowly imbricate in bud. Stamens 4 or 5, distinct. Male 

flowers: stamens to 1.5 × as long as petals; disk conic to cylindric or occasionally barrel-shaped; gynoecium rudimentary, of 4 or 5 

basally connate, divergent, and fingerlike carpels. Female flowers: stamens rudimentary, ligulate, much shorter than petals or some-


times lacking; disk pulvinate to barrel-shaped; gynoecium 4- or 5-carpelled; ovaries basally connate, otherwise contiguous, each with 1 

or 2 ovules; style apical, of 4 or 5 ± contiguous stylar elements; stigma peltate. Fruit of 1–5 basally connate follicles with abortive 

carpels, if any, persistent; outer part of pericarp (exocarp and mesocarp) dry or ± fleshy; endocarp cartilaginous. Seeds remaining 

attached in dehisced fruit; seed coat (except in Tetradium daniellii and T. calcicola) with thick inner layer of dense black sclerenchyma 

and spongy outer layer bounded externally by a shiny black or reddish pellicle; endosperm copious; embryo straight; cotyledons 

broadly elliptic, ± flattened; hypocotyl superior. 

Nine species: E, S, and SE Asia; seven species (one endemic) in China. 

Euodia lyi H. Léveillé (Bull. Géogr. Bot. 24: 142. 1914) is a synonym of Miliusa sinensis Finett & Gagnepain in the Annonaceae (see Fl. China 7). 

Euodia chaffanjonii H. Léveillé (Repert. Spec. Nov. Regni Veg. 13: 265. 1914) is a synonym of Euscaphis japonica (Thunberg) Dippel in the 

Staphyleaceae (see p. 498). 

Euodia was long confused with Tetradium and Melicope. Euodia comprises seven species native to NE Australia, New Guinea, and SW Pacific 

islands. 

1a. Plants flowering. 

2a. Flowers 4(or 5)-merous. 

3a. Leaflet blade secondary veins 13–22 on each side of midvein, margin crenulate or rarely entire; 

ovules 2 per carpel, subcollateral ............................................................................................................... 1. T. fraxinifolium 

3b. Leaflet blade secondary veins 11–14 on each side of midvein, margin entire; ovules 2 per carpel, 

collateral ..................................................................................................................................................... 5. T. trichotomum 

2b. Flowers (4 or)5-merous. 

4a. Ovules 1 per carpel; leaflet blades abaxially finely papillate ................................................................... 7. T. austrosinense 

4b. Ovules 2 per carpel; leaflet blades not abaxially papillate. 

5a. Ovules superposed. 

6a. Petals white or whitish, drying whitish or pale brown, outside glabrous; leaflet blade margins 

subentire to crenulate ............................................................................................................................ 2. T. daniellii 

6b. Petals purple, drying dull purplish red, outside sparsely to ± densely appressed pubescent; leaflet 

blade margins entire ............................................................................................................................ 3. T. calcicola 

5b. Ovules collateral or subcollateral. 

7a. Sepals ca. 0.5 mm; leaflet blades abaxially usually glaucous, reticulate veinlets dense and 

clearly defined ................................................................................................................................ 4. T. glabrifolium 

7b. Sepals 0.5–1.2 mm; leaflet blades abaxially rarely slightly glaucous, reticulate veinlets ± 

loose and ± inconspicuous ............................................................................................................... 6. T. ruticarpum 

1b. Plants fruiting. 

8a. Fruit (including abortive carpels, if any) 4(or 5)-carpelled. 

9a. Seeds 2 per follicle, subtrigonous, attached in dehisced follicle to a fleshy funiculate aril ..................... 1. T. fraxinifolium 

9b. Seeds 1 per follicle, paired with an abortive seed, subglobose to broadly ovoid, attached in dehisced 

follicle to an axile strip of pergamentaceous pericarp tissue ..................................................................... 5. T. trichotomum 

8b. Fruit (including abortive carpels, if any) (4 or)5-carpelled. 

10a. Fruit follicles 2-seeded. 

11a. Fruit follicles 5–11 mm, sparsely to densely spreading pubescent, apex beaked ................................. 2. T. daniellii 

11b. Fruit follicles ca. 5 mm, sparsely to densely appressed pubescent laterally, otherwise glabrate, 

apex not beaked .................................................................................................................................... 3. T. calcicola 

10b. Fruit follicles 1-seeded. 

12a. Seed not paired with an abortive seed; follicles ca. 3.5 mm ........................................................ 7. T. austrosinense 

12b. Seed paired with an abortive seed; follicles 3.5–6 mm. 

13a. Fruit follicles trigonous, sparsely to densely appressed pubescent laterally, glabrous 

abaxially, endocarp sparsely to ± densely pubescent ............................................................ 4. T. glabrifolium 

13b. Fruit follicles subglobose, glabrous or sometimes with sparse trichomes laterally and/or 

abaxially, endocarp glabrous ................................................................................................... 6. T. ruticarpum 

1. Tetradium fraxinifolium (Hooker) T. G. Hartley, Gard. 

Bull. Singapore 34: 102. 1981. 

无腺吴萸 wu xian wu yu 

Philagonia fraxinifolia Hooker, Icon. Pl. t. 710. 1845; 

Euodia fraxinifolia (Hooker) Bentham; E. impellucida Handel- 

Mazzetti; E. impellucida var. macrococca C. C. Huang; E. 

poilanei Guillaumin; E. subtrigonosperma C. C. Huang. 

Trees to 12 m tall. Leaves 23–67 cm, 5–15-foliolate; leaflet 

blades lanceolate or occasionally ovate, elliptic-oblong, or elliptic, 

9–25 × 3.5–8.5 cm, abaxially often glaucous and sometimes 

67

68 

subpapillate, secondary veins 13–22 on each side of midvein, 

reticulate veinlets abaxially usually inconspicuous, base in 

lateral leaflets rounded to narrowly cuneate and in terminal 

leaflet cuneate, margin crenulate or rarely entire, apex acuminate. 

Inflorescences 7–24 cm. Flowers 4(or 5)-merous. Sepals 

0.5–1.5 mm. Petals green to pale yellow but drying brown, 

3.5–6.5 mm, outside glabrous, inside glabrous to villous. Ovary 

glabrous or appressed pubescent; ovules 2 per carpel, subcollateral. 

Fruit (including abortive carpels, if any) usually 

4-carpelled; follicles compressed subglobose, 5.5–10 mm, glabrous 

to sparsely pubescent, apex not beaked; endocarp glabrous 

or sparsely pubescent. Seeds 2 per follicle, subtrigonous, 4–5 

mm, attached in dehisced follicle to a fleshy funiculate aril; seed 

coat with thick inner layer of dense black sclerenchyma and 

spongy outer layer bounded externally by a reddish brown to 

brownish black shiny pellicle. Fl. May and Nov, fr. Jul–Nov. 

Forests, thickets; 700–3000 m. SE Xizang, Yunnan [Bhutan, India, 

Myanmar, Nepal, Thailand, N Vietnam]. 

2. Tetradium daniellii (Bennett) T. G. Hartley, Gard. Bull. 

Singapore 34: 105. 1981. 

臭檀吴萸 chou tan wu yu 

Zanthoxylum daniellii Bennett, Ann. Mag. Nat. Hist., ser. 

3, 10: 201. 1862; Ampacus daniellii (Bennett) Kuntze; Euodia 

baberi Rehder & E. H. Wilson, p.p. as to E. H. Wilson 1164 p.p. 

Jul 1908 (see also synonymy of Tetradium ruticarpum); E. daniellii 

(Bennett) Hemsley; E. daniellii var. delavayi (Dode) C. C. 

Huang; E. daniellii var. henryi (Dode) C. C. Huang; E. daniellii 

var. hupehensis (Dode) C. C. Huang; E. daniellii var. labordei 

(Dode) C. C. Huang; E. daniellii var. villicarpa (Rehder & E. H. 

Wilson) C. C. Huang; E. delavayi Dode; E. henryi Dode; E. 

henryi var. villicarpa Rehder & E. H. Wilson; E. hupehensis 

Dode; E. labordei Dode; E. sutchuenensis Dode, p.p. as to R. P. 

Farges 1284 p.p. (see also synonymy of Phellodendron chinense 

var. chinense); E. velutina Rehder & E. H. Wilson; E. vestita 

W. W. Smith; Z. bretschneideri Maximowicz. 

Shrubs or trees, to 20 m tall. Leaves 15–44 cm, 5–9(or 

11)-foliolate; leaflet blades broadly ovate to lanceolate or rarely 

elliptic or elliptic-oblong, 5–18.5 × 2.5–10.5 cm, abaxially 

sometimes ± glaucous and not papillate, secondary veins 7–14 

on each side of midvein, reticulate veinlets abaxially usually 

inconspicuous, base in lateral leaflets narrowly cuneate, subtruncate, 

subrounded, or subcordate and in terminal leaflet narrowly 

cuneate, cuneate, or occasionally rounded, margin subentire 

to crenulate, apex acuminate. Inflorescences 3.5–19 cm. 

Flowers (4 or)5-merous. Sepals 0.5–1.5 mm. Petals white or 

whitish but drying whitish or pale brown, (2.5–)3–5 mm, outside 

glabrous, inside sparsely or sometimes densely pubescent or 

villous. Ovary pubescent between carpels, otherwise ± densely 

pubescent to glabrous; ovules 2 per carpel, superposed. Fruit 

(including abortive carpels, if any) usually 5-carpelled; follicles 

narrowly pyriform, 5–11 mm, sparsely to ± densely spreading 

pubescent, apex beaked; endocarp glabrous. Seeds 2 per follicle, 

superposed, black, ovoid to ellipsoid, shiny, attached in dehisced 

fruit to axile strip of pergamentaceous pericarp tissue; basal seed 

sterile, usually 1.5–3.5 mm, containing what appears to be en- 


dosperm but no embryo; apical seed fertile, 2.5–4 mm; seed coat 

sclerenchymatous, lacking spongy tissue, in fertile seed thick 

and resistant to breakage, in sterile seed thin and brittle. Fl. 

Jun–Aug, fr. Aug–Nov. 2n = 76, 78. 

Forests, forest margins, open slopes; near sea level to 3200 m. 

Anhui, S Gansu, Guizhou, Hebei, Henan, Hubei, Jiangsu, Liaoning, S 

Ningxia, E Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, SE Xizang, 

Yunnan [Korea]. 

3. Tetradium calcicola (Chun ex C. C. Huang) T. G. Hartley, 

Gard. Bull. Singapore 34: 108. 1981. 

石山吴萸 shi shan wu yu 

Euodia calcicola Chun ex C. C. Huang, Acta Phytotax. Sin. 

6: 120. 1957. 

Shrubs or trees, to 15 m tall. Leaves 9–19 cm, 5-foliolate; 

leaflet blades ovate to elliptic, 5–9.5 × 2.3–4.5 cm, abaxially ± 

glaucous and not papillate, secondary veins 10–14 on each side 

of midvein, reticulate veinlets abaxially usually inconspicuous, 

base in lateral leaflets and in terminal leaflet obtuse to rounded, 

margin entire, apex acuminate. Inflorescences 5.5–13 cm. 

Flowers 5-merous. Sepals 0.7–1 mm. Petals purple but drying 

dull purplish red, 3–4 mm, outside sparsely to ± densely appressed 

pubescent, inside villous. Ovary sparsely to ± densely 

appressed pubescent; ovules 2 per carpel, superposed. Fruit 

(including abortive carpels, if any) 5-carpelled; follicles 

pyriform, ca. 5 mm, laterally densely appressed pubescent, 

otherwise glabrate, apex not beaked; endocarp glabrous. Seeds 2 

per follicle, superposed, black, ellipsoid, shiny, attached in 

dehisced fruit to axile strip of pergamentaceous pericarp tissue; 

basal seed sterile, ca. 2 mm, containing what appears to be 

endosperm but no embryo; apical seed fertile, ca. 3 mm; seed 

coat sclerenchymatous, lacking spongy tissue, in sterile seed thin 

and brittle, in fertile seed thick and resistant to breakage. Fl. 

Jun–Sep, fr. Sep–Dec. 

● Forests, thickets; 600–800 m. N and W Guangxi, S Guizhou, SE 

Yunnan. 

4. Tetradium glabrifolium (Champion ex Bentham) T. G. 

Hartley, Gard. Bull. Singapore 34: 109. 1981. 

楝叶吴萸 lian ye wu yu 

Boymia glabrifolia Champion ex Bentham, Hooker’s J. 

Bot. Kew Gard. Misc. 3: 330. 1851; Ampacus meliifolia (Hance 

ex Walpers) Kuntze; Euodia ailantifolia Pierre; E. balansae 

Dode; E. fargesii Dode; E. glabrifolia (Champion ex Bentham) 

C. C. Huang; E. glauca Miquel; E. meliifolia (Hance ex Walpers) 

Bentham; E. taiwanensis T. Yamazaki; E. yunnanensis C. 

C. Huang; Eurycoma dubia Elmer; Megabotrya meliifolia Hance 

ex Walpers; Phellodendron burkillii Steenis; Tetradium glabrifolium 

var. glaucum (Miquel) T. Yamazaki; T. taiwanense (T. 

Yamazaki) T. Yamazaki. 

Shrubs or trees, to 20 m tall. Leaves 14–38 cm, (3 

or)5–19-foliolate; leaflet blades broadly ovate to lanceolate or 

less often elliptic or elliptic-oblong, 4–15 × 1.7–6 cm, abaxially 

usually glaucous and not papillate, secondary veins 8–18 on 

each side of midvein, reticulate veinlets abaxially clearly defined 

and dense, base in lateral leaflets narrowly cuneate to sub-

ounded to subtruncate and in terminal leaflet narrowly cuneate 

to cuneate, margin entire or ± crenulate, apex acuminate. Inflorescences 

9–19 cm. Flowers (4 or)5-merous. Sepals ca. 0.5 mm. 

Petals green, yellow, or white but drying whitish to brown, 2.7–4 

mm, outside glabrous or sometimes with sparse appressed trichomes, 

inside nearly glabrous to villous. Ovary ± densely pubescent 

between carpels, otherwise finely pubescent toward 

apex or glabrous; ovules 2 per carpel, collateral or subcollateral. 

Fruit (including abortive carpels, if any) usually 5-carpelled; 

follicles trigonous, 3.5–5 mm, laterally sparsely to densely appressed 

pubescent, otherwise glabrous, apex not beaked; endocarp 

sparsely to ± densely pubescent. Seeds 1 per follicle but 

paired with an abortive seed, subglobose to ovoid to broadly 

ellipsoid, 2.5–4 mm, attached in dehisced follicle to axile strip of 

pergamentaceous pericarp tissue; seed coat with thick inner layer 

of dense black sclerenchyma and spongy outer layer bounded 

externally by a shiny black pellicle. Fl. Jun–Sep, fr. Sep–Dec. 

Forests, thickets, open places; near sea level to 1200 m. Anhui, 

Fujian, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, 

Jiangxi, S Shaanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Bhutan, India, 

Indonesia, Japan (including Ryukyu Islands), Malaysia, Myanmar, 


5. Tetradium trichotomum Loureiro, Fl. Cochinch. 1: 91. 

1790. 

牛枓吴萸 niu dou wu yu 

Ampacus trichotoma (Loureiro) Kuntze; Brucea trichotoma 

(Loureiro) Sprengel; Euodia colorata Dunn; E. hainanensis 

Merrill; E. lenticellata C. C. Huang; E. trichotoma (Loureiro) 

Elmer; E. trichotoma var. pubescens C. C. Huang; E. viridans 

Drake. 

Shrubs or trees, to 8 m tall. Leaves 12–37 cm, (3 or)5–11(or 

13)-foliolate; leaflet blades elliptic-oblong, lanceolate, or occasionally 

ovate, 3–16 × 3–5 cm, abaxially neither glaucous nor 

papillate, secondary veins 11–14 on each side of midvein, reticulate 

veinlets abaxially inconspicuous, margin entire, base in 

lateral leaflets subrounded to cuneate and in terminal leaflet 

cuneate, apex acuminate. Inflorescences 5–30 cm. Flowers 4(or 

5)-merous. Sepals 0.5–1 mm. Petals green, yellow, or white but 

drying brown to whitish, 3–4 mm, outside glabrous, inside 

glabrous or sparsely villosulous. Ovary glabrous or with a few 

short trichomes between carpels toward base; ovules 2 per 

carpel, collateral. Fruit (including abortive carpels, if any) usually 

4-carpelled; follicles subglobose to obovoid, 4–7 mm, glabrous 

or laterally with sparse pubescence toward base, apex not 

beaked; endocarp glabrous. Seeds 1 per follicle but paired with 

an abortive seed, subglobose to broadly ovoid, 3.7–6 mm, 

attached in dehisced follicle to axile strip of pergamentaceous 

pericarp tissue; seed coat with thick inner layer of dense black 

sclerenchyma and spongy outer layer bounded externally by a 

shiny black pellicle. Fl. Apr–Aug, fr. Sep–Nov. 

Forests, thickets; 300–1900 m. SE Guangdong, Guangxi, S Guizhou, 

Hainan, Hubei, Shaanxi, S Sichuan, Yunnan [Laos, Thailand, N 

Vietnam]. 

6. Tetradium ruticarpum (A. Jussieu) T. G. Hartley, Gard. 

Bull. Singapore 34: 116. 1981. 


吴茱萸 wu zhu yu 

Boymia ruticarpa A. Jussieu, Mém. Mus. Hist. Nat. 12: 

507. 1825; Ampacus ruticarpa (A. Jussieu) Kuntze; Euodia 

baberi Rehder & E. H. Wilson, p.p. as to E. H. Wilson 1164 p.p. 

Oct 1908 (see also synonymy of Tetradium daniellii); E. 

bodinieri Dode; E. compacta Handel-Mazzetti; E. compacta var. 

meionocarpa Handel-Mazzetti; E. hirsutifolia Hayata; E. 

officinalis Dode; E. rugosa Rehder & E. H. Wilson; E. ruticarpa 

(A. Jussieu) Bentham; E. ruticarpa var. bodinieri (Dode) C. C. 

Huang; E. ruticarpa var. officinalis (Dode) C. C. Huang. 

Shrubs or trees, to 9 m tall. Leaves 15–40 cm, (3 or)5–13(or 

15)-foliolate; leaflet blades elliptic to ovate or sometimes 

lanceolate, oblanceolate, or obovate, 4.5–17 × 2–8 cm, abaxially 

rarely slightly glaucous and not papillate, secondary veins 9–17 

on each side of midvein, reticulate veinlets abaxially ± inconspicuous 

and ± loose, base in lateral leaflets obtuse to narrowly 

cuneate or sometimes rounded to cuneate or rarely attenuate, 

margin entire or irregularly crenulate, apex acuminate. Inflorescences 

2.5–18 cm. Flowers (4 or)5-merous. Sepals 0.5–1.2 

mm. Petals green, yellow, or white but drying brown to whitish, 

3–5 mm, outside glabrous or with sparse short trichomes, inside 

nearly glabrous to villous. Ovary glabrous or with sparse 

trichomes; ovules 2 per carpel, collateral or subcollateral. Fruit 

(including abortive carpels, if any) usually 5-carpelled; follicles 

subglobose, 3.5–6 mm, glabrous or sometimes with sparse 

trichomes, apex not beaked; endocarp glabrous. Seeds 1 per 

follicle but paired with an abortive seed, ovoid or sometimes 

ellipsoid or subglobose, 3.5–5.5 mm, attached in dehisced 

follicle to axile strip of pergamentaceous pericarp tissue; seed 

coat with thick inner layer of dense black sclerenchyma and 

spongy outer layer bounded externally by a shiny black pellicle. 

Fl. Apr–Jun, fr. Aug–Nov. 

Forests, thickets, open places; 100–3000 m. Anhui, Fujian, S 

Gansu, Guangdong, Guangxi, Guizhou, Hebei, S Henan, Hubei, Hunan, 

S Jiangsu, Jiangxi, S Shaanxi, Sichuan, Yunnan, Zhejiang [Bhutan, NE 


7. Tetradium austrosinense (Handel-Mazzetti) T. G. Hartley, 


华南吴萸 hua nan wu yu 

Euodia austrosinensis Handel-Mazzetti, Sinensia 5: 1. 

1934. 

Trees to 20 m tall. Leaves 20–35 cm, 7–11-foliolate; leaflet 

blades broadly elliptic or sometimes ovate, lanceolate, obovate, 

or oblanceolate, 5.5–14.5(–17) × 2.5–7(–8.5) cm, abaxially 

glaucous and finely papillate, secondary veins 9–12 on each side 

of midvein, reticulate veinlets abaxially usually ± conspicuous 

and ± loose, base in lateral leaflets obtuse to nearly rounded and 

in terminal leaflet cuneate, margin entire, apex acuminate. Inflorescences 

11–18 cm. Flowers (4 or)5-merous. Sepals ca. 0.5 

mm. Petals green to greenish yellow but drying brown, 2.5–3 

mm, outside glabrous to sparsely puberulent, inside sparsely to 

densely villous. Ovary glabrous; ovules 1 per carpel. Fruit (including 

abortive carpels, if any) usually 5-carpelled; follicles 

trigonous to subtrigonous, ca. 3.5 mm, glabrous or rarely with a 

69

70 

few scattered trichomes, apex not beaked; endocarp glabrous. 

Seeds 1 per follicle and not paired with an abortive seed, ellipsoid 

to subreniform, 2.3–2.5 mm, attached in dehisced follicle to 

axile strip of pergamentaceous pericarp tissue; seed coat with 

thick inner layer of dense black sclerenchyma and spongy outer 


layer bounded externally by a shiny black pellicle. Fl. Jun–Jul, 

fr. Sep–Nov. 

Forests; 300–1500 m. Guangdong, Guangxi, S Yunnan [N Vietnam]. 

4. MELICOPE J. R. Forster & G. Forster, Char. Gen. Pl. 28. 1775. 

蜜茱萸属 mi zhu yu shu 


Shrubs, trees, rarely scandent, evergreen, usually dioecious or sometimes monoclinous or andromonoecious. Leaves opposite [or 

whorled], digitately 3-foliolate or 1-foliolate. Inflorescences axillary or basal to leaves, cymulose to thyrsiform [or reduced to solitary 

flowers]. Sepals 4, connate at base or to nearly their full length. Petals 4, valvate or narrowly imbricate in bud. Stamens 4 or 8 [or 4–8], 

rudimentary in female flowers. Disk pulvinate to annular to cup-shaped. Gynoecium 4-carpelled, rudimentary or lacking in male 

flowers; ovaries connate at base, otherwise contiguous [or connate up to their full length]; ovules [1 or] 2 per locule; style apical or 

subapical, of 4 contiguous, coherent, or connate stylar elements; stigma usually punctiform, capitellate, or capitate. Fruit of 1–4 basally 

connate follicles [or grading to a syncarpous 4-loculed loculicidal capsule]; exocarp dry to fleshy; endocarp cartilaginous. Seeds 

remaining attached in dehisced fruit; seed coat with thick inner layer of dense black sclerenchyma and spongy outer layer bounded 

externally by a shiny black pellicle; endosperm copious; embryo straight or slightly curved; cotyledons elliptic, flattened; hypocotyl 

superior. 

About 233 species: E, S, and SE Asia, Australia, Indian Ocean islands (Mascarenes), Madagascar, Pacific islands; eight species (two endemic) in 

China. 

As discussed by Hartley (Allertonia 8: 19, f. 5. 2001), two types of attachment of mature seed are seen in dehisced fruit of Melicope. In Type A, the 

attachment is by a partially detached axile strip of pericarp tissue or by a partially detached raphe, or by both. In Type B, neither the axile pericarp tissue 

nor the raphe is detached, and the seed is connected to the axile placental region by a funiculus, which is simply the funiculus of the ovule enlarged. 

1a. Stamens 8; seed attachment Type A. 

2a. Leaves 1-foliolate; follicles ellipsoid to obovoid, ca. 6.5 mm ......................................................................... 1. M. patulinervia 

2b. Leaves mostly 3-foliolate; follicles subglobose, 3–4.5 mm ................................................................................... 2. M. triphylla 

1b. Stamens 4; seed attachment Type B. 

3a. Inflorescences 0.7–1.8(–2) cm wide; apex of staminal filaments at least in male and bisexual flowers obtuse 

or narrowly so or sometimes acute; petals usually persistent in fruit; leaves 3-foliolate and/or 1-foliolate. 

4a. Petioles sparsely puberulent to pubescent adaxially, otherwise glabrous or nearly so ..................................... 7. M. viticina 

4b. Petioles pubescent to velutinous all around .................................................................................................. 8. M. glomerata 

3b. Inflorescences 1.5–16 cm wide; apex of staminal filaments at least in male and bisexual flowers subulate 

to filiform; petals usually deciduous in fruit; leaves mostly 3-foliolate. 

5a. Young branchlets 4.5–10 mm wide in third internode; leaflet blades abaxially strigillose to ± 

appressed villosulous ............................................................................................................................. 4. M. semecarpifolia 

5b. Young branchlets 2–4.5 mm wide in third internode; leaflet blades abaxially glabrous to puberulent. 

6a. Larger primary branches of inflorescences usually spreading; leaflet blades ovate-elliptic, 

elliptic, or elliptic-obovate, or narrowly so; inflorescences 2–10 × 1.5–8 cm; pedicel sparsely 

puberulent to pubescent ........................................................................................................................... 6. M. pteleifolia 

6b. Larger primary branches of inflorescences usually ascending; leaflet blades usually elliptic or 

narrowly so; inflorescences 4.5–32 × 3–16 cm; pedicel glabrous to tomentulose. 

7a. Leaflet blades elliptic, sometimes ovate-elliptic, or rarely obovate; inflorescences 

8–32 × 3–16 cm; pedicel nearly glabrous to tomentulose ......................................................... 3. M. lunu-ankenda 

7b. Leaflet blades ± narrowly elliptic; inflorescences 4.5–9 × 3–4.5 cm; pedicel glabrous ....................... 5. M. chunii 

1. Melicope patulinervia (Merrill & Chun) C. C. Huang, Acta 

Phytotax. Sin. 6: 132. 1957. 

蜜茱萸 mi zhu yu 

Euodia patulinervia Merrill & Chun, Sunyatsenia 5: 87. 

1940. 

Shrubs 1–3 m tall. Young branchlets 2–3 mm wide in third 

internode. Leaves 1-foliolate, glabrous; petiole 0.2–3 cm; leaflets 

sessile; leaflet blades elliptic, narrowly elliptic, or elliptic-obovate, 

5–15 × 1.5–6 cm, apex acute to acuminate. Inflorescences 

axillary, to 3 cm. Sepals most likely in female flowers, 

ca. 0.5 mm, outside sparsely puberulent, basally connate, 

deciduous in fruit. Petals ca. 1.5 mm, deciduous in fruit. Stamens 

8, shorter than petals. Gynoecium glabrous; ovary globose; style 

very short. Infructescences 1–2 × 1.3–2.5 cm; peduncle 3–10

mm. Pedicel 2–3.5 mm, sparsely puberulent. Fruit follicles 

ellipsoid to obovoid, ca. 6.5 mm, glabrous. Seeds ellipsoid to 

obovoid, 4–5 mm, attachment Type A. Fl. Mar–Apr, fr. Sep– 

Oct. 

● Dense or ± open forests; 700–900 m. Hainan. 

2. Melicope triphylla (Lamarck) Merrill, Philipp. J. Sci., C, 7: 

375. 1912. 

三叶蜜茱萸 san ye mi zhu yu 

Fagara triphylla Lamarck, Encycl. 2: 447. 1788; Acronychia 

minahassae (Teijsmann & Binnendijk) Miquel; Ampacus 

incerta (Blume) Kuntze; A. triphylla (Lamarck) Kuntze; Bergera 

ternata Blanco; Euodia anisodora Lauterbach & K. Schumann; 

E. awadan Hatusima; E. glaberrima Merrill; E. incerta 

Blume; E. laxireta Merrill; E. microsperma F. M. Bailey; E. 

minahassae Teijsmann & Binnendijk; E. philippinensis Merrill 

& L. M. Perry; E. triphylla (Lamarck) Candolle; Melicope 

awadan (Hatusima) Ohwi & Hatusima; M. curranii Merrill; M. 

densiflora Merrill; M. gjellerupii Lauterbach; ?M. kanehirae 

Hatusima; M. luzonensis Engler ex Perkins; M. mahonyi F. M. 

Bailey; M. mindanaensis Elmer; M. monophylla Merrill; M. 

monophylla var. glabra Elmer; M. nitida Merrill; M. obtusa 

Merrill; M. odorata Elmer; M. rupestris Lauterbach; Zanthoxylum 

triphyllum (Lamarck) G. Don. 

Shrubs or trees, 1.5–6 m tall, dioecious or sometimes monoclinous 

or andromonoecious. Young branchlets 3–4 mm wide in 

third internode. Leaves 3-foliolate (occasional leaves 1-foliolate), 

glabrous; petiole 2–6.5 cm; leaflet blades obovate, oblanceolate, 

or rarely elliptic, in terminal leaflet 5–12 × 2–4.5 cm, 

apex acuminate or rarely acute. Inflorescences axillary, 3–7 × 

2–6 cm; peduncle obsolete or to 1.5 cm. Pedicel 1.5–3 mm and 

of similar length in fruit, glabrous. Sepals 0.5–0.8 mm, glabrous 

or with a few trichomes at apex, connate at base or to ± half their 

length, persistent in fruit. Petals 2.5–3.5 mm, deciduous in fruit. 

Stamens 8, in male and bisexual flowers those opposite sepals 

2–4 mm and in female flowers 0.5–3.5 mm; filaments at least in 

male and bisexual flowers acute to subulate at apex. Gynoecium 

glabrous, in female and bisexual flowers 0.8–1.6 mm, in male 

flowers 0.3–0.5 mm. Fruit follicles subglobose, 3–4.5 mm, glabrous. 

Seeds ellipsoid to obovoid, 3–4.5 mm, attachment Type 

A. 

Forests, thickets, open places; low elevations. Taiwan [Indonesia, 

Japan (Ryukyu Islands), New Guinea, Philippines; SW Pacific islands]. 

3. Melicope lunu-ankenda (Gaertner) T. G. Hartley, Sandakania 

4: 61. 1994. 

三刈叶蜜茱萸 san yi ye mi zhu yu 

Fagara lunu-ankenda Gaertner, Fruct. Sem. Pl. 1: 334. 

1788; Ampacus aromatica (Blume) Kuntze; Euodia arborea 

Elmer; E. arborescens D. D. Tao; E. aromatica Blume; E. 

concinna Ridley; E. lucida (Miquel) Miquel; E. lunu-ankenda 

(Gaertner) Merrill; E. lunu-ankenda var. tirunelvelica A. N. 

Henry & Chandrabose; E. marambong (Miquel) Miquel; E. 

obtusifolia Ridley; E. punctata Merrill; E. roxburghiana (Chamisso) 

Bentham; E. roxburghiana var. longipes Craib; E. triphylla 

(Lamarck) Candolle var. pubescens Ridley; Zanthoxylum 


aromaticum (Blume) Miquel; Z. lucidum Miquel; Z. marambong 

Miquel; Z. roxburghianum Chamisso. 

Trees to 30 m tall, dioecious. Young branchlets 2.5–4.5 

mm wide in third internode. Leaves 3-foliolate; petiole 2–12 cm, 

glabrous to tomentulose; leaflet blades elliptic, sometimes ovate- 

elliptic, or rarely obovate, in terminal leaflet 6–21 × 2.5–8.5 cm, 

glabrous or puberulent to sparsely puberulent on midvein especially 

abaxially and sometimes on secondary veins abaxially, 

apex acuminate. Inflorescences axillary, 8–32 × 3–16 cm; 

peduncle 4–15 cm; larger primary branches usually ascending. 

Pedicel 0.3–2.5 mm but 1–5 mm in fruit, nearly glabrous to 

tomentulose. Sepals 0.5–1 mm, connate at base or to ca. 1/4 their 

length, glabrous or outside puberulent, persistent in fruit. Petals 

1.5–2.5 mm, deciduous or subpersistent in fruit. Stamens 4, in 

male flowers 2–4 mm, in female flowers 0.5–2.5 mm; filaments 

at least in male flowers subulate to filiform at apex. Gynoecium 

in female flowers 1–3 mm, in male flowers 0.2–0.5 mm; ovary 

nearly glabrous to shortly pubescent or ± sparsely villous. Fruit 

follicles ellipsoid, 4.5–5 mm, glabrous to sparsely puberulent. 

Seeds subglobose, ovoid, or ellipsoid, 3.5–4 mm, attachment 

Type B. Fr. Aug–Sep. 

Montane forests; ca. 900 m. SE Xizang (Mêdog) [Bhutan, Cambodia, 

India, Indonesia, Malaysia, Myanmar, Nepal, Philippines, Sri 

Lanka, Thailand, Vietnam]. 

4. Melicope semecarpifolia (Merrill) T. G. Hartley, Fl. Taiwan, 

ed. 2, 3: 522. 1993. 

台湾蜜茱萸 tai wan mi zhu yu 

Euodia semecarpifolia Merrill, Publ. Bur. Sci. Gov. Lab. 

35: 23. 1906; E. camiguinensis Merrill; E. merrillii Kanehira & 

Sasaki; E. retusa Merrill. 

Small trees, dioecious. Young branchlets 4.5–10 mm wide 

in third internode. Leaves 3-foliolate; petiole 3.5–14 cm, ± 

appressed pilose or appressed villous when young; leaflet blades 

elliptic to obovate, in terminal leaflet 7.5–30 × 3–12 cm, both 

surfaces strigillose to ± appressed villosulous on midvein and 

often abaxially on secondary veins, apex acuminate or sometimes 

rounded or acute. Inflorescences axillary and sometimes 

basal to leaves, 3–9.5 × 2.5–8 cm; peduncle obsolete or to 3.5 

cm; larger primary branches usually spreading. Pedicel obsolete 

or to 2.5 mm, but 0.5–3 mm in fruit, glabrous to villosulous. 

Sepals 0.8–1 mm, connate at base or to ca. 1/4 their length, 

nearly glabrous to outside sericeous, persistent in fruit. Petals 

2–2.5 mm, deciduous or subpersistent in fruit. Stamens 4, in 

male flowers 3.5–4.5 mm, in female flowers 1–2 mm; filaments 

at least in male flowers subulate to filiform at apex. Gynoecium 

in female flowers ca. 1.5 mm, in male flowers 0.5–1 mm; ovary 

sericeous. Fruit follicles subglobose, 4–4.5 mm, nearly glabrous 

or sparsely pilose. Seeds subglobose to ovoid to ellipsoid, 3.5–4 

mm, attachment Type B. Fl. Jun–Jul, fr. Nov–Dec. 

Forests, thickets; lower and middle elevations. Taiwan [Philippines]. 

5. Melicope chunii (Merrill) T. G. Hartley, Allertonia 8: 237. 

2001. 

海南蜜茱萸 hai nan mi zhu yu 

71

72 

Euodia chunii Merrill, J. Arnold Arbor. 6: 132. 1925; E. 

lepta Merrill var. chunii (Merrill) C. C. Huang. 

Shrubs or trees, 1.5–12 m tall, dioecious. Young branchlets 

2–3 mm wide in third internode. Leaves 3-foliolate; petiole 2–6 

cm, glabrous or sparsely puberulent; leaflet blades ± narrowly 

elliptic, in terminal leaflet 5–12 × 1.5–3.5 cm, glabrous, apex 

acuminate or sometimes acute. Inflorescences axillary, 4.5–9 × 

3–4.5 cm; peduncle 2–3.5 cm; larger primary branches ascending. 

Pedicel 2–3.5 mm but 3–4 mm in fruit, glabrous. Sepals 

0.5–0.7 mm, connate at base or to ca. 1/4 their length, glabrous 

or margin irregularly ciliolate, persistent in fruit. Petals 2–2.5 

mm, deciduous in fruit. Stamens 4, in male flowers 2.5–3 mm, in 

female flowers 1.5–2.5 mm; filaments at least in male flowers 

subulate to filiform at apex. Gynoecium in female flowers 1.5–2 

mm, in male flowers 0.2–0.5 mm; ovary sparsely hirsutulous. 

Fruit follicles ellipsoid or obovoid, 5–6 mm, glabrate. Seeds 

subglobose to ellipsoid, 2.5–3.6 mm, attachment Type B. 

● Forests; below 400 m. Hainan. 

6. Melicope pteleifolia (Champion ex Bentham) T. G. Hartley, 

Fl. Taiwan, ed. 2, 3: 521. 1993. 

三桠苦 san ya ku 

Zanthoxylum pteleifolium Champion ex Bentham, Hooker’s 

J. Bot. Kew Gard. Misc. 3: 350. 1851; Euodia gracilis Kurz; 

E. lepta Merrill, nom. illeg. superfl.; E. lepta var. cambodiana 

(Pierre) C. C. Huang; E. oreophila Guillaumin (1945), not Guillaumin 

(1938); E. pteleifolia (Champion ex Bentham) Merrill; E. 

triphylla (Lamarck) Candolle var. cambodiana Pierre; Ilex lepta 

Sprengel, nom. illeg. superfl.; Lepta triphylla Loureiro. 

Shrubs or trees, rarely scandent, 1–14 m tall, dioecious or 

rarely monoclinous. Young branchlets 2.5–4 mm wide in third 

internode. Leaves 3-foliolate (occasional leaves 1-foliolate); 

petiole 1.5–14 cm, glabrous to pubescent; leaflet blades ovate- 

elliptic, elliptic, elliptic-obovate, or narrowly so, in terminal 

leaflet 6–22 × 2–8 cm, glabrous or puberulent on midvein especially 

abaxially, apex acuminate or subcaudate. Inflorescences 

axillary and sometimes basal to leaves, 2–10 × 1.5–8 cm; 

peduncle 0.4–5 cm; larger primary branches usually spreading. 

Pedicel 0.6–3 mm but 1.5–6 mm in fruit, sparsely puberulent to 

pubescent. Sepals 0.5–1(–1.5) mm, connate at base or to ca. 1/4 

their length, glabrous or outside pubescent, persistent in fruit. 

Petals 1.3–2(–2.5) mm, deciduous or rarely persistent in fruit. 

Stamens 4, in male and bisexual flowers 1.5–3(–4) mm, in female 

flowers 0.4–1.5 mm; filaments at least in male and bisexual 

flowers subulate to filiform at apex. Gynoecium in female and 

bisexual flowers 1–2 mm, in male flowers 0.2–0.5 mm; ovary 

glabrous to pubescent. Fruit follicles subglobose to ellipsoid to 

obovoid, (3–)3.5–7.5 mm, glabrous or sparsely puberulent. 

Seeds subglobose to ovoid to ellipsoid, 2.5–4 mm, attachment 

Type B. Fl. Apr–Jun, fr. Jul–Oct. 

Evergreen and mixed deciduous forests and forest margins, thickets, 

swamp forests, open places; near sea level to 2300(–2800) m. Fujian, 

Guangdong, SW Guangxi, Hainan, S Jiangxi, Taiwan, Yunnan, SE 

Zhejiang [Cambodia, Laos, Myanmar, Thailand, Vietnam]. 

7. Melicope viticina (Wallich ex Kurz) T. G. Hartley, Allertonia 

8: 262. 2001. 


单叶蜜茱萸 dan ye mi zhu yu 

Euodia viticina Wallich ex Kurz, J. Asiat. Soc. Bengal, Pt. 

2, Nat. Hist. 42: 64. 1873. 

Shrubs or trees, 0.8–6 m tall, dioecious or sometimes 

monoclinous. Young branchlets 1–3 mm wide in third internode. 

Leaves 1- or 3-foliolate. Leaves 3-foliolate: petiole 1.5–7 cm, 

adaxially sparsely puberulent to pubescent, otherwise glabrous 

or nearly so; leaflet blades elliptic or elliptic-obovate, or narrowly 

so, or rarely obovate or oblanceolate, in terminal leaflet 

5.5–21 × 1.5–6.5 cm, glabrous or sparsely puberulent to shortly 

pubescent on midvein, apex acuminate or rarely acute. Leaves 

1-foliolate: petiole 0.3–4.5 cm; leaflet blades elliptic or elliptic-obovate, 

or narrowly so, 2.2–22 × 0.7–7.5 cm, otherwise 

similar to 3-foliolate leaves. Inflorescences axillary and sometimes 

basal to leaves, 0.5–5(–8.5) × 0.7–1.8(–2) cm; peduncle 

obsolete or to 2.5 cm; larger primary branches spreading or 

ascending. Pedicel 0.5–1.5 mm but 1.5–3 mm in fruit, glabrous 

to sparsely puberulent. Sepals 1–1.8 mm, connate at base, 

glabrous or outside sparsely puberulent, persistent in fruit. Petals 

2.5–3 mm, persistent or rarely deciduous in fruit. Stamens 4, in 

male and bisexual flowers 2.2–3 mm, in female flowers 0.8–1.5 

mm; filaments at least in male and bisexual flowers obtuse or 

narrowly so at apex. Gynoecium in female and bisexual flowers 

1.5–2 mm, in male flowers 0.3–0.6 mm; ovary glabrous or rarely 

sparsely pubescent. Fruit follicles subglobose to ellipsoid, 5–7 

mm, glabrous. Seeds ovoid to ellipsoid, 4–6.5 mm, attachment 

Type B. Fl. Apr–May, fr. Aug–Oct. 

Open forests; 500–1300 m. S Yunnan [Cambodia, Laos, Myanmar, 

Thailand, Vietnam]. 

Plants mistakenly treated as Euodia simplicifolia Ridley var. simplicifolia 

by C. C. Huang (Fl. Reipubl. Popularis Sin. 43(2): 58. 1997) 

belong here. 

8. Melicope glomerata (Craib) T. G. Hartley, Allertonia 8: 263. 

2001. 

密果蜜茱萸 mi guo mi zhu yu 

Euodia glomerata Craib, Bull. Misc. Inform. Kew 1918: 

362. 1918; E. simplicifolia Ridley var. pubescens C. C. Huang. 

Shrubs 0.5–1.5 m tall or rarely trees to 11 m tall, dioecious 

or rarely monoclinous. Young branchlets 2–4.5 mm wide in 

third internode. Leaves 1- or 3-foliolate. Leaves 3-foliolate: 

petiole 1.5–7 cm, pubescent to velutinous; leaflet blades elliptic 

to obovate, or narrowly so, in terminal leaflet 3.5–16 × 1.3–6.5 

cm, pubescent or sparsely so at least abaxially on midvein and 

secondary veins and adaxially on midvein, apex obtuse to acuminate. 

Leaves 1-foliolate: petiole 0.8–1.5 cm; leaflet blade 

elliptic to obovate, 4–9.5 × 2–3.8 cm, otherwise similar to 

3-foliolate leaves. Inflorescences axillary and sometimes basal 

to leaves, 0.8–6 × 0.8–1.7 cm; peduncle 0.1–2.5 cm; larger 

primary branches spreading or ascending. Pedicel 0.6–2 mm 

but 2.5–4 mm in fruit, sparsely hispidulous to pubescent. Sepals 

1–2 mm, connate at base, outside hirsutulous or sparsely so or 

rarely nearly glabrous, persistent in fruit. Petals 2–3.5 mm, 

persistent in fruit. Stamens 4, in male and bisexual flowers

1.5–2.5 mm, in female flowers 1.5–2 mm; filaments at least in 

male and bisexual flowers narrowly obtuse or sometimes acute 

at apex. Gynoecium glabrous, in female and bisexual flowers 

1–1.5 mm, in male flowers ca. 0.3 mm. Fruit follicles obovoid, 


5–8 mm, glabrous. Seeds ovoid, 4–5 mm, attachment Type B. 

Fr. Aug. 

Open forests; 500–700 m. S Yunnan [Laos, Myanmar, Thailand]. 

5. BOENNINGHAUSENIA Reichenbach ex Meisner, Pl. Vasc. Gen. 1: 60; 2: 44. 1837, 

nom. cons., not Boenninghausia Sprengel (1826). 

石椒草属 shi jiao cao shu 


Herbs, perennial. Leaves alternate, pinnately to ternately decompound. Inflorescences terminal and axillary, paniculate, with 

simple foliar bracts. Flowers bisexual, actinomorphic. Sepals 4, connate at base or to nearly their full length. Petals 4, imbricate in bud, 

margin entire. Stamens (6–)8, distinct. Disk annular to cup-shaped, surrounding and separate from a slender manifestly accrescent 

gynophore which often becomes apically 4-parted. Gynoecium 4-carpelled; ovaries coherent in ± their basal half, otherwise 

contiguous; ovules several per locule; style lateral, of 4 coherent stylar elements; stigma punctiform or capitellate. Fruit of 4 distinct 

apically dehiscent follicles; endocarp thinly cartilaginous, adnate to mesocarp in dehisced fruit. Seeds reniform; seed coat leathery, 

tuberculate; endosperm ± copious; embryo curved; cotyledons elliptic, flattened; hypocotyl superior. 

One species: E, S, and SE Asia. 

1. Boenninghausenia albiflora (Hooker) Reichenbach ex 

Meisner, Pl. Vasc. Gen. 2: 44. 1837. 

臭节草 chou jie cao 

Ruta albiflora Hooker, Exot. Fl. 1: t. 79. 1823; Bodinieria 

thalictrifolia H. Léveillé; Boenninghausenia albiflora var. 

brevipes Franchet; B. albiflora var. pilosa Z. M. Tan; B. brevipes 

(Franchet) H. Léveillé; B. japonica Siebold ex Miquel; B. 

schizocarpa S. Y. Hu; B. sessilicarpa H. Léveillé; Podostaurus 

thalictroides Junghuhn. 

Plants to 1.2 m tall. Stems, branches, leaves, and inflorescences 

glabrous to pubescent. Larger leaves 4–16 cm; leaflet 

blades elliptic to obovate to broadly obovate to suborbicular, 

0.5–2.5 × 0.4–1.5 cm. Inflorescences to 60 cm. Flowers globose 

to ovoid to ellipsoid to oblong in bud. Sepals 0.5–1.3 mm. Petals 

elliptic to obovate, or broadly so, 3.5–6 mm, apex acute to 

broadly rounded. Disk 0.3–0.5 mm. Gynophore in flowers at 

anthesis 0.3–1.5 mm, in fruit 0.6–5.5 mm. Fruit follicles 2.5–5 

mm. Fl. and fr. May–Nov. 

Open forests, grassy slopes; 500–2800 m. Anhui, Fujian, Gansu, 

Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, 

Sichuan, Taiwan, SE Xizang, Yunnan, Zhejiang [Bhutan, India, Indonesia, 

Japan, Kashmir, Laos, Myanmar, Nepal, Pakistan, Philippines, 

Thailand, N Vietnam]. 

6. HAPLOPHYLLUM A. Jussieu, Mém. Mus. Hist. Nat. 12: 464. 1825 [“Aplophyllum”], 

nom. cons., not Aplophyllum Cassini (1824). 

拟芸香属 ni yun xiang shu 


Herbs, perennial [or sometimes low shrubs]. Leaves alternate, simple [occasional leaves 3- or 5-parted]. Inflorescences terminal, 

cymose and often corymbiform, (1- or) few to many flowered. Flowers bisexual, actinomorphic. Sepals 5, distinct or connate to ± half 

their length. Petals 5, imbricate in bud, margin entire. Stamens 10, distinct [or monadelphous]. Disk pulvinate. Gynoecium (2 

or)3–5-carpelled; ovaries connate in ± their basal half, otherwise contiguous; ovules few or several per locule; style lateral, of (2 or)3–5 

contiguous or connate stylar elements; stigma punctiform or capitellate. Fruit a (2 or)3–5-carpelled loculicidal capsule or rarely 

indehiscent, with carpels connate as in gynoecium; endocarp thinly cartilaginous, adnate to mesocarp in dehisced fruit. Seeds reniform; 

seed coat leathery, rugose; endosperm copious; embryo curved; cotyledons elliptic-oblong, flattened; hypocotyl superior. 

About 65 species: N Africa, Asia, S Europe; three species (one endemic) in China. 

1a. Leaf blade ovate to lanceolate, 5–20 mm wide, secondary veins prominent ........................................................... 2. H. acutifolium 

1b. Leaf blade elliptic, lanceolate, or linear, 1–5 mm wide, secondary veins not prominent. 

2a. Inflorescences many flowered; gynoecium 2- or 3-carpelled ............................................................................... 1. H. dauricum 

2b. Inflorescences 1-flowered; gynoecium 4- or 5-carpelled .......................................................................... 3. H. tragacanthoides 

1. Haplophyllum dauricum (Linnaeus) G. Don, Gen. Hist. 1: 

781. 1831. 

北芸香 bei yun xiang 

Peganum dauricum Linnaeus, Sp. Pl. 1: 445. 1753; Haplophyllum 

lineare (Candolle) G. Don; Ruta daurica (Linnaeus) 

Candolle. 

Herbs, perennial. Underground part woody, above-ground 

part many branched; branchlets slender, 10–20 cm, puberulent 

when young. Leaves subsessile; leaf blade lanceolate to linear, 

5–20 × 1–5 mm, grayish green, with oil glands, base attenuate, 

apex acute. Inflorescences many flowered or very rarely 

3-flowered; bracts linear, small. Sepals ca. 1 mm, basally connate, 

margin pubescent. Petals yellowish to pale yellowish 

73

74 

white, oblong, 6–8 mm, with scattered semipellucid oil glands. 

Staminal filaments enlarged in ± their basal half; anthers 

elongately ellipsoid; connective with an apical oil gland. Ovary 

globose, (2 or)3(or 4)-loculed; style slender; stigma slightly 

enlarged. Fruiting carpels 2-seeded. Seeds brownish black, 

2–2.5 × 1–1.5 mm. Fl. Jun–Jul, fr. Aug–Sep. 

Hill slopes; low elevations. Gansu, Hebei, Heilongjiang, Jilin, Nei 

Mongol, Ningxia, Shaanxi, Xinjiang [Mongolia, Russia]. 

2. Haplophyllum acutifolium (Candolle) G. Don, Gen. Hist. 1: 

780. 1831. 

大叶芸香 da ye yun xiang 

Ruta acutifolia Candolle, Prodr. 1: 711 1824; Haplophyllum 

flexuosum Boissier; H. perforatum Karelin & Kirilov; 

H. sieversii Fischer & C. A. Meyer; R. flexuosa (Boissier) 

Engler; R. perforata M. Bieberstein; R. sieversii (Fischer & C. 

A. Meyer) B. Fedtschenko. 

Herbs, perennial, to 80 cm tall. Stem many branched; 

branches stout, spreading, glabrous, with oil glands; branchlets 

angular. Leaves glabrous; petiole 4–6 mm or leaf subsessile; leaf 

blade ovate to rhombic but on basal part of branches lanceolate 

and on apical part of branches narrowly elliptic, 3–5 × 0.5–2 cm, 

thickly papery, with oil glands. Inflorescences many flowered; 

rachis pubescent when young; bracts linear to narrowly elliptic, 

margin ciliate. Flowers pedicellate. Sepals triangular-ovate, to 1 

mm, basally connate. Petals yellow, ca. 3 × 1.5 mm, with 

scattered semipellucid oil glands. Staminal filaments flattened in 

± their basal half. Ovary globose; style ca. 0.5 mm; stigma 

slightly enlarged. Capsule 4- or 5-carpelled, 3–4 mm in diam. 

when mature, with protruding oil glands when young, 2-seeded. 

Fl. May, fr. Sep. 


Grassy areas, riversides; ca. 700 m. Xinjiang [Afghanistan, 

Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Tajikistan, Turkmenistan, 

Uzbekistan; SW Asia]. 

C. C. Huang (Fl. Reipubl. Popularis Sin. 43(2): 87. 1997) mistakenly 

called this species Haplophyllum perforatum (M. Bieberstein) 

Karelin & Kirilov. The H. perforatum Karelin & Kirilov (1841) is based 

a different type than Ruta perforata M. Bieberstein (1800), and a combination 

of the R. perforata in Haplophyllum is a later homonym. The 

oldest specific epithet available for this species is R. acutifolia Candolle 

(1824). 

3. Haplophyllum tragacanthoides Diels, Notizbl. Bot. Gart. 

Berlin-Dahlem 9: 1028. 1926. 

针枝芸香 zhen zhi yun xiang 

Herbs, perennial, 10–15 cm tall. Branches dichotomous 

from stem base. Leaves sessile; leaf blade shortly linear to 

narrowly elliptic, 3–9 × 1–3 mm, thickly papery, grayish green, 

with scattered oil glands, margin denticulate. Flowers terminal, 

solitary. Sepals ovate, to 1 mm, basally connate, margin ciliate. 

Petals yellow, oblong, 7–8 × ca. 3 mm, with scattered semipellucid 

oil glands, margin irregular. Staminal filaments flattened in 

± their basal half, with trichomes. Ovary 4- or 5-carpelled; style 

ca. 2.5 mm; stigma slightly enlarged. Capsule 4- or 5-carpelled, 

with oil glands; carpels ca. 5 mm in diam., 1-seeded. Seeds 

reniform, 2–2.5 × ca. 1.5 mm; seed coat shrunken. Fl. May–Jun, 

fr. Jul–Aug. 

● Dry hill slopes; ca. 1500 m. Gansu, Nei Mongol, Ningxia. 

Townsend (Hooker’s Icon. Pl. 40: 35, t. 3902. 1986) placed Haplophyllum 

tragacanthoides in synonymy of H. dauricum (Linnaeus) G. 

Don. 

7. PSILOPEGANUM Hemsley, J. Linn. Soc., Bot. 23: 103. 1886. 

裸芸香属 luo yun xiang shu 


Herbs, perennial. Leaves alternate, digitately 3-foliolate (occasional leaves simple). Flowers bisexual, actinomorphic, axillary, 

solitary or in few-flowered racemes. Sepals 4, basally connate. Petals 4 or 5, imbricate in bud, margin entire. Stamens 8 or 10, distinct. 

Disk columnar. Gynoecium 2(or 3)-carpelled; ovaries connate in ± their basal 2/3, otherwise contiguous to divergent; ovules several 

per locule; style lateral, of 2 or rarely 3 coherent stylar elements; stigma capitellate. Fruit a loculicidal capsule, with carpels connate as 

in ovary; endocarp thinly cartilaginous, adnate to mesocarp in dehisced fruit. Seeds reniform; seed coat leathery, tuberculate; 

endosperm copious; embryo curved; cotyledons elliptic, flattened; hypocotyl superior. 

● One species: China. 

1. Psilopeganum sinense Hemsley, J. Linn. Soc., Bot. 23: 103. 

1886. 

裸芸香 luo yun xiang 

Plants 30–80 cm tall. Leaves 2.5–4.5 cm, glabrous, with 

citrus odor; petiole 0.8–1.5 cm; leaflet blades ovate-elliptic to 

elliptic-obovate, in lateral leaflets 4–10 × 2–6 mm, in terminal 

leaflet 1.5–2.8 × 0.6–1.2 cm. Sepals ca. 1 mm. Petals ovate- 

elliptic, 4–6 mm. Stamens slightly shorter than petals. Ovary 

usually obcordate; style to 2 mm. Capsule ca. 5 mm. Seeds ca. 

1.5 × 1 mm. Fl. and fr. May–Aug. 

● Hill slopes; ca. 800 m. Guizhou, Hubei, Sichuan. 

8. DICTAMNUS Linnaeus, Sp. Pl. 1: 388. 1753. 

白鲜属 bai xian shu 


Herbs, perennial. Leaves alternate, odd-pinnate. Inflorescences terminal. Flowers bisexual, zygomorphic. Sepals 5, basally


connate. Petals 5, 4 ascending and 1 descending, imbricate in bud. Stamens 10, distinct, declined. Disk thick, asymmetrically annular. 

Gynoecium 5-carpelled; ovaries basally connate, otherwise contiguous; ovules 3 per locule; style lateral, declined, of 5 basally 

coherent but apically connate stylar elements; stigma punctiform. Fruit of 5 basally connate follicles; endocarp cartilaginous, discharged 

elastically with seeds at dehiscence. Seeds black, subglobose, shiny; seed coat brittle, sclerenchymatous; endosperm copious; 

embryo straight; cotyledons broadly elliptic, flattened; hypocotyl partly included between cotyledons. 

One to five species: Asia, Europe; one species in China. 

1. Dictamnus dasycarpus Turczaninow, Bull. Soc. Imp. 

Naturalistes Moscou 15: 637. 1842. 

白鲜 bai xian 

Aquilegia fauriei H. Léveillé; Dictamnus albus Linnaeus 

subsp. dasycarpus (Turczaninow) L. Winter; D. albus var. dasycarpus 

(Turczaninow) T. N. Liou & Y. H. Chang. 

Plants 40–100 cm tall. Younger stems, leaves, and inflorescences 

pubescent or sparsely so; leaflet blades with scattered 

pellucid oil glands. Leaves 7–13-foliolate; rachis marginate to 

narrowly winged; leaflet blades ovate-elliptic, elliptic, or oblong, 

3–12 × 1–5 cm, lateral ones opposite and sessile but ter- 

minal one petiolulate, secondary veins ± inconspicuous, margin 

crenulate. Inflorescences, flowers, and follicles with opaque 

dark brown globose to ellipsoid sessile or stalked glands. Inflorescences 

compound racemose, to 30 cm. Pedicel 1–1.5 cm. 

Sepals 6–8 × 2–3 mm. Petals pinkish white to pink, with purplish 

stripes, oblanceolate, 2–2.5 × 0.5–0.8 cm. Stamens exserted. 

Fruit follicles 1–2 cm, usually with a stylar beak and apical 

abaxial beak. Fl. May, fr. Aug–Sep. 

Hill slopes, flat areas in thickets, grassy open forests. Anhui, 

Gansu, Hebei, Heilongjiang, Henan, Hubei, Jiangsu, Jiangxi, Jilin, Liaoning, 

Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, Xinjiang 

[Korea, Mongolia, Russia (Far East)]. 

9. TODDALIA Jussieu, Gen. Pl. 371. 1789, nom. cons. 

飞龙掌血属 fei long zhang xue shu 


Shrubs (usually sprawling) or woody climbers, dioecious, usually armed. Leaves alternate, digitately 3-foliolate (occasional 

leaves 1- or 2-foliolate). Inflorescences terminal and/or axillary, paniculate, racemose, or umbelliform. Sepals 4 or 5(or 6), connate at 

base or to ± half their length. Petals 4 or 5(or 6), valvate or narrowly imbricate in bud. Stamens 4 or 5(or 6), distinct, rudimentary in 

female flowers. Disk pulvinate. Gynoecium 4–7-loculed, syncarpous, rudimentary in male flowers; ovules 2 per locule; stigma sessile 

or nearly so. Fruit a 4–7-loculed drupaceous berry; exocarp fleshy; mesocarp undifferentiated; endocarp cartilaginous. Seeds brown to 

black, reniform, dull to lustrous; seed coat with thick inner layer of dense black sclerenchyma surrounded by outer layer of compact 

parenchymatous tissue; endosperm copious; embryo curved; cotyledons elliptic, flattened; hypocotyl superior. 

One species: Africa, E, S, and SE Asia, Madagascar, Mascarene Islands. 

1. Toddalia asiatica (Linnaeus) Lamarck, Tabl. Encycl. 2: 116. 

1797. 

飞龙掌血 fei long zhang xue 

Paullinia asiatica Linnaeus, Sp. Pl. 1: 365. 1753, typ. 

cons.; Aralia labordei H. Léveillé; Toddalia asiatica var. floribunda 

(Wallich) Kurz; T. asiatica var. gracilis Gamble; T. 

asiatica var. obtusifolia Gamble; T. floribunda Wallich; T. 

tonkinensis Guillaumin. 

Shrubs (usually sprawling) or woody climbers, usually 

armed. Petiole 1–4 cm; leaflet blades usually sessile or subsessile, 

elliptic or narrowly elliptic to obovate to oblanceolate, 3–10 

× 1–4 cm, base narrowly cuneate to attenuate, apex acuminate or 

rarely acute to obtuse or rounded. Inflorescences to 17 cm. 

Sepals 0.3–0.5 mm. Petals cream-white, ovate to elliptic, 1–3.5 

mm. Stamens in male flowers 3–4 mm, in female flowers 

ligulate and 0.2–0.8 mm. Disk 0.2–0.5 mm. Gynoecium in 

female flowers ovoid to ellipsoid and 1.5–2.5 mm, in male 

flowers subcylindric and 1–2 mm. Fruit 5–10 mm in diam. Seeds 

5–6.5 mm. Fl. year-round but mostly in spring and summer, fr. 

autumn and winter. 

Secondary forests, thickets; near sea level to 2000 m. Fujian, 

Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, 

Shaanxi, Sichuan, Taiwan, Xizang, Yunnan [Bangladesh, Bhutan, India, 

Indonesia, Japan (Ryukyu Islands), Laos, Malaysia, Myanmar, Nepal, 

Philippines, Sri Lanka, Thailand, Vietnam; Africa, Madagascar]. 

10. PHELLODENDRON Ruprecht, Bull. Cl. Phys.-Math. Acad. Imp. Sci. 

Saint-Pétersbourg 15: 353. 1857. 

黄檗属 huang bo shu 


Trees, deciduous, dioecious. Bark corky, fissured and gray or grayish brown at maturity; inner bark (phloem) yellow. Branchlets 

dark purplish. Leaves opposite, odd-pinnate; base of petiole excavated, concealing axillary bud; leaflet blades with oil glands restricted 

to margin, margin subentire to minutely crenulate or serrulate. Inflorescences terminal, thyrsiform. Sepals 5(–8), connate at base. Petals 

75

76 


5(–8), narrowly imbricate in bud. Stamens 5(–7), distinct. Disk columnar. Male flowers: stamens to 1.5 × as long as petals; gynoecium 

rudimentary, of 5(or 6) fingerlike carpels, basally connate, apically ± divergent. Female flowers: stamens rudimentary, ligulate, much 

shorter than petals; gynoecium 5(–10)-loculed, syncarpous; ovules 1 per locule; stigma peltate. Fruit a drupaceous berry, black or 

purplish black, 5(–10)-loculed; exocarp fleshy; mesocarp undifferentiated; endocarp thinly cartilaginous. Seeds brown or black, 

asymmetrically ellipsoid, dull, inconspicuously rugulose; seed coat with inner layer of dense black sclerenchyma and outer layer of 

compact parenchymatous tissue; endosperm ± scant; embryo straight; cotyledons elliptic, flattened; hypocotyl superior. 

Two to four species: E and SE Asia; two species (one endemic) in China. 

1a. Inflorescences and infructescences ± lax, rachis, branches, and pedicels slender; leaflet blades papery to thinly 

papery ........................................................................................................................................................................... 1. P. amurense 

1b. Inflorescences and infructescences ± compact, rachis, branches, and pedicels robust; leaflet blades papery ............. 2. P. chinense 

1. Phellodendron amurense Ruprecht, Bull. Cl. Phys.-Math. 

Acad. Imp. Sci. Saint-Pétersbourg 15: 353. 1857. 

黄檗 huang bo 

Trees 10–30 m tall, d.b.h. to 1 m. Leaves 7–13-foliolate; 

rachis glabrous to pubescent; leaflet blades ovate to ovate- 

lanceolate, 6–12 × 2.5–4.5 cm, papery to thinly papery, apex 

acuminate. Inflorescences and infructescences ± lax, rachis, 

branches, and pedicels slender. Fruit globose, ca. 1 cm in diam. 

Seeds ca. 6 × 3 mm. Fl. May–Jun, fr. Sep–Oct. 

Montane forests and thickets, river valleys. Anhui, Hebei, Heilongjiang, 

Henan, Jilin, Liaoning, Nei Mongol, Shandong, Shanxi, Taiwan 

[Japan, Korea, Russia (Far East)]. 

This species is widely cultivated. 

Phellodendron amurense var. wilsonii was accepted by C. E. 

Chang (Fl. Taiwan, ed. 2, 3: 527. 1993). 

2. Phellodendron chinense C. K. Schneider, Ill. Handb. 

Laubholzk. 2: 126. 1907. 

川黄檗 chuan huang bo 

Trees to 15 m tall. Leaves 7–15-foliolate; rachis glabrous, 

thinly pubescent, or rust-colored tomentose; leaflet blades ovate- 

elliptic to oblong-lanceolate, 8–15 × 3.5–6 cm, papery, base 

cuneate to attenuate and oblique, apex mucronate to acuminate. 

Inflorescences and infructescence ± compact, rachis, branches, 

and pedicels robust. Fruit subglobose to ellipsoid, 1–1.5 cm in 

diam. Seeds 6–7 × 3–5 mm. Fl. May–Jun, fr. Sep–Nov. 

● Mixed broad-leaved forests, open to dense forests; 800–1500 

(–3000) m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, 

Hubei, Hunan, Jiangsu, Shaanxi, Sichuan, Yunnan, Zhejiang. 

1a. Rachis of leaves rust-colored tomentose 

.................................................................. 2a. var. chinense 

1b. Rachis of leaves glabrous or thinly 

pubescent ....................................... 2b. var. glabriusculum 

2a. Phellodendron chinense var. chinense 

川黄檗(原变种) chuan huang bo (yuan bian zhong) 

Phellodendron fargesii Dode; P. macrophyllum Dode, p.p. 

as to R. P. Farges 1284 p.p. altitude 1400 m (see also synonymy 

of Tetradium daniellii). 

Rachis of leaves rust-colored tomentose. 

● Mixed broad-leaved forests; above 900 m. Anhui, Henan, Hubei, 

Hunan, Sichuan, Yunnan. 

2b. Phellodendron chinense var. glabriusculum C. K. 

Schneider, Ill. Handb. Laubholzk. 2: 126. 1907. 

秃叶黄檗 tu ye huang bo 

Phellodendron chinense var. falcatum C. C. Huang; P. 

chinense var. omeiense C. C. Huang; P. chinense var. yunnanense 

C. C. Huang. 

Rachis of leaves glabrous or thinly pubescent. 

● Open to dense forests; 800–1500(–3000) m. Fujian, Gansu, 

Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Shaanxi, Sichuan, 

Yunnan, Zhejiang. 

11. ACRONYCHIA J. R. Forster & G. Forster, Char. Gen. Pl. 27. 1775, nom. cons. 

山油柑属 shan you gan shu 


Shrubs or trees, evergreen. Leaves opposite, 1-foliolate [or digitately 3-foliolate]. Inflorescences axillary or basal to leaves, 

cymulose to thyrsiform [or reduced to solitary flowers]. Flowers bisexual. Sepals 4, distinct or basally connate. Petals 4, valvate in bud. 

Stamens 8, distinct, ones opposite sepals nearly as long as petals; filaments with indumentum toward base [or rarely glabrous], 

gradually tapering from ± broad base to subulate apex, becoming reflexed; anthers obtuse or obtusely mucronulate. Disk pulvinate. 

Gynoecium 4-loculed, ± completely syncarpous [or grading to subapocarpous]; ovules 2 per locule; stigma punctiform or capitellate. 

Fruit a ± completely syncarpous 4-loculed drupaceous berry [or grading to subapocarpous with 4 basally connate drupaceous berries]; 

exocarp ± fleshy or dry; mesocarp, if evident, spongy-crustaceous or ± woody; endocarp cartilaginous. Seeds ellipsoid to pyriform, 

dull to ± shiny; seed coat with thick [to thin] inner layer of dense black sclerenchyma surrounded by outer layer of compact 

parenchymatous tissue; endosperm copious; embryo straight or nearly so; cotyledons ovate to elliptic, flattened; hypocotyl superior. 

About 48 species: S and SE Asia, Australia, SW Pacific islands; one species in China.

1. Acronychia pedunculata (Linnaeus) Miquel, Fl. Ned. Ind., 

Eerste Bijv. 532. 1861. 

山油柑 shan you gan 

Jambolifera pedunculata Linnaeus, Sp. Pl. 1: 349. 1753; 

Acronychia apiculata Miquel; A. arborea Blume; A. barberi 

Gamble; A. elliptica Merrill & L. M. Perry; A. laurifolia Blume; 

Clausena simplicifolia Dalzell; Cyminosma pedunculata (Linnaeus) 

Candolle; Gela lanceolata Loureiro; J. arborea (Blume) 

Zollinger & Moritzi; J. rezinosa Loureiro; Laxmannia ankenda 

(Gaertner) Raeuschel; Melicope conferta Blanco; Selas lanceolatum 

(Loureiro) Sprengel; Ximenia lanceolata (Loureiro) Candolle. 

Shrubs or small to large trees, to 28 m tall. Petiole glabrous 

to finely pubescent, 0.5–5 cm; leaflet blades usually elliptic to 

elliptic-oblong but grading to obovate, oblanceolate, or nearly 

oblong, 3.5–24.5 × 2–8.5 cm, base cuneate or sometimes 

rounded or attenuate, apex obtusely acuminate with an acumen 


to 2 cm or sometimes obtuse or rounded. Inflorescences 2–25 

cm, few to many flowered. Pedicel 2–12 mm. Sepals 0.6–1.5 

mm. Petals 4–12 mm. Ovary completely pubescent or rarely 

pubescent only at apex, with or without apical septicidal fissures; 

style pubescent at base, otherwise glabrous. Fruit subglobose or 

sometimes grading to ellipsoid, pyriform, or broadly conic, 

0.5–1.5 cm in diam., with or without apical septicidal fissures, 

usually ± sparsely pubescent with ring of dense appressed trichomes 

at base but grading to tomentose or glabrous, apex often 

apiculate; outer part of pericarp (exocarp and mesocarp) drying 

0.5–3 mm thick; mesocarp woody or subwoody. Seeds reddish 

black to black, 3–7 mm. Fl. Apr–Aug, fr. Aug–Dec. 

Secondary forests, woods or thickets on lower hills; near sea level 

to 900 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan 

[Bangladesh, Bhutan, Cambodia, India, Indonesia, Laos, Malaysia, 

Myanmar, New Guinea, Philippines, Sri Lanka, Thailand, Vietnam]. 

Acronychia pedunculata is the dominant in some plant communities. 

12. MACLURODENDRON T. G. Hartley, Gard. Bull. Singapore 35: 4. 1982. 

贡甲属 gong jia shu 


Trees, evergreen, dioecious. Leaves opposite, 1-foliolate. Inflorescences axillary, thyrsiform or racemose. Sepals 4, basally 

connate. Petals 4, narrowly imbricate or valvate in bud. Stamens 8, distinct, ones opposite sepals nearly as long as petals in both male 

and female flowers; filaments sublinear, ± incurved, glabrous; anthers apiculate, lacking pollen in female flowers. Gynoecium 

4-loculed, rudimentary and minute in male flowers; ovary syncarpous; ovules 2 per locule; style of 4 coherent stylar elements; stigma 

capitate. Fruit a 4-loculed drupaceous berry; exocarp fleshy; mesocarp undifferentiated; endocarp thinly cartilaginous. Seeds ovoid to 

reniform; seed coat with thick inner layer of dense black sclerenchyma and spongy outer layer [rarely lacking] bounded externally by a 

shiny black pellicle; endosperm copious; embryo straight or slightly bent; cotyledons ovate to elliptic, flattened; hypocotyl superior. 

Six species: SE Asia; one species in China. 

1. Maclurodendron oligophlebium (Merrill) T. G. Hartley, 


贡甲 gong jia 

Acronychia oligophlebia Merrill, Philipp. J. Sci. 23: 246. 

1923. 

Trees to 13(–25) m tall. Petiole 0.8–2.5 cm, puberulent to 

finely pubescent especially adaxially, glabrescent; leaflet blades 

obovate, oblanceolate, or sometimes elliptic, 6–21.5 × 2.5–8.2 

cm, base cuneate to attenuate, apex acuminate with a 5–10 mm 

acumen but sometimes obtuse or rounded. Inflorescences 3.5–10 

cm. Pedicel 1.5–5 mm. Sepals 0.6–0.7 mm. Petals 2.5–3 mm. 

Gynoecium glabrous. Fruit globose or subglobose, 6–10 mm in 

diam., apex often apiculate. Seeds 4.5–5.5 mm; seed coat with 

spongy outer tissue bounded externally by a shiny black pellicle. 

Fl. Apr–Aug, fr. Aug–Dec. 

Primary and secondary forests; 200–1400 m. Guangdong, Hainan 

[N Vietnam]. 

13. SKIMMIA Thunberg, Nov. Gen. Pl. 3: 57. 1783, nom. cons. 

茵芋属 yin yu shu 


Shrubs or trees, evergreen, usually dioecious or monoclinous, unarmed. Leaves alternate, simple. Inflorescences terminal, 

thyrsiform. Sepals (3 or)4 or 5(–7), distinct or basally connate. Petals (3 or)4 or 5(–7), imbricate in bud. Stamens (3 or)4 or 5(–7), 

distinct, rudimentary in female flowers. Disk annular or pulvinate. Gynoecium 2–5-loculed, syncarpous, rudimentary in male flowers; 

ovules 1(or ?2) per locule; style short or ± as long as ovary. Fruit a fleshy drupaceous berry, with 1–5 1-seeded leathery pyrenes. Seeds 

ovoid to ellipsoid; seed coat membranous; endosperm copious; embryo straight; cotyledons oblong to suborbicular, flattened; 

hypocotyl superior. 

Five to six species: E, S, and SE Asia; five species (one endemic) in China. 

1a. Leaf blade midvein puberulent. 

77

78 


2a. Fruit red ................................................................................................................................................................. 1. S. reevesiana 

2b. Fruit bluish black .............................................................................................................................................. 2. S. melanocarpa 

1b. Leaf blade midvein glabrous. 

3a. Leaf blade secondary veins 12–20 on each side of midvein; rudimentary gynoecium in male flowers entire 

or nearly so; petals reflexed ................................................................................................................................. 5. S. multinervia 

3b. Leaf blade secondary veins fewer than 10 on each side of midvein; rudimentary gynoecium in male flowers 

3- or 4-lobed; petals erect to spreading. 

4a. Shrubs to 2 m tall; fruit red ................................................................................................................................ 3. S. laureola 

4b. Trees to 12 m tall; fruit bluish black ........................................................................................................... 4. S. arborescens 

1. Skimmia reevesiana (Fortune) Fortune, J. Tea Countr. China 

329. 1852. 

茵芋 yin yu 

Ilex reevesiana Fortune, Gard. Chron. 1851: 5. 1851; 

Skimmia arisanensis Hayata; S. distinctevenulosa Hayata; S. 

fortunei Masters; S. hainanensis C. C. Huang; S. japonica 

Thunberg var. distinctevenulosa (Hayata) C. E. Chang; S. 

japonica subsp. reevesiana (Fortune) N. P. Taylor & Airy Shaw; 

S. japonica var. reevesiana (Fortune) N. P. Taylor; S. orthoclada 

Hayata. 

Shrubs 1–2 m tall. Branches with hollow pith; bark pale 

grayish green, smooth, cracked when dry. Leaves clustered at 

branch apex, with a citrus-leaf odor; petiole 5–10 mm; leaf blade 

elliptic, lanceolate, ovate, or oblanceolate, 5–12 × 1.5–4 cm, 

leathery, midvein adaxially puberulent, base cuneate, apex mucronate 

to obtuse or acuminate. Inflorescences puberulent, with 

condensed flowers. Flowers male, female, or bisexual, (3 or)4or 

5-merous, fragrant. Sepals rounded, 1–1.5 mm, margin ciliate. 

Petals yellowish white, 3–5 mm. Stamens (3 or)4 or 5. Male 

flowers: rudimentary ovary compressed globose, entire or shallowly 

2–4-lobed, apex mucronate. Female flowers: ovary subglobose; 

style elongated; stigma capitate. Fruit red, globose to 

ellipsoid-obovoid, 0.8–1.5 cm, 2–4-seeded. Seeds compressed 

ovoid, 5–9 × 4–6 mm. Fl. Mar–May, fr. Sep–Nov. 2n = 60*. 

Montane moss forests; 1200–2600 m. S Anhui, Fujian, Guangdong, 

Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangxi, Sichuan, 

Taiwan, E Yunnan, Zhejiang [Myanmar, Philippines, S Vietnam]. 

Taylor (Kew Mag. 4: 182, 183. 1987) treated Skimmia reevesiana 

as S. japonica subsp. reevesiana and placed S. arisanensis, S. distinctevenulosa, 

S. hainanensis, and S. orthoclada in synonymy of S. japonica 

var. reevesiana. 

2. Skimmia melanocarpa Rehder & E. H. Wilson in Sargent, 

Pl. Wilson. 2: 138. 1914. 

黑果茵芋 hei guo yin yu 

Shrubs 1–2 m tall, monoecious, dioecious, or monoclinous. 

Branches with hollow pith; bark pale grayish green, smooth, 

cracked when dry. Petiole 5–10 mm; leaf blade elliptic to elliptic-lanceolate, 

3–7 × 1.5–2.5 cm, midvein adaxially pubescent, 

apex acuminate to acute. Inflorescences to 4 cm, with condensed 

flowers; rachis puberulent; bracts ca. 1 mm. Flowers subsessile. 

Sepals broadly ovate, 1–1.5 mm, margin ciliate. Petals 5, yellowish 

white, oblanceolate to oblong, 3–4 mm, subequal, 

reflexed in male flowers. Male flowers: stamens as long as or 

slightly longer than petals. Female and bisexual flowers: stamens 

shorter than petals; ovary subglobose; style cylindric. Fruit 

bluish black, subglobose, ca. 8 mm in diam., (4 or)5-loculed, 

apex with (4 or)5 stylar scars. Fl. Mar–May, fr. Sep–Nov. 

● Dense or open forests; 2000–3000 m. S Gansu, W Hubei, S 

Shaanxi, Sichuan, SE Xizang, Yunnan. 

Taylor (Kew Mag. 4: 189. 1987) placed Skimmia melanocarpa in 

synonymy of S. laureola subsp. laureola. 

3. Skimmia laureola (Candolle) Siebold & Zuccarini ex 

Walpers, Repert. Bot. Syst. 5: 405. 1845–1846. 

月桂茵芋 yue gui yin yu 

Limonia laureola Candolle, Prodr. 1: 536. 1824. 

Shrubs to 1.3 m tall. Branches grayish green, dichotomous 

or nearly whorled. Leaves nearly opposite or whorled, clustered 

near apex of branches, spreading, with a citrus-leaf odor; leaf 

blade oblong-lanceolate, (4–)7–10(–12) × 2–3 cm, adaxially 

dark green, shiny, and with translucent oil glands, midvein 

slender, secondary veins hardly distinguishable, base attenuate, 

margin entire, apex mucronate. Inflorescences 5–8 cm; rachis 

stout, angulate. Pedicel puberulent; bracteoles opposite near 

middle of pedicel, caducous. Sepals ovate, base pubescent, margin 

ciliate. Petals 5, yellowish, oblong, ca. 3 × as long as sepals, 

apex obtuse. Stamens 5, erect. Ovary ovoid, 2- or 3-loculed, 

glabrous, with oil glands; ovules 1(or ?2) per locule; style short; 

stigma small. Fruit ovoid, 8–10 mm in diam. Fl. Apr, fr. autumn. 

2n = 30*, 60. 

Evergreen forests; below 2800 m. S Xizang (Yadong) [Bhutan, NE 


4. Skimmia arborescens T. Anderson ex Gamble, J. Linn. Soc., 

Bot. 43: 491. 1916. 

乔木茵芋 qiao mu yin yu 

Skimmia euphlebia Merrill; S. japonica Thunberg var. euphlebia 

(Merrill) N. P. Taylor; S. japonica var. kwangsiensis (C. 

C. Huang) N. P. Taylor; S. kwangsiensis C. C. Huang. 

Trees to 8 m tall, d.b.h. to 20 cm. Branches with small but 

prominent pith. Petiole 1–2 cm; leaf blade elliptic, oblong, or 

obovate-elliptic, 5–18 × 2–6 cm, papery, glabrous, midvein 

adaxially prominent, secondary veins 7–10 on each side of 

midvein. Inflorescences 2–5 cm; rachis puberulent or glabrous; 

bracts broadly ovate, 1–1.5 mm. Sepals ovate, margin ciliate. 

Petals 5, yellowish, obovate to ovate-oblong, 4–5 mm. Male 

flowers: stamens longer than petals; filaments filiform; rudimentary 

gynoecium barlike, 3–4 mm, apex 3- or 4-lobed. Female 

flowers: rudimentary stamens shorter than petals; ovary 

globose; style ca. 1 mm; stigma capitate. Fruit bluish black, 

globose, 6–8 mm in diam., 1–3-seeded. Fl. Apr–Jun, fr. Jul–Sep. 

Shady moist montane areas; 1000–2800 m. Guangdong, Guangxi, 

Guizhou, Sichuan, E Xizang, Yunnan [Bhutan, NE India, Laos, Myanmar, 

Nepal, N Thailand, N Vietnam].

Taylor (Kew Mag. 4: 183–185. 1987) treated Skimmia arborescens 

as S. arborescens subsp. arborescens, S. euphlebia as S. japonica var. 

euphlebia, and S. kwangsiensis as S. japonica var. kwangsiensis. 

5. Skimmia multinervia C. C. Huang, Acta Phytotax. Sin. 7: 

348. 1958. 

多脉茵芋 duo mai yin yu 

Skimmia laureola (Candolle) Siebold & Zuccarini ex 

Walpers subsp. multinervia (C. C. Huang) N. P. Taylor & Airy 

Shaw. 

Trees to 13 m tall. Branches dark grayish, with scattered 

lenticels. Petiole 1–2 cm, stout; leaf blade oblanceolate to rarely 

narrowly oblong, 10–18 × 3–5 cm, leathery, midvein adaxially 

slightly prominent, secondary veins 12–20 on each side of mid- 


vein, margin slightly revolute. Inflorescences 2–6 cm; rachis 

stout when young, puberulent, elongated in fruit; bracts ovate, 

1–2 mm, margin ciliate. Pedicel stout. Sepals ovate, ca. 2 mm, 

margin ciliate. Petals 5, yellowish white, obovate-oblong to 

oblong, 4–5 mm, reflexed at anthesis. Stamens 5. Male flowers: 

stamens longer than petals; rudimentary gynoecium barlike, ca. 

1.5 mm, apex entire or shallowly lobed. Female and bisexual 

flowers: rudimentary stamens as long as petals; ovary globose, 

5-loculed; style ca. 1.5 mm; stigma capitate. Fruit bluish black, 

subglobose to slightly compressed, 6–8 mm in diam., 4- or 

5-seeded. Fl. Apr–Jun, fr. Jul–Sep. 

Montane forests; ca. 2000 m or above. SW Sichuan, NW Yunnan 

[Bhutan, NE India, Myanmar, Nepal, N Vietnam]. 

Taylor (Kew Mag. 4: 189. 1987) treated Skimmia multinervia as S. 

laureola subsp. multinervia. 

14. MICROMELUM Blume, Bijdr. 137. 1825, nom. cons. 

小芸木属 xiao yun mu shu 


Shrubs or trees, unarmed. Leaves alternate, odd-pinnate [or rarely digitately 3-foliolate and/or 1-foliolate]. Inflorescences 

terminal or terminal and axillary, paniculate. Flowers bisexual, ellipsoid or broadly ellipsoid to obovoid to oblong in bud. Calyx 

cup-shaped, shallowly 5-lobed or -toothed. Petals 5, valvate in bud. Stamens 10, distinct, alternately unequal in length; filaments 

sublinear, ± straight. Disk annular or columnar. Gynoecium 3–5-loculed, syncarpous; radial walls of locules becoming curved after 

anthesis; ovules 2 per locule; style to 1.5 × as long as ovary, deciduous in fruit. Fruit a berry with neither pulp nor pulp vesicles; 

endocarp membranous. Seeds ellipsoid; seed coat membranous; endosperm lacking; embryo straight; cotyledons broadly elliptic, thin, 

convolute and/or folded; hypocotyl superior. 

About ten species: S and SE Asia, Australia, SW Pacific islands; two species in China. 

1a. Flower buds ellipsoid or broadly ellipsoid; petals 3–4 mm, outside glabrous or pubescent; margin of leaflet blades 

serrate or repand ............................................................................................................................................................ 1. M. falcatum 

1b. Flower buds oblong; petals 5–10 mm, outside densely pubescent; margin of leaflet blades crenate, repand, or 

entire ...................................................................................................................................................................... 2. M. integerrimum 

1. Micromelum falcatum (Loureiro) Tanaka, Bull. Mus. Natl. 

Hist. Nat., sér. 2, 2: 257. 1930. 

大管 da guan 

Aulacia falcata Loureiro, Fl. Cochinch. 1: 273. 1790. 

Trees 1–3 m tall. Branchlets, petioles, and inflorescence 

rachises hirsute. Leaves 5–11-foliolate; petiolules 3–7 mm; 

leaflet blades alternate, ovate to lanceolate, inequilateral, 4–9 × 

1.2–3.5(–4.5) cm, margin serrate or repand, apex acuminate to 

caudate. Inflorescences terminal. Flowers ellipsoid or broadly so 

in bud. Calyx lobes broadly triangular, less than 1 mm. Petals 

white, oblong, 3–4 mm, outside glabrous or pubescent. Longer 

stamens subequal to petal length, shorter ones subequal to 

gynoecium length. Style cylindric, longer than ovary; stigma 

capitate. Fruit ellipsoid to obovoid, 8–10 × 7–9 mm, 1- or 

2-seeded. Fl. Jan–Apr, fr. Jun–Aug. 

Mountains; below 1200 m. SW Guangdong, Guangxi, Hainan, S 

Yunnan [Cambodia, Laos, Myanmar, Thailand, Vietnam]. 

2. Micromelum integerrimum (Buchanan-Hamilton ex Candolle) 

Wight & Arnott ex M. Roemer, Fam. Nat. Syn. Monogr. 

1: 47. 1846. 

小芸木 xiao yun mu 

Trees 6–8 m tall. Bark gray, smooth. Leaves 7–15-foliolate 

or 5–11-foliolate; petiolules 2–5 mm; leaflet blades alternate to 

subopposite, elliptic, lanceolate, or ovate, inequilateral, both 

surfaces dark green, margin entire, apex acuminate. Flower buds 

pale green, oblong. Calyx lobes ca. 1 mm. Petals pale yellow, 

5–10 mm, outside densely pubescent. Disk columnar. Stigma 

capitate. Fruit ellipsoid to obovoid, 1–1.5 × 0.6–1.2 cm, 1- or 

2-seeded. Seed coat membranous. Fl. Feb–Apr, fr. Jul–Sep. 2n = 

18. 

Moist montane forests, valley forests, maritime thickets in sandy 

soil; near sea level to 2000 m. SW Guangdong, Guangxi, S and SW 

Guizhou, Hainan, SE Xizang, S Yunnan [Bhutan, Cambodia, India, 

Laos, Myanmar, Nepal, Philippines, Thailand, Vietnam]. 

1a. Young parts rust-colored pubescent; 

longest leaflet blades to 20 cm ......... 2a. var. integerrimum 

1b. Young parts rust-colored hispid; 

longest leaflet blades to 24 cm .......... 2b. var. mollissimum 

2a. Micromelum integerrimum var. integerrimum 

小芸木(原变种) xiao yun mu (yuan bian zhong) 

Bergera integerrima Buchanan-Hamilton ex Candolle, 

Prodr. 1: 537. 1824. 

79

80 

Trees to 8 m tall. Young parts rust-colored pubescent. 

Leaves 7–15-foliolate; leaflet blades basally on rachis ca. 4 cm 

and apically to 20 × 8 cm. Fruit 1–1.5 × 0.7–1.2 cm. 

Moist montane forests, maritime thickets in sandy soil; near sea 

level to 2000 m. SW Guangdong, Guangxi, S and SW Guizhou, Hainan, 

SE Xizang, S Yunnan [Bhutan, Cambodia, India, Laos, Myanmar, 

Nepal, Thailand, Vietnam]. 

2b. Micromelum integerrimum var. mollissimum Tanaka, 

Bull. Mus. Natl. Hist. Nat., sér. 2, 2: 157. 1930. 


毛叶小芸木 mao ye xiao yun mu 

Trees 6–8 m tall. Young parts rust-colored hispid. Leaves 

5–11-foliolate; leaflet blades 10–24 × 4–8 cm. Young fruit 

ellipsoid, ca. 1 × 0.6–0.7 cm, pubescent, glabrescent when ripe, 

vermilion. Fl. Feb–Apr, fr. Jul–Sep. 

Valley forests; 100–600 m. SE Guangxi, S Yunnan [Cambodia, 

Laos, Philippines, Vietnam]. 

15. GLYCOSMIS Corrêa, Ann. Mus. Natl. Hist. Nat. 6: 384. 1805, nom. cons. 

山小橘属 shan xiao ju shu 


Shrubs or trees, unarmed, with rust-colored villosulous indumentum (sometimes becoming bleached) on terminal and axillary 

buds and usually on young inflorescences. Leaves alternate [rarely opposite], odd-pinnate, 1-foliolate, or simple. Inflorescences 

terminal and/or axillary, paniculate, compoundly racemose, or reduced to 1 or a few flowers. Flowers bisexual, globose to ellipsoid in 

bud. Sepals 4 or 5, basally connate. Petals 4 or 5, imbricate in bud. Stamens 8 or 10, distinct, alternately ± unequal in length; filaments 

± straight. Disk annular, pulvinate, columnar, conic, or bell-shaped. Gynoecium 2–5-loculed, syncarpous; radial walls of locules 

straight; ovules 1(or 2) per locule; style to nearly as long as ovary, persistent in fruit. Fruit a berry, with mucilaginous pulp or dry, 

without pulp vesicles; endocarp membranous. Seeds with membranous seed coat; endosperm lacking; embryo straight; cotyledons 

elliptic, plano-convex, neither convolute nor folded; hypocotyl partly included between cotyledons. 

About 50 species: E, S, and SE Asia, Australia; 11 species (two endemic) in China. 

1a. Leaves, or most of them, simple or 1-foliolate. 

2a. Leaves, or most of them, simple .................................................................................................................. 1. G. cochinchinensis 

2b. Leaves, or most of them, 1-foliolate. 

3a. Leaflet blades lustrous when dry; petiole (0.6–)1.5–3 cm .............................................................................. 2. G. montana 

3b. Leaflet blades not lustrous when dry; petiole 0.2–0.5(–1.6) cm. 

4a. Inflorescences less than 1 cm; anther connective with an apical oil gland and 2 or 3 abaxial oil 

glands ............................................................................................................................................ 3. G. pseudoracemosa 

4b. Inflorescences 2–5 cm; anther connective with only an apical oil gland ................................................ 4. G. longifolia 

1b. Leaves, or most of them, 2–7-foliolate. 

5a. Margin of leaflet blades serrate or dentate; inflorescences often longer than 10 cm. 

6a. Staminal filaments widest in ± their apical half; ovary glabrous ............................................................... 5. G. pentaphylla 

6b. Staminal filaments widest in ± their basal half; ovary rust-colored villosulous ............................................ 6. G. esquirolii 

5b. Margin of leaflet blades entire or repand; inflorescences rarely longer than 10 cm. 

7a. Ovary terete; petals ca. 5 mm, not caducous ....................................................................................................... 7. G. lucida 

7b. Ovary globose or ovoid; petals less than 5 mm, caducous. 

8a. Leaves (3–)5–7-foliolate .......................................................................................................................... 8. G. oligantha 

8b. Leaves (1 or)2–5-foliolate. 

9a. Leaflet blades concolorous; petals ca. 4 mm .................................................................................. 10. G. parviflora 

9b. Leaflet blades discolorous; petals ca. 1.5 or ca. 3 mm. 

10a. Petals ca. 1.5 mm; leaflet blades 5–9 cm wide ................................................................... 11. G. xizangensis 

10b. Petals ca. 3 mm; leaflet blades 2–3 cm wide ................................................................................ 9. G. craibii 

1. Glycosmis cochinchinensis (Loureiro) Pierre in Engler & 

Prantl, Nat. Pflanzenfam. 3(4): 185. 1896. 

山橘树 shan ju shu 

Toluifera cochinchinensis Loureiro, Fl. Cochinch. 1: 262. 

1790; Glycosmis cochinchinensis var. contracta Craib; G. 

parkeri Narayanaswamy; G. touranensis Guillaumin. 

Shrubs or trees, 1–4 m tall. Leaves, or most of them, 

simple; petiole 3–10 mm; leaf blade suborbicular, broadly 

elliptic, ovate, oblong, or lanceolate, 4–26 × 2–8 cm, papery to 

leathery, glabrous, base rounded, obtuse, cuneate, or attenuate, 

margin entire, apex rounded, obtuse, mucronate, or acuminate. 

Inflorescences axillary or terminal, with flowers conglomerate, 

rarely solitary, or in panicle ca. 5 cm. Sepals ovate, less than 1 

mm. Petals white, ca. 3 mm. Stamens 10; filaments widest in ± 

their basal half. Disk obvious. Ovary globose; style short, narrow; 

stigma slightly expanded. Fruit reddish, 0.8–1.4 cm in 

diam. Fl. and fr. year-round. 

Below 1000 m. Guangdong, S Guangxi, Hainan, S Yunnan [Cambodia, 

Laos, Myanmar, Thailand, Vietnam].

2. Glycosmis montana Pierre, Fl. Forest. Cochinch. 3: t. 285b. 

1893. 

海南山小橘 hai nan shan xiao ju 

Glycosmis hainanensis C. C. Huang; G. tonkinensis Tanaka 

ex Guillaumin; Tetracronia cymosa Pierre. 

Shrubs or trees, 1–3 m tall. Leaves, or most of them, 

1-foliolate; petiole (0.6–)1.5–3 cm; petiolule 2–10 mm; leaflet 

blades obovate-oblong to oblanceolate, 5–15 × 1.5–6.5 cm, 

papery to leathery, glabrous, base often obtuse, margin entire, 

apex abruptly acuminate to elongately caudate. Inflorescences 

paniculate, 1–3 cm. Flowers small, globose in bud. Sepals 

broadly ovate, less than 1 mm. Petals white, ca. 3 mm, caducous. 

Stamens 10; filaments widest in ± their basal half. Disk obviously 

uplifted. Ovary broadly ovoid to subglobose; style stout; 

stigma slightly expanded. Fruit pink, globose, ca. 8 mm in diam. 

Fl. Oct–Mar, fr. Jul–Sep. 

Woods on hillsides, near rivers; 200–500 m. Guangdong, Hainan, 

SE Yunnan (Funing) [Vietnam]. 

Stone (Proc. Acad. Nat. Sci. Philadelphia 137: 10. 1985) placed 

Glycosmis montana in synonymy of G. lanceolata (Blume) Sprengel ex 

Teijsmann & Binnendijk. 

3. Glycosmis pseudoracemosa (Guillaumin) Swingle, Notul. 

Syst. (Paris) 2: 162. 1912. 

华山小橘 hua shan xiao ju 

Atalantia pseudoracemosa Guillaumin, Notul. Syst. (Paris) 

1: 181. 1911; Glycosmis sinensis C. C. Huang. 

Shrubs or trees, 1.5–3 m tall. Leaves, or most of them, 

1-foliolate; petiole 2–5(–16) mm; petiolule 1–5 mm; leaflet 

blades oblong to lanceolate, 4–18 × 1.5–7 cm, papery, margin 

entire, apex acuminate. Inflorescences terminal or axillary, with 

flowers conglomerate or forming a short cyme, less than 1 cm. 

Sepals broadly ovate, less than 1 mm. Petals 5, white, ca. 4 mm. 

Stamens 10; filaments widest in ± their basal half. Ovary ovoid, 

becoming clavate; style extremely short. Fruit orange, broadly 

ellipsoid to subglobose, 1–1.2 × 0.8–1 cm. Fl. nearly year-round, 

fr. May–Dec. 

Hilly areas in woods and thickets; 400–1200 m. SW and W 

Guangxi, SE Yunnan (Malipo) [N Vietnam]. 

4. Glycosmis longifolia (Oliver) Tanaka, Bull. Soc. Bot. France 

75: 709. 1928. 

长叶山小橘 chang ye shan xiao ju 

Glycosmis pentaphylla (Retzius) Candolle subvar. longifolia 

Oliver, J. Linn. Soc., Bot. 5(Suppl. 2): 37. 1861; G. cyanocarpa 

(Blume) Sprengel var. simplicifolia Kurz; G. cymosa 

(Kurz) J. Narayanaswami ex Tanaka var. simplicifolia (Kurz) J. 

Narayanaswami. 

Trees to 5 m tall. Bark brownish gray. Leaves, or most of 

them, 1-foliolate; petiole less than 5 mm; leaflet blade oblong, 

10–18 × 4–5 cm, papery, base cuneate, margin entire, apex 

acuminate. Inflorescences axillary or terminal, paniculate, 2–5 

cm. Sepals broadly ovate, less than 1 mm. Petals 3–4 mm, 


caducous. Stamens 10; filaments widest in ± their basal half. 

Ovary globose; style short; stigma slightly expanded. Fruit 

oblong to obovoid, 10–15 × 8–10 mm, 1-seeded. Fl. Apr–Aug. 

Valley woods; ca. 1300 m. SW and W Yunnan (Lushui, Zhenkang) 

[India, NE Myanmar, Sri Lanka]. 


Glycosmis longifolia in synonymy of G. cyanocarpa (Blume) Sprengel. 

5. Glycosmis pentaphylla (Retzius) Candolle, Prodr. 1: 538. 

1924. 

山小橘 shan xiao ju 

Limonia pentaphylla Retzius, Observ. Bot. 5: 24. 1789; 

Glycosmis arborea (Roxburgh) Candolle; G. chylocarpa Wight 

& Arnott; G. quinquefolia Griffith; Limonia arborea Roxburgh; 

Myxospermum chylocarpum (Wight & Arnott) M. Roemer. 

Trees to 5 m tall. Leaves (3 or)5-foliolate; petiolules 2–10 

mm; leaflet blades oblong, 10–25 × 3–7 cm, papery, base 

cuneate, margin serrate, apex mucronate. Inflorescences axillary 

or terminal, paniculate. Flowers globose in bud. Sepals broadly 

ovate, less than 1 mm. Petals white or pale yellow, 3–4 mm, 

caducous. Stamens 10. Ovary globose to broadly ovoid; style extremely 

short; stigma slightly expanded. Fruit reddish, subglobose, 

8–10 mm in diam. Fl. Jul–Oct, fr. Jan–Mar. 2n = 36, 54. 

Hillside and valley woods; 600–1200 m. S and SW Yunnan [Bhutan, 

Cambodia, NE India, Indonesia, Laos, Malaysia, Myanmar, Nepal, 

Pakistan, Philippines, Sri Lanka, Thailand, NW Vietnam]. 

6. Glycosmis esquirolii (H. Léveillé) Tanaka, Bull. Soc. Bot. 

France 75: 709. 1928. 

锈毛山小橘 xiu mao shan xiao ju 

Clausena esquirolii H. Léveillé, Repert. Spec. Nov. Regni 

Veg. 9: 324. 1911; C. ferruginea C. C. Huang; Glycosmis ferruginea 

(C. C. Huang) C. C. Huang; G. winitii Craib. 

Trees 6–10 m tall. Leaves 4–7-foliolate; leaflet blades 

ovate to oblong, 10–16 × 4–7 cm, base cuneate, margin dentate, 

apex acuminate to obtuse. Inflorescences axillary or terminal, 

paniculate. Sepals broadly ovate, ca. 1 mm. Petals pale yellowish 

white, 3–4 mm. Stamens 10; filaments widest in ± their basal 

half; anthers elliptic, ca. 1 mm. Ovary subglobose, rust-colored 

villosulous; style extremely short. Fl. Oct–Mar, fr. Apr. 

Mountain woods and thickets; 400–1300 m. W Guangxi, S Guizhou, 

S and SE Yunnan [Myanmar, Thailand]. 

7. Glycosmis lucida Wallich ex C. C. Huang, Guihaia 7: 119. 

1987. 

亮叶山小橘 liang ye shan xiao ju 

Glycosmis cyanocarpa (Blume) Sprengel var. cymosa 

Kurz; G. cymosa (Kurz) J. Narayanaswami ex Tanaka (1941), 

not Zippelius ex Spanoghe (1841). 

Trees 3–5 m tall. Leaves (1 or)3- or 5(or 6)-foliolate; 

petiolules 2–8 mm; leaflet blades oblong to lanceolate, 7–18 × 

3–6 cm, margin entire or repand, apex acuminate. Inflorescences 

81

82 

axillary or terminal, paniculate, 1–4 cm, many flowered. Sepals 

broadly ovate, ca. 1 mm, margin ciliate. Petals ca. 5 × 2–3 mm. 

Stamens 10; filaments linear. Ovary terete. Fruit orange, broadly 

ellipsoid to ovoid, 1–1.4 × 0.8–1.2 cm, 1-seeded. Fl. and fr. 

year-round but flowering peak Jan–Mar. 

Mountain woods; 900–1400 m. Yunnan [Bhutan, India, NE 

Myanmar]. 


Glycosmis lucida in synonymy of G. cyanocarpa (Blume) Sprengel. 

8. Glycosmis oligantha C. C. Huang, Guihaia 7: 122. 1987. 

少花山小橘 shao hua shan xiao ju 

Shrubs or trees, to 3 m tall. Leaves (3 or)5- or 7-foliolate; 

petiolules 3–5 mm; leaflet blades narrowly lanceolate, 5–9 × 

1.5–2.5 cm, base attenuate, margin entire or repand, apex acuminate. 

Inflorescences axillary, 1–5-flowered. Sepals broadly 

ovate, ca. 1 mm wide. Petals ovate, ca. 3 mm, glabrous, apex 

obtuse. Stamens 10, longer ones ca. 2.5 mm, shorter ones ca. 1.5 

mm. Ovary globose; style extremely short. Fruit not seen. 

● Hillside woods; 200–500 m. SW Guangxi. 

9. Glycosmis craibii Tanaka, Bull. Mus. Natl. Hist. Nat., sér. 2, 

2: 159. 1930. 

毛山小橘 mao shan xiao ju 

Glycosmis puberula Lindley var. craibii (Tanaka) B. C. 

Stone. 

Trees to 5 m tall. Leaves (1–)3- or 5-foliolate; leaflet blades 

oblong, lanceolate, or ovate, 5–10 × 2–3 cm, papery or rigidly 

so, base attenuate to cuneate, margin entire, apex acuminate to 

mucronate. Inflorescences axillary or terminal, less than 3 cm or 

rarely ca. 4 cm. Sepals broadly ovate to ovate, less than 1 mm or 

ca. 1 mm. Petals ca. 3 mm. Filaments linear or widest in ± their 

basal half. Ovary terete in bud, becoming broadly conic, gourdshaped, 

or ovoid at anthesis, slightly pubescent or glabrous. Fruit 

reddish to orange, subglobose, 0.8–1.4 cm in diam., 1- or 

2-seeded. Fl. and fr. year-round. 

Hillside forests, woods, and thickets. Hainan, S Yunnan [N and NE 


Stone (Proc. Acad. Nat. Sci. Philadelphia 137: 18. 1985) reduced 

Glycosmis craibii to a variety of G. puberula Lindley. 

1a. Filaments linear; ovary slightly pubescent 

...................................................................... 9a. var. craibii 

1b. Filaments widest in ± their basal half; ovary 

glabrous ....................................................... 9b. var. glabra 

9a. Glycosmis craibii var. craibii 

毛山小橘(原变种) mao shan xiao ju (yuan bian zhong) 

Trees to 3 m tall. Leaflet blades oblong to lanceolate, 7–9 × 

2.5–3 cm, papery, base attenuate, apex acuminate. Inflorescences 

axillary, less than 3 cm, few flowered. Sepals broadly 

ovate, ca. 1 mm wide. Petals subelliptic. Stamens slightly longer 

than petals; filaments linear. Ovary conic to gourd-shaped at 

anthesis, slightly pubescent. Fruit reddish to orange, 0.8–1 cm in 

diam., 1-seeded. Fl. Apr–May, fr. Aug–Oct. 


Hillside forests. S Yunnan [NE Thailand]. 

9b. Glycosmis craibii var. glabra (Craib) Tanaka, Bull. Mus. 

Natl. Hist. Nat., sér. 2, 2: 159. 1930. 

光叶山小橘 guang ye shan xiao ju 

Glycosmis singuliflora Craib var. glabra Craib, Fl. Siam. 1: 

226. 1926. 

Trees to 5 m tall. Petiolules 2–6 mm; leaflet blades oblong, 

lanceolate, or ovate, 5–10 × 2–3 cm, rigidly papery, base attenuate 

to cuneate, apex acuminate to mucronate. Inflorescences 

axillary or terminal, rarely to 4 cm. Sepals ovate, less than 1 mm. 

Petals caducous. Stamens 10; filaments widest in ± their basal 

half. Ovary broadly ovoid at anthesis, glabrous. Fruit orange, 

1–1.4 cm in diam., 1- or 2-seeded. Fl. and fr. year-round. 

Hillside woods and thickets; 300–500 m. Hainan [N Thailand, 

Vietnam]. 


Glycosmis craibii var. glabra in synonymy of G. ovoidea Pierre. 

10. Glycosmis parviflora (Sims) Little, Phytologia 2: 463. 

1948. 

小花山小橘 xiao hua shan xiao ju 

Limonia parviflora Sims, Bot. Mag. 50: t. 2416. 1823; 

Citrus erythrocarpa Hayata; Glycosmis citrifolia Lindley; G. 

erythrocarpa (Hayata) Hayata; L. citrifolia Willdenow (1809), 

not Salisbury (1796). 

Shrubs or trees, 1–3 m tall. Leaves (1 or)2–4(or 

5)-foliolate; petiolules 1–5 mm; leaflet blades elliptic, oblong, or 

lanceolate, 5–19 × 2.5–8 cm, glabrous, base cuneate, margin 

entire, apex mucronate, acuminate, or obtuse. Inflorescences 

axillary or terminal, paniculate, 3–5 cm when axillary, to 14 cm 

when terminal. Sepals ovate, ca. 1 mm wide, apex obtuse. Petals 

white, ca. 4 mm, oblong. Stamens (8 or)10. Ovary broadly ovoid 

to globose; style extremely short; stigma slightly expanded. Fruit 

pale yellowish white but turning reddish to dark vermilion, 

globose to ellipsoid, 1–1.5 cm in diam., (1 or)2- or 3-seeded. Fl. 

Mar–May, fr. Jul–Sep. 2n = 54. 

Mountain woods; 200–1000 m. Fujian, Guangdong, Guangxi, 

Guizhou, Hainan, Taiwan, S and W Yunnan [Japan (including Ryukyu 

Islands), Myanmar, NE Vietnam]. 

11. Glycosmis xizangensis (C. Y. Wu & H. Li) D. D. Tao, Acta 

Phytotax. Sin. 32: 369. 1994. 

西臧山小橘 xi zang shan xiao ju 

Walsura xizangensis C. Y. Wu & H. Li, Acta Phytotax. Sin. 

18: 110. 1980; Glycosmis medogensis D. D. Tao ex C. C. Huang, 

nom. illeg. superfl.; G. motuoensis D. D. Tao, nom. illeg. 

superfl. 

Shrubs or trees, to 3 m tall. Leaves 3- or 4-foliolate; 

petiolules 4–10 mm; leaflet blades alternate to subopposite, 

broadly ovate to obovate-elliptic, 12–20 × 5–9 cm, subleathery, 

abaxially pale green, adaxially dark green, base cuneate, margin 

entire or repand, apex mucronate. Inflorescences axillary, 2.5–4

cm. Sepals broadly ovate, less than 2 mm, margin ciliate. Petals 

oblong, ca. 1.5 mm, glabrous, apex obtuse. Stamens 10; filaments 

flattened; anthers elliptic. Ovary subglobose; style terete. 

Fruit not seen. 


● Hillside woods; ca. 800 m. SE Xizang (Mêdog). 

The later names Glycosmis medogensis and G. motuoensis are 

based on the same type (Qinghai-Xizang Exped. 74–4540) as Walsura 

xizangensis. 

16. CLAUSENA N. L. Burman, Fl. Indica, 87, 243. 1768. 

黄皮属 huang pi shu 


Shrubs or trees, unarmed, without rust-colored villosulous indumentum on terminal and axillary buds or young inflorescences. 

Leaves alternate, odd-pinnate. Inflorescences terminal or axillary, paniculate or in loose racemes. Flowers bisexual or very rarely 

female, globose to pyriform or rarely ovoid in bud. Sepals 4 or 5, connate at base or to their full length. Petals 4 or 5, imbricate in bud. 

Stamens 8 or 10, distinct, alternately unequal in length; filaments ± abruptly dilated toward base, ± straight or geniculate. Disk 

columnar, conic, bell-shaped, or ± hourglass-shaped. Gynoecium (2–)4- or 5-loculed, syncarpous; radial walls of locules straight; 

ovules 2 per locule; style 0.5–2.5 × as long as ovary, deciduous in fruit. Fruit a berry, with neither pulp nor pulp vesicles; endocarp 

membranous. Seeds with membranous seed coat; endosperm lacking; embryo straight; cotyledons elliptic, plano-convex, neither 

convolute nor folded; hypocotyl partly included between cotyledons. 

Between ca. 15 and 30 species: Africa, E, S, and SE Asia, NE Australia, SW Pacific islands; ten species (five endemic) in China. 

This genus is especially diverse in Guangdong, Guangxi, and Yunnan. 

1a. Calyx lobes 4; petals 4(or 5); stamens 8. 

2a. Leaves 15–37-foliolate; fruit vermilion or pale yellow. 

3a. Fruit vermilion; leaflet blades 2–9 × 1–3 cm .................................................................................................. 1. C. excavata 

3b. Fruit pale yellow; leaflet blades not more than 2 × 1 cm ........................................................................... 2. C. hainanensis 

2b. Leaves not more than 15-foliolate; fruit bluish black. 

4a. Leaves 5–15-foliolate; leaflet blades asymmetric .......................................................................................... 3. C. dunniana 

4b. Leaves 5–7-foliolate; leaflet blades symmetric ................................................................................................... 4. C. vestita 

1b. Calyx lobes 5; petals (4 or)5; stamens 10. 

5a. Leaves 19–25-foliolate; stigma capitate or shield-shaped ....................................................................................... 5. C. odorata 

5b. Leaves not more than 15-foliolate; stigma punctiform to capitellate. 

6a. Fruit bluish black ................................................................................................................................................... 10. C. lenis 

6b. Fruit pale yellow to reddish. 

7a. Ovary and fruit pubescent; flower buds 5-ridged ....................................................................................... 6. C. lansium 

7b. Ovary pubescent or glabrous but fruit glabrous; flower buds not ridged. 

8a. Fruit longer than wide; leaflets more than 5 cm wide at middle of rachis ................................... 7. C. yunnanensis 

8b. Fruit ± as wide as long; leaflets rarely to 5 cm wide at middle of rachis. 

9a. Leaflet blades 2–6 × 1–3 cm, subsessile .................................................................................. 8. C. emarginata 

9b. Leaflet blades 5–12 × 2–4 cm, subtended by a 2–4 mm petiolule ....................................... 9. C. anisum-olens 

1. Clausena excavata N. L. Burman, Fl. Indica, 89. 1768. 

假黄皮 jia huang pi 

Amyris punctata Roxburgh; Clausena excavata var. lunulata 

(Hayata) Tanaka; C. forbesii Engler; C. lunulata Hayata; C. 

moningerae Merrill; C. punctata (Roxburgh) Wight & Arnott; 

C. tetramera Hayata; Lawsonia falcata Loureiro. 

Shrubs 1–2 m tall. Branchlets and leaf rachises pubescent, 

with oil glands. Leaves 21–27-foliolate but on young plants to 

41-foliolate; petiolules 2–5 mm; leaflet blades ovate, lanceolate, 

or rhomboid, asymmetric, 2–9 × 1–3 cm, both surfaces pubescent 

or only pubescent along veins, base oblique, margin repand. 

Inflorescences terminal; bracts opposite. Flowers globose in 

bud. Petals white or pale yellowish white, ovate to obovate, 2–3 

× 1–2 mm. Stamens 8; filaments basally dilated, geniculate at 

middle, apically linear. Style stout. Fruit ellipsoid, 1.2–1.8 × 

0.8–1.5 cm, 1- or 2-seeded. Fl. Apr–May and Jul–Aug(–Oct in 

Hainan), fr. Aug–Oct. 2n = 36. 

Below 1000 m. S Fujian, Guangdong, Guangxi, Hainan, Taiwan, S 

Yunnan [Bangladesh, Bhutan, Cambodia, India, Indonesia, Laos, Malaysia, 

Myanmar, Nepal, Philippines, Thailand, Vietnam]. 

2. Clausena hainanensis C. C. Huang & F. W. Xing, Guihaia 

12: 215. 1992. 

海南黄皮 hai nan huang pi 

Shrubs or trees, to 5 m tall. Leaves 25–37-foliolate; petiolules 

ca. 1 mm; leaflet blades alternate or opposite, asymmetrically 

elliptic, ca. 2 × 1 cm, midvein slightly depressed, base 

oblique and obtuse, margin repand, apex obtuse. Flowers not 

known. Infructescences terminal, narrowly paniculate, ca. 5 cm. 

Fruit pale yellow, ellipsoid, ca. 8 × 5 mm, pubescent. Fr. 

Jul–Aug. 

● Limestone mountains; 900–1000 m. Hainan (Changjiang). 

3. Clausena dunniana H. Léveillé, Repert. Spec. Nov. Regni 

Veg. 11: 67. 1912. 

齿叶黄皮 chi ye huang pi 

83

84 

Trees 2–5 m tall, deciduous. Leaves 5–15-foliolate; petiolules 

4–8 mm; leaflet blades ovate to lanceolate, 4–10 × 2–5 cm, 

glabrous or villous, base asymmetric, margin serrate or rarely 

repand, apex acute to acuminate. Inflorescences terminal. Flowers 

4(or 5)-merous, globose in bud. Stamens 8(or 10); filaments 

geniculate at middle, subulate at apex. Disk small. Ovary globose; 

style shorter than ovary. Fruit bluish black when ripe, globose, 

1–1.5 cm in diam., 1- or 2-seeded. Fl. Jun–Jul, fr. Oct– 

Nov. 

Montane forests, moist areas in mountains; 300–1500 m. Guangdong, 

Guangxi, Guizhou, W Hubei, Hunan, E and SE Sichuan, S Yunnan 

[NE Vietnam]. 

Molino (Bull. Mus. Natl. Hist. Nat., B, Adansonia 16: 130. 1994) 

placed Clausena dunniana in synonymy of C. anisata (Willdenow) J. D. 

Hooker ex Bentham. 

1a. Leaflet blades glabrous ........................... 3a. var. dunniana 

1b. Leaflet blades villous ................................. 3b. var. robusta 

3a. Clausena dunniana var. dunniana 

齿叶黄皮(原变种) chi ye huang pi (yuan bian zhong) 

Clausena dentata (Willdenow) M. Roemer var. dunniana 

(H. Léveillé) Swingle. 

Leaflet blades glabrous. 

Montane forests; 300–1500 m. Guangdong, Guangxi, Guizhou, 

Hunan, SE Sichuan, S Yunnan [NE Vietnam]. 


placed Clausena dentata var. dunniana in synonymy of C. anisata. 

3b. Clausena dunniana var. robusta (Tanaka) C. C. Huang, 


毛齿叶黄皮 mao chi ye huang pi 

Clausena dentata var. robusta Tanaka, J. Bot. (Morot) 66: 

228. 1930; C. dentata var. henryi Swingle; C. henryi (Swingle) 

C. C. Huang. 

Leaflet blades villous. Leaflet blades and fruit slightly 

larger than those of autonym variety. 

● Moist areas in mountains; 300–1300 m. W Hubei, Hunan, 

Guangxi, Guizhou, E Sichuan, S Yunnan. 


placed Clausena dunniana var. robusta, C. dentata var. robusta, C. 

dentata var. henryi, and C. henryi in synonymy of C. anisata. 

4. Clausena vestita D. D. Tao, Acta Bot. Yunnan. 6: 73. 1984. 

毛叶黄皮 mao ye huang pi 

Trees to 4 m tall. Leaves 5–7-foliolate; petiolules 2–4 mm; 

leaflet blades alternate, broadly ovate, elliptic, or orbicular, 3–11 

× 1.5–8 cm, apex rounded, obtuse, or acute. Calyx 4-parted, 

persistent in fruit. Fruit bluish black, globose to broadly ellipsoid, 

1.2–1.6 cm in diam., glabrous, 1–3-seeded. Seeds ovoid to 

broadly ellipsoid, 8–12 × 6–9 mm. Fr. May. 

● Thickets in dry hot river valleys; ca. 1900 m. NW Yunnan 

(Lijiang). 



placed Clausena vestita in synonymy of C. anisata (Willdenow) J. D. 


5. Clausena odorata C. C. Huang, Acta Phytotax. Sin. 8: 92. 

1959. 

香花黄皮 xiang hua huang pi 

Trees to 2 m tall. Shoots dark purplish red. Leaves 19– 

25-foliolate; petiolules 1–2 mm; leaflet blades oblong to lanceolate, 

asymmetric, 4–7 × 2–3.5 cm, base oblique, margin crenulate 

to subentire, apex acuminate and often retuse. Inflorescences 

terminal, paniculate. Flowers 5-merous, fragrant. Petals white, 

3–4 mm. Stamens 10. Stigma capitate to shield-shaped. Fl. Apr. 

● Thickets; ca. 1800 m. Yunnan (Mojiang). 


placed Clausena odorata in synonymy of C. anisata (Willdenow) J. D. 


6. Clausena lansium (Loureiro) Skeels, Bull. Bur. Pl. Industr. 

U.S.D.A. 168: 31. 1909. 

黄皮 huang pi 

Quinaria lansium Loureiro, Fl. Cochinch. 1: 272. 1790; 

Clausena wampi (Blanco) Oliver; Cookia wampi Blanco. 

Trees to 12 m tall. Leaves 5–11-foliolate; petiolules 4–8 

mm; leaflet blades ovate to ovate-elliptic, 6–14 × 3–6 cm, 

midvein often pubescent, base rounded to broadly cuneate, 

margin repand to crenulate. Inflorescences terminal, paniculate. 

Flowers globose in bud. Calyx lobes broadly ovate, ca. 1 mm. 

Petals oblong, ca. 5 mm. Stamens 10; filaments linear, basal 

portion slightly expanded. Disk short. Ovary hirsute. Fruit pale 

yellow, globose, ellipsoid, or broadly ovoid, 1.5–3 × 1–2 cm, 

1–4-seeded. Fl. Apr–May, fr. Jul–Aug (fl. and fr. ca. 1 month 

earlier in Hainan). 2n = 18. 

Fujian, Guangdong, Guangxi, S Guizhou, Hainan, Sichuan, SE 

Yunnan [Vietnam]. 

This species is cultivated for its fruit in China and elsewhere in 

warm areas of the world. 

7. Clausena yunnanensis C. C. Huang, Acta Phytotax. Sin. 8: 

91. 1959. 

云南黄皮 yun nan huang pi 

Trees 3–8 m tall. Bark gray. Branchlets stout; pith large. 

Leaves 5–11-foliolate; petiolules 4–6 mm; leaflet blades oblong 

to ovate-elliptic, 10–40 × 5–16 cm, papery, margin denticulate. 

Inflorescences terminal, paniculate, to 40 cm; bracts narrowly 

ovate, apex acute. Flowers globose in bud. Pedicel 1.5–3 mm. 

Sepals ovate, ca. 1 mm. Petals 2–3 mm. Stamens 10. Fruit 

orange, ellipsoid, ca. 3 × 2 cm, 1- or 2-seeded. Fl. Jun, fr. Sep– 

Oct. 

● Mountain forests, forests on limestone; 500–1300 m. SW and W 

Guangxi, SE Yunnan. 


placed Clausena yunnanensis in synonymy of C. engleri Tanaka. 

1a. Branchlets glabrate; leaflet blades 

abaxially shortly pubescent on 

venation .............................................. 7a. var. yunnanensis

1b. Branchlets densely pubescent; leaflet 

blades abaxially with dense tufts of 

long weak trichomes, especially on 

midvein ........................................... 7b. var. longgangensis 

7a. Clausena yunnanensis var. yunnanensis 

云南黄皮(原变种) yun nan huang pi (yuan bian zhong) 

Branchlets glabrate. Leaf rachises minutely pubescent; 

petiolules glabrous or minutely pubescent; leaflet blades abaxially 

shortly pubescent on venation, adaxially glabrous or 

glabrate. 

● Mountain forests. W Guangxi, SE Yunnan. 

7b. Clausena yunnanensis var. longgangensis C. F. Liang & 

Y. X. Lu, Guihaia 10: 104. 1990. 

弄岗黄皮 nong gang huang pi 

Clausena yunnanensis var. dolichocarpa C. F. Liang & Y. 

X. Lu ex C. C. Huang. 

Branchlets, leaf rachises, and petiolules densely pubescent. 

Leaflet blades abaxially with dense tufts of long weak trichomes, 

especially on midvein. 

● Forests on limestone. SW Guangxi (Ningming). 


placed Clausena yunnanensis var. longgangensis in synonymy of C. 

engleri. Clausena yunnanensis var. dolichocarpa was not mentioned. 

8. Clausena emarginata C. C. Huang, Acta Phytotax. Sin. 8: 

93. 1959. 

小黄皮 xiao huang pi 

Trees 4–15 m tall. Branchlets grayish black. Leaves 

5–11-foliolate; leaflet blades subsessile, obliquely lanceolate to 

ovate, 2–6 × 1–3 cm, dark brownish black when dry, base 

oblique, margin crenate, apex obtuse. Inflorescences terminal or 

axillary, 3–7 cm; bracts subulate. Calyx lobes broadly ovate. 

Petals ca. 4 mm at anthesis. Stamens 10; filaments slightly 

expanded in ± their basal half, geniculate, longer than anthers. 

Disk elongated. Ovary globose, glabrous. Fruit pale yellow, 


globose, 8–10 mm in diam., 1- or 2-seeded. Fl. Mar–Apr, fr. 

Jun–Jul. 

● Valley forests; 300–800 m. W Guangxi, SE Yunnan. 


placed Clausena emarginata in synonymy of C. sanki (Perrottet) J. F. 

Molino, which he later (Taxon 44: 427. 1995) decided should be called 

C. anisum-olens. 

9. Clausena anisum-olens (Blanco) Merrill, Publ. Bur. Sci. 

Gov. Lab. 17: 21. 1904. 

细叶黄皮 xi ye huang pi 

Cookia anisum-olens Blanco, Fl. Filip. 359. 1837; Clausena 

grandifolia Merrill; C. laxifolia Quisumbing & Merrill; C. 

loheri Merrill; C. todayensis Elmer; C. warburgii Perkins. 

Trees 3–6 m tall. Leaves 5–11-foliolate; petiolules 2–4 

mm; leaflet blades falcately lanceolate to obliquely ovate, 5–12 

× 2–4 cm, margin repand, apex acuminate to slightly obtuse. 

Inflorescences terminal. Flowers globose in bud, fragrant. Calyx 

lobes ovate, ca. 1 mm. Petals white, oblong, ca. 3 mm. Stamens 8 

or 10; filaments slightly expanded in ± their basal half, geniculate. 

Style slightly shorter than ovary. Fruit pale yellow, globose, 

1–2 cm in diam., 1- or 2-seeded. Fl. Apr–May, fr. Jul–Aug. 

Taiwan (Lan Yu); cultivated in Guangdong, Guangxi, Yunnan 

[Philippines]. 

10. Clausena lenis Drake, J. Bot. (Morot) 6: 276. 1892. 

光滑黄皮 guang hua huang pi 

Clausena kerrii Craib. 

Trees 2–3 m tall. Branchlet pith fairly large. Leaves 

9–15-foliolate; leaflet blades ovate to lanceolate, asymmetric, 

2–5 × 1.5–3.5 cm at base of rachis, ca. 18 × 11 cm at middle and 

apex of rachis, margin crenate. Inflorescences terminal. Flowers 

ovoid in bud. Calyx lobes 5. Petals (4 or)5, white but basally 

reddish to dark yellow. Stamens (8 or)10; filaments short; anthers 

oblong. Fruit globose, ca. 1 cm in diam., 1–3-seeded. Fl. 

Apr–Jun, fr. Sep–Oct. 

Mountain forests; 500–1300 m. Guangdong, SW Guangxi, Hainan, 

S Yunnan [Laos, Thailand, NE Vietnam]. 

17. MURRAYA J. Koenig ex Linnaeus, Mant. Pl. 2: 554, 563. 1771 [“Murraea”], nom. cons. 

Bergera J. Koenig ex Linnaeus; Camunium Adanson. 

九里香属 jiu li xiang shu 


Shrubs or trees, unarmed, without rust-colored villosulous indumentum on terminal and axillary bud or young inflorescences. 

Leaves alternate, odd-pinnate (occasional leaves even-pinnate or 1-foliolate). Inflorescences terminal or terminal and axillary, 

paniculate or reduced to cymes or few to several-flowered racemes. Flowers bisexual, ellipsoid to obovoid, or narrowly so or 

subcylindric in bud. Sepals 4 or 5, connate at base or to ± half their length. Petals 4 or 5, imbricate in bud. Stamens (5–)8 or 10, distinct, 

alternately unequal in length; filaments linear or sublinear, ± straight. Disk annular, pulvinate, or columnar. Gynoecium 2–5-loculed, 

syncarpous; radial walls of locules straight; ovules 1 or 2 per locule; style 3–7 × as long as ovary, deciduous in fruit or sometimes basal 

portion persistent. Fruit a berry, with mucilaginous pulp and without pulp vesicles; endocarp fleshy. Seeds with membranous or fleshy 

seed coat; endosperm lacking; embryo straight; cotyledons elliptic, plano-convex, neither convolute nor folded; hypocotyl partly 

included between cotyledons. 

85

86 


About 12 species: E, S, and SE Asia, Australia, SW Pacific islands; nine species (five endemic) in China. 

In a cladistic analysis of selected species of subfamily Aurantioideae based on plastid DNA sequences and phytochemical features, R. Samuel et 

al. (Pl. Biol. 3: 77–87. 2001) proposed that Murraya is heterogeneous and that plants of M. sect. Bergera (J. Koenig ex Linnaeus) But & Y. C. Kong 

(including species 4–9 in the present enumeration) are generically distinct from those of M. sect. Murraya. However, as they pointed out, it is not clear 

yet whether Bergera is distinct from Clausena. 

1a. Petals 1–2 cm; seeds villous. 

2a. Leaf rachises winged .................................................................................................................................................... 1. M. alata 

2b. Leaf rachises not winged. 

3a. Leaflet blades mostly suborbicular to ovate to elliptic, 1.5–6 cm wide ....................................................... 2. M. paniculata 

3b. Leaflet blades elliptic-obovate or obovate, 0.5–3 cm wide ............................................................................... 3. M. exotica 

1b. Petals not more than 0.8 cm; seeds glabrous. 

4a. Flowers 4(or 5)-merous; stamens 8(or 10). 

5a. Leaflet blades lanceolate, 0.8–2 cm wide, tertiary veins inconspicuous ...................................................... 4. M. tetramera 

5b. Leaflet blades ovate to elliptic or rarely lanceolate or obovate, 2–4 cm wide, tertiary veins adaxially 

slightly prominent .................................................................................................................................... 5. M. euchrestifolia 

4b. Flowers 5-merous; stamens 10. 

6a. Leaves 11–31-foliolate. 

7a. Leaflet blades 3–6 mm wide, apex rounded to obtuse ........................................................................ 6. M. microphylla 

7b. Leaflet blades 5–20 mm wide, apex mucronate to acuminate ................................................................... 7. M. koenigii 

6b. Leaves 3–11-foliolate. 

8a. Leaflet blades pubescent abaxially ..................................................................................................... 9. M. kwangsiensis 

8b. Leaflet blades glabrous. 

9a. Leaflet blades 5–6 × 2–3 cm ............................................................................................................ 8. M. crenulata 

9b. Leaflet blades 7–18 × 4–10 cm ................................................................................................... 9. M. kwangsiensis 

1. Murraya alata Drake, J. Bot. (Morot) 6: 276. 1892. 

翼叶九里香 yi ye jiu li xiang 

Murraya alata var. hainanensis Swingle. 

Shrubs 1–2 m tall. Branches yellowish gray to grayish 

white. Leaves 5–9-foliolate; petiolules ± short or leaflets subsessile; 

leaflet blades obovate to obovate-elliptic, 1–3 × 0.6–1.5 

cm, margin crenulate or entire, apex rounded or rarely obtuse. 

Inflorescences axillary, cymose. Flowers 5-merous. Sepals 

1.5–2 mm. Petals white, 10–15 × 3–5 mm. Stamens 10. Ovary 

2-loculed; ovules 1 per locule; style ca. 2 × as long as ovary; 

stigma capitate. Fruit vermilion, ovoid to globose, ca. 1 cm in 

diam., 2–4-seeded. Fl. May–Jul, fr. Oct–Dec. 

Thickets in sandy areas; near sea level. SW Guangdong (Leizhou 

Bandao), SW Guangxi (Beihai), S Hainan [NE Vietnam]. 

2. Murraya paniculata (Linnaeus) Jack, Malayan Misc. 1: 31. 

1820. 

千里香 qian li xiang 

Chalcas paniculata Linnaeus, Mant. Pl. 1: 68. 1767; C. 

exotica (Linnaeus) Millspaugh; Murraya omphalocarpa Hayata; 

M. paniculata var. exotica (Linnaeus) C. C. Huang; M. paniculata 

var. omphalocarpa (Hayata) Tanaka. 

Shrubs or trees, 1.8–12 m tall. Older branchlets grayish 

white to pale yellowish gray. Leaves 2–5-foliolate; petiolules 

less than 1 cm; leaflet blades mostly suborbicular to ovate to 

elliptic, 2–9 × 1.5–6 cm, margin entire or crenulate, apex 

rounded to acuminate. Inflorescences terminal or terminal and 

axillary. Flowers 5-merous, fragrant. Sepals ovate to lanceolate, 

to 2 mm, persistent in fruit. Petals white, narrowly elliptic to 

oblanceolate, to 2 cm. Stamens 10. Fruit orange to vermilion, 

narrowly ellipsoid or rarely ovoid, 1–2 × 0.5–1.4 cm. Seeds 

villous. Fl. Apr–Oct, fr. Apr–Feb. 2n = 18. 

Thickets, montane forests; near sea level to 1300 m. Fujian, 

Guangdong, Guangxi, S Guizhou, Hainan, S Hunan, Taiwan, Yunnan 

[Bhutan, Cambodia, India, Indonesia, Japan (Ryukyu Islands), Laos, 

Malaysia, Myanmar, Nepal, New Guinea, Pakistan, Philippines, Sri 

Lanka, Thailand, Vietnam; Australia, SW Pacific islands]. 

Murraya paniculata var. omphalocarpa was accepted by Chang 

(Fl. Taiwan, ed. 2, 3: 527. 1993). 

3. Murraya exotica Linnaeus, Mant. Pl. 2: 563. 1771. 

九里香 jiu li xiang 

Chalcas exotica (Linnaeus) Millspaugh; Murraya paniculata 

(Linnaeus) Jack var. exotica (Linnaeus) C. C. Huang. 

Trees to 8 m tall. Older branchlets grayish white to pale 

yellowish gray. Leaves 3–7-foliolate; petiolules rather short; 

leaflet blades elliptic-obovate or obovate, 1–6 × 0.5–3 cm, margin 

entire, apex rounded or obtuse. Inflorescences terminal or 

terminal and axillary. Flowers 5-merous, fragrant. Sepals ovate, 

ca. 1.5 mm. Petals white, oblong, 1–1.5 cm. Stamens 10. Fruit 

orange to vermilion, broadly ovoid, 8–12 × 6–10 mm. Seeds 

villous. Fl. Apr–Aug, fr. Sep–Dec. 2n = 18*. 

● Thickets; near sea level. S Fujian, Guangdong, Guangxi, S 

Guizhou, Hainan, Taiwan. 

This species is widespread in cultivation in tropical and subtropical 

areas. 

4. Murraya tetramera C. C. Huang, Acta Phytotax. Sin. 8: 102. 

1959. 

四数九里香 si shu jiu li xiang


mm; leaflet blades lanceolate, 2–5 × 0.8–2 cm, dark brownish 

black when dry, apex acuminate. Inflorescences paniculate. 

Flowers 4-merous. Sepals ovate, less than 1 mm, connate at 

base. Petals white, oblong, 4–5 mm. Stamens 8. Ovary ellipsoid, 

ca. 1 mm. Fruit reddish, globose, 1–1.2 cm in diam., with many 

oil glands, 1–3-seeded. Seed coat membranous, smooth. Fl. 

Mar–Apr, fr. Jul–Aug. 

● Often on limestone mountains. W Guangxi (Bose, Debao), SE 

Yunnan. 

5. Murraya euchrestifolia Hayata, Icon. Pl. Formos. 6: 11. 

1916. 

豆叶九里香 dou ye jiu li xiang 

Chalcas euchrestifolia Tanaka; Clausena euchrestifolia 

(Hayata) Kanehira. 

Shrubs or trees, 1.5–7 m tall. Leaves 5–9-foliolate; leaflet 

blades ovate to elliptic or rarely lanceolate or obovate, 5–8 × 2–4 

cm, margin entire, apex mucronate to acuminate. Inflorescences 

paniculate. Flowers 4(or 5)-merous. Sepals pale yellowish green, 

ovate, 0.6–1.5 mm. Petals obovate-elliptic, 3–5 mm. Stamens 

8(or 10); anthers subglobose. Ovary pale yellowish green. Fruit 

red to dark red, globose, 1–1.5 cm in diam., 1- or 2-seeded. 

Seeds glabrous. Fl. Apr–Jul, fr. Sep–Dec. 

● Well-drained forests and thickets; 500–1400 m. Guangdong 

(Fengkai, Nan’ao), Guangxi (Fangcheng), S and SW Guizhou (Wangmo, 

Xingyi), Hainan (Changjiang), Taiwan, Yunnan. 

6. Murraya microphylla (Merrill & Chun) Swingle, J. Wash. 

Acad. Sci. 32: 26. 1942. 

小叶九里香 xiao ye jiu li xiang 

Clausena microphylla Merrill & Chun, Sunyatsenia 2: 251. 

1935. 

Shrubs or small trees. Leaves 11–21-foliolate; petiolules 

extremely short; leaflet blades suborbicular to ovate to elliptic, 

3–20 × 3–6 mm, glabrous, base usually inequilateral, margin 

crenate, apex rounded to obtuse. Inflorescences terminal, 10– 

30-flowered. Flowers 5-merous. Sepals ovate-elliptic, ca. 1 mm, 

persistent in fruit. Petals in bud 4–5 mm. Fruit ellipsoid to 

ovoid-ellipsoid, to 1 cm. Fl. Apr–May and Jul–Oct, fr. Nov– 

Dec. 

● Thickets in sandy areas. Guangdong, Hainan. 

7. Murraya koenigii (Linnaeus) Sprengel, Syst. Veg. 2: 315. 

1817. 

调料九里香 tiao liao jiu li xiang 

Bergera koenigii Linnaeus, Mant. Pl. 2: 555, 563. 1771; 

Chalcas koenigii (Linnaeus) Kurz. 

Shrubs or trees, to 4 m tall. Leaves 17–31-foliolate; leaflet 

blades ovate, 2–5 × 0.5–2 cm, base obtuse to rounded and 

oblique, margin entire or crenulate. Inflorescences terminal, 

paniculate, many flowered. Flowers 5-merous, ellipsoid in bud. 

Sepals ovate, less than 1 mm. Petals white, oblanceolate to 


oblong, 5–7 mm. Stamens 10. Stigma capitate. Fruit bluish 

black, ovoid to oblong, 1–1.5 cm, 1- or 2-seeded. Seed coat 

membranous. Fl. Mar–Apr, fr. Jul–Aug. 

Moist forests; 500–1600 m. Guangdong, S Hainan, S Yunnan 

(Xishuangbanna) [Bhutan, India, Laos, Nepal, Pakistan, Sri Lanka, 


8. Murraya crenulata (Turczaninow) Oliver, J. Linn. Soc., Bot. 

5(Suppl. 2): 29. 1861. 

兰屿九里香 lan yu jiu li xiang 

Glycosmis crenulata Turczaninow, Byull. Moskovsk. 

Obshch. Isp. Prir., Otd. Biol. 30: 250. 1858; Chalcas crenulata 

(Turczaninow) Tanaka. 

Small trees. Branchlets and leaves glabrous. Leaves 7– 

11-foliolate; petiolules ca. 3 mm; leaflet blades ovate-elliptic, 

5–6 × 2–3 cm, base obtuse and oblique, apex acuminate. Inflorescences 

terminal, ca. 8 cm. Flowers 5-merous. Sepals suborbicular. 

Petals oblong, ca. 6 mm. Stamens 10. Ovary ca. 1.5 

mm; style ca. 2.5 mm, stout, basal portion often persistent in 

fruit. Fruit broadly ovoid to subglobose, ca. 6 mm. 

Taiwan (Lan Yu) [Indonesia, New Guinea, Philippines; SW Pacific 

islands]. 

9. Murraya kwangsiensis (C. C. Huang) C. C. Huang, Acta 

Phytotax. Sin. 16(2): 85. 1988. 

广西九里香 guang xi jiu li xiang 


mm; leaflet blades alternate, ovate-oblong, 7–18 × 3–10 cm, 

leathery, both surfaces glabrous or abaxially pubescent, margin 

crenulate, apex obtuse to rounded. Flowers 5-merous, ellipsoid 

in bud. Sepals broadly ovate, ca. 1 mm. Petals ca. 4 mm. Stamens 

10. Fruit globose, ca. 1 cm in diam. Fl. May, fr. Oct. 

● Forests and thickets in valleys; 200–800 m. SW and W Guangxi, 

SE Yunnan. 

1a. Leaflet blades abaxially pubescent 

........................................................... 9a. var. kwangsiensis 

1b. Leaflet blades glabrous ...................... 9b. var. macrophylla 

9a. Murraya kwangsiensis var. kwangsiensis 

广西九里香(原变种) guang xi jiu li xiang (yuan bian zhong) 

Clausena kwangsiensis C. C. Huang, Acta Phytotax. Sin. 8: 

90. 1959. 

Leaflet blades 3–10 × 2.5–6.5 cm, abaxially pubescent. 

● Forests and thickets in valleys; 200–800 m. SW and W Guangxi, 

SE Yunnan. 

9b. Murraya kwangsiensis var. macrophylla C. C. Huang, 


大叶九里香 da ye jiu li xiang 

Leaflet blades 7–18 × 4–10 cm, glabrous. Fruit red. 

● Valley forests. SW Guangxi. 

87

88 


18. LUVUNGA Buchanan-Hamilton ex Wight & Arnott, Prodr. Fl. Ind. Orient. 90. 1834. 

三叶藤橘属 san ye teng ju shu 


Woody climbers, with straight or recurved axillary spines, without rust-colored villosulous indumentum on terminal and axillary 

buds or young inflorescences. Leaves alternate, digitately 3-foliolate; leaflets petiolulate (occasional leaves 1-foliolate, including 

sometimes all of those on new shoots). Inflorescences terminal, axillary, or basal to leaves, paniculate or racemose. Flowers bisexual. 

Calyx cup-shaped, 3–5-lobed (often irregularly) or truncate. Petals 3–5, imbricate in bud. Stamens 6–10, equal in length, distinct or 

monadelphous. Disk annular, pulvinate, or columnar. Gynoecium 2–4-loculed, syncarpous; ovules 1 or 2 per locule; style to 4 × as long 

as ovary, deciduous in fruit. Fruit a berry, with mucilaginous pulp and without pulp vesicles; endocarp membranous or fleshy. Seeds 

with membranous seed coat; endosperm lacking; embryo straight; cotyledons elliptic, plano-convex; hypocotyl partly included 

between cotyledons. 

About ten species: S and SE Asia; one species in China. 

1. Luvunga scandens (Roxburgh) Buchanan-Hamilton ex 

Wight & Arnott, Prodr. Fl. Ind. Orient. 90. 1834. 

三叶藤 san ye teng 

Limonia scandens Roxburgh, Fl. Ind., ed. 1832, 2: 380. 

1832; Luvunga nitida Pierre. 

Heavy woody climbers. Leaves digitately 3-foliolate; 

petiole 2–9 cm; petiolules 3–10 mm; leaflet blades elliptic to 

obovate, 6–20 × 3–9 cm. Flowers ellipsoid in bud. Calyx 2.5–4 

mm, shallowly 4-lobed. Petals 4.6–10 mm. Stamens 8 or fewer. 

Fruit yellow, globose or obpyriform, 3–5 cm in diam., surface 

smooth, 1–4-seeded; outer part of pericarp (exocarp and 

mesocarp) thick. Seeds broadly ovoid, 2–3 cm. Fl. Mar–Apr, fr. 

Oct–Dec. 

Riverbanks, valleys of evergreen broad-leaved forests; below 600 

m. Guangdong, Hainan, S Yunnan [Cambodia, India, Laos, Malaysia, 

Myanmar, Thailand, Vietnam]. 

19. PARAMIGNYA Wight, Ill. Ind. Bot. 1: 108. 1838. 

单叶藤橘属 dan ye teng ju shu 


Woody climbers or erect or scrambling shrubs. Branchlets with straight or recurved spines or rarely unarmed, without 

rust-colored villosulous indumentum on terminal and axillary buds or young inflorescences. Leaves alternate, 1-foliolate or simple; 

petiole 0.4–2.5 cm, usually bent and/or twisted and ± swollen apically. Flowers bisexual, axillary, solitary or in few-flowered fascicles. 

Sepals 4 or 5, connate at base or to ca. 2/3 their length. Petals 4 or 5, imbricate in bud. Stamens 8 or 10, equal in length, distinct. Disk 

cup-shaped, conic, or columnar. Gynoecium 3–5-loculed, syncarpous; ovules 1 or 2 per locule; style to 4 × as long as ovary, deciduous 

in fruit. Fruit a berry, with copious mucilaginous pulp and without pulp vesicles; endocarp fleshy. Seeds with membranous seed coat; 

endosperm lacking; embryo straight; cotyledons elliptic, plano-convex; hypocotyl partly included between cotyledons. 

About 15 species: S and SE Asia, N Australia; one species in China. 

1. Paramignya confertifolia Swingle, J. Arnold Arbor. 21: 17. 

1940. 

单叶藤橘 dan ye teng ju 

Woody climbers or erect or scrambling shrubs, with recurved 

spines 2–8 mm or unarmed. Leaves 1-foliolate or simple; 

petiole 5–12 mm; leaf blade ovate to elliptic to obovate, 5–12 × 

2–4.5 cm, glabrous or abaxially grading from sparsely pubescent 

to ± densely pubescent on midvein, base rounded to obtuse, 

margin crenate toward apex or entire, apex acuminate to shortly 

so. Pedicel 3–5 mm. Flowers 5-merous, ellipsoid to obovoid in 

bud. Sepals 0.5–1.5 mm, pubescent and/or margin ciliate, per- 

sistent in fruit. Petals 6–10 mm, glabrous, deciduous in fruit. 

Stamens 10; filaments pubescent. Disk thickly columnar, ca. 1 

mm, glabrous, ± accrescent and forming a stipe in fruit. Ovary 

hirsute; style ca. 5 × as long as ovary, pubescent basally or 

glabrous. Fruit reportedly green, ± globose, larger ones 1.5–2 cm 

in diam., subtended by a 1.3–2 mm stipe. Fl. Jul–Sep, fr. 

Oct–Dec. 

Forests, streamsides, trailsides; 300–900 m. Guangdong, S 

Guangxi, Hainan, S Yunnan (Xishuangbanna) [N Vietnam]. 

Plants mistakenly treated as Paramignya rectispinosa Craib by C. 

C. Huang (Fl. Reipubl. Popularis Sin. 43(2): 155. 1997) belong here. 

20. ATALANTIA Corrêa, Ann. Mus. Natl. Hist. Nat. 6: 383, 385, 386. 1805, nom. cons. 

Severinia Tenore ex Endlicher. 

酒饼簕属 jiu bing le shu 

Zhang Dianxiang (张奠湘); Thomas G. Hartley


Shrubs or trees, unarmed or with straight axillary spines, without rust-colored villosulous indumentum on terminal and axillary 

buds or young inflorescences. Leaves alternate, 1-foliolate or simple; petiole 2–13 mm, usually not bent, twisted, or swollen. 

Inflorescences terminal and/or axillary, fasciculate, racemose, or paniculate. Flowers bisexual. Sepals 3–5, connate at base to nearly 

their full length, or calyx splitting irregularly into 2 or 4 segments. Petals 3–5, imbricate in bud. Stamens 6–10, equal or alternately 

unequal in length, distinct, monadelphous, filaments coherent in phalanges, or filaments irregularly coherent. Disk annular, 

cup-shaped, or columnar. Gynoecium 2–5-loculed, syncarpous; ovules 1 or 2 per locule; style to 1.5 × as long as ovary, deciduous in 

fruit. Fruit a berry, usually with pulp vesicles and with or without mucilaginous pulp; endocarp membranous. Seeds with membranous 

seed coat; endosperm lacking; embryo straight; cotyledons elliptic, plano-convex; hypocotyl partly included between cotyledons. 

About 17 species: S and SE Asia; seven species (two endemic) in China. 

1a. Leaves simple; petiole not articulated with leaf blade. 

2a. Branchlets with spines ca. 4 cm or rarely unarmed; leaf blade 2–6(–10) × 1–5 cm, apex rounded or rarely 

narrowly obtuse, retuse to emarginate at tip; stamens 10; ripe fruit bluish black ................................................. 1. A. buxifolia 

2b. Branchlets unarmed or rarely with sparse short spines; leaf blade 11–21 × 3–6(–10) cm, apex acuminate, 

sometimes retuse at tip; stamens 8; ripe fruit red ........................................................................................ 2. A. kwangtungensis 

1b. Leaves 1-foliolate; petiole articulated with leaf blade. 

3a. Fruit ellipsoid to ovoid but pyriform when young, ca. 3 cm in diam. .................................................................. 7. A. fongkaica 

3b. Fruit globose or subglobose, 1.2–3 cm in diam. 

4a. Flowers 5–6 mm in diam.; petals 3–4 mm; fruit smooth ..................................................................................... 3. A. henryi 

4b. Flowers 8–15 mm in diam.; petals 6–10 mm; fruit scabrous. 

5a. Young branchlets and leaves glabrous .................................................................................................... 6. A. acuminata 

5b. Young branchlets and leaves with indumentum. 

6a. Leaflet blade apex acuminate; petiole 6–10 mm; fruit 1.5–2 cm or more in diam. ........................ 4. A. dasycarpa 

6b. Leaflet blade apex mucronate to obtuse; petiole 10–12 mm; fruit 2.5–3 cm in diam. ................. 5. A. guillauminii 

1. Atalantia buxifolia (Poiret) Oliver ex Bentham, Fl. Hongk. 

51. 1861. 

酒饼簕 jiu bing le 

Citrus buxifolia Poiret, Encycl. 4: 580. 1797; Dumula 

sinensis Loureiro ex B. A. Gomes; Limonia monophylla 

Loureiro (1790), not Linnaeus (1767); Severinia buxifolia 

(Poiret) Tenore; S. monophylla Tanaka. 

Shrubs ca. 2.5 m tall. Branches grayish brown; branchlets 

green, with spines or rarely unarmed; spines ca. 4 cm, apex red. 

Leaves simple; petiole 1–7 mm; leaf blade ovate, obovate, elliptic, 

or suborbicular, 2–6(–10) × 1–5 cm, leathery, with oil glands 

and an orange smell, midvein slightly ridged, secondary veins 

joined in an arched marginal vein, apex rounded to obtuse and 

retuse to emarginate at tip. Inflorescences axillary, fasciculate, 

(1- or) several flowered. Flowers 5-merous, subsessile. Calyx 

persistent. Petals white, 3–4 mm, with oil glands. Stamens 10; 

filaments white, distinct or sometimes a few basally connate. 

Style green, ± as long as ovary. Fruit bluish black when ripe, 

globose, slightly oblate, or subellipsoid, 0.8–1.2 cm in diam., 

smooth, 1- or 2-seeded. Seeds with 1(or 2) embryos; cotyledons 

green, with many oil glands. Fl. May–Dec, fr. Sep–Dec. 2n = 36. 

Forests or thickets near ocean; below 300 m. S Fujian, S Guangdong, 

S Guangxi, Hainan, S Taiwan, Yunnan [Malaysia, Philippines, 

Vietnam]. 

2. Atalantia kwangtungensis Merrill, Philipp. J. Sci. 21: 496. 

1922. 

广东酒饼簕 guang dong jiu bing le 

Atalantia hainanensis Merrill & Chun ex Swingle; A. roxburghiana 

J. D. Hooker var. kwangtungensis (Merrill) Swingle. 

Shrubs 1–2 m tall. Young branchlets green, slightly flat, 

ridged. Leaf blade elliptic, lanceolate, or rarely obovate-elliptic, 

11–21 × 3–6(–10) cm, pale green, abaxially grayish yellow 

when dry, oil glands pellucid and conspicuous, base cuneate, 

margin sinuate. Inflorescences axillary, 5 mm or less, with 3 to 

several flowers in a fasciclelike raceme. Flowers 4-merous. 

Petals white, 3–5 mm. Stamens 8, monadelphous or filaments 

coherent in phalanges. Style ± as long as ovary; stigma slightly 

clavate. Fruit red, ovoid, ellipsoid, or rarely globose, 1.3–1.8 × 

0.7–1(–1.5) cm, smooth, with large oil glands, 1–3-seeded; 

exocarp ca. 0.5 mm thick. Seeds narrowly ovoid, 1–1.5 cm; 

embryo solitary. Fl. Jun–Jul, fr. Nov–Jan. 

● Moist and shady places in evergreen broad-leaved forests; 

100–400 m. W Guangdong, SE Guangxi, Hainan. 

3. Atalantia henryi (Swingle) C. C. Huang, Guihaia 11: 5. 

1991. 

薄皮酒饼簕 bo pi jiu bing le 

Atalantia racemosa Wight & Arnott var. henryi Swingle, J. 

Arnold Arbor. 21: 127. 1940. 

Trees usually 3–7 m tall. Branchlets brown, slightly flat, 

glabrous or very sparsely pilose; spines few, 2 mm or less. 

Leaves 1-foliolate; petiole 4–8 mm; leaflet blade broadly ovate, 

ovate-elliptic, or sublanceolate, 5–11 × 2.5–5 cm, thickly papery 

to leathery, oil glands slightly sunken when dry, midvein slightly 

ridged, secondary veins very numerous and joined in an arched 

marginal vein, apex shortly acuminate to obtuse and emarginate 

at tip. Inflorescences axillary, racemose, 1–3 cm, ca. 30-flowered. 

Pedicel 1–5 mm, with 1 bracteole. Sepals 4(or 5), 1–1.5 

mm, pubescent, basally connate, margin ciliate. Petals white, ca. 

4 mm. Stamens 8; filaments 1–2 mm, distinct or sometimes in 

basally coherent phalanges. Gynoecium nearly as long as filaments; 

ovary subglobose, with 4 large oil glands, 2- or 3-loculed, 

89

90 

with 1 ovule per locule; stigma capitate. Fruit reddish orange, 

globose, 1.5–2 cm in diam., smooth, oil glands inconspicuous. 

Seeds broadly ovoid, 1–1.4 cm; seed coat thinly crustose, 

smooth; embryo solitary. Fl. Apr–May, fr. Nov–Dec. 

Moist forests in limestone mountains; 300–1100 m. W Guangxi, 

Yunnan [NE Vietnam]. 

4. Atalantia dasycarpa C. C. Huang, Acta Phytotax. Sin. 16(2): 

85. 1978. 

厚皮酒饼簕 hou pi jiu bing le 

Trees 2–5 m tall. Branchlets, leaves, and petioles with short 

crisped trichomes but glabrescent. Young branchlets slightly 

flat, with or without spines. Petiole 6–10 mm, adaxially narrowly 

canaliculate; leaflet blade narrowly elliptic to elliptic- 

lanceolate, 10–17 × 3–6 cm, dark green to yellowish green when 

dry, secondary veins numerous and conspicuous, base rounded, 

margin sinuate, apex acuminate and rarely emarginate at tip. 

Inflorescences axillary, racemose, 1–2 cm, 3–9-flowered; 

peduncle thinly pubescent. Flowers 4-merous. Pedicel thinly 

pubescent. Calyx lobes ovate-triangular, apex acute. Petals 

white, 6–8 mm. Stamens 8, slightly shorter than petals; filaments 

broad, distinct or sometimes cohering in phalanges, apex acute. 

Disk 1/4–1/3 as long as ovary. Ovary ellipsoid, with 4 large pale 

oil glands, 3- or 4-loculed, with 1 or 2 ovules per locule, apex 

narrow; style 4–5 mm. Fruit globose, 1.5–2 cm or more in diam., 

oil glands numerous, large, and conspicuously concave when 

dry; exocarp ca. 1.5 mm thick. Seeds broadly ellipsoid, 10–12 × 

6–9 mm, narrowed at both ends; embryo solitary. Fl. Apr–May, 

fr. Oct–Dec. 

Forests by rivers in valleys; 200–400 m. SW Guangxi [NE Vietnam]. 

5. Atalantia guillauminii Swingle, Notul. Syst. (Paris) 2: 159. 

1911. 

大果酒饼簕 da guo jiu bing le 

Trees to 6 m tall. Young branchlets sparsely pubescent, 

unarmed or with few spines. Petiole 1–1.2 cm, stout, adaxially 

with short pubescence when young; leaflet blade narrowly 

elliptic, 14–18 × 5–6 cm, thickly papery, secondary veins 

numerous and conspicuous, base broadly cuneate, margin entire, 

apex narrowly obtuse to shortly acuminate. Ovary 3- or 

4-loculed, with 1 or 2 ovules per locule. Infructescences axillary, 

racemose, 1–1.5 cm, thinly pubescent. Fruit subglobose, 2.5–3 

cm in diam., with oil glands conspicuously sunken when dry; 


outer part of pericarp (exocarp and mesocarp) ca. 1 mm thick. 

Seeds broadly ellipsoid and slightly flat, 1.6–2.2 × ca. 1.5 cm; 

embryo solitary. Fr. Sep. 

Moist forests in valleys; 200–300 m. SE Yunnan (Hekou) [N 

Vietnam]. 

6. Atalantia acuminata C. C. Huang, Guihaia 11: 6. 1991. 

尖叶酒饼簕 jian ye jiu bing le 

Trees 2–6 m tall, whole plant glabrous. Branchlets slender, 

with 2–5 mm spines. Petiole 3–6(–8) mm, adaxially shallowly 

canaliculate; leaflet blade lanceolate, 6–12 × 2–4 cm, secondary 

veins numerous and conspicuous, base narrowly cuneate, margin 

entire, apex acuminate. Inflorescences axillary, racemose, 

3–9-flowered. Flowers 1–1.5 cm in diam. Calyx lobes 4, broadly 

ovate, ca. 2 mm. Petals white, 8–10 mm. Stamens 8, slightly 

shorter than petals; filaments broad, distinct or sometimes 

cohering in phalanges, apex acute. Ovary ellipsoid, without large 

oil glands, 3- or 4-loculed, with 1 or 2 ovules per locule, apex 

narrow; style 4–5 mm; stigma capitate. Fruit globose, 1.2–1.5 

cm in diam., oil glands dense, large, and conspicuously concave 

when dry. Fl. May, fr. Oct. 

Forests and thickets on limestone mountains; 700–900 m. W 

Guangxi, SE Yunnan [N Vietnam]. 

7. Atalantia fongkaica C. C. Huang, Guihaia 11: 5. 1991. 

开封酒饼簕 kai feng jiu bing le 

21. CITRUS Linnaeus, Sp. Pl. 2: 782. 1753. 

柑橘属 gan ju shu 

Zhang Dianxiang (张奠湘); David J. Mabberley 

Shrubs to 1 m tall. Branchlets green, slightly flat, glabrous, 

with very short sometimes slightly flat spines. Leaves 1-foliolate; 

petiole 4–7 mm, articulated with blade; leaflet blade variable 

in shape and size, some obovate and 7–9 × 3–4 cm with 

shortly acuminate apex, others oblanceolate to narrowly elliptic 

and 10–16 × 4–6 cm with acuminate apex, thickly papery to 

leathery, glabrous, abaxially grayish yellow to yellowish gray 

when dry, midvein ridged, secondary veins numerous, tertiary 

veins conspicuous, base broadly cuneate to obtuse, margin 

sinuate or apically with minute obtuse teeth. Flowers axillary, 

solitary. Ovary 5-loculed, with 1 ovule per locule. Fruit pyriform 

when young but ellipsoid to ovoid when mature, ca. 3.8 × 3 cm, 

smooth, with slightly convex oil glands when dry; outer part of 

pericarp (exocarp and mesocarp) ca. 2 mm thick. Seeds 1.5–1.8 

× 1–1.3 cm; seed coat smooth; embryo solitary; cotyledons 

green. Fr. Aug–Oct. 

● Forests by rivers in valleys; below 200 m. Guangdong (Fengkai). 

Aurantium Tournefort ex Miller; Citreum Tournefort ex Miller; ×Citrofortunella J. Ingram & H. E. Moore; ×Citroncirus J. Ingram 

& H. E. Moore; +Citroponcirus H. Wu et al.; Fortunella Swingle; Limon Tournefort ex Miller; Papeda Hasskarl; Poncirus 

Rafinesque; Pseudaegle Miquel; Sarcodactilis C. F. Gaertner. 

Shrubs or small trees, evergreen, rarely deciduous. Young branches often flat and angled, usually with solitary (rarely paired) 

spines at axils. Leaves 1-foliolate, rarely 3-foliolate or simple; petiole usually articulated with base of leaf blade, usually conspicuously 

winged; leaf blade subleathery to leathery, with dense pellucid fragrant oil dots, margin crenulate or rarely entire. Flowers axillary,


hermaphrodite or male, solitary or in small fascicles, fragrant. Calyx cup-shaped; lobes 3–5, subglabrous. Petals (3 or)4 or 5(–8), white 

or outside pinkish red, imbricate, thick. Stamens usually 4(–10) × as many as petals, free or basally coherent. Disk annular or short, 

with nectary glands. Ovary (3–)5–14(–18)-loculed, each locule with 2–8 or more ovules; stigma large. Fruit a berry (hesperidium) with 

sarcocarp segments of pulp vesicles and adaxially attached seeds. Seed coat smooth or ridged; embryo(s) 1 to many, like cotyledons 

milky white, green, or rarely yellowish, germination hypogeous. 

Between 20 and 25 species: E, S, and SE Asia, Australia, SW Pacific islands, with many cultivated taxa widely naturalized in warm countries; 11 

species and hybrid species (three endemic) native, naturalized, or extensively cultivated in China, plus five hybrid species cultivated to a limited extent. 

In China, many early hybridizations appear to have taken place, and many cultivated taxa have become naturalized (these are listed at the end of 

the generic account). For a discussion of the status of several taxa formerly considered species see Nicolosi et al. (Theor. Appl. Genetics 100: 

1155–1166. 2000) and Mabberley (Blumea 49: 481–498. 2004). Hybrids readily form between species, and as apomixis is common, such hybrids can be 

fixed as cultivars, with occasional outcrossings leading to yet more. Because these hybrids can thereby span, through backcrossing, the spectrum of 

variation between the original, probably geographically isolated, species, it is impossible to provide a key to cover all plants that may be found. The key 

here therefore covers the apparently wild species and some of the most common cultivar groups now referred to as hybrid taxa. Doubtful taxa are not 

included in the key, but descriptions of two are included at the end of the treatment. 

Because of the enormous worldwide economic importance of the genus, Citrus is treated more fully, particularly with regard to synonymy, than is 

the norm in this flora. The following treatment is the first floristic one to take account of current advances in the understanding of the genus. 

1a. Plants deciduous; leaves of mature plants 3(–5)-foliolate ............................................................................................ 1. C. trifoliata 

1b. Plants evergreen; leaves 1-foliolate or rarely simple. 

2a. Ovary with 3 or 4(–6) locules, each with 3 or 4 ovules ......................................................................................... 4. C. japonica 

2b. Ovary with (6 or)7–15 locules, each with many ovules. 

3a. Leaves with a winged petiole much more than half as long as leaf blade. 

4a. Leaf blade margin entire to finely crenulate, apex acuminate ................................................................ 2. C. cavaleriei 

4b. Leaf blade margin conspicuously crenulate, apex obtuse to subrounded and sometimes mucronate ......... 3. C. hystrix 

3b. Leaves with a winged petiole less than half as long as leaf blade or absent. 

5a. Petiole not winged; fruit pericarp thicker than sarcocarp ............................................................................ 5. C. medica 

5b. Petiole winged although sometimes only narrowly; fruit pericarp thinner than sarcocarp. 

6a. Cotyledons green. 

7a. Fruit oblong or at least with a marked apical mammilla; petals outside pinkish or reddish ......... 9. C. ×limon 

7b. Fruit globose, oblate, pyriform, or broadly obconic and without a marked apical mammilla; 

petals outside white or purplish. 

8a. Young branches, calyx lobes, and ovaries with trichomes; fruit 10 cm or more in diam. ..... 6. C. maxima 

8b. Young branches, calyx lobes, and ovaries glabrous; fruit to 10 cm in diam. but often 

much smaller ................................................................................................................... 10. C. ×aurantium 

6b. Cotyledons milky white. 

9a. Flowers solitary or in small fascicles; pericarp easily removed. 

10a. Fruit pale yellow, orange, red, or carmine; sarcocarp usually sweet ................................ 7. C. reticulata 

10b. Fruit pale yellow; sarcocarp acid and bitter ........................................................................ 11. C. ×junos 

9b. Flowers usually in fascicles; pericarp not easily removed. 

11a. Fruit greenish yellow, with an apical papilla; sarcocarp very acid ........................... 8. C. ×aurantiifolia 

11b. Fruit orange, reddish, or sometimes yellow, without an apical papilla; sarcocarp 

sour or sweet ................................................................................................................ 10. C. ×aurantium 

1. Citrus trifoliata Linnaeus, Sp. Pl., ed. 2, 2: 1101. 1763. 

枳 zhi 

Aegle sepiaria Candolle; Citrus trifolia Thunberg; C. trifoliata 

subf. monstrosa (T. Itô) Hiroe; C. trifoliata var. monstrosa 

T. Itô; C. triptera Desfontaines; Poncirus trifoliata (Linnaeus) 

Rafinesque; P. trifoliata var. monstrosa (T. Itô) Swingle; 

Pseudaegle sepiaria (Candolle) Miquel. 

Trees or treelets, 1–5 m tall. Branches green, flat, ridged 

when young; spines ca. 4 cm, base flat, apex rufous. Foliage 

spurs unarmed, developed from dormant buds on twigs of 

previous year, with very short internodes bearing 1–5 leaves. 

Leaves palmately 3(–5)-foliolate, in juveniles usually simple or 

1-foliolate; petiole narrowly winged; leaflet blades 2–5 × 1–3 

cm, central one as long or longer than laterals, midvein with 

short trichomes when young, margin finely crenulate or entire. 

Flowers solitary or paired, 3.5–8 cm in diam. Calyx lobes 5–7, 

basally connate. Petals (4 or)5(or 6), white, obovate, 1.5–3 cm, 

imbricate. Stamens usually 20; filaments of different lengths. 

Ovary 6–8-loculed, hairy; ovules 4–8, in 2 rows per locule; style 

short, thick; stigma clavate. Fruit dark yellow, subglobose to 

pyriform, usually 3–4.5 × 3.5–6 cm, with coarse ring-shaped 

furrows or sometimes smooth. Seeds 20–50, broadly ovoid, 

0.9–1.2 cm; embryo(s) solitary or several; seed coat smooth or 

with inconspicuous fine ridges. Fl. May–Jun, fr. Oct–Nov. 

● Anhui, Chongqing, S Gansu, N Guangdong, N Guangxi, Guizhou, 

Henan, Hubei, Hunan, Jiangsu, NW Jiangxi, S Shaanxi, Shandong, 

S Shanxi, Zhejiang. 

2. Citrus cavaleriei H. Léveillé ex Cavalerie, Bull. Géogr. Bot. 

21: 211. 1911. 

宜昌橙 yi chang cheng 

91

92 

Citrus ×aurantium Linnaeus subsp. ichangensis (Swingle) 

Guillaumin; C. hongheensis Y. M. Ye et al.; C. ichangensis 

Swingle; C. macrosperma T. C. Guo & Y. M. Ye. 

Trees or shrubs, 2–10 m tall. Branchlets subglabrous; 

spines straight, stout, smaller on flowering branches. Leafy 

petiole 1–3 × as long as blade, narrowly elliptic, 6–16 × 2.5–4 

cm, base cuneate, margin finely crenulate, apex rounded; leaf 

blades ovate-lanceolate, ca. 2(–8) × 0.7–1.5(–4.5) cm, margin 

entire to finely crenate, apex acuminate. Flowers solitary or to 9 

in fascicles, 3–3.5 cm in diam.; buds pale purplish red, broadly 

ellipsoid. Calyx lobes 5. Petals 4 or 5, white or pink, 1–1.8 × 

0.5–0.8 cm. Stamens 16–30; filaments distinct or cohering in 

bundles, ciliate. Ovary pale green, subellipsoid; style ca. 6.5 

mm; stigma pale yellow, with fine shallow grooves. Fruit pale 

yellow, oblate, globose, or pyriform, usually 3–5 × 4–6 cm but 

when pyriform to 9–10 × 7–8(–12) cm, with narrow longitudinal 

grooves, oil dots large and conspicuously prominent, base 

rounded, apex rounded, dimpled, and with or without a papilla; 

pericarp to 2 cm thick but usually much less; sarcocarp in 7–13 

segments, yellowish white, very sour. Seeds 30 or more, subglobose 

to irregularly pyramidal, ca. 1.5 × 1.5 × 1.2 cm; seed 

coat yellowish white, chalaza dark brown, large, and covering 

almost half of coat; embryo(s) solitary to numerous; cotyledons 

milky white. Fl. Mar–Jun, fr. Oct–Dec. 

● Mountains, hills, valleys; below 2500 m. S Gansu, N Guangxi, 

Guizhou, W Hubei, NW and W Hunan, S Shaanxi, Sichuan, Yunnan. 

This species is a parent with Citrus ×aurantium of C. ×webberi 

Wester, and possibly a parent with C. reticulata of C. ×junos. 

3. Citrus hystrix Candolle, Cat. Pl. Horti Monsp. 19, 97. 1813. 

箭叶橙 jian ye cheng 

Citrus auraria Michel; C. echinata Saint-Lager; C. hyalopulpa 

Tanaka; C. kerrii (Swingle) Tanaka; C. macroptera Montrouzier 

var. kerrii Swingle; C. papeda Miquel; Fortunella 

sagittifolia F. M. Feng & P. I Mao; Papeda rumphii Hasskarl. 

Trees 3–6 m tall. Branchlets with spines. Leaves dark red 

when young; petiole winged, apex rounded to truncate; leaf 

blade ovate, 5–8 × 2.5–4.5 cm, 1–2.5 cm longer (rarely same 

length) and 0.5–1 cm wider than winged petiole, tertiary veins 

conspicuous, margin apically conspicuously and sparsely crenate, 

apex narrowly obtuse. Inflorescences with (1 or)3–5 

flowers; peduncle 1–5 mm. Flower buds globose. Calyx lobes 4 

or 5, broadly triangular, ca. 4 × 6 mm. Petals white but pinkish 

red outside, 7–10 mm. Stamens ca. 30; filaments distinct. Style 

short, thick. Fruit lemon yellow, ellipsoid to subglobose, 5–7 × 

3–5 cm, slightly coarse or smooth, oil dots numerous and 

prominent, apex rounded; pericarp thick; sarcocarp in 11–13 

segments, very acidic and slightly bitter. Seeds numerous 

1.5–1.8 × 1–1.2 cm, ridged; embryo solitary; cotyledons milky 

white. Fl. Mar–May, fr. Nov–Dec. 

N Guangxi, Yunnan [Indonesia, Myanmar, New Guinea, Philippines, 

Thailand]. 

Although apparently native to S China into SE Asia and Malesia, 

the natural distribution of this species is obscured by cultivation. Selected 

forms are cultivated throughout the warm parts of the world for their 


culinary (leaves) and medicinal (fruit) uses. All named taxa (save perhaps 

some from central Malesia) seem to have been based on cultivated 

plants as discussed by Mabberley (Gard. Bull. Singapore 54: 173–184. 

2002). Commonly seen in China are cultivated plants (the “lime leaves” 

of commerce) with the following characteristics: leaf blade broadly 

elliptic, apex obtuse to rounded; fruit subglobose, ca. 4 × 3.5 cm, smooth, 

apex with a papilla; pericarp ca. 2 mm thick; sarcocarp in 6 or 7 segments, 

6–8-seeded but 1 or 2 seeds undeveloped; seeds pyramidal, 

1.5–1.8 × 1–1.4 cm, 0.8–1.2 mm thick, with alveolate ridges. 

4. Citrus japonica Thunberg, Nova Acta Regiae Soc. Sci. 

Upsal. 3: 208. 1780. 

金柑 jin gan 

Atalantia hindsii (Champion ex Bentham) Oliver ex Bentham; 

Citrus ×aurantium Linnaeus var. globifera Engler; C. 

×aurantium subsp. japonica (Thunberg) Engler; C. ×aurantium 

var. japonica (Thunberg) Hooker; C. ×aurantium subvar. madurensis 

(Loureiro) Engler; C. ×aurantium subvar. margarita 

(Loureiro) Engler; C. ×aurantium var. oliviformis Risso ex Loiseleur; 

C. ×aurantium subvar. spinosa Siebold & Zuccarini ex 

Engler; C. hindsii (Champion ex Bentham) Govaerts; C. inermis 

Roxburgh; C. japonica subf. crassifolia (Swingle) Hiroe; C. 

japonica subf. hindsii (Champion ex Bentham) Hiroe; C. 

japonica var. madurensis (Loureiro) Guillaumin; C. japonica 

subf. margarita (Loureiro) Hiroe; C. japonica var. margarita 

(Loureiro) Guillaumin; C. kinokuni Tanaka; C. madurensis 

Loureiro; C. margarita Loureiro; C. ×nobilis Loureiro var. inermis 

(Roxburgh) Sagot; Fortunella chintou (Swingle) C. C. 

Huang; F. crassifolia Swingle; F. hindsii (Champion ex Bentham) 

Swingle; F. hindsii var. chintou Swingle; F. japonica 

(Thunberg) Swingle; F. margarita (Loureiro) Swingle; F. obovata 

Tanaka; F. venosa (Champion ex Bentham) C. C. Huang; 

Sclerostylis hindsii Champion ex Bentham; S. venosa Champion 

ex Bentham. 

Trees to 5 m tall, d.b.h. to 20 cm. Branchlets numerous; 

spines variable, to 5 cm on young growth but some only a few 

mm on flowering shoots. Leaves 1-foliolate or sometimes mixed 

with simple leaves; petiole 6–9 mm, narrowly winged; leaf blade 

elliptic to obovate-elliptic, 4–6 × 1.5–3 cm, base rounded to 

broadly cuneate, margin dentate near apex or rarely entire, apex 

rounded and rarely mucronate. Flowers solitary or fascicled, 

subsessile. Calyx 4- or 5-lobed. Petals 5, ca. 5 mm or less. Stamens 

ca. 20; filaments cohering into 4 or 5 bundles. Ovary as 

long as style, 3- or 4-loculed, with 3 or 4 ovules per locule. Fruit 

bright orange to red, globose to slightly oblate, 9–10 mm in 

diam., smooth, 3- or 4-seeded; pericarp sweet and edible; sarcocarp 

acidic. Seeds broadly ovoid, apex mucronate; seed coat 

smooth; embryos at least sometimes numerous; cotyledons 

green. Fl. Apr–May, fr. Oct–Dec. 2n = 18*, 20*, 36*. 

● Evergreen broad-leaved forests; 600–1000 m. S Anhui, Fujian, 

Guangdong, SE Guangxi, Hainan, Hunan, Jiangxi, Zhejiang. 

This appears to be the truly wild kumquat from which cultivars 

with greatly increased fruit-sizes of different shapes have been selected. 

However, recent field studies in China show a considerable amount of 

variation within some wild populations of what is treated here as Citrus 

japonica. Future comprehensive field and cytological studies throughout

the complex may show that geographical subspecies or cytological races 

of the wild form could be formally recognized. 

None of the cultivars, formerly recognized as species, has been 

found in truly “natural” habitats. They are cultivated on a large scale in S 

China and have sometimes become naturalized. Citrus japonica and C. 

reticulata are parents of C. ×microcarpa. Many of the above synonyms 

can perhaps best be referred to cultivar groups, corresponding to the 

subformae of Hiroe. Commonly seen cultivar groups in China have the 

following characteristics, but some cultivars are intermediate. 

Round Kumquat Group 

Trees 2–5 m tall. Petiole 6–10 mm or rarely longer, wings 

narrow; leaf blade ovate-elliptic to elliptic-lanceolate, 4–8 × 

1.5–3.5 cm, base broadly cuneate, apex obtuse and sometimes 

mucronate. Flowers 1–3 per fascicle; peduncle 6 mm or less. 

Petals 6–8 mm. Stamens 15–25. Ovary globose, ± as long as 

style, 4–6-loculed. Fruit orangish yellow to orangish red, globose, 

1.5–2.5 cm in diam., 2–5-seeded; pericarp 1.5–2 mm thick, 

sweet. Seeds ovoid, base rounded; embryo solitary. Fl. Apr– 

May, fr. Nov–Feb. 

Oval Kumquat Group 

Trees to 3 m tall. Petiole ca. 1.2 cm, wings very narrow; 

leaf blade ovate-lanceolate to long elliptic, 5–11 × 2–4 cm, base 

broadly cuneate to nearly rounded, apex obtuse to slightly acute. 

Flowers 1–3 per fascicle; peduncle 3–5 mm. Petals 6–8 mm. 

Stamens 20–25. Ovary elliptic; style usually ca. 1.5 × as long as 

ovary, slender; stigma slightly clavate. Fruit orangish yellow to 

orangish red, ellipsoid to ovoid-ellipsoid, 2–3.5 cm in diam., oil 

dots usually slightly expanded, 2–5-seeded; pericarp ca. 2 mm 

thick, sweet; sarcocarp in 4 or 5 segments, acidic. Seeds ovoid, 

apex acute; embryo solitary or rarely numerous. Fl. Mar–May, 

fr. Oct–Dec. 

5. Citrus medica Linnaeus, Sp. Pl. 2: 782. 1753. 

香橼 xiang yuan 

Aurantium medicum (Linnaeus) M. Gómez; Citreum vulgare 

Tournefort ex Miller; Citrus alata (Yu. Tanaka) Tanaka; C. 

×aurantium Linnaeus subvar. amilbed Engler; C. ×aurantium 

subvar. chakotra Engler; C. cedra Link; C. cedrata Rafinesque; 

C. fragrans Salisbury; ?C. kwangsiensis Hu; C. ×limon (Linnaeus) 

Osbeck var. digitata Risso; C. medica var. alata Yu. 

Tanaka; C. medica var. digitata Risso; C. medica var. ethrog 

Engler; C. medica f. monstrosa Guillaumin; C. medica var. 

proper J. D. Hooker; C. medica var. sarcodactylis (Hoola van 

Nooten) Swingle; C. odorata Roussel; C. sarcodactylis Hoola 

van Nooten; C. tuberosa Miller; Sarcodactilis helicteroides C. F. 

Gaertner. 

Shrubs or small trees. Branches, leaf buds, and flower buds 

purplish when young. Branches with ca. 4 cm spines. Leaves 

simple or rarely 1-foliolate; petiole short, not winged; leaf blade 

elliptic to ovate-elliptic, 6–12 × 3–6 cm or larger, margin serrate, 

apex rounded, obtuse, or rarely mucronate. Inflorescences axillary, 

ca. 12-flowered or sometimes flowers solitary. Flowers 

bisexual or sometimes male by ± complete abortion of pistil. 

Petals 5, 1.5–2 cm. Stamens 30–50. Ovary cylindric; style long 

and thick; stigma clavate. Fruit pale yellow, elliptic to sub- 


globose, to 2 kg, surface coarse; pericarp white to pale yellow 

and soft within, thicker than sarcocarp, removed with difficulty; 

sarcocarp with 10–15 segments, colorless, nearly pellucid to pale 

milky yellow, acidic to slightly sweet, fragrant. Seeds small; 

seed coat smooth; embryo(s) solitary to several; cotyledons 

milky white. Fl. Apr–May, fr. Oct–Nov. 2n = 18, 20. 

Cultivated and sometimes naturalized. Guangxi, SW Guizhou, 

Hainan, Sichuan, E Xizang, Yunnan [native to NE India and possibly 

Myanmar]. 

This species is a parent with Citrus ×aurantium of C. ×limon and 

with ?C. hystrix of C. ×aurantiifolia. 

The Buddha-hand Citron (佛手 fo shou), with separated segments 

± surrounded by pericarp, is best treated as a cultivar, correctly Citrus 

medica ‘Fingered.’ 

6. Citrus maxima (Burman) Merrill, Interpr. Herb. Amboin. 

296. 1917. 

柚 you 

Aurantium maximum Burman in Rumphius & Burman, 

Herb. Amboin. Auctuar. 7: Index [16]. 1755; A. decumanum 

(Linnaeus) Miller; Citrus ×aurantium Linnaeus subsp. decumana 

(Linnaeus) Tanaka; C. ×aurantium var. decumana Linnaeus; 

C. ×aurantium f. grandis (Linnaeus) Hiroe; C. ×aurantium 

var. grandis Linnaeus; C. costata Rafinesque; C. decumana 

(Linnaeus) Linnaeus; C. grandis (Linnaeus) Osbeck; C. grandis 

var. pyriformis (Hasskarl) Karaya; C. grandis var. sabon (Siebold 

ex Hayata) Hayata; ?C. kwangsiensis Hu; C. medica Linnaeus 

subf. pyriformis (Hasskarl) Hiroe; C. obovoidea Yu. Tanaka; 

C. pampelmos Risso; C. pompelmos Risso; C. pyriformis 

Hasskarl; C. sabon Siebold ex Hayata. 

Trees. Young branches, abaxial surface of leaves, peduncles, 

and ovaries pilose. Branches usually purplish, flat with 

ridges when young. Petiole 2–4 × 0.5–3 cm or less, winged; leaf 

blade broadly ovate or elliptic, 9–16 × 4–8 cm or larger, thick, 

dark green, base rounded, apex rounded to obtuse and sometimes 

mucronate. Flowers solitary or in racemes; flower buds 

purplish or rarely milky white. Calyx 3–5-lobed. Petals 1.5–2 

cm. Stamens 25–35, some undeveloped. Style long and thick. 

Fruit pale yellow and yellowish green, globose, oblate, pyriform, 

or broadly obconic, usually more than 10 cm in diam., with large 

prominent oil dots, to 200-seeded or seedless; pericarp spongy; 

sarcocarp with 10–15(–19) segments, white, pink, reddish, or 

rarely milky yellow. Seeds irregularly shaped, with conspicuous 

ridges, undeveloped seeds numerous; embryo solitary; cotyledons 

milky white. Fl. Apr–May, fr. Sep–Dec. 2n = 18, 36. 

Cultivated and naturalized in S China [probably native to SE Asia]. 

Pomelo or shaddock includes cultivars with round to obovoid fruit 

much favored for festival decoration as well as eating. No truly wild 

plants (presumably with much smaller fruit) have been seen. Citrus 

maxima is a parent with C. reticulata of C. ×aurantium. 

7. Citrus reticulata Blanco, Fl. Filip. 610. 1837. 

柑橘 gan ju 

Citrus ×aurantium Linnaeus f. deliciosa (Tenore) Hiroe; C. 

×aurantium var. tachibana Makino; C. daoxianensis S. W. He 

93

94 

& G. F. Liu; C. deliciosa Tenore; C. depressa Hayata; C. 

erythrosa Yu. Tanaka; C. madurensis Loureiro var. deliciosa 

(Tenore) Sagot; C. mangshanensis S. W. He & G. F. Liu; C. 

×nobilis Loureiro subf. deliciosa (Tenore) Hiroe; C. ×nobilis 

var. deliciosa (Tenore) Guillaumin; C. ×nobilis subf. erythrosa 

(Yu. Tanaka) Hiroe; C. ×nobilis var. major Ker Gawler; C. 

×nobilis var. ponki Hayata; C. ×nobilis subf. reticulata (Blanco) 

Hiroe; C. ×nobilis var. spontanea Ito; C. ×nobilis subf. succosa 

(Tanaka) Hiroe; C. ×nobilis var. sunki Hayata; C. ×nobilis 

subf. tachibana (Makino) Hiroe; C. ×nobilis var. tachibana 

(Makino) Ito; C. ×nobilis subf. unshiu (Marcowicz) Hiroe; C. 

×nobilis var. unshiu (Marcowicz) Tanaka ex Swingle; C. ×nobilis 

var. vangasy (Bojer) Guillaumin; C. ponki Yu. Tanaka; C. 

poonensis Yu. Tanaka; C. reticulata var. austera Swingle; C. 

reticulata subsp. deliciosa (Tenore) Rivera et al.; C. reticulata 

subsp. tachibana (Tanaka) Rivera et al.; C. reticulata subsp. 

unshiu (Marcowicz) Rivera et al.; C. succosa Tanaka; C. suhuiensis 

Hayata; C. sunki Tanaka; C. tachibana (Makino) Yu. Tanaka; 

C. tachibana subf. depressa (Hayata) Hiroe; C. tachibana 

subf. ponki (Hayata) Hiroe; C. tachibana subf. suhuiensis (Hayata) 

Hiroe; C. tachibana subf. sunki (Hayata) Hiroe; C. tangerina 

Yu. Tanaka; C. tankan Hayata; C. unshiu Marcowicz; C. 

vangasy Bojer. 

Small trees. Branchlets numerous, with few spines. Leaves 

1-foliolate; leaf blade lanceolate, elliptic, or broadly ovate, basal 

articulated part to leaf blade usually narrow or only a remnant, 

midvein furcate near apex, margin apically obtusely crenulate or 

rarely entire, apex emarginate. Flowers solitary to 3 in a fascicle. 

Calyx irregularly 3–5-lobed. Petals usually 1.5 cm or less. Sta- 

8. Citrus ×aurantiifolia (Christmann) Swingle, J. Wash. Acad. 

Sci. 3: 465. 1913 [“aurantifolia”], pro sp. 

来檬 lai meng 

Limonia ×aurantiifolia Christmann, Vollst. Pflanzensyst. 

1: 618. 1777 [“aurantifolia”], pro sp.; Citrus ×acida Persoon; 

C. ×aurantiifolia subsp. murgetana Garcia Lidón et al.; C. ×aurantium 

Linnaeus subsp. aurantiifolia (Christmann) Guillaumin; 

C. ×aurantium var. proper Guillaumin; C. hystrix Candolle 

subsp. acida Engler; C. ×javanica Blume; C. ×lima Macfadyen; 

C. medica Linnaeus subf. aurantiifolia (Christmann) Hiroe. 

Small trees. Branchlets numerous and irregular, with short 

stout spines. Leaves slightly stiff with a short, conspicuous 

petiole; leaf blade broadly ovate to elliptic, 5–8 × 2–4 cm, base 

rounded, margin crenulate, apex obtuse and sometimes mucronate. 

Inflorescences with ca. 7 flowers or rarely flowers solitary. 

Calyx cup-shaped; lobes 4 or 5. Petals (4 or)5, white, 1–1.2 cm, 

rather thick. Stamens 20–25. Ovary globose; style short, ± as 

long as stigma; stigma large. Fruit greenish yellow, globose, 

ellipsoid, or obovoid, usually 4–5 cm in diam., smooth, with 

prominent oil glands, apex with a papilla; pericarp thin; sarcocarp 

with 9–12 segments, very acidic. Seeds few, ovoid; seed 

coat smooth; cotyledons milky white. Fl. Apr–May, fr. Sep– 

Dec. 2n = 18, 27. 

Trailsides; low elevations. Occasionally naturalized in Yunnan. 


Hybrid taxa sometimes naturalized in China 

mens 20–25. Style long, slender; stigma clavate. Fruit pale yellow, 

orange, red, or carmine, oblate to subglobose, smooth or 

coarse; pericarp very thin to thick, easily removed; sarcocarp 

with 7–14 segments or rarely more, sweet to acidic and sometimes 

bitter, with few to many seeds or rarely seedless; pulp 

vesicles plump, short, rarely slender and long. Seeds usually 

ovoid, base rounded, apex narrow and acute; embryos numerous, 

rarely solitary; cotyledons dark green, pale green, or milky 

white; chalaza purple. Fl. Apr–May, fr. Oct–Dec. 2n = 18, 27, 36. 

Extensively cultivated in China S of the Qin Ling [possibly native 

to SE China and/or S Japan (see below)]. 

Tangerine or mandarin orange is the parent with Citrus maxima of 

C. ×aurantium, with C. japonica of C. ×microcarpa, and possibly with 

C. cavaleriei of C. ×junos. The recently recognized subspecies are perhaps 

better considered as cultivar groups to which most of the synonyms 

would be referred. However, some names in the synonymy may be 

referable to those cultivars of C. ×aurantium that are repeated backcrosses 

with C. reticulata. Trees referred to C. tachibana may represent 

true wild forms and have the following characteristics: 

Trees to 3 m tall. Branchlets numerous, with short spines. Petiole 

8–10 mm, very narrowly winged; leaf blade elliptic, 6–7 × 3.5–4 cm, 

secondary veins inconspicuous, base broadly cuneate, margin crenulate, 

apex narrow, obtuse, and conspicuously emarginate. Flowers solitary, 

1.2–1.4 cm in diam.; flower buds subglobose. Pedicel ca. 2 mm. Petals 

white. Stamens ca. 20. Fruit yellow, oblate, 2–2.5 × 2.5–3.4 cm, smooth; 

pericarp 1.5–2 mm thick; sarcocarp with 7–9 segments, yellow, very 

acidic and bitter, 5- or 6-seeded. Seeds broadly ovoid, ca. 1 cm; seed coat 

smooth; embryos numerous; cotyledons greenish. 

Hillside forests; low elevations. Taiwan [Japan (Ryukyu Islands)]. 

As a common name, lime covers a number of different Citrus 

species and hybrid species. Those referred to as C. ×aurantiifolia are 

hybrids involving C. medica and possibly C. hystrix. 

9. Citrus ×limon (Linnaeus) Osbeck, Reis Ostindien China, 

250. 1765 [“limonia”], pro sp. 

柠檬 ning meng 

Citrus medica Linnaeus var. limon Linnaeus, Sp. Pl. 2: 782. 

1753; C. ×aurantium Linnaeus subsp. bergamia (Risso) Engler; 

C. ×aurantium var. bergamia (Risso) Brandis; C. ×aurantium 

var. mellarosa (Risso) Engler; C. ×bergamia Risso; C. ×bergamia 

subsp. mellarosa (Risso) Rivera et al.; C. ×bergamota 

Rafinesque; C. ×limodulcis Rivera et al.; C. ×limonum Risso; C. 

medica Linnaeus f. limon (Linnaeus) Hiroe; C. medica subsp. 

limonum (Risso) J. D. Hooker; C. medica var. limonum (Risso) 

Brandis; C. ×mellarosa Risso; C. ×meyeri Yu. Tanaka; Limon 

×vulgaris Ferrarius ex Miller (1754), not C. ×vulgaris Risso 

(1813). 

Small trees. Branches ± spiny. Young leaves and flower 

buds reddish purple. Leaf blade ovate to elliptic, 8–14 × 4–6 cm, 

margin conspicuously crenulate, apex usually mucronate. Flowers 

solitary or several in fascicles. Flowers bisexual or male by ± 

complete abortion of pistil. Calyx cup-shaped; lobes 4 or 5. 

Petals 1.5–2 cm, outside purplish, inside white. Stamens 20–25 

or more. Ovary subcylindric or barrel-shaped; stigma clavate.

Fruit yellow, ellipsoid to ovoid, narrowed at both ends, surface 

usually coarse and lemon scented, apex usually with a mammilla; 

pericarp thick, difficult to remove; sarcocarp in 8–11 segments, 

pale yellow, acidic. Seeds ovoid, small, apex acute; seed 

coat smooth; embryo usually solitary but sometimes numerous; 

cotyledons milky white. Fl. Apr–May, fr. Sep–Nov. 2n = 18, 36. 

Cultivated and sometimes naturalized in S China. 

The parents of the lemon are Citrus ×aurantium and C. medica. 

Backcrosses with either parent give a range of sour to sweet lemons 

which go under various names and perhaps would best be considered as 

forming cultivar groups, e.g., Bergamot Group. The rough lemon, C. 

×taitensis Risso (C. ×aurantium subsp. jambhiri Engler; C. ×jambhiri 

Lushington; C. ×sinensis subsp. jambhiri (Lushington) Engler), sometimes 

included here, is perhaps C. medica × C. reticulata. 

The name “Citrus limonia” has been misapplied to other Citrus 

taxa. 

10. Citrus ×aurantium Linnaeus, Sp. Pl. 2: 782. 1753, pro sp. 

酸橙 suan cheng 

Aurantium ×acre Miller; A. ×corniculatum Miller; A. 

×distortum Miller; A. ×humile Miller; A. ×myrtifolium Descourtilz; 

A. ×sinense Miller; A. ×vulgare (Risso) M. Gómez; 

Citrus ×amara Link; C. ×aurantium subsp. amara Engler; C. 

×aurantium var. bigaradia (Loiseleur) Brandis; C. ×aurantium 

var. crassa Risso; C. ×aurantium var. daidai Makino; C. 

×aurantium var. dulcis Hayne; C. ×aurantium var. fetifera Risso; 

C. ×aurantium var. lusitanica Risso; C. ×aurantium var. 

myrtifolia Ker Gawler; C. ×aurantium subf. nobilis (Loureiro) 

Hiroe; C. ×aurantium var. sanguinea Engler; C. ×aurantium 

subf. sinensis (Linnaeus) Hiroe; C. ×aurantium subsp. sinensis 

(Linnaeus) Engler; C. ×aurantium var. sinensis Linnaeus; C. 

×aurantium subsp. suntara Engler; C. ×aurantium var. vulgaris 

(Risso) Risso & Poiteau; C. ×aurata Risso; C. ×bigaradia 

Loiseleur; C. ×changshan-huyou Y. B. Chang; C. ×communis 

Poiteau & Turpin; C. decumana (Linnaeus) Linnaeus var. 

paradisi (Macfadyen) H. H. A. Nicholls; C. ×dulcis Persoon; 

C. ×florida Salisbury; C. ×humilis (Miller) Poiret; C. maxima 

(Burman) Merrill var. uvacarpa Merrill; C. ×myrtifolia (Ker 

Gawler) Rafinesque; C. ×paradisi Macfadyen; C. ×sinensis 

(Linnaeus) Osbeck; C. ×sinensis var. brassiliensis Tanaka; C. 

×sinensis subsp. crassa (Risso) Rivera et al.; C. ×sinensis 

subsp. fetifera (Risso) Rivera et al.; C. ×sinensis subsp. lusitanica 

(Risso) Rivera et al.; C. ×sinensis var. sanguinea (Engler) 

Engler; C. ×sinensis var. sekkan Hayata; C. ×sinensis 

subsp. suntara (Engler) Engler; C. ×taiwanica Tanaka & Shimada; 

C. ×tangelo Ingram & H. E. Moore; C. ×vulgaris Risso. 

Small trees. Branches with spines up to ca. 8 cm. Petiole 

obovate, 1–3 × 0.6–1.5 cm, base narrow; leaf blades dark green, 

thick. Inflorescences racemes, with few flowers or flowers solitary. 

Flowers perfect or male by ± complete abortion of pistil; 

buds ellipsoid to subglobose. Calyx lobes 4 or 5. Petals 2–3.5 

mm in diam. Stamens 20–25, usually basally connate into bundles. 

Fruit orange to reddish, globose to oblate, surface coarse; 

pericarp thick, sometimes difficult to remove; sarcocarp with 

10–13 segments, acidic and sweet or sometimes bitter. Seeds 

numerous, large, with ridges; embryo(s) solitary to numerous; 

cotyledons milky white. Fl. Apr–May, fr. Sep–Dec. 2n = 18. 


Cultivated and sometimes naturalized in most of China S of the Qin 

Ling. 

Oranges and grapefruit are Citrus maxima × C. reticulata and the 

backcrosses with the parental species. Many of these hybrids arose in 

China, although others were synthesized, especially in the U.S.A., and 

introduced in China. The cultivars can be arranged in groups: Sour 

Orange Group (the sour or bitter orange most like the original cross), 

Sweet Orange Group (the commercially most important being 

backcrosses with C. reticulata), and Grapefruit Group (representing 

backcrosses with C. maxima and first being made in the Caribbean). 

Sweet Orange Group 

Petiole long and narrow; leaf blade ovate, ovate-elliptic, or 

rarely lanceolate, 6–10 × 3–5 cm or larger. Calyx lobes 3–5. 

Petals white or rarely purplish, 1.2–1.5 cm. Style stout; stigma 

large. Fruit orangish yellow to orangish red, globose, depressed 

globose, or ellipsoid; sarcocarp with 9–12 segments, yellow, 

orange, or purplish, sweet or slightly acidic. Seeds few or absent; 

seed coat slightly ridged; embryos numerous; cotyledons milky 

white. Fl. Mar–Jun, fr. Oct–Dec but some cultivars Feb–Apr. 2n 

= 18, 27, 36, 45. 

Below 1500 m. Cultivated S of the Qin Ling, as far NW as SE Gansu 

and as far SW as SE Xizang. 

Grapefruit Group 

Branches glabrous. Leaves similar to those of Citrus 

maxima but leaf blade smaller and narrower, midvein ciliate. 

Flowers smaller than those of C. maxima. Calyx lobes glabrous. 

Fruit yellow, depressed globose to globose, smaller than those of 

C. maxima; pericarp thin; sarcocarp with 12–15 segments, yellowish 

white or pink, tender, juicy, slightly fragrant and acidic. 

Seeds few or absent; embryos numerous. Fr. Oct–Nov. 2n = 18, 

20, 27, 36. 

11. Citrus ×junos Siebold ex Tanaka, Sieb. Sens. Tor. Hy-akun. 

Kin. Ronbunshu, 65. 1924. 

香橙 xiang cheng 

Citrus ×aurantium Linnaeus subsp. junos (Siebold ex Tanaka) 

Makino; C. ×hsiangyuan Tanaka; C. medica subf. junos 

(Siebold ex Tanaka) Hiroe; ?C. sechen Kokaya; C. sechen subsp. 

sjanshen Kokaya; ?C. ×wilsonii Tanaka. 

Small trees. Branches often with long stout spines; 

branchlets, leaves, and petioles sparsely pubescent when young. 

Petiole obovate-elliptic, 1–2.5 × 0.4–1.5 cm, base cuneate, apex 

rounded to obtuse; leaf blade ovate to lanceolate, 2.5–8 × 1–4 

cm, thickly papery, base rounded to obtuse, margin apically 

finely dentate or rarely entire, apex acuminate, mucronate, or 

often obtuse and emarginate. Flowers solitary, shortly pedicellate. 

Calyx cup-shaped; lobes 4 or 5, broadly ovate, apex acute. 

Petals white, sometimes outside purplish, 1–1.3 cm. Stamens 

20–25. Style ca. 2 × as long as ovary. Fruit pale yellow, oblate to 

nearly pyriform, 4–8 cm in diam., surface coarse and with large 

oil dots, apex grooved; pericarp 2–4 mm thick, easily removed; 

sarcocarp with 9–11 segments, pale yellowish white, very acidic 

and bitter. Seeds ca. 40, broadly ovoid, smooth; cotyledons 

milky white; embryo(s) solitary to numerous. Fl. Apr–May, fr. 

Oct–Nov. 

95

96 

Cultivated and sometimes naturalized. S Anhui, S Gansu, Guizhou, 

Hubei, Hunan, S Jiangsu, S Shaanxi, Sichuan, Yunnan, Zhejiang. 

Citrus ×microcarpa Bunge, Enum. Pl. China Bor. 10. 1833. 

×Citrofortunella microcarpa (Bunge) Wijnands; ×C. mitis 

(Blanco) J. Ingram & H. E. Moore; Citrus ×mitis Blanco. 

Calamondin or calamansi is hybrid between Citrus reticulata and 

C. japonica. 

Citrus ×latifolia (Tanaka ex Yu. Tanaka) Tanaka, Kwaju Bunruigaku 

[Systematic Pomology], 140. 1951. 

Citrus ×aurantiifolia (Christmann) Swingle var. latifolia 

Tanaka ex Yu. Tanaka, Iconogr. Jap. Citrus Fruits 1: 57. 1946; 

C. ×aurantiifolia subsp. latifolia (Tanaka ex Yu. Tanaka) S. 

Ríos et al. 

Seedless lime is a hybrid between Citrus ×aurantiifolia and possibly 

C. ×limon. 

Citrus ×floridana (J. Ingram & H. E. Moore) Mabberley, Telopea 

7: 337. 1998. 

Citrus ×polytrifolia Govaerts, World Checkl. Seed Pl. 3(1): 15. 

1999. 

富民枳 fu min zhi 

Poncirus ×polyandra S. Q. Ding et al., Acta Bot. Yunnan. 

6: 292. 1984, not Citrus polyandra Tanaka (1928). 

Trees to 2.5 m tall, evergreen. Young branches green, triangular 

but becoming cylindric with age. Leaves palmately 

3-foliolate; petiole 1–2 cm, narrowly winged; lateral leaflet 

blades 2.7–3.8 × 0.7–1.7 cm; central leaflet blade 3.5–5 × 

0.9–1.4 cm, dark green, base cuneate, margin sinuate crenulate, 

apex mucronate. Flower solitary, 6.4–7 cm in diam. Pedicel 3–7 

cm, ca. 2 mm in diam. Calyx lobes 5, broadly ovate, ca. 7 × 5 

mm. Petals white, 5–9, 3.2–3.4 × 1.6–1.9 cm, broadly elliptic, 

lanuginous with more trichomes especially at margins. Stamens 

35–43; filaments ca. 4 mm, distinct; anthers yellow with milky 

white subpellucid dots. Ovary oblate, ca. 6 mm in diam., lanuginous, 

10-loculed; style ca. 2 mm; stigma green, clavate, ca. 2 

mm, apex emarginate. Fruit green, oblate, lanuginous when 

young. Fl. Mar–Apr, fr. Aug–Sep. 


The parents of this hybrid species are possibly Citrus cavaleriei 

and C. reticulata. 

Additional Hybrid Taxa Cultivated to a Limited Extent in China 

● Forests on mountain slopes; ca. 2400 m. SE Yunnan (Funing). 

This appears to be a Citrus trifoliata hybrid with another Citrus 

Doubtful Taxa 

×Citrofortunella floridana J. W. Ingram & H. E. Moore, 

Baileya 19: 170. 1975. 

Limequat is a hybrid between Citrus japonica and C. ×aurantiifolia. 

Citrus ×georgiana Mabberley, Blumea 49: 490. 2004. 

Citrangequat is a hybrid between Citrus ×insitorum and C. 

japonica. 

Citrus ×insitorum Mabberley, Gard. Bull. Singapore 54: 193. 

2002. 

×Citroncirus webberi J. Ingram & H. E. Moore, Baileya 

19: 171. 1975, not Citrus ×webberi Wester (1915). 

Citrange, a hybrid between Citrus ×aurantium and C. trifoliata, is 

widely grown in China as a rootstock. 

species. Such hybrids are commonly found where the parents are grown 

together. 

Fortunella bawangica C. C. Huang, Guihaia 11: 8. 1991. 

霸王金橘 ba wang jin ju 

Trees to 4 m tall. Young branches flat; spines ca. 4 cm. 

Petiole 3–5(–17) mm; leaf blade elliptic to ovate, (2–)4–7(–10) × 

(1–)2–3 cm, base rounded to obtuse, margin conspicuously 

crenulate on basal half, apex rounded. Flowers solitary. Pedicel 

ca. 5 mm but ca. 1 cm in fruit. Calyx lobes ca. 1 mm. Petals 

elliptic to lanceolate, ca. 7 mm. Stamens 20–25; filaments cohering 

into bundles; anthers mostly fertile. Ovary ovoid, 

5–7-loculed, with 1 or 2 ovules per locule; style short; stigma 

clavate. Fruit pyriform, 2.2–2.5 × 1.8–2.2 cm, 1- or 2-seeded; 

carpopodium ca. 2 mm thick. Seeds ovoid, base rounded, apex 

acute; seed coat smooth; embryo solitary; cotyledons green. 

● Scrub; ca. 1200 m. Hainan. 

Recent field studies show that populations morphologically in accordance 

with Fortunella bawangica are widely distributed in Hainan 

Island. The only character that can be used to distinguish this entity from 

typical wild populations of Citrus japonica is its pear-shaped fruit. Further 

study is needed to ascertain whether F. bawangica is a separate species 

or only a geographical race of C. japonica. 

22. AEGLE Corrêa, Trans. Linn. Soc. London 5: 222. 1800, nom. cons. 

木橘属 mu ju shu 


Trees, deciduous, with straight axillary spines. Leaves alternate, odd-pinnately (1 or)3(or 5)-foliolate. Inflorescences terminal and 

axillary, loosely fasciculate or racemose and few flowered or flowers solitary. Flowers bisexual, fragrant. Calyx cup-shaped, 4- or 

5-lobed. Petals 4 or 5, imbricate in bud. Stamens 30–50 or more; filaments short, subulate, distinct or irregularly coherent at base; 

anthers linear-lanceolate. Disk columnar or bell-shaped. Gynoecium 8–20-loculed, syncarpous; ovules many per locule, in 2 rows;


style short and thick; stigma capitate, cylindric, or bluntly conic, longitudinally grooved. Fruit a berry, globose to ellipsoid to pyriform; 

exocarp thin, parenchymatous; mesocarp woody; endocarp fleshy, soft and pulpy, becoming hard and reddish orange when dry, 

composed largely of elongate sessile pulp vesicles which are lacking within seed locules. Seeds depressed ovoid, woolly when ripe, 

embedded in a clear glutinous substance that becomes hard when dry; seed coat fleshy; endosperm lacking; embryo solitary, straight; 

cotyledons ovate, plano-convex; hypocotyl partly included between cotyledons. 

One species: native to India, cultivated in China. 

1. Aegle marmelos (Linnaeus) Corrêa, Trans. Linn. Soc. London 

5: 223. 1800. 

木橘 mu ju 

Crateva marmelos Linnaeus, Sp. Pl. 1: 444. 1753. 

Trees to 10 m tall; spines to 3 cm. Leaflet blades ovate to 

elliptic, 4–12 × 2–6 cm, base rounded to narrowly cuneate, 

margin crenulate, apex acuminate or sometimes acute. Calyx 

lobes ca. 1 mm. Petals white, ca. 1 cm. Stamens nearly as long as 

petals. Gynoecium ca. 6 mm. Fruit greenish yellow, 10–12 × 6–8 

cm; mesocarp ca. 3 mm thick. Seeds ca. 8 mm. Fr. Oct. 2n = 18, 

36. 

Cultivated in forests on slightly dry hillsides; 600–1000 m. S and 

SW Yunnan [native to India]. 

This species is used medicinally. 

97

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