CAMPANULACEAE [Draft]

桔梗科  jie geng ke

Hong Deyuan (洪德元)[1], Ge Song (葛颂)[2]; Thomas G. Lammers[3]

Herbs perennial with rhizomes or caudexes (caulorhiza), or annual, erect, climbing or twining, rarely (absent in China) shrubs or small trees, lactiferous. Leaves simple, alternate, less frequently opposite, rarely whorled. Inflorescences of diverse racemes or cymes, paniculate, capitate or pseudo-racemose, or sometimes flowers solitary. Flowers perfect, or rarely unisexual, mostly (3–)5(or 6 or 10)-merous, epigynous, semiepigynous, or rarely (Cyananthus) hypogynous. Calyx tube (hypanthium) adnate to whole ovary or up to different parts of ovary (Codonopsis), or free from ovary (Cyananthus), or sometimes calyx totally free and thus without calyx tube; calyx lobes valvate or imbricate. Corolla sympetalous but in some groups (e.g. Codonopsis, Wahlenbergia) corolla cleft nearly to base and rotate; lobes valvate, rarely imbricate, actinomorphic (Campanuloideae), or zygomorphic due to division to nearly base at posterior side (Lobelioideae). Stamens free from each other, or connivent into a tube via villous hairs on filaments (Lobelioideae) or totally connate (Lobelioideae); filaments usually dilated into lamellar at base, glabrous or densely villous on margins; anthers introrse, rarely lateral, tetrasporangiate and dithecal, often unequal in size in Lobelioideae, usually 2 or more with setose hairs at top. Disk absent, or present, discrete or annular, epigynous. Ovary inferior, semi-inferior, or superior for calyx but inferior or semi-inferior for corolla (Codonopsis), or rarely entirely superior (Cyananthus), 2–5(or 6 or 10)-locular; style single, often with collecting hairs at top; stigma 2–5(or 6 or 10)-fid; ovules numerous, on axile (rarely parietal) placentas, anatropous. Fruit commonly a capsule, loculicidal at superior part or poricidal at inferior part, rarely a berry, or a dry fruit. Seeds numerous, with endosperm; embryo straight.

About 70 genera and ca. 2000 species: cosmopolitan; 15 genera (two endemic) and 161 species (82 endemic) in China, mainly in the southwest.

Pentaphragma Wallich ex G. Don and Sphenoclea Gaertner were included in the Campanulaceae in FRPS but are treated in this volume as Pentaphragmataceae (p. xxx) and Sphenocleaceae (p. xxx), respectively. See the notes under those two families.

[Note: “Papua New Guinea” has been used throughout this manuscript. Actually, in some or most cases “New Guinea” is meant. The whole island is New Guinea; the E half of the island is Papua New Guinea; the W half of the island is Irian Jaya, which is part of Indonesia. Therefore it is wrong to treat “Papua New Guinea” and “New Guinea” as if they are the same.]

Hong De-yuan, Lian Yong-shan & Shen Lian-dai. 1983. Campanulaceae (Campanuloideae and Lobelioideae). In: Hong De-yuan, ed., Fl. Reipubl. Popularis Sin. 73(2): 1–173.

1a.       Corolla zygomorphic; stamens entirely connate; ovary 2-locular; inflorescence of raceme (Lobelioideae)                                                                                                                               15. Lobelia

1b.       Corolla actinomorphic; stamens free, rarely connivent via villous hairs on filaments; ovary 3–5(–6)-locular; inflorescences of diverse cymes, paniculate, pseudo-racemose (Campanuloideae).

2a.       Ovary superior for both calyx and corolla; flowers usually solitary and terminal; calyx often brown- or black hirsute or setose ...........................................................................................  1. Cyananthus

2b.       Ovary inferior or semi-inferior for both calyx and corolla, or at least for corolla; flowers in inflorescences, rarely solitary; calyx glabrous or with other types of hairs.

3a.       Fruit a capsule poricidal at inferior part, or a dry fruit, irregularly lacerate at inferior part.

4a.       Fruit a dry fruit, pericarp thin, membranous, irregularly lacerate; plants annual or perennial, roots thin, not carrot-shaped.

5a.       Flowers solitary and axillary, long pedicellate; plants perennial, with rhizomes often (?) tuberous at ends; seed-coat smooth ..................................................................................  12. Peracarpa

5b.       Flowers 1–3, fascicled on very short branches, sessile; plants annual, prostrate; seed-coat reticulate                                                                                                              13. Homocodon

4b.       Fruit a capsule, poricidal; plants mostly perennial and roots carrot-shaped, rarely annual.

6a.       Corolla lobed, campanulate.

7a.       Disk absent; capsule poricidal at base, middle or upper part .................  9. Campanula

7b.       Disk present, annular or tubular; capsule poricidal at base .................  10. Adenophora

6b.       Corolla divided nearly to base, corolla lobes petaloid.

8a.       Perennials with carrot-shaped roots; corolla lobes linear .......................  11. Asyneuma

8b.       Annuals with fibrous roots; corolla lobes lanceolate ................................  14. Triodanis

3b.       Fruit a capsule loculicidal at superior part, or a berry; ovary inferior, or inferior or semi-inferior only for corolla; stigma segments short, not circinate.

9a.       Fruit a berry; ovary and fruit apically flat.

10a.     Herbaceous vines; calyx lobes ovate-deltoid or ovate-lanceolate, entire
                                                                                                          7. Campanumoea

10b.     Erect herbs; calyx lobes linear or linear-lanceolate, dentate, rarely entire
                                                                                                                8. Cyclocodon

9b.       Fruit a capsule; ovary and fruit apically conical.

11a.     Glands 5, orbicular and compressed, alternate with stamens; herbaceous twiners  2. Leptocodon

11b.     Glands absent; erect herbs, rarely climbers or twiners.

12a.     Stigma segments ovate or oblong; calyx lobes and corolla lobes sometimes inserted at different positions; flowers often solitary; stems erect, diffuse, climbing or twining  3. Codonopsis

12b.     Stigma segments linear; calyx lobes and corolla inserted at same position; flowers in cymes or lax panicles; stems erect or ascending.

13a.     Leaves whorled or opposite, rarely alternate; ovary and capsule 5-locular; capsule valves opposite to persistent calyx lobes .....................................................................  5. Platycodon

13b.     Leaves alternate; ovary and capsule 2–5-locular; capsule valves alternate with persistent calyx lobes.

14a.     Calyx lobes echinate; leaves pinnatifid ...........................................  4. Echinocodon

14b.     Calyx lobes not echinate; leaves entire or serrate ..........................  6. Wahlenbergia

1. CYANANTHUS Wallich ex Bentham in Royle, Ill. Bot. Himal. Mts. 309. 1836, nom. cons., not Rafinesque (1815).

蓝钟花属  lan zhong hua shu

Hong Deyuan (洪德元); Thomas G. Lammers

Herbs perennial or annual; perennials with caudexes; caudexes thick, lignified, covered with numerous scales, usually multibranched and thus stems caespitose. Leaves alternate, but sometimes 4 or 5 leaves below pedicels aggregated, whorl-like. Flowers solitary, terminal on main stems and branches, rarely in a dichasium and dichasia aggregated on top of stems and branches, pedicellate. Calyx tubular or tubular-campanulate, 5-, rarely 4-lobed. Corolla tubular-campanulate, (3–)5-lobed. Stamens 5, often aggregated and surrounding ovary at upper part. Ovary superior, conical, mostly 5-locular, less frequently 4- or 3-locular. Fruits a capsule, loculicidal. Seeds numerous, brown-red or brown-black.

About 22 species: Himalayas and Hengduan Mountains; 19 species (ten endemic) in China.

1a.       Annuals.

2a.       Plants dwarf, less than 20 cm tall; flowers almost sessile, (3 or)4(or 5)-merous; corolla usually less than 10 mm, rarely up to 15 mm ................................................................................  19. C. hookeri

2b.       Plants taller; flowers pedicellate, usually 5-merous; corolla usually more than 15 mm.

3a.       Calyx covered with red-brown setae, base of setae swollen, black-verrucose; calyx lobes obovate-oblong, broadest at middle or above middle; corolla pale yellow or white  16. C. lichiangensis

3b.       Calyx villous; calyx lobes nearly linear or triangular, broadest below middle; corolla blue or blue-purple.

4a.       Leaves glabrous or sparsely puberulent; flowers in dichasia, fascicled at top of stems; calyx lobes linear .........................................................................................................  17. C. fasciculatus

4b.       Leaves hairy; flowers solitary, terminal on top of stems and branches, rarely 3–5-fascicled; calyx lobes triangular ..................................................................................................  18. C. inflatus

1b.       Perennials; caudexes robust, densely clothed with persistent and straw-colored scales.

5a.       Calyx densely covered only with brown-black setae, or mixed with such hairs; corolla lobes nearly orbicular, broadly ovate or oblong, as long as or slightly longer than broad.

6a.       Calyx covered with brown-black and white setae; stems sparsely arachnoid.

7a.       Leaf blades cordate to triangular-ovate, base cordate or rounded, margin obviously crenulate     14. C. cordifolius

7b.       Leaf blades linear-elliptic, base cuneate, margin entire or sinuous .............  15. C. sherriffii

6b.       Calyx covered with only brown-black setae; stems villous, scaberulose or sericeous.

8a.       Leaves less than 10 mm ...................................................................  13. C. microphyllus

8b.       Leaves more than 10 mm.

9a.       Leaf blades usually oblanceolate, obovate-lanceolate or obovate, margin with 3–9 gross and obtuse teeth above middle .................................................................................  10. C. lobatus

9b.       Leaf blades elliptic or ovate-lanceolate, margin entire or sinuous-crenulate above middle.

10a.     Leaves hispidulous on both surfaces; pedicels 12–32 mm; no several leaves aggregated below pedicels .....................................................................................  11. C. pedunculatus

10b.     Leaves adaxially glabrous, abaxially densely sericeous; pedicels 5–12 mm; several leaves aggregated below pedicels, whorl-like ..................................................................  12. C. sericeus

5b.       Calyx glabrous or covered with hairs other than brown-black setae, often ± swollen into saclike after anthesis; corolla lobes usually oblong, longer than broad.

11a.     Flowers several on a stem in racemelike; leaves ovate-lanceolate, abaxially densely sericeous; margin strongly recurved .....................................................................................  9. C. longiflorus

11b.     Flowers solitary and terminal on a stem; leaves rhomboid, flabellate, spatulate, or ovate, rarely lanceolate, rarely sericeous, margin spread or slightly recurved.

12a.     Leaves usually rhomboid, flabellate or spatulate, (2–)2.5–4.8(–5.5) mm, base usually truncate or broad-cuneate, obviously petiolate.

13a.     Flowers 1.6–2.4 cm; leaves rhomboid, base broad-cuneate ....................  1. C. delavayi

13b.     Flowers 2.5–4 cm.

14a.     Middle cauline leaves flabellate, with length/width ratio 0.6–1, base truncate, margin obviously serrate ...........................................................................................................  2. C. formosus

14b.     Middle cauline leaves spatulate or rhomboid, with length/width ratio more than 1.25, base truncate, margin entire or sinuous-crenulate ..............................................  3. C. chungdianensis

12b.     Leaves usually spatulate, elliptic, ovate or obovate-lanceolate, (3–)4.5–16 mm, base cuneate or rounded, attenuated into shortly petiolate, or leaves sessile.

15a.     Plants densely covered with yellow-brown arachnoid hairs except corolla; scales on caudexes 10–15 × ca. 3 mm; corolla 3–4 mm, lobed to 1/3 .....................................................  5. C. wardii

15b.     Plants glabrous or covered with other types of hairs, leaves adaxially white hirsute or white hispidulous; scales on caudexes much smaller; corolla 2–2.5(–3.8) cm, lobed to middle.

16a.     Leaves spatulate, with decurrent short petioles (0.5–)1.3–5 mm
.......................................................................................................  4. C. macrocalyx

16b.     Leaves elliptic, ovate, long ovate or obovate-lanceolate, sessile or with extremely short petioles less than 1(–1.7) mm.

17a.     Leaves sessile, broad-ovate or obovate-lanceolate, base rounded; flowers yellow or white        8. C. flavus

17b.     Leaves shortly petiolate or sessile, elliptic, narrowly elliptic or oblanceolate, base cuneate; flowers blue or blue-purple.

18a.     Pedicels (6–)10–32 mm; calyx tube mostly glabrous, rarely sparsely hairy outside  7. C. dolichosceles

18b.     Pedicels 2–10 mm; calyx tube densely hairy, rarely glabrous ..............  6. C. incanus

1. Cyananthus delavayi Franchet, J. Bot. (Morot) 1: 280. 1887.

细叶蓝钟花  xi ye lan zhong hua

Cyananthus barbatus Franchet (1885), not Edgeworth (1846); C. microrhombeus C. Y. Wu; C. microrhombeus var. leiocalyx C. Y. Wu.

Herbs perennial. Caudexes numerously branched, scales deltoid-lanceolate, ca. 2 × 1 mm. Stems caespitose, slender, 10–20 cm, multibranched, prostrate or ascending, white villous. Leaves alternate, 3–5 leaves below flowers aggregated and whorl-like; petioles ca. 25 mm, white-villous; leaf blades nearly orbicular or ovate-deltoid, 3–5 × 2–4 mm, adaxially hispidulous, abaxially silver-sericeous, base nearly rounded or truncate, apex rounded or obtuse, margin slightly recurved, entire or sinuous. Flowers solitary, terminal on main stems and some branches; pedicels 3–7 mm. Calyx tubular, 8–10 × 3–5 mm, base rounded, below enlarged in fruit, brown hirsute or glabrescent; lobes narrowly triangular, 2–4 × 1.5–2.5 mm, hirsute both outside and inside. Corolla blue, tubular-campanulate 1.8–3 cm, glabrous outside, densely villous at throat inside; lobes linear-oblong, 6–10 mm. Ovary 5-locular, nearly equal to calyx in length at anthesis; style extended to corolla throat; stigma 5-fid. Capsules conical, longer than calyx. Seeds oblong, ends pointed, ca. 1.8 × 0.3 mm, brown. Fl. Aug–Sep.

* Calcareous grassy slopes, forests, forest margins; 1900–4000 m. SW Sichuan, C and NW Yunnan.

2. Cyananthus formosus Diels, Notes Roy. Bot. Gard. Edinburgh 5: 172. 1912.

美丽蓝钟花  mei li lan zhong hua

Herbs perennial. Roots carrot-shaped, up to 10 mm wide. Caudexes robust, often numerously branched; scales persistent, linear-lanceolate, ca. 5 mm. Stems slender, caespitose, 10–25 cm, pale purple, prostrate to ascending, simple or with short branches. Leaves alternate, 4 or 5 below flowers aggregated and whorl-like; lower leaves scalelike; middle and upper ones larger; petioles 3–7 mm; leaf blades rhomboid-flabellate, 4–9 × 2–6 mm, adaxially sparsely white hirsute or glabrous, abaxially densely white hirsute, base broad-cuneate or nearly truncate, apex truncate, margin recurved, with 3–5 obtuse teeth. Flowers large, solitary and terminal on main stems and branches; pedicels ca. 3 mm. Calyx tubular-campanulate; tube 8–12 mm, densely covered with pale-brown hirsute hairs; lobes narrowly triangular, 4–6 × 2–3 mm, hirsute both inside and outside. Corolla dark blue or purple-blue, 2.5–3.5 cm, glabrous outside, densely villous at throat inside; lobes obovate-oblong, 1/2–1/3 as long as tube, apex with a tuft of hairs outside. Ovary 5-locular, nearly as long as calyx tube; style extended up to corolla throat; stigma 5-fid. Fl. Aug–Sep.

* Grassy slopes, forest glades, forest margins; 2800–4300 m. SW Sichuan (Muli), NW Yunnan (Heqing, Lijiang).

3. Cyananthus chungdianensis C. Y. Wu in Wu & Li, Rep. Stud. Pl. Trop. Subtrop. Reg. Yunnan, 1: 85. 1965.

中甸蓝钟花  zhong dian lan zhong hua

Perennial herbs Caudexes robust, multibranched, densely covered with scales; scales ovate-lanceolate, ca. 3 mm, scarious. Stems caespitose, with slender branches above, prostrate or ascending, 5–10 cm, below glabrous, with subulate, scalelike leaves, above densely white-villous, with ordinary leaves. Leaves small, rhomboid, oblanceolate or obovate, 3–11 × 1–5 mm including cuneate petioles, adaxially sparsely villous, abaxially densely white-villous, apex acute, margin recurved, entire or serrate above middle. Flowers solitary and terminal; pedicels ca. 4 mm. Calyx tubular, densely brown-yellow or white hispidulous outside; tube 8–10 × 6–7 mm; lobes narrowly triangular, ca. 5 mm. Corolla purple-blue, 2.3–3.2 cm, lobed to 1/3; lobes obovate-oblong, densely yellow-villous near base inside. Fl. Aug–Sep.

* Alpine meadows with shrubs, grassy places at forest margins; 3500–4300 m. NW Yunnan (Xianggelila).

4. Cyananthus macrocalyx Franchet, J. Bot. (Morot) 1: 297. 1887.

大萼蓝钟花  da e lan zhong hua

Herbs perennial. Caudexes robust, lignified, multibranched, above with persistent ovate-lanceolate scales; scales 3–4 mm. Stems caespitose, ascending, simple, 5–20 cm, base often sparsely white-villous, above sparsely white-villous or glabrous. Leaves alternate, gradually becoming larger up stems, 4 or 5 below flowers aggregated and whorl-like; petioles 2–4 mm; leaf blades rhomboid, nearly orbicular or spatulate, 5–8 × 3–6 mm, adaxially sparsely hispidulous, abaxially densely white hirsute, base suddenly attenuated into petioles, apex obtuse or acute, margin recurved, entire or sinuous-serrate. Flowers solitary, terminal; pedicels 4–10 mm. Calyx tubular at anthesis, 7–13 mm, yellow-green or purplish, enlarged in fruit, below globose, obviously veined, glabrous, brown hirsute or glabrescent; lobes narrowly triangular, glabrous or brown hirsute outside, brown hirsute inside. Corolla yellow, sometimes purplish or red-veined, or less frequently purple below but yellow at exserted part, tubular-campanulate, 2–3 cm, glabrous outside, densely villous at throat inside; lobes linear-obovate, 2/5 as long as total length of corolla. Ovary extended up to throat. Capsules exserted above calyx. Seeds oblong, ca. 1.3 mm, smooth. Fl. Jul–Aug.

Alpine meadows, grassy slopes; 2500–5300 m. SE Gansu, S Qinghai, W Sichuan, SE Xizang, Yunnan [Bhutan, NE India (Assam, Sikkim), Myanmar, E Nepal].

1a. Leaf blade ovate to suborbicular, margin entire; pedicel less than 5 mm; calyx more than 11 mm  4a. subsp. macrocalyx

1b. Leaf blade obovate-spatulate, crenulate; pedicel ca. 10 mm; calyx less than 11 mm .......  4b. subsp. spathulifolius

4a. Cyananthus macrocalyx subsp. macrocalyx

大萼蓝钟花(原亚种)  da e lan zhong hua (yuan ya zhong)

Cyananthus incanus J. D. Hooker & Thomson var. leiocalyx Franchet; C. leiocalyx (Franchet) Cowan; C. leiocalyx subsp. lucidus K. Shrestha; C. macrocalyx var. flavopurpureus C. Marquand; C. macrocalyx var. pilosus C. Marquand; C. neurocalyx C. Y. Wu.

Leaf blade ovate to suborbicular, margin entire. Pedicel less than 5 mm. Calyx more than 11 mm.

* Alpine meadows, grassy slopes; 2500–5000 m. SE Gansu (Zhugqu), S Qinghai (Nangqên), W Sichuan (Barkam, Daocheng, Hongyuan, Jinchuan, Kangding, Lixian, Muli, Qianning, Sêrtar, Xiangcheng), Yunnan (Dali, Dêqên, Dongchuan, Eryuan, Gongshan, Heqing, Huize, Lijiang, Qiaojia, Weixi, Xianggelila).

4b. Cyananthus macrocalyx subsp. spathulifolius (Nannfeldt) K. Shrestha, Acta Phytotax. Sin. 35: 412. 1997.

匙叶蓝钟花  ci ye lan zhong hua

Cyananthus spathulifolius Nannfeldt, Acta Horti Gothob. 5: 30. 1929.

Leaf blade obovate-spatulate, margin crenulate. Pedicel ca. 10 mm. Calyx less than 11 mm.

Alpine meadows; 3000–5300 m. SE Xizang (Cona, Mainling, Nyingchi, Yadong, Zayü) [Bhutan, NE India (Assam, Sikkim), Myanmar, E Nepal].

5. Cyananthus wardii C. Marquand, J. Linn. Soc., Bot. 48: 196. 1929.

棕毛蓝钟花  zhong mao lan zhong hua

Herbs perennial. Roots ca. 5 mm in diam. Caudexes with 2 or 3 grades of branches, densely covered with persistent and straw-colored scales; scales large, elliptic, 10–15 × 3–4 mm. Stems several from each branch of caudex, caespitose, 10–18 cm, lower half with small, straw-colored and elliptic scales, sparsely yellow-arachnoid. Leaves at upper half of stems, larger up stems, alternate but 4 or 5 below flowers aggregated and whorl-like, sessile or nearly sessile, oblanceolate or elliptic, 5–11 × 3–5 mm, densely covered with brown-yellow arachnoid hairs on both surfaces, base broad-cuneate, apex rounded or obtuse, margin entire. Flowers solitary and terminal; pedicels 2–4 mm, covered with brown-yellow arachnoid hairs. Calyx short-tubular, 10–12 × 6–8 mm, densely covered with brown-yellow arachnoid hairs; lobes as long as tube, ovate-lanceolate, apex obtuse. Corolla tubular, 3–4 cm, glabrous outside but densely villous inside, lobed to 1/3; lobes long obovate, with a tuft of yellow villous hairs at top outside. Stamens 1/2 as long as corolla tube. Style glabrous, slightly exceeding corolla tube.

* 3400–5000 m. SE Xizang.

6. Cyananthus incanus J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 2: 20. 1857.

灰毛蓝钟花  hui mao lan zhong hua

Herbs perennial. Caudexes robust, numerously branched, top with persistent ovate-lanceolate scales; scales ca. 4 mm. Stems caespitose, simple or below branched, white-villous. Leaves alternate, but 4 or 5 below flowers aggregated and whorl-like, gradually larger upward stems, shortly petiolate; leaf blades ovate-elliptic to linear-elliptic, 4–12 × 1.5–4 mm, white hirsute on both surfaces, base cuneate, apex obtuse, margin recurved, nearly entire or sinuous-serrate. Flowers solitary, terminal on main stems and branches; pedicels 4–13 mm, hirsute. Calyx short-tubular, narrower below than above at anthesis, but broader than above in fruit, densely with yellow-brown hirsute hairs to glabrous; tube 5–8 mm; lobes triangular, 2–3 mm, slightly longer than broad, densely white hirsute inside. Corolla blue purple or dark blue, 2.5–3 times longer than calyx, glabrous outside, densely villous at throat inside; lobes obovate-oblong, ca. 2/5 as long as total length of corolla. Ovary nearly equal to calyx tube in length at anthesis; style extended up to corolla throat. Capsule exserted above calyx, 5-loculicidal, 10–13 mm. Seeds oblong, pale brown. Fl. Jul–Sep, fr. Sep–Oct.

Grassy slopes, forest glades, grassy places at forest and thicket margins, alpine meadows; 2700–5300 m. S Qinghai, W Sichuan, E Xizang, NW Yunnan [Bhutan, N India (Sikkim), Nepal].

1a. Calyx tubes glabrous or densely white hirsute outside; leaves usually white hirsute and glaucous; cauline leaves elliptic, with length/width ratio 1.5–2.5
.......................................................................................................................  6a. subsp. incanus

1b. Calyx tubes usually densely yellow-brown hirsute outside, occasionally glabrous; leaves yellow hirsute; cauline leaves linear-elliptic, with length/width ratio 2.5–6.5 ..........................  6b. subsp. petiolatus

6a. Cyananthus incanus subsp. incanus

灰毛蓝钟花(原亚种)  hui mao lan zhong hua (yuan ya zhong)

Cyananthus incanus var. decumbens Y. S. Lian; C. incanus subsp. orientalis K. Shrestha; C. incanus var. parvus C. Marquand; C. incanus var. trichocalyx Franchet.

Cauline leaves ovate or ovate-elliptic, 4–8 × 1.5–4 mm, usually white hirsute and glaucous. Calyx tubes glabrous or densely white hirsute outside.

Grassy slopes, forest glades, alpine meadows; 3100–5300 m. S Qinghai (Nangqên), W Sichuan (Daocheng, Derong, Sêrtar, Xiangcheng), E Xizang (W to Namling and Nyalam) [Bhutan, N India (Sikkim), Nepal].

6b. Cyananthus incanus subsp. petiolatus (Franchet) D. Y. Hong & L. M. Ma, Acta Phytotax. Sin. 29: 44. 1991.

毛叶蓝钟花  mao ye lan zhong hua

Cyananthus petiolatus Franchet, Bull. Soc. Philom. Paris, sér. 8, 3: 147. 1891; C. neglectus C. Marquand; C. petiolatus var. pilifolius (C. Y. Wu) Y. S. Lian; C. pilifolius C. Y. Wu; C. pilifolius var. minor C. Y. Wu; C. pilifolius var. pallidocoeruleus C. Y. Wu.

Cauline leaves elliptic to linear-elliptic, 5–12 × 2–3 mm, yellow hirsute; calyx tubes usually densely yellow-brown hirsute, occasionally glabrous outside.

* Alpine meadows, grassy slopes, grassy places at forest and thicket margins; 2700–5000 m. W Sichuan, NW Yunnan (Dêqên, Xianggelila).

7. Cyananthus dolichosceles C. Marquand, Bull. Misc. Inform. Kew 1924: 250. 1924.

川西蓝钟花  chuan xi lan zhong hua

Herbs perennials. Caudexes robust, multibranched, top covered with persistent scales; scales ovate-lanceolate, 5–10 × 2–3 mm. Stems caespitose, prostrate or ascending, simple, 5–15 cm, base covered with brown-yellow or white villous hairs, above sparsely white-villous. Leaves alternate, only 4 or 5 below flowers aggregated and whorl-like, becoming gradually larger up stems, sessile or almost sessile, obovate-lanceolate or long oblong, 5–13 × 3–5 mm, with length/width ratio 2–3, sparsely covered with appressed white hirsute hairs, base long cuneate, apex acute, margin recurved, entire or sinuous-crenulate. Flowers solitary and terminal; pedicels (6–)10–32 mm, sparsely brown-yellow villous or glabrescent. Calyx tubular, base rounded, below swollen and nearly globose in fruit, glabrous or sparsely brownish hirsute, 10–15 mm; lobes triangular, brown-yellow hirsute inside, 1/4–1/3 as long as tube. Corolla purple-blue, 2.5–3 times as long as calyx, outside glabrous, inside densely villous at throat; lobes obovate-oblong, 2/5 as long as total length of corolla. Ovary about equal to calyx in length at anthesis; style extended up to throat. Capsules much longer than calyx at maturity, 5-loculicidal. Seeds oblong, obtuse at two ends, 1.5–1.7 × ca. 0.8 mm. Fl. Jul–Aug.

* Grassy slopes, alpine meadows, alpine meadows with scrub; 2800–4800 m. W Sichuan (Daocheng, Dawu, Garzê, Jinchuan, Kangding, Qianning, Muli, Xiangcheng, Xiaojin).

8. Cyananthus flavus C. Marquand, Bull. Misc. Inform. Kew 1924: 247. 1924.

黄钟花  huang zhong hua

Herbs perennial. Roots carrot-shaped, up to 5 mm in diam. Caudexes robust, multibranched, top covered with persistent scales; scales long ovate, ca. 3 × 1 mm. Stems caespitose, ascending, 7–16 cm, simple or with short branches, densely white-villous to nearly glabrous. Leaves alternate, only 4 or 5 below flowers aggregated and whorl-like, becoming gradually larger up stems, sessile or nearly sessile, elliptic or ovate-orbicular, 5–16 × 3–8 mm, gray-white hirsute or glabrous on both surfaces, both base and apex acute, margin entire, recurved. Flowers solitary and terminal; pedicels 1–2 cm, usually glabrous, rarely sparsely hirsute. Calyx short-tubular, base rounded, 8–12 mm, glabrous, enlarged in fruit; lobes triangular, 3–4 × ca. 3 mm, inside yellow hirsute. Corolla yellow or white, 2.5–3 cm, outside glabrous, inside densely white-villous at throat; lobes obovate-oblong or obovate-elliptic, 14–16 × 5–7 mm, apex often very sparsely brown-villous. Ovary nearly equal to calyx in length; style exceeding corolla tube. Fl. Jul–Aug.

* Grassy slopes; 2700–3600 m. SW Sichuan, NE and NW Yunnan.

1a. Flowers yellow; leaves broad-ovate, apex often rounded ....................................  8a. subsp. flavus

1b. Flowers white; leaves obovate-lanceolate, apex often acute ..........................  8b. subsp. montanus

8a. Cyananthus flavus subsp. flavus

黄钟花(原亚种)  huang zhong hua (yuan ya zhong)

Flowers yellow, pale yellow or yellowish green. Cauline leaves usually ovate or broad-ovate, apex obtuse.

* Grassy slopes; 3100–3600 m. NW Yunnan (Lijiang, Xianggelila).

8b. Cyananthus flavus subsp. montanus (C. Y. Wu) D. Y. Hong & L. M. Ma, Acta Phytotax. Sin. 29: 46. 1991.

白钟花  bai zhong hua

Cyananthus montanus C. Y. Wu in Wu & Li, Rep. Pl. Trop. Subtrop. Reg. China 1: 89. 1965; Atropanthe mairei (H. Léveillé) H. Léveillé; C. albiflorus D. F. Chamberlain; C. mairei (H. Léveillé) Cowan; Wahlenbergia mairei H. Léveillé.

Flowers often white. Cauline leaves obovate-lanceolate, apex acute.

* Grassy slopes; 2700–3400 m. SW Sichuan (Xichang, Zhaojue), NE Yunnan (Huize).

9. Cyananthus longiflorus Franchet, J. Bot. (Morot) 1: 280. 1887.

长花蓝钟花  chang hua lan zhong hua

Cyananthus argenteus C. Marquand; C. obtusilobus C. Marquand.

Perennial herbs. Caudexes robust, ± lignified, multibranched, top with some ovate scales; scales ca. 2 mm. Stems erect, 4–22 cm tall, caespitose, below lignified, multibranched, densely covered with gray-white lanate hairs; branches very short. Leaves alternate but those below flowers aggregated; petioles 2–4 mm; leaf blades elliptic or ovate-elliptic, 5–10 × 2–3 mm, adaxially sparsely puberulent or glabrescent, abaxially densely silvery-sericeous, base cuneate, apex acute, margin recurved, entire. Flowers solitary, terminal on main stems and branches, nearly sessile. Calyx tubular, ca. 10 × 6–8 mm at anthesis, outside densely brown-yellow villous; lobes lanceolate, 5–7 × ca. 3 mm, two surfaces both hairy. Corolla tubular-campanulate, purple-blue or blue-purple, 3.5–5 cm outside glabrous, inside densely villous at throat; lobes obovate-oblong, 1/3–1/2 as long as total length of corolla, apex with a tuft of setose hairs. Style almost extending up to corolla throat. Capsules slightly longer than calyx at maturity. Seeds oblong, ca. 1.3 mm. Fl. Jul–Sep.

* Pinus forests, dry slopes, sandy dunes; 2800–4300 m. W Yunnan.

10. Cyananthus lobatus Wallich ex Bentham in Royle, Ill. Bot. Himal. Mts. 309. 1839.

裂叶蓝钟花  lie ye lan zhong hua

Cyananthus insignis R. E. Grahame [Gard. Chron., ser. 3, 107: 10. 1940, w/o Latin descr.??]; C. lobatus var. farreri C. Marquand.

Herbs perennial. Roots carrot-shaped, up to 1.3 cm in diam. Caudexes robust, multibranched, top with persistent scales; scales ovate-lanceolate, ca. 4 × 2.5 mm. Stems caespitose, prostrate or ascending, 20–50 cm, simple or branched, above sparsely villous. Leaves alternate, sessile or with petioles less than 5 mm; leaf blades nearly leathery, various in shape, size and degree of division, usually oblanceolate, obovate-lanceolate, obovate or rhomboid, 7–30 × 5–16 mm, adaxially sparsely hispidulous, glabrescent, abaxially villous, base cuneate, margin slightly recurved, with 3–9 gross teeth at upper part. Flowers solitary, terminal on main stems and branches; pedicels 1–3 cm, brown hirsute. Calyx tubular-campanulate; tube 1.2–1.5 cm, densely covered with brown-red to brown-black hispid hairs; lobes triangular to lanceolate-triangular, 1/4–1/2 as long as tube, brown-hispid both inside and outside. Corolla tubular-campanulate, purple-blue to pale blue, 3–4 cm, outside glabrous, inside densely villous at throat; lobes nearly orbicular, slightly more than 1/3 of total length of corolla, apex with a tuft of short brown hairs in center. Ovary conical, nearly equal to calyx tube in length; style extending up to middle part of corolla; stigma large, capitate, 5-fid. Fl. Aug–Sep.

Grassy slopes, forests; 2800–4500 m. SE Xizang (Nyingchi, Yadong, Zayü), NW Yunnan (Bijiang, Gongshan, Weixi) [Bhutan, India, NE Myanmar, Nepal].

11. Cyananthus pedunculatus C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 434. 1881.

有梗蓝钟花  you geng lan zhong hua

Cyananthus linifolius Wallich ex J. D. Hooker & Thomson, p.p.

Herbs perennial. Roots carrot-shaped, up to 1.2 cm in diam. Caudexes robust, multibranched, top covered with persistent scales; scales elliptic-lanceolate, straw-colored. Stems caespitose, ascending or diffuse, 10–20 cm, usually simple, hispidulous. Leaves alternate, becoming gradually larger up stems, almost sessile; leaf blades ovate-elliptic, ovate-lanceolate to linear-lanceolate, 8–16 × 2–6 mm, hispidulous on both surfaces, base rounded, margin entire or sinuous above middle. Flowers solitary, terminal; pedicels 1.2–4 cm, covered with brown-black villous hairs. Calyx tubular, 10–15 × 7–10 mm, densely covered with brown-black hirsute hairs, lobes lanceolate-triangular, 1/2–2/5 as long as tube. Corolla purple-blue, funnelform-campanulate, 2.5–4.5 cm, throat glabrous or sparsely villous; lobes nearly orbicular, 1/3 as long as total length of corolla, apex with a tuft of brown hirsute hairs. Ovary nearly equal to calyx in length; style extending up to corolla throat; stigma 5-fid. Fl. Aug–Sep.

Alpine scrub; 3600–4900 m. S Xizang (Yadong) [Bhutan, N India, Nepal].

12. Cyananthus sericeus Y. S. Lian, Acta Phytotax. Sin. 17(3): 122. 1979.

绢毛蓝钟花  juan mao lan zhong hua

Herbs perennial, 20–30 cm tall. Caudexes robust, multibranched, top densely covered with long ovate or ovate-oblong persistent scales; scales 3–5 mm. Stems slender, densely sericeous when young, glabrescent. Leaves alternate, only 3 or 4 below flowers aggregated and whorl-like; lower leaves scalelike, ca. 2 × 1 mm, sparsely villous; middle and upper ones long elliptic, 8–12 × 2–4 mm, adaxially glabrous or sparsely hairy, abaxially densely covered with retrorse and appressed sericeous hairs. Flowers solitary and terminal; pedicels 5–12 mm, with brown-black villous hairs. Calyx tubular-campanulate, broader above than below, 15–20 × ca. 10 mm, densely covered with brown-black hirsute hairs; lobes lanceolate-triangular, ca. 1/2 as long as tube, margin white-villous. Corolla blue, with dark blue veins, 3.5–4 cm, throat glabrous; lobes broad-obovate, about 1/3 as long as total length of corolla. Ovary conical, slightly shorter than calyx. Style almost extending up to corolla throat; stigma capitate. Fl. Sep.

* Rocky crevices; 3500–3600 m. S Xizang (Cona).

13. Cyananthus microphyllus Edgeworth, Trans. Linn. Soc. London 20: 81. 1846.

小叶蓝钟花  xiao ye lan zhong hua

Cyananthus linifolius Wallich ex J. D. Hooker & Thomson, p.p.; C. nepalensis Kitamura.

Herbs perennial. Caudexes robust, multibranched, top densely covered with scales; scales elliptic-lanceolate, ca. 5 × 2 mm. Stems slender, 5–25 cm, below branched, brown-red, glabrous or sparsely puberulent. Leaves alternate, almost sessile, ovate, ovate-lanceolate to linear-elliptic, 5–13 × 1.5–5 mm, adaxially glabrous, abaxially sericeous, base rounded to shallowly cordate, apex obtuse or acute, margin recurved, entire or sinuous. Flowers solitary and terminal; pedicels 3–10 mm, covered with brown-black setae. Calyx tubular-campanulate, 7–10 mm, with brown-black setae, base truncate; lobes triangular, 1/3–2/5 as long as whole calyx, hairy on both sides. Corolla tubular-campanulate, blue-purple or blue, 1.8–2.8 cm, outside glabrous, inside densely covered with fimbriate white-villous hairs; lobes obovate-oblong, nearly equal to tube in length. Ovary conical, nearly as long as calyx; style extending up to corolla throat. Seeds bright-brown, oblong, ca. 2 × 0.5–0.7 mm. Fl. Sep.

Mountain slopes; 3300–4300 m. S Xizang (Nyalam) [India, Nepal].

14. Cyananthus cordifolius Duthie, Bull. Misc. Inform. Kew 1912: 37. 1912.

心叶蓝钟花  xin ye lan zhong hua

Herbs perennial. Caudexes robust, multibranched. Stems slender, 14–27 cm, base prostrate, sparsely arachnoid-villous, middle with slender and sterile branches. Leaves alternate, nearly sessile, cordate to triangular-ovate, 10–15 × 5–9 mm, adaxially glabrous or sparsely puberulent, abaxially sparsely villous, base cordate or nearly rounded, apex obtuse or acute, margin serrate; leaves on sterile branches much smaller, elliptic-ovate, almost entire, base rounded, rarely cordate. Flowers solitary and terminal; pedicels 5–18 mm, with brown-black setae. Calyx tubular-campanulate, 11–16 × ca. 8 mm, base truncate, densely covered with brown-black setae; lobes lanceolate-triangular, slightly shorter than tube, inside both black- and white-villous. Corolla blue or pale blue, tubular-campanulate, 2.5–3.5 cm, densely villous at throat; lobes oblong, 12–15 × 5–6 mm. Ovary conical-ovoid, slightly shorter than calyx; style extending up to corolla throat. Fl. Aug.

Mountain thickets; 3000–4000 m. S Xizang (Gyirong) [NW India, Nepal].

15. Cyananthus sherriffii Cowan, New Fl. Silva 10: 181. 1938.

杂毛蓝钟花  za mao lan zhong hua

Herbs perennial. Caudexes robust, nearly lignified, multibranched, top densely covered with persistent oblong-lanceolate scales; scales up to 10 × 2.5 mm. Stems caespitose, slender, terete, dark brown, 5–15 cm, sparsely arachnoid. Leaves alternate, but several below flowers aggregated and whorl-like, gradually larger up stems, sessile or shortly petiolate; leaf blades linear-elliptic, 4–7 × 2–3 mm, but those below flowers larger, 9–12 × 3–3.5 mm, adaxially sparsely arachnoid-villous along middle vein, abaxially silvery-sericeous. Flowers solitary, terminal on longer stems; pedicels 5–8 mm. Calyx tubular, 10–15 × ca. 7 mm, densely covered with glaucous and brown-black setae, base truncate; lobes triangular-lanceolate, slightly shorter than tube, hairy on both sides. Corolla blue or gray-blue, tubular-campanulate, ca. 2.5 cm, outside glabrous, inside villous at throat; lobes oblong, apex with a tuft of brown villous hairs. Ovary conical, nearly as long as calyx; style extending up to corolla throat; stigma 5-fid. Fl. Jul–Aug.

Alpine meadows or thickets; 3200–5000 m. S Xizang (Lhünzê, Nangxian) [Bhutan, India, Nepal].

16. Cyananthus lichiangensis W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 8: 109. 1913.

丽江蓝钟花  li jiang lan zhong hua

Herbs annual. Stems caespitose, 10–30 cm tall, glabrous, simple or with slender branches. Leaves sparse, alternate but 4 or 5 below flowers aggregated and whorl-like; petioles 2–5 mm, villous; leaf blades ovate-triangular or rhomboid, 5–13 × 5–11 mm, both surfaces hispidulous, base long cuneate, apex obtuse, margin recurved, entire or sinuous-crenulate. Flowers solitary, terminal on both main stems and branches; pedicels 2–5 mm, glabrous. Calyx tubular, tube 8–10 × 6–8 mm, outside covered with red-brown setae, base of setae swollen, often black-verrucose; lobes obovate-oblong, ca. 1/3 as long as tube, broadest part at middle or above middle, outside sparsely covered with red-brown setae, inside with appressed red-brown villous hairs. Corolla pale yellow or green-yellow, sometimes blue- or purple-striate, tubular-campanulate, 2 times as long as calyx tube, outside glabrous, inside densely villous at throat; lobes oblong, 1/3–1/4 as long as whole corolla, apex acute. Ovary nearly as long as calyx tube; style extending up to corolla throat. Capsules exserted above calyx at maturity. Seeds oblong, obtuse at both ends, ca. 1 × 0.5 mm. Fl. Aug.

* Grassy slopes, grassy places at forest margins; 3000–4000 m. SW Sichuan, N Yunnan (Dêqên, Lijiang, Zhenxiong).

17. Cyananthus fasciculatus C. Marquand, Bull. Misc. Inform. Kew 1924: 247. 1924.

束花蓝钟花  shu hua lan zhong hua

Herbs annuals, 30–100 cm tall. Stems multibranched, glabrous or sparsely villous, branches long and patent. Leaves alternate, scattered, but several below flowers aggregated and whorl-like; petioles slender, 5–10 mm, glabrous or sparsely villous; leaf blades cordate or triangular-ovate, 4–16 × 4–15 mm, adaxially sparsely hispidulous, abaxially usually glabrous, base shallowly cordate to cuneate, apex rounded. Flowers in dichasia and aggregated at top of main stems and branches; pedicels 2–4 mm, slender, glabrous. Calyx tubular, broader below than above, 5–7 × 3–5 mm, rather densely brown-setose, bottom rounded; lobes usually 5, but some flowers on lower branches with 4 lobes, linear-triangular, ciliate. Corolla pale blue, tubular-campanulate, 14–17 mm, outside glabrous, inside villous near throat; lobes usually 5, but rarely 4, obovate-oblong, ca. 5 × 2–3 mm. Ovary nearly equal to calyx tube in length; style exserted above corolla tube. Capsules exceeding calyx in height at maturity. Seeds ellipsoid, two ends pointed, ca. 0.5 × 0.2 mm. Fl. Sep–Oct.

* Forests, thickets, grassy slopes; 2400–3500 m. W Sichuan (Lixian, Muli, Yanbian, Yanyuan), N Yunnan (Kunming, Lijiang, Luquan, Xianggelila).

18. Cyananthus inflatus J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 2: 21. 1857.

胀萼蓝钟花  zhangn e lan zhong hua

Cyananthus forrestii Diels; C. pseudoinflatus P. C. Tsoong.

Herbs annual, up to 80 cm tall. Stems erect or ascending, branched, sparsely villous. Leaves alternate, sparse, but 3 or 4 below flowers aggregated and whorl-like; petioles slender, 2–7 mm, villous; leaf blades rhomboid, orbicular-rhomboid, or fan-shaped, 5–15 × 3–10 mm, adaxially hispidulous, abaxially hirsute, base rounded or cuneate, apex obtuse, margin entire or obscurely crenulate. Flowers usually solitary, terminal on main stems and branches; pedicels 2–5 mm, slender, brown hirsute. Calyx urceolate at anthesis, 8–12 mm, obviously inflated after flowering, outside densely brown hirsute; lobes 5, lanceolate-triangular, 1/4–2/5 as long as calyx tube, both sides brown hirsute. Corolla pale blue, tubular-campanulate, nearly 2 times as long as calyx, outside glabrous, inside densely villous at throat; lobes 5, obovate-oblong, nearly 1/3 as long as total length of corolla. Ovary slightly shorter than calyx; style extending up to corolla throat. Capsules ovoid, exceeding calyx in height at maturity, 5-loculicidal. Seeds brown-red, ellipsoid, obtuse at two ends, ca. 0.5 × 0.2 mm. Fl. Aug–Sep. 2n = 14.

Alpine meadows, grassy and shrubby slopes; 1900–4900 m. W Guizhou (Weining), W Sichuan, SE Xizang (Cona, Mainling, Yadong), N Yunnan [Bhutan, India, N Myanmar, Nepal].

19. Cyananthus hookeri C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 435. 1881.

蓝钟花  lan zhong hua

Cyananthus cronquistii Shrestha; C. hookeri var. densus C. Marquand; C. hookeri var. grandiflorus C. Marquand; C. hookeri var. hispidus Franchet; C. hookeri var. levicalyx Y. S. Lian; C. hookeri var. levicaulis Franchet.

Herbs annual. Stems usually caespitose, nearly erect or ascending, glabrous to rather densely villous, 4–20 cm, short-branched, branches 1.5–10 cm. Leaves alternate but several below aggregated and whorl-like; petioles 1–3 mm; leaf blades rhomboid, rhomboid-triangular or ovate, 3–7 × 2–6 mm, adaxially sparsely to rather densely hispidulous, abaxially sparsely to rather densely hirsute, base cuneate, suddenly attenuated into petioles, apex obtuse, margin sparsely crenulate, sometimes entire. Flowers small, solitary, terminal on main stems and branches, almost sessile. Calyx ovoid, 3–7 mm, outside densely clothed with yellow-brown hirsute hairs, to nearly glabrous; lobes usually 4, less frequently 3 or 5, triangular, 1/2–1/3 as long as tube, both sides hirsute. Corolla purple-blue, tubular, 7–15 mm, outside glabrous, inside densely villous at throat; lobes usually 4, sometimes 3 or 5, obovate-oblong, apex with 3 or 4 brown-yellow villous hairs. Stamens 4. Style extending above corolla throat; stigma 4-fid. Capsules ovoid, exserted above calyx at maturity. Seeds long ovoid, ca. 1.2 × 0.4 mm. Fl. Aug–Sep.

Thickets, grasslands; 2700–4500 m. SE Gansu (Tao He basin), S Qinghai (Tongren, Yushu), W Sichuan (W of Kangding and Lixian), E Xizang (E of Namling), N Yunnan (Dali, Dêqên, Gongshan, Heqing, Huize, Kunming, Qiaojia, Weixi, Xianggelila) [Bhutan, N India (Sikkim), Nepal].

2. LEPTOCODON (J. D. Hooker) Lemaire, Ill. Hort. 3: Misc. 49. 1856.

细钟花属  xi zhong hua shu

Hong Deyuan (洪德元); Thomas G. Lammers

Codonopsis sect. Leptocodon J. D. Hooker, Ill. Himal. Pl. t. 16, f. a. 1855.

Perennial vines. Stems slender. Leaves alternate, leaves on branches often opposite or nearly opposite. Flowers solitary, outside axils, less frequently in axils, or opposite leaves. Calyx 5-lobed, tube short-obconical. Corolla tubular, 5-lobed. Stamens 5; filaments long, base slightly dilated, alternate with 5 free, lamellar glands. Ovary semi-inferior, with superior part long conical, 3-locular; style long; stigma 3-fid. Capsule 3-loculicidal at superior part. Seeds numerous.

Two species: E Himalayas to SW China; both species in China.

1a.       Plants glabrous throughout except young leaves sparsely villous; calyx lobes linear-oblong, often reflexed, distant from each other, glabrous, base clawlike
                                                                                                                                                       1. L. gracilis

1b.       Plants ± hairy, rarely sparsely villous only on leaf margin; calyx lobes ovate to long ovate, not reflexed, overlapping each other, at least villous on margin, base obtuse, truncate or cordate, but not clawlike        2. L. hirsutus

1. Leptocodon gracilis (J. D. Hooker) Lemaire, Ill. Hort. 3: Misc. 49. 1856.

细钟花  xi zhong hua

Codonopsis gracilis J. D. Hooker, Ill. Himal. Pl. t. 16, f. a. 1855.

Plants glabrous throughout except sparsely villous young leaves, awfully odorous. Roots unknown. Stems long, slender, branched. Leaves alternate, rarely those on branchlets opposite, long petiolate; leaf blades membranous, ovate-orbicular, 1–2.5 × 1–2.5 cm, adaxially green, abaxially gray, base and apex both obtuse, margin sinuate-crenate. Pedicels slender, 1–5 cm, upright or curved. Calyx glabrous, tube ca. 2 mm; lobes linear-oblong, 6–8 × 1.5–2 mm, pendulous from anthesis, claw 1–2 mm, margin with 1 or 2 pairs of serratures, serratures pointed upwards, callose. Corolla blue or purple, 3–3.5 cm; tube ca. 4 mm in diam., limb ca. 1 cm in diam., lobes triangular, 4–5 mm. Filaments ca. 2 cm. Style ca. 2 cm. Inferior part of capsule semiglobose, ca. 5 mm, superior part ca. 1 cm, conical. Seeds unknown. Fl. Aug–Oct.

Forests, thickets; 2000–2500 m. SW Sichuan (Huili, Muli, Yanbian), NW Yunnan (Binchuan, Dali, Heqing, Lijiang) [N India (Sikkim), E Nepal].

2. Leptocodon hirsutus D. Y. Hong, Acta Phytotax. Sin. 18: 246. 1980.

毛细钟花  mao xi zhong hua

Codonopsis hirsuta (D.Y. Hong) K. E. Morris & Lammers (1999), not (Handel-Mazzetti) D. Y. Hong & L. M. Ma (1992); C. hongii Lammers.

Perennials awfully odorous. Roots slender, not thickened. Stems hirsute on young parts. Leaves on main stems alternate, those on branchlets almost opposite, all leaves long petiolate; leaf blades abaxially villous, rarely extremely sparsely. Flowers solitary, nearly opposite to leaves. Calyx lobes hirsute on abaxial side and margin, ovate, overlapping each other at anthesis, separated at fruiting, upright, ca. 5 × 3–4 mm, base cordate or nearly truncate, not clawlike, margin sinuate-crenate. Inferior part of capsule semiglobose, 8–10 mm in diam., superior part conical, ca. 10 mm. Seeds ellipsoid, ca. 1 mm. Fl. Jul–Aug, fr. Oct–Nov.

Mixed forests, thickets, grasslands; 2000–2700 m. SE Xizang (Cona, Bomi, Zayü), NW Yunnan (Gongshan, Weixi) [Bhutan, N Myanmar].

3. CODONOPSIS Wallich in Roxburgh, Fl. Ind. 2: 103. 1824.

党参属  dang shen shu

Hong Deyuan (洪德元); Thomas G. Lammers

Perennial herbs. Caudexes often short. Roots usually thickened, various in shape, mostly fleshy, rarely lignified, carrot-shaped, fusiform or tuberous. Stems erect, ascending, climbing, or twining. Leaves alternate, opposite, fascicled or pseudo-verticillate. Flowers solitary, terminal on main stems and branches, sometimes opposite to leaves, rarely axillary. Calyx tube adnate to ovary up to different parts of ovary, often 10-ribbed; calyx lobes 5. Corolla epigynous, broad-campanulate, campanulate, funnel-shaped, tubular-campanulate, or tubular, shallowly 5-lobed or 5-fid nearly to base, and thus rotate, red-purple, blue-purple, blue-white, yellow-green or green. Stamens 5; filaments often dilated at base, glabrous or hairy; anthers basifixed, connective glabrous or setose. Ovary inferior or at least inferior for corolla, 3-locular; ovules numerous; style glabrous or hairy; stigma usually 3-fid, lobes broad. Capsule with persistent calyx, half-globose or obconical in lower part, rostrate in upper part, 3-loculicidal in superior part. Seeds numerous, ellipsoid, oblong or ovoid, winged or wingless, smooth or reticulate; embryo straight, embedded in copious endosperm.

About 45 species: C and E Asia; 41 species (28 endemic) in China.

Where belongs Codonopsis limprichtii Lingelsheim & Borza, Repert. Spec. Nov. Regni Veg. 13: 391. 1914, described from Yunnan?

1a.       Corolla divided mostly to near base, rotate, rarely deep lobed and campanulate; inferior part of capsule obconical, superior part short and obtuse; roots often tuberous, ovoid, or globose.

2a.       Stems robust, base up to 1 cm in diam.; calyx lobes linear; corolla campanulate, divided to 1/3       36. C. dicentrifolia

2b.       Stems slender, base less than 5 mm in diam.; calyx lobes not linear; corolla divided to near base, rotate.

3a.       Stems erect or ± prostrate, scapiform; leaves rosulate ................................  37. C. rosulata

3b.       Stems twining or nearly erect; leaves opposite or alternate, not rosulate.

4a.       Leaves usually aggregated at one part of stems, mostly at lower part; stems usually less than 1 m, erect or twining only at apex.

5a.       Stems and adaxial side of leaves hirsute; leaves ovate, serrate ..................  39. C. hirsuta

5b.       Stems and leaves glabrous; leaves linear or pinoid, rarely lanceolate or ovate
                                                                                                        40. C. graminifolia

4b.       Leaves generally evenly distributed on stems; stems much longer, twining.

6a.       Leaves ovate-lanceolate to linear-lanceolate, slightly cordate, truncate to cuneate at base, nearly sessile to shortly petiolate, with petioles less than 1.2 cm ........................  38. C. convolvulacea

6b.       Leaves cordate-ovate or cordate-lanceolate, cordate at base, with petioles 1.5–3 cm  41. C. efilamentosa

1b.       Corolla shallowly 5-lobed, campanulate, tubular-campanulate or tubular; capsule nearly semiglobose at lower inferior half, conical at superior part; roots carrot-shaped or fusiform (tuberous only in C. ussuriensis).

7a.       Stems twining.

8a.       Leaves fascicled of 3–4 at top of lateral branches, pseudo-verticillate.

9a.       Roots usually fusiform; seeds winged .....................................................  1. C. lanceolata

9b.       Roots tuberous; seeds not winged .........................................................  2. C. ussuriensis

8b.       Leaves alternate or opposite, not pseudo-verticillate.

10a.     Corolla tubular; calyx lobes toothed or crenulate on margins.

11a.     Leaves deeply incised, gross-serrate, base cordate to broad-cuneate
                                                                                                             11. C. deltoidea

11b.     Leaves shallowly incised, serrate or nearly entire, base cuneate to rounded
                                                                                                              13. C. tubulosa

10b.     Corolla campanulate or tubular-campanulate; calyx lobes entire; leaves entire or shallowly incised.

12a.     Lower cauline leaves cordate, vary rarely truncate or rounded at base.

13a.     Stems almost simple; leaves nearly entire or slightly sinuous on margins, acute at apex  3. C. cordifolioidea

13b.     Stems branched; leaves serrate on margins, obtuse at apex.

14a.     Calyx adnate to ovary up to middle, sinus between calyx lobes narrow and acute.

15a.     Corolla large, more than 1.5 × 1.7 cm, usually longer than calyx lobes
                                                                                                           4. C. pilosula

15b.     Corolla small, less than 1 × 1.5 cm, shorter than or nearly equal to calyx lobes  5. C. micrantha

14b.     Calyx adnate to ovary up to top, sinus between calyx lobes broad and obtuse.

16a.     Leaves less than 3 × 3 cm; calyx hirsute; calyx lobes ovate-orbicular or rhomboid-ovate, dentate and hirsute; corolla globose-campanulate
                                                                                                     6. C. subglobosa

16b.     Leaves much more than 3 × 3 cm; calyx tube sparsely hairy, calyx lobes narrow-oblong or lanceolate, nearly entire, densely puberulent; corolla broad-campanulate  7. C. affinis

12b.     Lower cauline leaves cuneate or rounded, occasionally cordate at base.

17a.     Calyx adnate to ovary only up to lowest part, and thus ovary almost superior for calyx  8. C. tangshen

17b.     Calyx adnate to ovary up to middle, and thus ovary semi-inferior for calyx.

18a.     Stems and leaves glabrous; leaves entire.

19a.     Leaves 0.5–2 × 0.5–1.5 cm, apex acute or obtuse ..............................  9. C. farreri

19b.     Leaves 6–7 × 2–2.5 cm, apex short-acuminate ............................  10. C. longifolia

18b.     Stems and leaves ± hairy; leaves crenulate or serrate on margins.

20a.     Lateral branches 5–15 cm; leaves crenulate or entire; calyx lobes broad, with sinus between them narrow and acute, not reflexed ..................................................  14. C. kawakamii

20b.     Lateral branches no more than 3 cm; leaves serrate; calyx lobes narrower, with sinus between them broad and obtuse, often reflexed
                                                                                                           12. C. henryi

7b.       Stems usually erect, scapiform, sometimes climbing or procumbent, but not twining.

21a.     Stems without branches like pinnate leaves and usually sterile.

22a.     Corolla tubular or tubular-campanulate; filaments villous; stems mostly procumbent or climbing.

23a.     Corolla 2–4 cm, yellow-green, base with brown-red shade, glabrous
                                                                                                            15. C. benthamii

23b.     Corolla only ca. 1.7 cm, yellowish, with purple-blue spots, densely puberulent outside  16. C. pianmaensis

22b.     Corolla campanulate; filaments without villous hairs; stems mostly erect or scapiform.

24a.     Leaves located at lower part of stems, main stems scapiform.

25a.     Petioles 2–7 cm, leaf blades 2–13 × 1.5–5 cm; main stems with 1–4 flowers; calyx lobes 5–7 × ca. 3 mm, with sinus between them broad and obtuse; corolla less than 1.5 cm wide  18. C. subscaposa

25b.     Petioles less than 1 cm, leaf blades 5–8 × 1.5–3 cm; main stems with 1–2 flowers; calyx lobes 8–12 × 4–6 mm, with sinus between them narrow; corolla up to 3 cm wide  19. C. meleagris

24b.     Leaves evenly arranged on main stems, not scapiform.

26a.     Leaves opposite; plants glabrous throughout ......................................  20. C. purpurea

26b.     Leaves alternate.

27a.     Plants ± hairy; corolla less than 2 cm ............................................  17. C. subsimplex

27b.     Plants glabrous; corolla more than 3.5 cm.

28a.     Leaves lanceolate, 1.5–6.5 cm, entire; calyx lobes narrowly lanceolate, 18–20 × 2–3 mm      . 22. C. gombalana

28b.     Leaves broadly lanceolate, 5–14 × 3–7 cm, serrate; calyx lobes 20–30 × 6–10 mm  23. C. chimiliensis

21b.     Stems with branches like pinnate leaves and usually sterile.

29a.     Veins on leaves conspicuous; leaf margins recurved forming thick periphery.

30a.     Corolla short-tubular, ca. 1.2 cm; lower slender branches sometimes fertile
                                                                                                          35. C. xizangensis

30b.     Corolla campanulate, more than 1.5 cm; lower slender branches sterile.

31a.     Leaves nearly entire; flowers entirely glabrous.

32a.     Leaves not more than 1.8 × 1.5 cm, sparsely puberulent beneath; corolla purple-green, with purple veins ...................................................................................................  30. C. alpina

32b.     Leaves up to 3.2 × 2.6 cm, glabrous on both surfaces; corolla pale blue, with red-brown spots inside .................................................................................................  31. C. cardiophylla

31b.     Leaves crenulate; flowers ± hairy on some parts.

33a.     Calyx-lobes 5–7 × 2–3 mm, with sinus between them broad and obtuse
.................................................................................................  34. C. chlorocodon

33b.     Calyx lobes more than 10 mm, with sinus between them narrow and acute.

34a.     Corolla villous inside; calyx tube mostly hairy; calyx lobes narrowly triangular, broadest at base, mostly entirely hispidulous ........................................................  33. C. tsinlingensis

34b.     Corolla totally glabrous; calyx tube mostly glabrous; calyx lobes ovate or ovate-lanceolate, broadest at middle, hispidulous only at apex outside
                                                                                                     32. C. viridiflora

29b.     Veins on leaves inconspicuous; leaf margins not recurved.

35a.     Corolla tubular or tubular-campanulate, at least lower part tubular.

36a.     Corolla tubular; connective setose ................................................  24. C. thalictrifolia

36b.     Corolla tubular-campanulate, lower half tubular but suddenly expanded at middle; connective glabrous ..........................................................................................  25. C. bulleyana

35b.     Corolla broad-campanulate.

37a.     Plants densely with white hairs and plants white; leaf blades less than 1.5 × 1 cm  28. C. canescens

37b.     Plants sparsely hairy and plants green; leaf blades usually larger.

38a.     Calyx lobes 15–20 × 6–8 mm, hairy only at apex; leaf blades 1–5 × 0.8–3.2 cm  21. C. clematidea

38b.     Calyx lobes much smaller, entirely hairy; leaf blades mostly smaller.

39a.     Corolla villous both inside and outside ...........................................  29. C. argentea

39b.     Corolla glabrous or hairy only above outside.

40a.     Leaves 5–8 mm; calyx lobes ca. 8 mm, two lateral sides recurved
.....................................................................................................  26. C. foetens

40b.     Leaves more than 10 mm; calyx lobes generally larger, recurved or not
....................................................................................................  27. C. nervosa

1. Codonopsis lanceolata (Siebold & Zuccarini) Bentham & J. D. Hooker, Gen. Pl. 2: 558. 1876.

羊乳  yang ru

Campanumoea lanceolata Siebold & Zuccarini, Fl. Jap. 1: 174. 1835; C. japonica Siebold & &/ex “Morr.” [see Belgique Hort. 337] (1863), not Maximowicz (1868); Codonopsis bodinieri H. Léveillé.

Plants glabrous throughout or occasionally sparsely villous on stems and leaves. Caudexes nearly cylindrical. Roots usually fusiform-thickened, 10–20 × 1–6 cm. Stems twining, more than 100 cm, yellow-green but with purplish shade, often branched. Leaves on main stems alternate, lanceolate or narrowly rhomboid-ovate, 8–14 × 3–7 mm; usually leaves 2–4 fascicled on top of branchlets, nearly opposite or verticillate, petioles 1–5 mm; leaf blades rhomboid-ovate, narrowly ovate or elliptic, 3–10 × 1.3–4.5 cm, base attenuate, apex acute or obtuse, margin usually entire or sparsely waved-dentate, adaxially green, abaxially gray-green, veins obvious. Flowers solitary or twin on top of branchlets; pedicels 1–9 cm. Calyx adnate to ovary by half; calyx tube semiglobose; calyx lobes ovate-deltoid, 1.3–3 × 0.5–1 cm, acute, entire; sinus between calyx lobes acute or gradually becoming broader after anthesis. Corolla broad-campanulate, 2–4 × 2–3.5 cm, shallowly lobed; lobes deltoid, recurved, 0.5–1 cm, yellow-green or milk-white, with purple spots. Disk fleshy, dark green. Filaments subulate, slightly dilated at base, 4–6 mm; anthers 3–5 mm; ovary inferior. Capsule semiglobose below, rostate at top, 2–2.5 cm in diam. Seeds numerous, ovoid, winged, brown. Fl. and fr. Jul–Aug. 2n = 16.

Shrublands in ravines, broad-leaved forests. N, E, and SC China [Japan, Korea, Russia (Far East)].

2. Codonopsis ussuriensis (Ruprecht & Maximowicz) Hemsley, J. Linn. Soc., Bot. 26: 6. 1889.

雀斑党参  qio ban dang shen

Glossocomia ussuriensis Ruprecht & Maximowicz, Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 15: 209. 1857.

Plants glabrous throughout, or stems and leaves sparsely villous. Roots tuberous or oblong, 1–3 cm in diam., gray-yellow. Stems twining, slender, green, white or dark-purple, glabrous or internodes sparsely villous. Leaves on main stems alternate, lanceolate or rhomboid-ovate, smaller, those on top of branches usually 3–5-fascicled, pseudo-verticillate, petioles short; leaf blades lanceolate or oblong, 3–5 × 1.5–2.5 cm, base attenuate, apex acute or obtuse, margin entire, adaxially green, glabrous, abaxially gray-green, glabrous or sparsely villous. Flowers solitary, terminal on slender branches; pedicels 2–5 cm, bracts single, small, lanceolate or narrowly rhomboid-ovate. Calyx tube adnate to ovary up to middle of ovary, half-globose; calyx lobes narrowly lanceolate or ovate-deltoid, 10–20 × 6–8 mm, apex acute, margin entire. Corolla campanulate, 2–3 × 1.5–2.5 cm, shallowly 5-lobed; lobes deltoid, dark purple, insides with remarkable black-purple stripes or black spots. Filaments 3–5 mm, slightly dilated at base, anthers 2–4 mm. Capsules half-globose in inferior part, rostrate in superior part. Seeds numerous, ovoid, wingless, dark brown, lucid. Fl. Jul–Aug. 2n = 16.

Ravines, moist meadows, mostly in sandy soil; ca. 800 m. E Heilongjiang, E Jilin [Japan, Korea, Russia (Far East)].

3. Codonopsis cordifolioidea P. C. Tsoong, Contr. Inst. Bot. Nat. Acad. Peiping 3(3): 95. 1935.

心叶党参  xin ye dang shen

Plants glabrous or nearly glabrous or only leaves sparsely short-setose. Stems twining, more than 1 m, with a few very short branches. Leaves on main stems lax, alternate, ca. 10 cm distant between two leaves; petioles filiform, 5–9 cm; leaf blades broad-ovate, nearly entire, up to 10 × 7 cm, base nearly cordate, with a square sinus, apex short-acuminate or acute, veins conspicuous, lateral veins from base, adaxially green, abaxially gray-green; leaves on branches usually only 2 and opposite, petioles less than 1 cm, similar to those on main stems in shape, but sinus not square. Flowers solitary, opposite leaves, pedicels 3–6 cm. Calyx tube adnate to ovary up to middle, semiglobose; calyx lobes deltoid-lanceolate, ca. 10 × 5–6 mm, entire, apex acuminate, sinus between calyx lobes narrow. Corolla campanulate, 1.7–1.8 × 1–1.2 cm, 5-lobed to middle, lobes lanceolate-deltoid, dark-blue. Filaments dilated at base, ca. 5 mm, anthers ca. 5 mm. Capsule semiglobose in lower half, rostrate above, ca. 1.5 cm in diam. Seeds numerous, ellipsoid, brown, obscurely reticulate. Fl. and fr. Sep–Oct.

* Forests; 1700–2200 m. NW Yunnan (Fugong, Gongshan).

4. Codonopsis pilosula (Franchet) Nannfeldt, Acta Horti Gothob 5: 29. 1930.

党参  dang shen

Roots carrot-shaped or fusiform-cylindrical, often branched, 15–30 × 1–3 cm. Stems twining, glabrous, 1–2 m, branched, lateral branches 15–30 cm, branchlets 1–5 cm, sterile or fertile. Leaves on main stems and branches alternate, those on branchlets opposite; petioles 0.5–2.5 cm, sparsely short-setose; leaf blades ovate or narrowly ovate, 1–6.5 × 0.8–5 cm, base nearly cordate, apex obtuse or acute, margin crenate, adaxially green, abaxially gray-green, sparsely or densely appressed hirsute or villous, rarely glabrous. Flowers solitary and terminal on branches, pedicellate. Calyx adnate to ovary up to middle; calyx lobes broad-lanceolate or narrowly oblong, 10–20 × 6–8 mm, apex obtuse or acute, margin nearly entire or slightly waved, sinus between calyx lobes narrow and pointed. Corolla broad-campanulate, 1.8–2.3 × 1.8–2.5 cm, yellow-green, with purple spots inside, shallowly lobed; lobes deltoid, apex acute. Filaments slightly dilated at base, ca. 5 mm; anthers elongate, 5–6 mm. Stigma white-setose. Capsule semiglobose in inferior part, conical at superior part. Seeds numerous, ovoid, wingless, brown-yellow. Fl. and fr. Jul–Oct. 2n = 16.

Forests, thickets, meadows at forest margins; 900–3900 m. E Gansu, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, E Qinghai, Shaanxi, Shandong, Shanxi, N and W Sichuan, NW Yunnan [Korea, Mongolia, Russia (Far East)].

1a. Leaves glabrous or often hispidulous ...............................................................  4a. subsp. pilosula

1b. Leaves usually hirsute ................................................................................  4b. subsp. handeliana

4a. Codonopsis pilosula subsp. pilosula

党参(原亚种)  dang shen (yuan ya zhong)

Campanumoea pilosula Franchet, Pl. David. 1: 192. 1884; Codonopsis glaberrima Nannfeldt; C. microtubulosa Z. T. Wang & G. J. Xu; C. modesta Nannfeldt; C. pilosula var. glaberrima (Nannfeldt) P. C. Tsoong; C. pilosula var. modesta (Nannfeldt) L. T. Shen; C. pilosula var. volubilis (Nannfeldt) L. T. Shen; C. silvestris Komarov; C. volubilis Nannfeldt.

Leaves glabrous 1–6.5 × 0.8–5 cm, or often hispidulous. Calyx lobes 1.4–1.8 cm. Corolla 2–2.3 cm.

Forests, thickets; 900–2900 m. E Gansu, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, E Qinghai, Shaanxi, Shandong, Shanxi, N Sichuan (Heishui, Pingwu, Songpan) [Korea, Mongolia, Russia (Far East)].

4b. Codonopsis pilosula subsp. handeliana (Nannfeldt) D. Y. Hong & L. M. Ma in Fl. Sichuanica, 10:532. 1992.

闪毛党参  shan mao dang shen

Codonopsis handeliana Nannfeldt in Handel-Mazzetti, Symb. Sin. 7: 1078. 1936; C. pilosula var. handeliana (Nannfeldt) L. T. Shen.

Leaves 1–3 × 0.8–2.5 cm, usually hirsute on both surfaces. Calyx lobes 1.5–2 cm. Corolla [text missing].

* Forests, thickets, meadows at forest margins; 2300–3900 m. W Sichuan, NW Yunnan.

5. Codonopsis micrantha Chipp, J. Linn. Soc., Bot. 38: 382. 1908.

小花党参  xiao hua dang shen

Campanumoea violifolia H. Léveillé; Melothria violifolia H. Léveillé.

Roots long cylindrical, 20–30 × 0.5–1 cm. Stems twining, more than 1 m, branched, yellow-green or green, sparsely villous or nearly glabrous. Leaves alternate, or opposite on branches, long petiolate; petioles 2–5 cm; leaf blades ovate, to broadly ovate, 2–5.5 × 2.4–4 cm, base cordate, leaf ears slightly curved toward inside, sinus broad and obtuse, or nearly square, apex obtuse or acute, margin crenate or serrate, green above, gray-green, sparsely villous or nearly glabrous beneath. Flowers axillary or opposite to leaves; pedicels 1–2.5 cm, glabrous. Calyx tube adnate to ovary to up to middle, semiglobose; calyx lobes triangular-lanceolate, 10–14 × 4–5 cm, apex acute, glabrous or sparsely ciliate, sinus narrow and pointed. Corolla campanulate, ca. 8 mm, 7–9 mm in diam., white, glabrous or ciliate, 5-divided to middle, lobes triangular. Filaments slightly dilated at base, ca. 3 mm; anthers ca. 2 mm. Ovary almost entirely inferior for corolla. Capsule semiglobose below, conical and rostrate above, 8–10 mm in diam., superior part very short, 2.5–3.5 mm. Seeds numerous, ovoid, slightly compressed, shortly tailed, ca. 1 × 0.5 mm, smooth, brown-yellow. Fl. and fr. Jul–Oct.

* Mountain scrub, sunny forests; 1900–2600 m. SW Sichuan, N Yunnan.

6. Codonopsis subglobosa W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 8: 108. 1913.

球花党参  qiu hua dang shen

Roots fusiform, carrot-shaped or cylindrical, 30–50 × 1.5–8 cm. Stems twining, more than 2 m, branched, yellow-green or green, sparsely white hirsute. Leaves on mains and branches alternate, those on branchlets nearly opposite; petioles 0.5–2 cm, sparsely white hirsute; leaf blades broadly ovate, ovate to narrowly ovate, 0.5–3 × 0.5–2.5 cm, base shallowly cordate to rounded, apex obtuse or acute, margin slightly waved or shallowly crenate, adaxially green, hispidulous, abaxially gray-green, hirsute. Flowers solitary, terminal on branchlets, or opposite to leaves; pedicels hirsute. Calyx tube adnate to whole ovary, semiglobose, with 10 conspicuous radiate ribs, ribs sparsely white hirsute; calyx lobes distant to each other, nearly orbicular or rhomboid-ovate, 9–13 × 6–8 mm, veins conspicuous, margins serrate, often recurved, abaxially white hirsute. Corolla globose-campanulate, ca. 2 × 2.5 cm, pale yellow-green, apex dark red-purple; lobes broadly triangular. Filaments slightly dilated; anthers ellipsoid, ca. 5 mm. Capsule semiglobose below, conical or rostrate above. Seeds numerous, ellipsoid or ovoid, wingless, smooth, yellow-brown. Fl. and fr. Jul–Oct.

* Stony mountain meadows, scrub by streams; 2500–3500 m. W Sichuan, NW Yunnan (Dêqên, Lijiang).

7. Codonopsis affinis J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 2: 12. 1857.

大叶党参  da ye dang shen

Roots much thickened, fusiform, branched. Stems twining, more than 2 m, branched, yellow green or green, nearly glabrous or sparsely white hirsute. Leaves on main stems and major branches alternate, those on branchlets nearly opposite; petioles 1–6 cm, shortly white hirsute; leaf blades ovate or ovate-oblong, 2.5–15 × 1–9 cm, base deeply cordate to rounded, apex short-acuminate, margin entire to shallowly crenate, adaxially green, sparsely white hirsute, abaxially gray-green, densely white hirsute. Flowers solitary, terminal, or opposite leaves; pedicels 1–4 cm, hispidulous. Calyx tube adnate to whole ovary, semiglobose, obscurely 10-ribbed, hispidulous; calyx lobes distant from each other, linear-oblong or linear-lanceolate, 8–17 × 2–3 mm, white-lanate on both surfaces, apex acute, or ± obtuse, margin waved, veins obscure. Corolla broadly campanulate, ca. 2 × 2 cm, yellow-green at lower and middle parts, purple-red at upper part, apically hirsute outside, shallowly 5-lobed; lobes triangular. Filaments slightly dilated at base; anthers ellipsoid, ca. 5 mm, glabrous. Capsule semiglobose or nearly globose below, conical and rostrate, above, 1.5–5 cm in diam., rostrum ca. 4 mm. Seeds numerous, ellipsoid or ovoid, wingless, yellow-brown, smooth. Fl. and fr. Jul–Oct.

Forests; 2300–3200 m. S Xizang (Nyalam) [N India (Sikkim), Myanmar, Nepal].

8. Codonopsis tangshen Oliver, Hooker’s Icon. Pl. 20: t. 1966. 1891.

川党参  chuan dang shen

Plants glabrous throughout except leaves. Roots fusiform or fusiform-cylindrical, branched, 15–30 × 1–1.5 cm. Stems twining, up to 3 m, major branches 15–50 cm, branchlets 1–5 cm, leafy, sterile or fertile with a terminal flower. Leaves on main stems and major branches alternate, those on branchlets nearly opposite; petioles 0.7–2.4 cm; leaf blades ovate, shallowly ovate or lanceolate, 2–8 × 0.2–3.5 cm, base cuneate or slightly rounded, rarely cordate, apex obtuse or acute, margin crenate, adaxially green, densely puberulent, abaxially gray-green, densely puberulent. Flowers solitary, terminal on branches and branchlets, pedicellate. Calyx nearly entirely free from ovary, 5-segmented; calyx lobes oblong-lanceolate, 14–17 × 5–7 mm, entire or slightly waved, apex acute. Corolla distant from insertion of calyx lobes by ca. 3 mm, campanulate, 1.5–2 × 2.5–3 cm, pale yellow-green, with purple spots inside, shallowly lobed; lobes nearly deltoid. Filaments slightly dilated at base, 7–8 mm; anthers 4–5 mm. Ovary inferior for corolla, 0.5–1.4 cm in diam. Capsule nearly globose below, conical above, 2–2.5 cm in diam. Seeds numerous, ellipsoid, wingless, brown-yellow. Fl. and fr. Jul–Oct.

* Scrub at forest margins; 900–2300 m. Chongqing, N Guizhou, W Hubei, NW Hunan, S Shaanxi, NE Sichuan; also cultivated on a large scale.

9. Codonopsis farreri J. Anthony, Notes Roy. Bot. Gard. Edinburgh 15: 181. 1926.

秃叶党参  tu ye dang shen

Codonopsis farreri var. grandiflora S. H. Huang, Acta Bot. Yunnan. 6: 394. 1984.

Stems twining, 60–120 cm, branched. Leaves on main stems and major branches alternate or nearly opposite, those on branchlets opposite or pseudo-verticillate; petioles 1–1.5 cm; leaf blades ovate, lanceolate or elliptic, 0.5–2 × 0.5–1.5 cm, base cuneate or rounded, apex acute or slightly obtuse, margin entire, glabrous on both surfaces. Flowers solitary, terminal; pedicels ca. 5 cm. Calyx tube adnate to ovary only up to middle; calyx lobes linear-lanceolate, nearly entire, ca. 2 × 0.5 cm, apex acute or slightly obtuse. Corolla campanulate, 3.3–5.5 × ca. 3 cm, pale green or greenish yellow, with brown-red or purple reticulate veins; tube 2–3.5 cm; lobes broad-triangular, obtuse, 1.2–1.6 × 1.2–2 cm. Filaments slightly dilated at base, ca. 6 mm; anthers elongate, 5–7 mm.

Alpine forests; 3600–4000 m. NW Yunnan (Bijiang) [N Myanmar].

10. Codonopsis longifolia D. Y. Hong, Acta Phytotax. Sin. 18: 245. 1980.

长叶党参  chang ye dang shen

Twining perennials, glabrous throughout. Roots carrot-shaped, not branched, ca. 18 × 1 cm. Stems purple, branched. Leaves on main stems and major branches alternate, larger; petioles 1–2 cm; leaf blades ovate-lanceolate, thin, entire, 6–7 × 2–2.5 cm, base rounded to cuneate, apex short-acuminate; those on branchlets similar but much smaller. Flowers solitary, terminal; pedicels up to 13 cm. Calyx tube adnate to ovary up to middle, semiglobose; calyx lobes broad-linear, ca. 22 × 8 mm, entire, apex obtuse. Corolla yellow-green. Fl. and fr. Aug–Sep.

* Mountain slopes; 3600–3700 m. S Xizang (Nyalam).

Similar to Codonopsis rotundifolia Bentham in Kashmir, but this species is entirely glabrous, with leaves entire and thin, base rounded to cuneate, not cordate, apex short-acuminate, veins not concave, and calyx lobes entire, and thus distinct from C. rotundifolia.

11. Codonopsis deltoidea Chipp, J. Linn. Soc., Bot. 38: 387. 1908.

三角叶党参  san jiao ye dang shen

Roots conical or cylindrical, ± branched, 15–30 ×1–1.4 cm. Stems twining, more than 1 m, branches and branchlets both short, no more than 5 cm, leafy, sterile or with terminal flowers, sparsely villous or gradually glabrescent. Leaves alternate or opposite; petioles 0.2–8 cm, villous or hirsute; leaf blades deltoid-ovate or broadly ovate, but those on branchlets ovate to narrowly ovate, 3–9.5 × 4–10 cm, base shallowly cordate, truncate, rounded or broadly cuneate, apex acuminate, margin dentate, adaxially green, sparsely puberulent or hirsute, abaxially gray-green, sparsely puberulent or hirsute. Flowers solitary, terminal on main stems, major branches and branchlets, sometimes in cymes; pedicels less than 2 cm, villous. Calyx tube adnate to ovary up to middle, semiglobose, glabrous; calyx lobes ovate, 11–13 × 5–7 mm, apex acute, margin dentate, often white-ciliate. Corolla campanulate, ca. 2.5 × 1–1.5 cm, pale yellow-green, with purple veins, shallowly lobed; lobes deltoid. Filaments slightly dilated at base, ca. 6 mm, glabrous; anthers ca. 3 mm. Capsules semiglobose below, short-conical above. Seeds numerous, void, wingless, smooth, yellow-white. Fl. and fr. Jul–Oct.

* Margins of mountain forests, scrub; 1800–2800 m. W Sichuan (Emei Shan).

12. Codonopsis henryi Oliver, Hooker’s Icon. Pl. 20: t. 1967. 1891.

川鄂党参  chuan e dang shen

Codonopsis levicalyx L. T. Shen; C. levicalyx var. hirsuticalyx L. T. Shen.

Stems twining, more than 1 m, branches short, less than 3 cm, with 2–4 leaves, sterile or with terminal flowers, sparsely puberulent or glabrescent at internodes. Leaves on main stems alternate, those on branches nearly opposite; petioles 0.2–2 cm, puberulent; leaf blades ovate-lanceolate or lanceolate, 3–15 × 1–7 cm, base decurrent or cuneate, apex acuminate, margin conspicuously dentate, adaxially green, puberulent, abaxially gray-green, appressed-puberulent. Flowers solitary, terminal on branches; pedicels ca. 1 cm, puberulent. Calyx tube adnate to ovary up to middle, semiglobose, puberulent or gradually glabrescent, calyx lobes distant from each other, triangular, 6–10 × 3–7 mm, apex acute, puberulent and ciliate, sinus between calyx lobes broad and obtuse. Corolla campanulate, or ± tubular-campanulate, 1.5–3 cm, corolla lobes triangular, glabrous. Stamens glabrous; filaments slightly dilated at base, ca. 7 mm; anthers ca. 5 mm. Capsule ca. 1.5 cm in diam. Fl. and fr. Jul–Sep.

* Margins of forests, scrub. W Hubei, N and W Sichuan, SE Xizang (Bomi, Mainling, Nyingchi).

13. Codonopsis tubulosa Komarov, Trudy Imp. S.-Petersburgsk. Bot. Sada, 29: 112. 1908.

管花党参  guan hua dang shen

Codonopsis accrescenticalyx H. Léveillé; C. macrocalyx Diels var. coerulescens Handel-Mazzetti; C. pilosa Chipp.

Roots carrot-shaped, simple or branched below middle, 10–20 × 0.5–2 cm. Stems usually 1–3 m, twining, rarely ascending, 50–75 cm, branched; branches and branchlets both leafy, sterile or with a terminal flower, nearly glabrous or sparsely villous. Leaves opposite or tending to alternate at top of stems and branches; petioles short, 1–7 mm, puberulent; leaf blades ovate to lanceolate, 3–8 × 1–4 cm, base cuneate or obtuse, apex acute or obtuse, margins crenate or nearly entire, adaxially green, hispidulous, abaxially gray-green, sparsely or densely puberulent. Flowers terminal; pedicels 1–6 cm, villous. Calyx tube adnate to ovary up to middle, semiglobose, nearly glabrous to densely villous; calyx lobes broadly ovate, ciliate, adaxially glabrous, abaxially sparsely villous, 10–18 × 5.5–12 mm, apex obtuse, margins crenate or verrucose-toothed. Corolla tubular, 2–3.7 × 0.5–1.6 cm, yellow-green, glabrous, shallowly lobed, corolla lobes deltoid, acute at apex. Filaments dilated at base, ciliate, ca. 10 mm; anthers 3–5 mm. Capsule globose below, conical above. Seeds ovoid, wingless, smooth, brown-yellow. Fl. and fr. Jul–Oct.

Mountain scrub, grasslands; 1900–3000 m. W Guizhou (Nayong, Panxian), S Sichuan, Yunnan (Dali, Lanping, Mengzi) [N Myanmar].

14. Codonopsis kawakamii Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30(1): 165. 1911.

台湾党参  tai wan dang shen

Roots carrot-shaped. Main stems scrambling or twining, fertile, sparsely villous; branches sterile or fertile, with nearly opposite leaves. Leaves on main stems alternate, those on branches nearly opposite; petioles 5–12 mm, hirsute; leaf blades ovate or deltoid-ovate, 7–23 × 5–17 mm, base usually rounded, rarely shallowly slightly cordate, apex acute or obtuse, margins mostly entire or sinuous, or very obscurely crenate, adaxially hispidulous, abaxially hispid. Flowers solitary, terminal on main stems and some branches, slightly pendent; pedicels 1–2.5 cm, sparsely hispid. Calyx tube adnate to ovary up to middle, semiglobose, glabrous or rarely sparsely hairy; calyx lobes ovate-lanceolate, 5–9 × 3–5 mm, nearly entire, sparsely hispid, apex acute; sinus between calyx lobes narrow and acute at anthesis. Corolla broadly campanulate, 1.5–2 cm, greenish yellow or pale blue, nearly glabrous, shallowly lobed. Stamens glabrous; filaments slightly dilated at base, ca. 3 mm; anthers ca. 3 mm. Capsule ca 10–15 × 6–12 mm. Seeds ellipsoid wingless, brown, ca. 1.5 × 1 mm. Fl. Aug–Sep, fr. Oct.

* Grassy slopes; 2500–3100 m. Taiwan (Nantou).

15.Codonopsis benthamii J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 2: 14. 1857.

大萼党参  da e dang shen

Codonopsis macrocalyx Diels; C. macrocalyx var. parviloba J. Anthony.

Roots carrot-shaped or cylindrical, 20–30 × 1.5–2.5 cm. Stems erect, climbing or ascending, up to 2 m, branched, branches and branchlets leafy, sterile or with a terminal flower, sparsely puberulent. Leaves alternate or those on branches nearly opposite; petioles 1–6 cm, sparsely villous; leaf blades broadly ovate, triangular-ovate, ovate or ovate-lanceolate, 3–9 × 1.5–7 cm, base cuneate or rounded, apex acute or obtuse, margin irregularly pinnatifid to shallowly lobed, or dentate, crenate, or rarely nearly entire, adaxially green, sparsely puberulent, abaxially gray-white, sparsely or densely puberulent. Flowers terminal, long pedicellate; pedicels villous. Calyx tube adnate to ovary up to middle; 10-ribbed, glabrous or sparsely villous; calyx lobes ovate, 13–25 × 5–15 mm, glabrous on both surfaces, apex acute or obtuse, margin crenate, ciliate, sinus between calyx lobes narrow and pointed. Corolla tubular, 2–4 × 1.5–2 cm, yellow-green, pale brown-red at base, glabrous, shallowly lobed; lobes triangular, apex acute. Stamens villous; filaments slightly dilated at base, 6–10 mm; anthers 5–6 mm. Capsule 1.2–1.6 cm in diam. Seeds ovoid, wingless, brown-yellow. Fl. and fr. Jul–Oct.

Grassy slopes, forest margins, scrub, by streams; 2800–3700 m. W Sichuan (Baoxing, Muli), SE Xizang (Markam, Yadong, Zayü), NW Yunnan [Bhutan, N India (Sikkim), N Myanmar].

16. Codonopsis pianmaensis S. H. Huang, Acta Bot. Yunnan. 6: 393. 1984.

片马党参  pian ma dang shen

Herbs perennial. Stems decumbent or climbing, 80–120 cm, branched, nearly glabrous. Leaves alternate; petioles 0.8–3.5 cm; leaf blades ovate-deltoid, 2–5.5 × 1.5–4 cm, adaxially sparsely puberulent, abaxially sparsely or densely puberulent, base cuneate or obtuse, apex acuminate, margin irregularly serrate; leaves on branches smaller. Flowers solitary and terminal; pedicels 1–6.5 cm. Calyx adnate to ovary up to middle, puberulent, semiglobose; calyx lobes ovate-deltoid, ca. 12 × 8 mm, apex acute, margin repand, sinus between calyx lobes acute. Corolla tubular-campanulate, ca. 17 × 10 mm, yellow, with purple-blue spots inside, densely puberulent outside; corolla lobes triangular. Filaments dilated at base, densely villous, ca. 7 mm; anthers oblong, ca. 4.5 mm, abaxially densely puberulent. Ovary semi-inferior. Fl. Jul.

* Grassy slopes with shrubs; ca. 3500 m. W Yunnan (Lushui).

17. Codonopsis subsimplex J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 2: 16. 1857.

藏南党参  zang nan dang shen

Roots carrot-shaped, 15–20 × 0.5–1.2 cm. Stems erect or ascending, few-branched, 35–70 cm, nearly glabrous or sparsely white-villous. Leaves on main stems alternate, those on branches nearly opposite; petioles 0.5–1.5 cm, sparsely white-villous; leaf blades broadly ovate, ovate or nearly elliptic, 3–7 × 2–4.5 cm, adaxially hispidulous, abaxially hispidulous but hirsute along veins, base cuneate or rounded, apex acute or slightly obtuse, margin crenate. Calyx tube adnate to whole ovary; calyx lobes triangular-ovate or ovate, 13–16 × 6–8.5 mm, apex acute or obtuse, margin dentate, sparsely ciliate, sinus between calyx lobes narrow and pointed. Corolla broadly campanulate, 1.5–2 × 1.5–2 cm, pale yellow-green or pale blue, glabrous, shallowly lobed; lobes ovate-deltoid. Filaments slightly dilated at base, ca. 2–3 mm; anthers ca. 5 mm. Capsules semiglobose below, short-conical above, ca. 8 mm in diam., rostrum 4–6 mm. Seeds numerous, ellipsoid, wingless, ca. 1.5 mm, yellow. Fl. Jul–Aug, fr. Sep–Oct.

Mountain forests or scrub; ca. 3100 m. S Xizang (Mainling, Yadong) [Bhutan, N India, Nepal].

18. Codonopsis subscaposa Komarov, Trudy Imp. S.-Petersburgsk. Bot. Sada 29: 114. 1908.

Chinese  Pinyin

Roots carrot-shaped, 15–20 × 0.5–1 cm. Stems erect, 40–100 cm, simple or branched below, sparsely pubescent, late glabrescent. Leaves on main stems alternate, those on branches opposite; petioles 2–10 cm, sparsely villous; leaf blades ovate, linear-elliptic or lanceolate, 2–13 × 1.5–5 cm, base cuneate, apex acute or obtuse, margin sparsely dentate or crenate, or nearly entire, adaxially green, sparsely puberulent, abaxially gray-green, glabrous or sparsely villous along veins. Flowers terminal or axillary, often 1–4 on tops of stems and branches, long pedicellate. Calyx tube adnate to ovary up to middle, semiglobose, 10-ribbed, sparsely villous; calyx lobes deltoid, 5–7 × ca. 3 mm, glabrous, apex acute, margin entire or crenulate. Corolla broadly campanulate, 1.5–3 × 2–4 cm, yellow but with red-purple veins or red-purple with yellow spots, 5-fid to near middle, glabrous on both sides or sparsely villous at apex of lobes. Stamens glabrous; filaments slightly dilated at base, ca. 7 mm; anthers ca. 5 mm. Capsule semiglobose below, conical above. Seeds ovoid, wingless, brown-yellow, smooth. Fl. and fr. Jul–Oct.

* Grassy slopes, open woods; 2500–4200 m. W Sichuan, NW Yunnan (Xianggelila).

19. Codonopsis meleagris Diels, Notes Roy. Bot. Gard. Edinburgh 5: 172. 1912.

珠鸡斑党参  zhu ji ban dang shen

Roots carrot-shaped, 15–20 × 0.5–1 cm. Stems erect, simple or with elongate branches in lower part, 40–90 cm, sparsely hairy, late glabrescent. Leaves on main stems alternate, those on branches opposite, sessile or with short petioles less than 1 cm; leaf blades ovate, long elliptic or lanceolate, 5–8 × 1.5–3 cm, base cuneate, apex acute or obtuse, margin crenate or nearly entire, adaxially green, glabrous or sparsely hispidulous, abaxially gray-green, densely hirsute. Flowers terminal or axillary, long pedicellate. Calyx tube adnate to ovary up to middle; semiglobose, 10-ribbed, glabrous or sparsely scaberulose; calyx lobes ovate or ovate-triangular, 8–12 × 4–6 mm, glabrous, apex acute, margins nearly entire or serrate. Corolla broadly campanulate, yellow with red-purple veins, or red-purple with yellow spots, 3–4 × 3–4 cm, glabrous on both side, 5-fid to middle; lobes often sparsely villous at apex. Stamens glabrous; filaments slightly dilated at base, ca. 7 mm; anthers ca. 5 mm. Capsules semiglobose below, conical above, ca. 10 ×8 mm. Seeds numerous, ovoid, wingless, brown-yellow. Fl. and fr. Jul–Oct.

* Grassy slopes, open forests; 3000–4000 m. NW Yunnan (Lijiang).

20. Codonopsis purpurea Wallich in Roxburgh, Fl. Ind. 2: 105. 1824.

紫花党参  zi hua dang shen

Campanula purpurea (Wallich) Sprengel; Wahlenbergia purpurea (Wallich) A. Candolle.

Plants glabrous throughout. Roots fusiform or carrot-shaped. Stems or nearly climbing, few-branched, 30–50 cm. Leaves opposite or those at top sometimes alternate; petioles 1–7 mm; leaf blades long elliptic, ovate or ovate-lanceolate, 4–9 × 2–3.5 cm, base rounded or cuneate, apex acute or obtuse, margin nearly entire or sinuous. Flowers terminal or opposite leaves at top. Calyx tube adnate to whole ovary, semiglobose; calyx lobes triangular-ovate, 10–20 × 6–9 mm, entire, apex acute. Corolla broadly campanulate or funnelform-campanulate, 2–3 × 3–4 cm, dark red-purple, 5-fid to middle; lobes triangular. Filaments slightly dilated, ca. 6 mm; anthers ca. 5 mm. Capsules semiglobose below, conical above, 1.5–2 cm in diam.; valves 5–8 mm. Seeds numerous, oblong, narrow-winged, ca. 2 × 1 mm, yellow. Fl. and fr. Sep–Oct.

Grassy slopes, scrub, epiphytic on trunks in forests; 2000–3300 m. S Xizang (Nyalam), W Yunnan (Jingdong) [N India, Nepal].

21. Codonopsis clematidea (Schrenk) C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 433. 1881.

新疆党参  xinjiang dang shen

Wahlenbergia clematidea Schrenk, Enum. Pl. Nov. 1: 38. 1841; Codonopsis ovata Bentham var. cuspidata Chipp; C. ovata var. obtusa Chipp.

Roots carrot-shaped, 25–45 × 1–3 cm. Stems single or caespitose, erect or ascending, or slightly procumbent, 50–100 cm tall, branches slender, erect or slightly spreading, mostly at lower part but those at upper part usually fertile, hispidulous when young. Leaves on main stems smaller and alternate, those on branches opposite; petioles up to 2.5 cm, sparsely hispidulous; leaf blades ovate, ovate-oblong, broadly lanceolate or lanceolate, 1–2.8(–5.2) × 0.8–1.5(–3.2) cm, adaxially sparsely or densely hispidulous, abaxially densely hispidulous, base slightly cordate or rounded, apex acute, margin entire. Flowers solitary, terminal on main stems and branches, long pedicellate; pedicels sparsely white hispidulous. Calyx tube adnate to ovary up to middle, semiglobose, 10-ribbed, glabrous or rarely sparsely white-scaberulose; calyx lobes ovate, elliptic or ovate-lanceolate, apex acute, 15–20 × 6–8 mm, entire, glabrous or apex sparsely puberulent; sinus between calyx lobes narrow and pointed. Corolla broadly campanulate, 2.8–2.6 cm, pale blue with dark blue veins and purple spots inside, glabrous. Stamens glabrous; filaments slightly dilated at base, 5–6 mm; anthers oblong, 5–6 mm