ASTERACEAE Tribe HELIANTHEAE [Draft]

向日葵族  xiang ri kui zu

Chen Yousheng (陈又生)[1]; D. J. Nicholas Hind[2]

Annuals, perennials, subshrubs. Leaves mostly opposite or mostly alternate, rarely whorled; petiolate or sessile. Heads usually radiate, sometimes discoid. Involucres cylindric to hemispheric or rotate. Phyllaries persistent, herbaceous, rarely membranous, in 1–6 series. Receptacles usually flat or convex, sometimes hemispheric or conic, usually paleate; paleae usually falling, sometimes persistent, distinct or rarely connate, herbaceous to scarious, apices sometimes 3-lobed or -toothed. Ray florets neuter, or female and fertile; ligule entire or 2–3-toothed. Disc florets bisexual, fertile or sterile; 4–5-lobed; anther thecae dark or pale; anther tails obtuse and entire, or sagittate, auriculate. Cypselae often compressed or angled; pappi 0, or usually of (1–)2(–8) scales and/or awns, sometimes coroniform.

About 209 genera, mostly in America; 30 genera in China, most are introduced (one endemic).

[Ed. note: Genera and species that are only cultivated in China, except for large-scale crops (and of course naturalized taxa), should have their treatments deleted, in accordance with the FOC Guidelines. However, these purely cultivated species should be mentioned in a comment here at the start of the tribe.]

1a.       Plant generally wind-pollinated or self-pollinated, heads small and not showy; all flowers imperfect; ray florets absent.

2a.       Phyllaries in staminate heads in 1–2 series, distinct to bases; phyllaries in pistillate heads in 6–12 series, their distinct tips mostly ± hooked (the distal 1–3 usually longer, stouter, and not hooked), the whole becoming a hard, prickly perigynium; florets 2 ..............................................  1. Xanthium

2b.       Phyllaries in staminate heads in 1 series, connate; phyllaries in pistillate heads in 1–8 series, usually with free tips forming tubercles, spines, or wings; florets 1(–5) .............................  2. Ambrosia

1b.       Plant generally adapted for attracting pollinating insects, heads colorful and attractive; some or all flowers perfect; ray florets present.

3a.       Only ray florets fertile, these achenes much longer than those of the sterile disk florets.

4a.       Ray achenes thick, rounded or weakly compressed, without wings; achenes thick, not much flattened; pappus none.

5a.       Achenes embraced and enclosed by the prickly inner bracts of the involucre
..................................................................................................  3. Acanthospermum

5b.       Achenes merely subtended by the unarmed bracts.

6a.       Phyllaries in several series, the inner completely clasping the achenes
                                                                                                       4. Melampodium

6b.       Phyllaries in 1–2 series, subtending, not clasping the achenes .............  5. Smallanthus

4b.       Ray achenes strongly flattened; pappus present or absent.

7a.       Leaves opposite; ray achenes in 2–3 series, even at flowering time, each achene completely free from the nearby bracts and falling separately .....................................................  6. Silphium

7b.       Leaves alternate; ray achenes in a single series, each achene more or less enclosed by and often attached to the subtending involucral bract and 2–3 adjacent receptacular bracts, all falling as a unit ...........................................................................................................  7. Parthenium

3b.       Disc florets fertile; ray florets present and fertile or sterile or absent.

8a.       Ray florets with or without short tubes, persistent.

9a.       Receptacles conic; achenes 3-angled or flattened; phyllaries in 3–4 series
....................................................................................................................  8. Zinnia

9b.       Receptacles convex to conic; achenes terete or obscurely 3–4-angled to compressed or flattened; phyllaries in 2–3 series ...........................................................................  9. Sanvitalia

8b.       Ray florets caducous.

10a.     Pappus absent, or awned.

11a.     Achenes all plumpy, or 3-angled in ray florets and compressed in disc florets.

12a.     Achenes enclosed by inner phyllaries or outer paleae; pappus absent or scale-like.

13a.     Female florets in one row; phyllaries 5, spreading; heads in sparse panicles; peduncles long       10. Sigesbeckia

13b.     Female florets in many rows; phyllaries 4, broad, in two pairs; heads axillary, peduncles very short ..........................................................................................................  11. Enydra

12b.     Achenes not enclosed by inner phyllaries.

14a.     Paleae narrowly long, even; ray florets in two series, ligules small; pappus absent or with two short awns.

15a.     Ligules white ...................................................................................  12. Eclipta

15b.     Ligules yellow ...............................................................................  13..Guizotia

14b.     Paleae concave or folio, more or less enclose florets.

16a.     Achenes in bisexual florets 4–5 angled, or compressed.

17a.     Pappus absent or coroniform, or of 2–8 unequal scales; receptacles subspheric to ovoid, or conic to columnar; leaves usually alternate
................................................................................................  14. Rudbeckia

17b.     Pappus scale-like, spricky, awnlike or absent; receptacles even or raised; leaves usually opposite.

18a.     Ray florets fertile.

19a.     Pappus 2–5, unequal, spine like or squama like, base connate; ray florets female, ligules short or very short, apex 2–4-dentate; heads small ..............  15. Blainvillea

19b.     Pappus absent, scale-like, cyathiform, coroniform or 1–2 bristles; heads larger  16. Wedelia

18b.     Ray florets sterile.

20a.     Receptacles paleate, each palea completely investing and falling with a cypsela, each forming a hardened perigynium ........................................  17. Sclerocarpus

20b.     Receptacle paleate, paleae sometimes conduplicate, ± enfolding cypselae, not forming perigynia.

21a.     Pappi 0, or coroniform, of connate scales, 1–2 scales sometimes subulate to aristate; peduncles usually distally dilated, fistulose
                                                                                              18. Tithonia

21b.     Pappi 0 or readily falling, of 2(–3) usually lanceolate, aristate, or erose scales plus 0–8 usually shorter scales ........................................................  19. Helianthus

16b.     Achenes in ray florets broadly ovate or elliptic, 3-angled; disc achenes ellipsoid, strongly compressed; pappus absent or of up to 10 awnlike bristles .............  20. Acmella

11b.     Achenes compressed.

22a.     Pappus 0, or persistent, of 2 bristly cusps or scales; leaves opposite.

23a.     Phyllaries free, outer ones coriaceous, subequal, inner ones shorter, similar with paleae; ray achenes narrowly oval and winged, with 2 triangular scales

24a.     Achenes markedly dimorphic, those of the rays with conspicuous, lacerate wing margins, others wingless .....................................................................................  21. Synedrella

24b.     Achenes all about alike, wingless or somewhat thick-winged distally, not lacerate  22. Calyptocarpus

23b.     Phyllaries in two row, outer ones few and smaller, inner ones membranous, connate at base; pappus of 2–4 awns or scales or absent.

25a.     Slender annual; ligules ca. 11 Χ 8 mm ..........................................  23. Coreopsis

25b.     Robust perennial or subshrub; ligules 35–60 Χ 15–25 mm .................  24. Dahlia

22b.     Pappus of retrorsely barbed awns; leaves opposite or upper ones alternate.

26a.     Pappus of 2–4 scabrid awns; style branches with short minute papillae.

27a.     Achenes apex beaked; ray florets red or purple ..............................  25. Cosmos

27b.     Achenes apex narrow, not beaked; ray florets yellow, white or absent
                                                                                                           26. Bidens

26b.     Pappus of 2 scabrid awns; style branches with long hairs ..........  27. Glossocardia

10b.     Pappus of subulate to acerose scales, or spatulate, entire to erose, fimbriate, or laciniate, sometimes aristate, scales in 1 series, or plumose, setiform scales (or flattened bristles) in 1 series.

28a.     Annuals; ray cypselae often each shed together with subtending phyllary and 2 adjacent paleae      28. Galinsoga

28b.     Perennials; cypselae shed separate from paleae ........................................  29. Tridax

1. XANTHIUM Linnaeus, Sp. Pl. 987. 1753.

苍耳属  cang er shu

Annuals. Stems erect, branched. Leaves cauline; mostly alternate (proximal 2–6 sometimes opposite); petiolate; blades lanceolate, linear, ovate, rounded-deltate, or suborbiculate, often palmately or pinnately lobed, ultimate margins entire or ± toothed, faces hirtellous or ± strigose, usually gland-dotted as well. Heads discoid, either pistillate (proximal) or functionally staminate (distal), in racemiform to spiciform arrays or borne singly (in axils). Pistillate heads: involucres ± ellipsoid, 2–5 mm diam. at anthesis; phyllaries in 6–12 series, outer 5–8 distinct, the rest (sometimes interpreted as paleae) proximally connate, their distinct tips mostly ± hooked (the distal 1–3 usually longer, stouter, and not hooked), the whole becoming a hard, prickly perigynium (a bur); florets 2, corollas 0. Staminate heads: involucres saucer-shaped, 3–5 mm diam.; phyllaries in 1–2 series, distinct to bases; receptacles conic to columnar; paleae spatulate to cuneiform or linear, membranous, distally villous or hirtellous; florets 20–50, corollas whitish, funnelform, lobes 5, erect or reflexed (filaments connate, anthers distinct or weakly coherent). Cypselae (black) fusiform, enclosed in obovoid to ellipsoid, hard, prickly, 2-chambered burs. Pappus absent. x = 18.

About 2–3 species, New World, introduced nearly worldwide; two species introduced in China.

1a.       Nodal spines 0; leaf blades suborbiculate to pentagonal or deltate ...........  1. X. strumarium

1b.       Nodal spines (1–)3-lobed, 15–30+ mm; leaf blades ± lanceolate to ovate or lance-linear  2. X. spinosum

1. Xanthium spinosum Linnaeus, Sp. Pl. 987. 1753.

刺苍耳  ci cang er

Xanthium ambrosioides Hooker & Arnott; X. spinosum var. inerme Bel.

Annual herbs, 10–60(–120) cm; nodal spines usually in pairs, simple or 2–3-partite, 15–30 mm. Leaves: petioles 1–15(–25) mm; blades ± ovate to lanceolate or lance-linear, 4–8(–12) Χ 1–3(–5) cm, often pinnately 3(–7)-lobed, abaxial faces gray to white, densely strigose. Burs 10–12(–15) mm. Fl. Jul–Oct. 2n = 36.

Damp or seasonally wet, alkaline soils, waste places, margins of agriculture. Beijing, Henan [native to North America and South America].

2. Xanthium strumarium Linnaeus, Sp. Pl. 987. 1753.

苍耳  cang er

Xanthium americanum Walter; X. chasei Fernald; X. chinense Miller; X. curvescens Millspaugh & Sherff; X. cylindricum Millspaugh & Fernald; X. echinatum Murray; X. echinellum Greene ex Rydberg; X. globosum C. Schull; X. inaequilaterum Candolle; X. inflexum Mackenzie & Bush; X. italicum Moretti; X. japonicum Widder; X. mongolicum Kitagawa; X. orientale Linnaeus; X. oviforme Wallroth; X. pensylvanicum Wallroth; X. sibiricum Patrin ex Widder; X. speciosum Kearney, X. strumarium Linnaeus subsp. sibiricum (Widder) Greuter; X. sibiricum Patrin ex Widder var. jingyuanense H. G. Hou & Y. T. Lu; X. sibiricum Patrin ex Widder var. subinerme (C. Winkler) Widder; X. strumarium var. canadense (Miller) Torrey & A. Gray, X. strumarium var. glabratum (Candolle) Cronquist; X. strumarium var. japonica (Widder) Hara; X. varians Greene; X. wootonii Cockerell.

Annuals, 20–120 cm; nodal spines 0. Median cauline leaves chartaceous, ovate-deltoid, 9–25 cm long, apex acute, base shallowly cordate to broadly cuneate, margins irregularly dentate, often obsoletely 3-lobed, densely scabrous on surfaces; petiole 3.5–10 cm long, not winged. Heads monoecious. Staminate heads in terminal umbels, involucral bracts in 1 series, oblong-lanceolate, 2.2 mm long; outer paleae oblong-lanceolate, inner paleae lanceolate, 2.2 mm long; corolla white, tubular, 2.5 mm long, 5-dentate. Pistillate heads axillary, involucral bracts in 1 series, oblong-lanceolate, 3 mm long, inner bracts connate with outer paleae. Utricles sessile, oblong, ellipsoid or ovoid, 10–18 Χ 6–12 mm, densely puberulent, 2-beaked, burs 10–30 mm. Fl. Jul–Aug, fr. Sep–Oct. 2n = 36.

Damp or seasonally wet, often alkaline, soils, waste places, margins of agriculture, common weeds in China; Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hebei, Henan, Heilongjiang, Hubei, Hunan, Jilin, Liaoning, Jiangsu, Jiangxi, Nei Mongol, Ningxia, Qinghai, Shandong, Shanxi, Shaanxi, Sichuan, Taiwan, Xizang, Xinjiang, Yunnan, Zhejiang [India, Iran, Japan, Korea, Mongolia, Russia; North America].

2. AMBROSIA Linnaeus, Sp. Pl. 987. 1753; Gen. Pl. ed. 5, 425. 1754.

豚草属  tun cao shu

Annuals, perennials, or shrubs. Stems erect, decumbent, or prostrate, branched. Leaves usually cauline; opposite throughout or opposite (proximal) and alternate or mostly alternate; sessile or petiolate; blades deltate, elliptic, filiform, lanceolate, linear, obovate, ovate, or rhombic, usually pinnately, sometimes palmately lobed, ultimate margins entire or toothed, faces hairy or glabrate, usually gland-dotted or stipitate-glandular. Heads discoid (unisexual, pistillate proximal to or intermixed with staminates, staminates usually in racemiform to spiciform arrays; rarely, single plants all or mostly staminate or pistillate). Pistillate heads: phyllaries in 1–8 series, outer (1–)5–8 distinct or connate, herbaceous, the rest connate, usually with free tips forming tubercles, spines, or wings; florets 1(–5), corollas 0. Staminate heads: involucres cup-shaped to saucer-shaped, 1.5–6 mm diam.; phyllaries in 1 series, connate; receptacles flat or convex; paleae spatulate to linear, membranous, sometimes villous, hirtellous, and/or gland-dotted or stipitate-glandular, sometimes none; florets 5–60; corollas whitish or purplish, funnelform, lobes 5, erect or incurved; staminal filaments connate, anthers distinct or weakly coherent. Cypselae (black) ovoid or fusiform, enclosed within globose to obovoid, pyramidal, pyriform, obconic, or fusiform, hard, smooth, tuberculate, spiny, or winged "burs"; pappi 0. x = 18.

About species, tropical to subtropical and temperate New World, mostly North America, two species introduced in China.

1a.       Leaves one or more commonly twice pinnatifid .....................................  1. A. artemisiifolia

1b.       Leaves palmately 3- to 5-lobed, sometimes undivided .....................................  2. A. trifida

1. Ambrosia artemisiifolia Linnaeus, Sp. Pl. 988. 1753.

豚草  tun cao

Ambrosia artemisiifolia var. elatior (Linnaeus) Descourtilz; A. elatior Linnaeus.

Annuals, 20–150 cm. Stems erect. Leaves opposite and alternate; petioles 25–35(–60) mm; blades deltate to lanceolate or elliptic, 25–55(–90) Χ 20–30(–50) mm, 1–2-pinnately lobed, bases cuneate, ultimate margins entire or toothed, abaxial faces sparsely pilosulous to strigillose, adaxial faces strigillose, both gland-dotted. Pistillate heads clustered, proximal to staminates; florets 1. Staminate heads: peduncles 0.5–1.5 mm; involucres shallowly cup-shaped (usually without black nerves), 2–3 mm diam., glabrous or hispid to pilosulous; florets 12–20. Burs: bodies ± globose to pyriform, 2–3 mm, ± pilosulous, spines or tubercles 3–5, near middles or distal, conic to acerose, 0.1–0.5 mm, tips straight. Fl. Jul–Oct, fr. Sep–Oct. 2n = 34, 36.

Introduced obnoxious weeds of wet to dry soils; below 1000 m. widely distributed in China [native to North and Central America; widely distributed as an introduction in Asia and Europe].

2. Ambrosia trifida Linnaeus, Sp. Pl. 987. 1753.

三裂叶豚草  san ye tun cao

Annuals, 30–150 cm. Stems erect. Leaves mostly opposite; petioles 10–30(–70) mm; blades rounded-deltate to ovate or elliptic, 40–150(–250) Χ 30–70(–200) mm, usually some blades palmately 3(–5)-lobed, bases truncate to cuneate, sometimes decurrent onto petioles, margins usually toothed, rarely entire, abaxial and adaxial faces scabrellous and gland-dotted. Pistillate heads clustered, proximal to staminates; florets 1. Staminate heads: peduncles 1–3 mm; involucres saucer-shaped, 2–4 mm diam., scabrellous; florets 3–25. Burs: bodies pyramidal, 3–5(–7) mm, glabrous or glabrate, spines 4–5, distal, acerose, 0.5–1 mm, tips straight. Fl. Jul–Aug, fr. Sep–Nov. 2n = 24, 48.

Disturbed sites, waste places, damp soils; up to 1600 m [native to North America].

3. ACANTHOSPERMUM Schrank, Pl. Rar. Hort. Monac. 2: plate 53. 1820.

刺苞果属  chi bao guo shu

Annuals. Leaves cauline; opposite; petiolate or sessile; blades mostly elliptic to deltate, rhombic, or ovate, sometimes lyrate, ultimate margins entire or toothed, faces usually pilosulous to sericeous or scabrellous, sometimes glabrate or glabrescent, usually gland-dotted. Heads radiate, 1(–3) in "forks" of branches. Involucres hemispheric. Phyllaries persistent (outer) or falling, 10–13 in 2 series. Ray florets 5–8, pistillate, fertile; corollas yellowish (tubes shorter than to equaling laminae, laminae ovate to elliptic or linear). Disc florets 3–8(–12), functionally staminate; corollas yellowish, tubes shorter than funnelform or campanulate throats, lobes 5, deltate. Cypselae each enclosed within and shed with an often hardened, prickly perigynium (the ultimate "fruits" plumply ellipsoid to fusiform, or ± compressed); pappi 0 or rudimentary. x = 11.

About six species, mostly tropical to warm-temperate New World; one species introduced in China.

1. Acanthospermum australe (Loefling) O. Kuntze, Revis. Gen. Pl. 1: 303. 1891.

刺苞果  chi bao guo

Melampodium australe Loefling, Iter Hispan., 268. 1758.

Annuals, 10–60(–120) cm high. Stems erect, ascending or procumbent. Leaf blades deltate to rhombic or ovate, 13–37 Χ 7–32 mm, faces sparsely scabrellous to glabrate or glabrescent, gland-dotted. Fruits plumply ellipsoid to fusiform, weakly compressed, 7–9+ mm, 5–7-ribbed, lacking terminal spines, prickles uncinate, mostly along ribs. Fl. Jun–Jul, fr. Aug–Sep. 2n = 22.

Streamside, plains; 300–1900 m. Naturized inYunnan [native to South America].

4. MELAMPODIUM Linnaeus, Sp. Pl. 2: 921. 1753.

皇帝菊属  huang di ju shu

Annuals, perennials, or subshrubs. Stems erect or prostrate. Leaves cauline; opposite; petiolate or sessile; blades deltate, lance-elliptic, lanceolate, lance-linear, linear, linear-oblong, ovate, or rhombic, sometimes pinnately lobed, ultimate margins entire or toothed, faces usually hairy, usually gland-dotted (at least abaxial). Heads radiate, borne singly (from forks of branches, peduncles often uncinate). Involucres mostly hemispheric. Phyllaries persistent (outer) or falling with cypselae, 8–20+ in 2 series; outer ones often connate, herbaceous; inner each investing a ray ovary, forming a perigynium, shed with enclosed cypsela. Receptacles flat or convex to conic; paleae lanceolate to linear, scarious, conduplicate, often with dilated, erose tips. Ray florets pistillate, fertile; corollas yellow, orange, or cream-white, sometimes purplish abaxially (tubes often wanting). Disc florets functionally staminate; corollas ochroleucous, yellow, or orange, tubes shorter than funnelform throats, lobes 5, ± deltate. Cypselae each enclosed within and shed with a smooth, sculpted, or tuberculate, scarious, coriaceous, or hardened perigynium. Pappus absent. x = 12.

About 36 species: United States, Mexico, West Indies, Central America, South America; one species introduced in China.

1. Melampodium divaricatum (Richard) Candolle, Prodr. 5: 520. 1836.

皇帝菊  huang di ju

Dysodium divaricatum Richard in C. H. Persoon, Syn. Pl. 2: 489. 1807; Melampodium paludosum Kunth, Nov. Gen. Sp. 4: 273. 1820.

Annuals, 15–100 cm high. Leaf blades rhombic to lanceolate, 40–150 Χ 25–95 mm, lengths –2(–3) times widths, margins coarsely toothed or entire. Peduncles 1–12 cm. Outer phyllaries 5, connate 1/4–1/3 their lengths, ovate to orbiculate, 3.5–6 mm. Ray florets 8–13; corollas yellow-orange, laminae oblong-elliptic, 3.5–7 Χ 1.6–3 mm. Disc florets 40–70, yellow-orange. Achenes 2.8–4 mm long. Fl. Jun–Oct. 2n = 24.

Widely cultivated in China (Beijing, Guangdong, Guangxi, Taiwan) [native to Mexico, West Indies (Antilles), Central America, South America].

5. SMALLANTHUS Mackenzie ex Small, Man. S. E. Fl. 1406, 1509. 1933.

包果菊属  bao guo ju shu

Perennials, annuals or shrubs, 100–300(–1200) cm. Stems erect. Leaves cauline; opposite; petiolate (petioles usually winged), sessile; blades mostly deltate to ovate, usually palmately lobed, ultimate margins dentate to denticulate, faces hirtellous, pilosulous, or puberulent, gland-dotted (at least abaxially). Heads radiate, borne singly or 2–5 in crowded, corymbiform arrays. Involucres hemispheric, 8–15 mm diam. Phyllaries persistent, 12–13(–25) in 2 series, herbaceous, inner as many as rays, more membranous to scarious, narrower and shorter). Receptacles flat to convex, paleate (paleae obovate to spatulate, scarious). Ray florets 7–13(–25+), pistillate, fertile; corollas yellow, white or orange; tubes hairy, laminae linear to elliptic [ovate]. Disc florets (20–)40–80(–150), functionally staminate; corollas yellow or orange, tubes shorter than abruptly campanulate [funnelform] throats, lobes 5, deltate. Cypselae (obliquely inserted on receptacles, each shed separate from subtending phyllary) obovoid [quadrangular], somewhat compressed, finely 30–40-ribbed or -striate (not narrowed at bases, not apically beaked); pappi 0 (cypselae sometimes hairy at apices). x = 16.

About 23 species; United States, Mexico, South America. Two species introduced in China.

1a.       Leaf blade usually lobed; underground stems without tubers ..........................  1. S. uvedalia

1b.       Leaf blade unlobed; underground stems tuberous ....................................  2. S. sonchifolius

1. Smallanthus uvedalia (Linnaeus) Mackenzie ex Small, Man. S. E. Fl. 1509. 1933.

包果菊  bao guo ju

Osteospermum uvedalia Linnaeus, Sp. Pl. 2: 923. 1753; Polymnia uvedalia (Linnaeus) Linnaeus.

Perennial herbs, 1–3 m tall. Stems erect and hollow, purple-spotted. Leaves opposite; petioles 3–12 cm, blades ovate to deltoid, 10–35(–60) Χ 10–35 cm, usually palmately 3–5-lobed, sessile or with broad conspicuous wings to base of petiole. Heads clusteed in loose leafy cymes; phyllaries 4–6, 10–20 mm long, 10–12 mm wide, ovate to ovate-lanceolate; ray florets 7–13, pistillate, fertile, ligules yellow, 12–30 mm. Achenes 5–6 mm long, ca. 4 mm wide; pappus absent. Fl. Jun–Aug(–Oct). 2n = 32.

Thickets, fields, naturized in Anhui, Jiangsu [native to North America and Central America].

Yellow-flowered leaf-cup is primarily a weed of pastures, hay fields, fencerows, and roadsides in North America.

2. Smallanthus sonchifolius (Poeppig & Endlicher) H. Robinson, Phytologia 39(1): 51. 1978.

菊薯  ju shu

Polymnia sonchifolia Poeppig & Endlicher, Nov. Gen. Sp. Pl. 3: 47. 1843.

Perennial herbs, 1–3 m tall. Stems cylindrical and hollow; underground part very irregular, very branched mass; numerous roots grow from this part as well as above-ground shoots, some of the roots increase from the central part already when they reach as short as 10 mm length and continue to be longer and wider until they create a spindle-shaped tuber 100–200 mm long and 30–80 mm in diam. Lower leaves broadly ovate and hastate or subhastate, connate and auriculate et the base; upper leaves ovate-lanceolate, without lobes and hastate base; upper and lower surfaces densely pubescent. Infloroscence terminal, composed of 1–5 axes, each one whith 3 heads; penduncles densely pilose; phyllaris 5, uniseriate and ovate. Flowers yellow to bright orange; ray flowers 2- or 3-toothed, depending on the clone, to 12 Χ 7 mm, pistillate; disc flowers about 7 mm long, staminate. Immature cypselas purple, and turn dark brown or black at maturity. Fl. Jun–Sep.

Cultivated in Fujian, Guizhou, Hebei, Hubei, Hunan, Hainan, Shandong, Taiwan, Yunnan, Zhejiang [native to Bolivia, Colombia and Ecuador].

Named as yacon, it is an ancient crop of the South American Andes with tuberous roots that can be eaten raw or cooked.

6. SILPHIUM Linnaeus, Sp. Pl. 919. 1753.

松香草属  song xiang cao shu

Perennials, 20–250+ cm. Stems usually erect, usually branched. Leaves basal and cauline, whorled, opposite, subopposite, or alternate; petiolate or sessile; blades deltate, elliptic, linear, ovate, or rhombic, sometimes 1–2-pinnately lobed or -pinnatifid, bases cordate or truncate to cuneate, margins entire or toothed, faces glabrous or hairy (sometimes stipitate-glandular). Heads radiate, in paniculiform or racemiform arrays. Involucres campanulate to hemispheric, 10–30 mm diam. Phyllaries persistent, 11–45 in 2–4 series. Receptacles flat to slightly convex, paleate. Ray florets 8–35+ in 1–4 series, pistillate, fertile; corollas yellow or white. Disc florets 20–200+, functionally staminate; corollas yellow or white, tubes much shorter than narrow, cylindric throats, lobes 5, deltate. Cypselae obflattened. Pappus absent, or persistent, of 2 awns. x = 7.

About 12 species: North America; one species introduced in China.

1. Silphium perfoliatum Linnaeus, Syst. Nat. ed. 10. 2: 1232. 1759.

串叶松香草  chuan ye song xiang cao

Plants caulescent, 75–300 cm; fibrous rooted. Stems square, glabrous, hispid, or scabrous. Leaves: basal caducous; cauline usually opposite, rarely whorled (in 3’s), petiolate or sessile; blades deltate, lanceolate, or ovate, 2–41 Χ 0.5–24 cm, bases attenuate or truncate (distal connate-perfoliate), margins entire, dentate, or bidentate, apices acuminate to acute, faces scabrous to hispid. Phyllaries 25–37 in 2–3 series, outer appressed, apices acute to acuminate, abaxial faces scabrous or hispid. Ray florets 17–35; corollas yellow. Disc florets 85–150 (–200); corollas yellow. Cypselae 8–12 Χ 5–9 mm; pappus 0.5–1.5 mm. Fl. Jun–Sep, Fr. Sep–Oct.

Cultivated in Beijing, Hebei, Liaoning, Sichuan [native to Eastern North America].

7. PARTHENIUM Linnaeus, Sp. Pl. 988. 1753; Gen. Pl. ed. 5, 426. 1754.

银胶菊属  yin jiao ju shu

Annuals, biennials, perennials, subshrubs, or shrubs. Stems erect, usually branched. Leaves usually cauline, sometimes in rosettes; alternate; petiolate or sessile; blades elliptic, lanceolate, linear, lyrate, oblanceolate, obovate, ovate, rounded-deltate, spatulate, sometimes (1–)2-pinnately lobed, ultimate margins entire or toothed, faces usually hairy and gland-dotted (at least the abaxial). Heads usually radiate, sometimes ± disciform. Involucres hemispheric. Phyllaries falling, in 2 series, outer 5(–8) herbaceous to scarious, inner 5–8 scarious to membranous. Receptacles flat to conic; paleae cuneate to flabelliform, scarious or membranous, distally papillate and/or fimbrillate, all or the peripheral each ± enfolding a disc floret. Ray (pistillate) florets 5(–8), fertile; corollas ochroleucous, tubes stout, glandular. Disc florets functionally staminate; corollas ochroleucous, funnelform, lobes 5. Cypselae oblanceoloid, obovoid, or pyriform, often obcompressed. Pappus 0. x = 9.

About 16 species: North America to South America, West Indies; two species introduced in China.

1a.       Leaves usually 2-pinnately lobed; heads 3–4 mm diameter; bisexual florets corolla 4-lobed, stamens 4     1. P. hysterophorus

1b.       Leaves entire or sharply toothed; heads ca. 6 mm diameter; bisexual florets corolla 5-lobed, stamens 5    2. P. argentatum

1. Parthenium hysterophorus Linnaeus, Sp. Pl. 988. 1753.

银胶菊  yin jiao ju

Annuals, 30–120 cm high. Leaf blades ovate to elliptic, 30–180 Χ 10–50(–90) mm, (1–)2-pinnately lobed, ultimate lobes lanceolate to linear, 3–50 Χ 2–15 mm, faces sparsely to densely scabrellous and gland-dotted. Heads obscurely radiate, borne in open, paniculiform arrays. Peduncles 1–8(–15+) mm. Phyllaries: outer 5(–6), lance-elliptic, 2–4 mm, inner 5(–6) ovate to orbiculate, 2.5–4 mm. Pistillate florets 5(–6); corolla laminae reniform or orbiculate to oblong, 0.3–1 mm. Disc florets 12–30(–60). Cypselae obovoid, 1.5–2(–3.5) mm; pappus-like enations erect, deltate to ovate, 0.5–1 mm. Fl. Apr–Aug. 2n = 34.

Fields, roadsides; up to 1500 m. Guangdong, Guangxi, Guizhou, Yunnan [Vietnam; tropical America].

2. Parthenium argentatum A. Gray in W. H. Emory, Rep. U. S. Mex. Bound. 2(1): 86. 1859.

灰白银胶菊  hui bai jin jiao ju

Shrubs, 30–100 cm. Leaf blades lanceolate to oblanceolate, 15–25(–40+) Χ 6–15(–25) mm, margins mostly entire, some with 1–2(–5) sharp teeth, faces densely strigillose (gray to white) and obscurely or not at all gland-dotted. Heads radiate, in glomerules of 3–5+ at ends of ± ebracteate stalks 8–15(–20) cm. Peduncles 1–2(–6) mm. Phyllaries: outer 5 oval-elliptic, 2.5–3 mm, inner 5 orbiculate, 3.5–4 mm. Pistillate florets 5; corolla laminae ovate, 1.2–1.5 mm. Disc florets 20–30. Cypselae ± obovoid, 2.5–3 mm; pappus-like enations 2(–4), erect to spreading, subulate, 0.3–0.8 mm. Fl. Apr–Aug. 2n = 36.

Cultivated in S China [native to Mexico and the United States].

Guayule has sometimes been used for commercial production of natural rubber.

8. ZINNIA Linnaeus, Syst. Nat. ed. 10. 2: 1189, 1221, 1377. 1759.

百日菊属  bai ri ju shu

Annual or perennial herbs or subshrubs. Stems prostrate or erect. Leaves cauline; opposite or subopposite; sessile or shortly petiolate; blades acerose, elliptic, lance-linear, lanceolate, linear, oblong, or ovate, bases rounded to cuneate, sheathing the stem, margins entire, faces hairy, usually gland-dotted. Heads usually radiate, borne singly. Involucres campanulate, cylindric, to hemispheric or broader. Phyllaries persistent, in 3–4 series. Receptacles conic, paleate. Ray florets pistillate, fertile; corollas yellow, orange, red, maroon, purple, or white. Disc florets bisexual, fertile; corollas usually yellow to reddish, sometimes purple-tinged, tubes much shorter than cylindric throats, lobes 5, lance-ovate. Cypselae 3-angled or flattened (disc; not winged); pappi 0, or persistent, of 1–3(–4) awns or toothlike scales.

About 17 species: United States, Mexico, Central America, South America; three species introduced in China.

1a.       Heads 5–6 cm diameter ................................................................................  1. Z. elegans

1b.       Heads less than 3 cm diameter.

2a.       Heads 2.5–3 m diameter; scarlet red or maroon, sometimes yellow; paleae apex rounded, margin fimbriacte ...............................................................................................  2. Z. peruviana

2b.       Heads 1–2 cm diameter; florets orange; paleae entire ..............................  3. Z. haageana

1. Zinnia elegans Jacquin, Coll. Bot. 3: 152. 1789.

百日菊  bai ri ju

Annuals, to 100(–200) cm. Stems greenish, becoming yellowish to purplish, unbranched or sparingly branched distal to bases, hirsute to strigose or scabrous. Leaf blades 3–5-nerved, ovate to oblong, mostly 60–100 Χ 20–60 mm, scabrellous to glabrate. Peduncles to 85 mm. Involucres ± hemispheric or broader, 10–15 Χ 5–25 mm. Phyllaries obovate, becoming scarious, glabrous or sparsely hairy, apices rounded, erose or fimbriate. Paleae red to purple, apices rounded to acute, fimbriate. Ray florets 8–21 (more in "double" cultivars); corollas usually red (white, yellow, or purple in cultivars), laminae spatulate to obovate, 10–35 mm. Disc florets 100–150+; corollas yellow, 7–9 mm, lobes 1–2.5 mm. Cypselae 6–10 mm, 3-angled (ray) or ± compressed (disc), not or faintly ribbed, ciliolate; pappi 0. Fl. Jun–Sep, fr. Jun–Oct. 2n = 24.

Introduced and widely cultivated in China [native to North America].

2. Zinnia peruviana (Linnaeus) Linnaeus, Syst. Nat. ed. 10. 2: 1221. 1759.

多花百日菊  duo hua bai ri ju

Chrysogonum peruvianum Linnaeus, Sp. Pl. 920. 1753; Zinnia multiflora Linnaeus; Z. pauciflora Linnaeus.

Annuals, mostly 30–50(–100) cm. Stems greenish, becoming purplish or yellowish, unbranched or sparingly branched distal to bases, strigose. Leaf blades 3–5-nerved, ovate to elliptic or broadly lanceolate, 25–70 Χ 8–35 mm, scabrellous. Peduncles 10–50(–70) mm. Involucres narrowly to broadly campanulate, 9–18 Χ 10–20 mm. Phyllaries obovate to oblong, becoming scarious, glabrous, apices rounded, usually entire or erose, sometimes ciliate. Paleae red to purple or yellow, apices obtuse, erose or subentire. Ray florets 6–15(–21); corollas usually scarlet red or maroon, sometimes yellow, laminae linear to spatulate, 8–25 mm. Disc florets 12–50; corollas yellow, 5–6 mm, lobes ca. 1 mm. Cypselae 7–10 mm, 3-angled (ray) or compressed (disc), ribbed, ciliate; pappi usually of 1 stout awn 4–6 mm. Fl. Jun–Nov, fr. Jul–Nov. 2n = 24.

Roadsides, grasslands, mountain slopes; up to 1300 m. Introduced and naturized in Gansu, Hebei, Henan, Sichuan, Yunnan [native to Mexico].

3. Zinnia haageana Regel, Gartenfl. 10: 355. 1861.

小百日菊  xiao bai ri ju

Annual herbs. Stems up to 60 cm high, becoming purple, strigose-pilose. Leaves ca. 3.5 Χ 0.7 cm, lanceolate, sessile, hispid. Heads radiate, 1–2 cm in diam.; peduncle up to 6 cm. Involucre broadly campanulate to hemispheric. Receptacle conic, paleae yellow, tipped with black. Phyllaries obovate, with wide, dark apical band; ray florets 8–9, oblong, ca. 17 mm long, bright orange and velvety above, dull orange to yellow and hirsute below; disc florets numerous, orange, darker below. Achenes ca. 4 mm long, 3-angled. Pappus of 2 unequal awns, or absent. Fl. Jun–Sep. 2n = 24.

Introduced and cultivated in China [native to Mexico].

9. SANVITALIA Lamarck, J. Hist. Nat. 2: 176, plate 33. 1792.

蛇目菊属  she mu ju shu

Annuals or perennials, 10–30 cm. Stems prostrate to erect, branched from bases or ± throughout. Leaves cauline; opposite; petiolate or sessile; blades obovate or spatulate to linear, bases rounded to cuneate, margins entire [toothed or lobed], faces hairy. Heads radiate, borne singly. Involucres hemispheric to ± rotate, 4–12+ mm diam. Phyllaries persistent, 8–21 in 2–3 series (lanceolate to linear, outer distally herbaceous, others each with stiff, subulate appendage). Receptacles convex to conic, paleate (paleae conduplicate, scarious). Ray florets 5–20, pistillate, fertile; corollas white or yellow (laminae sessile, persistent, becoming papery). Disc florets 15–60, bisexual, fertile; corollas distally yellow to orange (sometimes drying white), tubes much shorter than funnelform throats, lobes 5, deltate. Cypselae terete or obscurely 3–4-angled to compressed or flattened (all usually tuberculate and usually bearing uncinate hairs; none, some, or all in each head winged); pappi persistent, of 3–4 awns. x = 8, 11.

Five species: SW United States, Mexico, Central America, South America; one species introduced in China.

1. Sanvitalia procumbens Lamarck, J. Hist. Nat. 2: 176, plate 33. 1792.

蛇目菊  she mu ju

Stems procumbent to erect, 3–15 cm. Leaf blades ovate to lance-linear, 10–60 Χ 4–31 mm. Phyllaries 13–21, unequal. Ray corollas 2–9 mm. Cypselae: rays 2.5–3.5 mm with awns 1–3 mm, clearly 3-faced, adaxial faces often 2–3-nerved; discs strongly dimorphic within single heads: outer 4-angled, wingless, inner ± flattened, 1–2-winged. Flowering summer–fall. 2n = 16, 32.

Introduced in China [native to North and Central America].

10. Sigesbeckia Linnaeus, Sp. Pl. 900. 1753.

豨莶属  xi qian shu

Annual herbs. Leaves opposite, petiolate, densely pubescent. Heads apical and axillary; polyanthus, on peduncles, arranged usually in small corymbs. Involucre biseriate, companulate; outer phyllaries 5 in number, linear-spathulate, divaricate, beset with thick glandular hairs, longer than inner ones. Flower yellow, marginal florets unseriate, short-ligulate pistillate; disc florets tubular, bisexual. Receptacle plane with scarious bracts, embracing achenes. Achenes elongate-obovate, tetragonate, apex truncate. Pappus absent. x = 15.

About four species: Tropical and subtropical areas; three species in China.

Sigesbeckia esquirolii H. Lιveillι & Vaniot (Repert. Spec. Nov. Regni Veg. 8: 59. 1910) was described from Guizhou but its identity cannot be ascertained here because we have not seen any specimens.

1a.       Branches forked in upper part; leaves characeous, irregularly lobulate ..........  1. S. orientalis

1b.       Branches not forked; leaves thinly membranaceous, usually toothed.

2a.       Stems and both surfaces of leaves uniformly soft pubescent; peduncle not glandular; achenes ca. 2 mm long .....................................................................................................  2. S. glabrescens

2b.       Stems and lower surfaces of leaves densely white pubescent; peduncle usually glandular pilose; achenes 2.5–3.5 mm long ....................................................................................  3. S. pubescens

1. Sigesbeckia orientalis Linnaeus, Sp. Pl. 900. 1753.

豨莶  xi qian

Sigesbeckia brachiata Roxburgh; S. gracilis Candolle; S. humilis Koidzumi; S. iberica Willdenow; S. jorullensis Kunth; S. microcephala Candolle.

Annuals. Stems erect, simple or dichotomously pranched, branches opposite, obtusely angulate, 30–75(100) cm high, more or less crisp pubescent to densely pubescent, especially in upper part. Leaves ovate-triangular, ovate or oblong-ovate, cuneate or rounded to subcordiform at base, largely and usually unequally toothed (to sinuato-dentate) at margin, sometimes resembling the reduced leaves of sunflower, finely and appressed pilose (especially beneath), acute or acuminate, sometimes with sparse small glands and hairs. Heads small (about 5 mm wide). External bracts of involucre beset densely with capitate hairs on pedicels, oblong-obovate or linear-spathulate, considerably longer than the internal ones, sometimes broader and with small number of glands. [var. caspica (Fisch. Et Mey) Grossh] or shorter; internal bracts shorter than the external ones. Achenes obpyramidal, dark-gray or dull-black, sometimes with sparse, more lucid, levigate irregular tubercles, ca. 3 Χ 1.2 mm, with white annular vallicule at apex, marginal achenes slightly curvate. Fl. Apr–Sep, fr. Jun–Nov. 2n = 30, 60.

Fields, thickets, forest margin, forests, 100–2800 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejaing [India, Japan, Laos, Malesia, Nepal, Russia, Thailand, Vietnam; Oceania Australia; Africa; Tropic America].

2. Sigesbeckia glabrescens Makino, J. Jap. Bot. 1: 25. 1917.

毛梗豨莶  mao geng xi qian

Sigesbeckia orientalis f. glabrescens Makino, Bot. Mag. Tokyo 18: 100. 1904; S. formosana Kitamura; S. glabrescens var. leucoclada Nakai; S. orientalis subsp. glabrescens (Makino) H. Koyama.

Annuals. Stems 35–100 high, short appressed pilose. Median cauline leaves ovate-deltoid, 5–13 Χ 3.5–11 cm, irregularly toothed, upper surface short appressed pubescent, petiole winged; upper leaves oblong, sessile; uppermost leaves linear. Heads radiate, ca. 12 mm across; peduncle 1–3 cm long, densely short pubescent. Phyllaries spathulate, densely glandular pilose. Ligulate corolla 1.5–2.5 mm long, 3-toothed, tube 0.5–1 mm long, pilose; disc corolla ca. 1.5 mm long, 3-toothed, tube ca. 0.5 mm long, densely pilose. Achenes ca. 2 mm long. Fl. Apr–Sep, fr. Jun–Sep. 2n = 30.

Roadside, fields, thickets; 300–2500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunnan, Jiangsu, Jiangxi, Liaoning, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea].

3. Sigesbeckia pubescens Makino, J. Jap. Bot. 1: 21. 1917.

腺梗豨莶  xian geng xi qian

Annuals. Stems 60–120 cm high, densely white pubescent, especially on upper part. Median cauline leaves ovate to deltoid-ovate, 7–19 Χ 6–18 cm, short sppressed pubescent on both surfaces, veins on lower surface densely white pubescent. Heads radiate, ca. 20 mm across; peduncle 15–35 mm long, densely glandular pilose. Phyllaries linear, rounded at apex, glandular pilose at base; ligulate corolla ca. 3.5 mm long, shallowly 2- or 3-toothed, tube ca. 1.5 mm long; disc corolla 2–2.5 mm long, 5-toothed, tube 0.5–1 mm long, pilose. Achenes 2.5–3.5 mm long. Fl.May–Aug, fr. Jun–Oct. 2n = 30.

Mountain slopes, forest margin, thickets, grassland; up to 3400 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Hebei, Henan, Hubei, Hunan, Jilin, Liaoning, Jiangsu, Jiangxi, Nei Mongol, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [India, Japan, Korea].

11. ENYDRA Loureiro, Fl. Cochinch. 510. 1790

沼菊属  zhao ju shu

Herbs. Leaves opposite, sessile, entire or toothed. Heads nearly sessile, solitary, terminal or axillary. Phyllaries 4, foliaceous. Receptacle convex to conic, convex to conic, paleate, paleae persistent, apex glandulose. Ray florets female, fertile; ligules small, apex 3–4-toothed. Disc florets bisexual, tubular, limbs campanulate, 5–6-toothed. Anther tails obtuse, entire, or unconspicuously auriculate. Achenes oblong, glabrous. Pappus absent.

About 10 species: Tropical and subtropical regions; one species in China.

1. Enydra fluctuans Loureiro, Fl. Cochinch. 511. 1790.

沼菊  zhao ju

Herbs. Stems stout, cylindrical, slightly fleshy, prostrate in the lower part, 50–80 cm long. Leaves nearly sessile, oblong, linear-oblong, 2–6 cm Χ 4–14 mm, apex obtuse or acute, base amplexicaul, margin sparsely serrate, both surfaces glabrous. Heads 8–10 mm in diam. Involucre 4, dorsally glabrous, outer pair larger, ovate-oblong, 10–11 mm long, apex rounded. Receptacle ca. 3 mm in diam.; paleae rigid, ca. 5 mm long, apex toothed and sparsely pubescent. Ray florets ca. 3 mm long, ligules 3–4-lobed in apex. Disc florets 5-lobed in apex, stamens 5, rarely 6. Achenes obovoid-cylindrical, ca. 3.5 mm long. Fl. Nov–Apr.

Marshes, streamside. Hainan, Yunnan [India, Indonesia, Malaysia, Myanmar, Thailand, Vietnam; Australia].

12. Eclipta Linnaeus, Mant. Pl. 157, 286. 1771.

鳢肠属  li chang shu

Erect annual or perennial herbs, branched, strigose. Leaves opposite, toothed. Heads terminal on stems and branches or axillary, peduncled, heterogamous. Involucre campanulate, bracts imbricate, in ca. 2 series. Receptacle flat or convex, paleaceous, paleae awn-like. Florets whitish. Ray florets bisexual, mostly fertile, corolla tubular, white, 4- or rarely 5-lobed. Anthers entire or very shortly bifid at base. Style branches obtuse, mammillate at apex. Achenes thick, those of ray florets 3-angled, those of disc florets compressed, 4-angled, apex truncate and depressed, margins with 1–3 minute teeth, coarsely hairy, tuberculate. Pappus none, or of a few short teeth. x = 11.

About four species: mostly warm-temperate to tropical New World; introduced in Old World, two species in China.

1a.       Plants less than 60 cm tall, strigose-pilose; leaves lanceolate to oblong, entire or slightly undulate; pedicel 2–4 cm long, receptacle ca. 1 cm wide ...........................................................  1. E. prostrata

1b.       Plants more than 100 cm tall, velutinous; leaves ovate, oblong to oblanceolate, irregularly undulate; pedicel less than 2 cm long, receptacle ca. 0.5 cm wide
                                                                                                                                             2. E. zippeliana

1. Eclipta prostrata (Linnaeus) Linnaeus, Mant. Pl. 286. 1771.

鳢肠  li chang

Verbesina prostrata Linnaeus, Sp. Pl. 902. 1753; Eclipta alba (Linnaeus) Hasskarl; E. thermalis Bunge.

Annual herbs. Stems erect, ascending or prostrate, to 60(–100) cm tall, strigose-pilose, branched at base. Leaves chartaceous, lanceolate, 3–10 Χ 0.5–2.5 cm, apex gradually acuminate, base narrowed, sessile or short petiolate, serrulate, densely strigose-pubescent on surfaces. Heads ca. 6 mm across; peduncle slender, 2–4.5 cm long; involucre globose-campanulate, ca. 5 Χ 6–7 mm, enlarging to 11 mm broad in fruit, bracts 5 or 6, 2-seriate, oblong, acute, outer ones longer. Ray florets in 2 series; corolla white, 2.5–3 Χ ca. 0.4 mm, bifid or entire. Disc florets many, corolla white, ca, 1.5 mm long, 4-lobed. Achenes ca. 2.8 Χ 1.5 mm, ribbed on margins. Fl. Jun–Sep.

Riverside, fields; up to 1600 m. Anhui, Gansu, Fujian, Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jilin, Jiangsu, Jiangxi, Liaoning, Shandong, Shanxi, Shaanxi, Sichuan, Taiwan, Yunnan, Zhejiang [North America, Central America; South America; introduced in Europe, Asia, Africa, Pacific islands, Australia].

2. Eclipta zippeliana Blume, Bijdr. 914. 1825

毛鳢肠  mao li chang

Eclipta prostrata var. zippeliana (Blume) J. Koster.

Plants coarse, densely hirsute, erect, branched, 30–100 cm tall; stems often reddish. Leaves 3–9 cm long, sessile, oblong obovate to lanceolate, hirsute, acuminate, base narrowed, margins irregularly and rather coarsely spinulose-toothed. Heads ovoid, 6–7 Χ 5–6 mm, larger in fruit; peduncle hirsute, ca. 1 cm long; bracts green, hairy, as long as or exceeding florets; ligule of ray florets ca. 1.5 mm long. Achenes black, ca. 3 mm long, apically hairy.

Abandoned ponds and roadsides; probably introduced from southeast Asia. Taiwan [Malaysia, Philippines].

13. Guizotia Cassin,i Bull. Soc. Philom. Paris 127. 1827.

小葵子属  xiao kui zi shu

Annuals, perennials, subshrubs, or shrubs. Stems erect or creeping, branched. Leaves mostly cauline; opposite, sessile; blades rhombic to lanceolate or oblanceolate, margins entire or serrate, faces glabrous or puberulent to pilose, gland-dotted at least abaxial. Heads radiate, in corymbiform arrays or borne singly. Involucres campanulate or hemispheric. Phyllaries persistent, 10–13 in 2 series. Receptacles conic to hemispheric; paleae oblong to lanceolate, membranous to scarious. Ray florets 6–18, pistillate, fertile; corollas yellow. Disc florets numerous, bisexual, fertile; corollas yellow, tubes cylindric, hairy, shorter than campanulate throats, lobes 5, deltate. Achenes weakly compressed, 3–4-angled, glabrous. Pappus absent. x = 15.

Six species: Africa; one species introduced in China.

1. Guizotia abyssinica (Linnaeus f.) Cassini, Dict. Sci. Nat. 59: 248. 1829.

小葵子  xiao kui zi

Polymnia abyssinica Linnaeus f., Suppl. Pl. 383. 1782.

Annuals. Stems 30–50 cm to 1–2 m high, subglabrous, more or less, pilose above. Leaves sessile (lower ones petiolate), oblong-ovate or lanceolate, semiamplexicaul, acuminate, glabrous above, more or less, short-haired beneath, especially along nerves. Heads 2–6 cm in diam., on long or sometimes short pubescent peduncles. External involucral bracts ovate or latioval, herbaceous; internal ones narrower paleaceous. Ligulate florets with short tube and tridentate limb, pubescent along all surface or only at the base of limb (ligule) and in its lower part; central florets tubular, pubescent as the ligulate ones, quinquelobate. Achenes deprived of pappus, external achenes triquetrous; internal ones, more or less, tetragonate, 3–6 mm long, 1.5–3.0 mm wide, obpyramidal, in transverse section subrhombical or triangular, brown or black, with thin pericarp.

Cultivated in Fujian, Sichuan, Yunnan [native to Africa].

Grown for its edible oil and seed, originated in the Ethiopian highlands.

14. Rudbeckia Linnaeus, Sp. Pl. 906. 1753.

金光菊属  jin guang ju shu

Annuals, biennials, or perennials. Stems erect, branched distally, glabrous or hairy, sometimes glaucous. Leaves basal and cauline; alternate; petiolate or sessile; blades elliptic, lanceolate, linear, oblanceolate, ovate, or spatulate, often pinnately lobed to 1–2-pinnatifid, ultimate margins entire, dentate, serrate, or coarsely toothed, faces glabrous or hairy, sometimes glaucous, sometimes gland-dotted. Heads radiate or discoid, borne singly or in ± corymbiform or paniculiform arrays. Involucres hemispheric to rotate. Phyllaries persistent, 5–20 in 1–2(–3) series, narrowly triangular to lanceolate or narrowly elliptic, subequal, herbaceous, distally or throughout. Receptacles subspheric to ovoid, or conic to columnar, paleate (paleae mostly tan proximally, green to maroon distally, obovate, concave, each ± clasping a floret, apices acute to cuspidate or truncate to rounded, abaxial tips glabrous or hairy, sometimes gland-dotted, resin ducts 2–3, maroon, 1 medial and 1 near each margin; receptacles plus paleae and florets equaling discs, 8–80 Χ 5–30 mm). Ray florets 0 or 5–25, neuter; corollas usually yellow to yellow-orange or bicolor (laminae often proximally maroon or each with a maroon splotch, distally yellow). Disc florets numerous, bisexual, fertile; corollas yellow, yellowish green, or brown-purple (often bicolor), tubes shorter than cylindric to funnelform throats, lobes 5, triangular. Cypselae black, obpyramidal and 4-angled, faces glabrous, angles sometimes hairy. Pappus 0, coroniform, or of 2–8 unequal scales. x = 16, 18, 19.

About 23 species: North America; about six species introduced in China.

1a.       Stems glabrous; leaves sessile, blades bases auriculate and clasping, faces glabrous, glaucous; pappus absent .................................................................................................  1. R. amplexicaulis

1b.       Stems usually hairy; leaves blade bases not auriculate and clasping; pappus usually coroniform or of 2–6+ scales, sometimes 0.

2a.       Leaves often bluish green, glaucous; receptacles usually conic to columnar, receptacles sometimes hemispheric to ovoid; ray florets 0 or 8–15+, corollas bright yellow; disc corollas proximally yellow to yellowish green, distally yellow or greenish to brown-purple; cypselae (3–)3.5–7.5 mm; pappi coroniform, or of 2–6 scales 0.1–2.5 mm.

3a.       Blades of all but distalmost leaves usually 1–2-pinnatifid or pinnately lobed; receptacles hemispheric or globose to ovoid; disc corolla lobes yellow
                                                                                                                 2. R. laciniata

3b.       Blades of basal leaves ± pinnatifid to pinnately lobed (distal leaves sometimes not lobed); receptacles ovoid to conic .........................................................................................  3. R. maxima

2b.       Leaves green, not glaucous; receptacles usually conic to hemispheric, rarely columnar; ray florets 6–25+, corollas usually yellow-orange proximally, yellow distally, sometimes with basal maroon splotch; disc corollas proximally yellow to yellowish green, distally usually brown-purple, lobes sometimes yellowish or greenish; cypselae 1.5–3.5(–4) mm; pappi coroniform, or of 8+ unequal scales 0.1–2 mm, or absent.

4a.       Annuals or biennials; pappi absent .................................................................  4. R. hirta

4b.       Perennials; pappi coroniform, 0.1–0.5 mm long.

5a.       Proximal cauline leaves elliptic or ovate, usually 3(–5)-lobed; paleae cuspidate, tips awnlike, ca. 1.5 mm .........................................................................................................  5. R. triloba

5b.       Proximal cauline leaves elliptic, linear, spatulate, or ovate, rarely lobed; paleae acute, obtuse, or rounded .................................................................................................  6. R. fulgida

1. Rudbeckia amplexicaulis Vahl, Skr. Naturhist. Selsk. 2(2): 29, plate 4. 1793.

抱茎金光菊  bao jing jin guang ju

Dracopis amplexicaulis (Vahl) Cassini.

Annual herbs. Stems 30–60 cm high, glabrous. Leaf blades entire or serrate, sessile, 3–15 Χ 0.5–4 cm, lower leaves oblong to spathulate, upper leaves ovate, ovate-oblong, lanceolate, apex acute. Involucres 1–5 cm diam. Phyllaries spreading to reflexed, green, linear to lanceolate, herbaceous. Ray laminae spreading, eventually reflexed, elliptic to obovate, 12–30 Χ 7–15 mm, abaxially hirsute. Disc corollas 2.8–3.5 mm. Cypselae: each face 4–5-striate and minutely cross-rugose, glabrous; pappi 0. Fl. Jun–Sep. 2n = 32.

Cultivated ornamental in China [native to North America].

2. Rudbeckia laciniata Linnaeus, Sp. Pl. 906. 1753.

金光菊  jin guang ju

Perennials, 50–300 cm (rhizomes often elongate, slender, plants colonial, roots fibrous). Leaves green, blades broadly ovate to lanceolate, all but distalmost 1–2-pinnatifid or pinnately compound, leaflets/lobes 3–11, bases cuneate to attenuate or cordate, margins entire or dentate, apices acute to acuminate, faces glabrous or hairy (sometimes with translucent patches); basal (often withering before flowering) petiolate, 15–50 Χ 10–25 cm; cauline petiolate or sessile, mostly lobed to pinnatifid, sometimes not lobed, 8–40 Χ 3–20 cm. Heads (2–25) in loose, corymbiform arrays. Phyllaries to 2 cm (8–15, ovate to lanceolate, margins mostly ciliate, glabrous or hairy). Receptacles hemispheric or ovoid to globose; paleae 3–7 mm, apices (at least of proximal) truncate or rounded, abaxial tips densely hairy. Ray florets 8–12; laminae elliptic to oblanceolate, 15–50 Χ 4–14 mm, abaxially hairy. Discs 9–30 Χ 10–23 mm. Disc florets 150–300+; corollas yellow to yellowish green (lobes yellow), 3.5–5 mm; style branches 1–1.5 mm, apices acute to rounded. Cypselae 3–4.5 mm; pappi coroniform or of 4 scales, to 1.5 mm. Fl. Jul–Oct.

Widely cultivated ornamental in China [native to North America].

3. Rudbeckia maxima Nuttall, Trans. Amer. Philos. Soc., n. s. 7: 354. 1840.

大金光菊  da jin guang ju

Perennials, to 250 cm (rhizomatous, roots fibrous). Leaves bluish green (heavily glaucous, dried blades white under UV light, at least abaxially), blades elliptic, ovate to obovate or pandurate (not lobed), leathery, margins crenate, dentate, or entire, apices acute to rounded, faces glabrous; basal petiolate, elliptic to ovate, 15–65 Χ 4–15 cm, bases attenuate to cuneate; cauline petiolate or sessile, 7–50 Χ 4–14 cm, blades ovate to pandurate, bases cuneate to auriculate (and clasping). Heads borne singly or (3–15) in ± corymbiform arrays. Phyllaries to 1.5 cm (ovate to lanceolate, margins ciliate). Receptacles ovoid to conic; paleae 6–8 mm, apices mostly rounded, abaxial tips glabrous (hairy subapically). Ray florets 10–20; laminae elliptic to oblanceolate, 30–80 Χ 8–15 mm, abaxially sparsely hairy. Discs 40–80 Χ 15–35 mm. Disc florets 300–600+; corollas maroon (at least distally), 4–6 mm; style branches ca. 2.2 mm, apices acute. Cypselae 6.5–7 mm; pappi of 4–6 scales to 1.5 mm. Fl. May–Aug. 2n = 36.

Cultivated ornamental in China [native to North America].

4. Rudbeckia hirta Linnaeus, Sp. Pl. 907. 1753.

黑心金光菊  hei xin jin guang ju

Annuals, biennials, or perennials, to 100 cm. Stems hispid to hirsute (hairs spreading, 1+ mm). Leaves: blades elliptic, lanceolate, or ovate (not lobed), bases attenuate to cuneate, margins entire or serrate, apices acute, faces hispid to hirsute; basal petiolate, blades 8–30 Χ 0.5–7 cm; cauline petiolate or sessile, blades (sometimes pandurate) 3–20 Χ 0.4–4 cm. Heads borne singly or (2–5) in loose, corymbiform arrays. Phyllaries to 3 cm (faces hispid to hirsute). Receptacles hemispheric to ovoid; paleae 4–6 mm, apices acute, often attenuate, abaxial tips hirsute to hispid. Ray florets 8–16; laminae elliptic to oblong or oblanceolate, 15–45 Χ 5–10 mm, abaxially hispid to hirsute. Discs 12–22 Χ 10–20 mm. Disc florets numerous; corollas proximally yellowish green, distally brown-purple, 3–4.2 mm; style branches ca. 1.5 mm, apices subulate. Cypselae 1.5–2.7 mm; pappi 0. Fl. Jun–Oct.

Widely cultivated ornamental in China [native to North America].

5. Rudbeckia triloba Linnaeus, Sp. Pl. 2: 907. 1753.

三裂叶金光菊  san lie ye jin guang ju

Perennials, to 150 cm. Stems glabrate to hirsute or strigose. Leaves: blades ovate to subcordate or elliptic (not lobed), margins serrate, apices acute to acuminate, faces hirsute to strigose; basal petiolate, 10–30 Χ 2–8 cm, bases truncate or rounded to cordate; cauline petiolate or sessile, ovate to elliptic, proximal usually 3–5-lobed, 2–20 Χ 1.5–8 cm (smaller, fewer lobed distally), bases rounded to attenuate, sometimes clasping. Heads (10–30) in paniculiform arrays. Phyllaries to 1.5 cm (faces moderately hirsute). Receptacles conic to subhemispheric; paleae 5–6.5 mm, apices cuspidate (tips awnlike, 1.5+ mm), glabrous. Ray florets 8–15; laminae (corollas yellow to yellow-orange with basal maroon splotches) linear to oblanceolate, 8–30 Χ 3–8 mm, abaxially sparsely strigose. Discs 8–15 Χ 10–20 mm. Disc florets 150–300+; corollas yellowish green basally, otherwise brown-purple, 3–4 mm; style branches ca. 1.2 mm, apices obtuse to rounded. Cypselae 1.9–2.8 mm; pappi coroniform, to 0.2 mm. Fl. Jul–Sep.

Cultivated ornamental in China [native to North America].

6. Rudbeckia fulgida Aiton, Hort. Kew. 3: 251. 1789.

全缘金光菊  quan yuan jin guang ju

Perennials, to 120 cm high, stoloniferous, rosettes forming at stolon apices. Stems glabrous or moderately hirsute (branches spreading). Leaves: blades lanceolate to broadly ovate or elliptic (not lobed), herbaceous, bases attenuate to cordate, margins usually entire or serrate, sometimes lacerate, apices acute, faces glabrous or hirsute to strigose; basal petiolate, 5–30 Χ 1–8 cm; cauline petiolate, 2–25 Χ 0.5–7 cm, bases attenuate to cordate or auriculate. Heads borne singly or (2–7) in corymbiform arrays. Phyllaries to 2 cm. Receptacles hemispheric to ovoid; paleae 2.5–4 mm, (apical margins usually ciliate) apices obtuse to acute, abaxial tips usually glabrous. Ray florets 10–15; laminae elliptic to oblanceolate, 15–25 Χ 3–6 mm, abaxially strigose. Discs 12–16 Χ 10–18 mm. Disc florets 50–500+; corollas proximally yellowish green, brown-purple distally, 3–4.2 mm; style branches ca. 1.3 mm, apices rounded. Cypselae 2.2–4 mm; pappi coroniform, to 0.2 mm. Fl. Aug–Oct.

Cultivated in China [native to North America].

15. Blainvillea Cassini, Dict. Sci. Nat. 29: 493. 1823.

百能葳属  bai neng wei shu

Annual or perennial herbs. Leaves opposite or alternate in upper part, petiolate, margin serrate. Heads small, terminal or axillary, with slender and long petioles. Corolla light yellow, yellow or rarely white. Involucre ovoid, ovoid-campanulate or hemispherical. Phyllaries few, outer ones papery. Receptacle convex, paleae rigid and dry membranous. Ray florets female, 1–2 series, ligules short or very short, apex 2–4-dentate. Disc florets tubular, limbs 5-dentate. Anther tails obtuse, entire or unconspicously auriculate. Achenes glabrous or pubescent, apex truncate, those in female florets 3-ribbed, dorsally compressed, those in bisexual florets 3–4-ribbed, or laterally compressed. Pappus 2–5, unequal, spine like or squama like, base connate.

About 10 species: Tropics; one species in China.

1. Blainvillea acmella (Linnaeus) Philipson, Blumea 6: 350. 1950.

百能葳  bai neng wei

Verbesina acmella Linnaeus, Fl. Zeyl. 309. 1748; Blainvillea latifolia (Linnaeus filius) Candolle; Eclipta latifolia Linnaeus filius.

Annual herbs. Stems erect, 40–60 cm high, branched. Lower leaves opposite, with up to 1 cm long petioles, blade ovate to ovate-lanceolate, 3–6 Χ 2–3 cm, apex acuminate, base cuneate, margin sparsely serrate, both surface scabrous; upper leaves smaller, usually alternate, ovate to ovate-oblong, 2–3 Χ 1.3–1.5 cm, base usually rounded. Heads axillary or terminal, ca. 1 cm in diam., peduncles slender, 15–40 mm long, with spreading hairs. Phyllaries in two series; outer ones papery, green, ovate-oblong, ca. 6 mm long, apex shortly acute or obtuse, dorsally densely hairy; inner ones ovate to oblong-linear, ca. 5 mm long, apex acute, sparsely pubescent. Paleae oblong-lanceolate, ca. 5 mm long, apex aristiform, dorsally pubescent. Ray florets one series, yellow or yellowish-white, ligules ca. 3 mm long, apex 2–4-toothed. Disc florets campanulate, limbs 5-toothed. Achenes densely pubescent, those in female florets trigonous, ca. 4 mm long, those in bisexual florets compressed, ca. 5 mm long. Pappus short, unequal, 2–5, spine like. Fl. Apr–Jun.

Sparsely broadleaf forests, grassy slopes. Hainan, Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam; Africa, America, Australia].

16. Wedelia Jacquin, Enum. Syst. Pl. 8, 28. 1760.

蟛蜞菊属  peng qi ju

Annuals, perennials, subshrubs or shrubs. Stems mostly erect, branched. Leaves opposite, petiolate or sessile, margin toothed or rarely entire. Heads discoid, borne singly in corymbiform arrays. Involucres obconic to hemispheric, 4–8(–15) mm diameter. Phyllaries persistent, 8–16+ in 2–3+ series, outer ones usually larger and/or more herbaceous than inner ones. Receptacles convex, paleate, paleae conduplicate, chartaceous to scarious. Ray florets 0 or 4–18, pistillate, fertile; corollas yellow to orange, purplish or white. Disc florets 8–150+, bisexual, fertile; corollas yellow to orange or purplish, sometimes marked with purple, tubes shorter than or equaling funnelform or cylindric throats, lobes 5, deltate. Cypselae ± dimorphic; peripheral sometimes obcompressed and weakly 3-angled, inner compressed and biconvex or flattened, or somewhat 4-angled; pappi persistent, cyathiform. x = 15

About 25 species: tropical and subtropical New World; five species in China.

1a.       Leaf base extended into an abruptly terminating petiolar wing; leaves often 3-lobed  1. W. trilobata

1b.       Petioles usually distinct; leaf blade entire or dentate, but not lobed.

2a.       Leaf blade margin 1–3 toothed or entire, lower leaves no more than 1.3 cm broad, petioles indistinct.

3a.       Phyllaries longer than paleae; achenes apex rounded; pappus distinct, cup shaped; achenes not angled                                                                                                                2. W. chinensis

3b.       Phyllaries shorter than paleae or rarely subequal; achenes apex truncate; pappus nearly obsolete, bristles 1 or 2 ..................................................................................................  3. W. prostrata

2b.       Leaf blade margin densely toothed, lower leaves broader than 3 cm; petioles distinct.

4a.       Paleae apex obtuse or shortly acute; achenes apex truncate; pappus absent
                                                                                                                   4. W. biflora

4b.       Paleae apex acuminate; achenes apex rounded; pappus usually 2–3, distinct, cup-shaped  5. W. wallichii

1. Wedelia trilobata (Linnaeus) Hitchcock, Rep. (Annual) Missouri Bot. Gard. 4: 99. 1893.

南美蟛蜞菊  nan mei peng qi ju

Silphium trilobatum Linnaeus, Syst. Nat., ed. 10. 2: 1233. 1759.

Perennial herbs, procumbent; stems stout, glabrous or pubescent, seldom scabrous. Leaves opposite, somewhat succulent, to 18 cm long, elliptic or lanceolate, often with triangular lobes and conspicuous marginal teeth, apex acute, base cuneate, glabrous or sparsely pubescent, sometimes scabrous; petiole distinct but less than 5 mm long. Heads solitary on elongate peduncle, radiate; involucre green, bracts lanceolate, ciliate, obscurely nerved, 10–15 mm long, innermost bracts narrower; ray florets 4–8, showy, yellow, 15–20 mm long, 3 or 4 denticulate, fertile; disc florets yellow, numerous, ca. 2 cm long, corolla 5–6 mm long. Achenes blackish, sometimes mottled, clavate, angled, ca. 5 mm long. Pappus of fused scales, coroniform.

Cultivated as an ornamental ground cover and escaped in many countries, occasionally naturalized on slopes and along roadsides at lower elevations. Taiwan [native of the New World tropics].

2. Wedelia chinensis (Osbeck) Merrill, Philipp. J. Sci. 12: 111. 1917.

蟛蜞菊  peng qi ju

Solidago chinensis Osbeck, Dagb. Ostind. Resa 241. 1757; Verbesina calendulacea Linnaeus; Wedelia calendulacea (Linnaeus) Lessing, Syn. Gen. Compos. 222. 1832, non Richard (1807).

Stems prostrate with ascending tips, elongate, rooting from lower nodes, appressed pilose. Leaves chartaceous, linear-oblong to lanceolate, 2–10 cm long, 6–20 mm wide, apex acute, base narrowed, margins sparsely mucronulate-serrulate, sessile or short petiolate, appressed pilose on surfaces. Heads 2–2.5 cm across, solitary on erect branches, peduncle 6–12 cm long; involucre hemispheric, 8–9 mm long, bracts 5, nearly equal in length, in 1 series, oblong, acute or sometimes obtuse, short appressed pubescent. Ray florets in 1 series, yellow, corolla 9–11 Χ 3–3.5 mm, 2- to 3-dentate. Disc corolla 4–4.5 mm long, 5-lobed. Achenes obovoid, 3.5 Χ 1.5–2 mm, coarsely hairy at tip. Pappus vase-or cup-shaped. Fl. Mar–Sep.

On paddy ridges, in grassy fields and moist lowland depressions in the north, also common in littoral areas. Liaoning, Guangdong, Taiwan [India, Japan, Myanmar, southeast Asia].

3. Wedelia prostrata (Hooker & Arnott) Hemsley, J. Linn. Soc. Bot. 23: 434. 1888.

卤地菊  lu di ju

Stems long creeping, rooting at nodes, coarsely hirsute; flowering branches ascending, leafy, densely strigose. Cauline leaves thick coriaceous, oblong, sometimes ovate or lanceolate, acute, base cuneate, loosely dentate, 3-nerved, coarsely strigose on surfaces, petiole 2–8 mm long. Heads 5–22 mm long, 16–22 mm across, usually solitary, terminal; peduncle 1–7 cm long; involucre hemispheric, 10–12 Χ 5–6 mm, bracts in 1 series, ovate to ovate-oblong, 2–3.5 mm wide, acute to obtuse, coarsely strigose. Ray corollas yellow, 8–11 Χ ca. 4 mm. Disc corollas tubular, 5–6 mm long, 5-dentate. Achenes 3.5–4 Χ ca. 2 mm, apically strigillose, 3- or 4-angled. Pappus bristles 1 or 2, or obsolete. 2n = 30.

Littoral sand dunes, sandy seashores. Taiwan [Japan, Korea; SE Asia].

1a. Leaves oblong, 1.5–4.5 cm, coriaceous; heads 1.6–2.2 cm across, usually solitary  3a. var. prostrata

1b. Leaves ovate, 3–12 cm, chartaceous; heads 2.–2.5 cm across, in 3’s or sometimes solitary  3b. var. robusta

3a. Wedelia prostrata var. prostrata

卤地菊(原变种)  lu di ju (yuan bian zhong)

Verbesina prostrata Hooker & Arnott, Bot. Beechey Voy. 195. 1836; Eclipta dentata H. Lιveillι & Vaniot; Wollastonia prostrata Hooker & Arnott.

Insert diagnosis.

Littoral sand dunes, sandy seashores. Taiwan [Japan, Korea; SE Asia].

3b. Wedelia prostrata var. robusta Makino, J. Jap. Bot. 1: 23, fig. 2. 1917; Koyama, Fl. Jap. 3b: 34. 1995.

大天蓬草舅  da tian peng cao ju

Wedelia robusta (Makino) Kitamura.

Stems much elongate; leaves thick chartaceous to somewhat coriaceous, ovate, 3–12 Χ 1.5–6 cm, apex acute, base cuneate, loosely obtuse-toothed, petiole 3–28 mm long; heads 20–25 mm across, in 3’s or sometimes solitary.

Littoral regions. Taiwan [Japan].

4. Wedelia biflora (Linnaeus) Candolle in Wight, Contr. Bot. India 18. 1834.

孪花蟛蜞菊  luan hua peng qi ju

Subshrubs; stems elongate, branched, scandent, coarsely appressed-strigose. Cauline leaves thick chartaceous, long-petiolate, ovate, 7–14 Χ 3–8 cm, acuminate at apex, rounded at base, margins much-toothed, appressed-strigose on surfaces; petioles 12–23 mm long. Heads 3–6, terminal, 2–3 cm wide, peduncles 1.5–5.5 cm, slender; involucre 10–13 Χ 5–7 mm, bracts in 1 series, ovate-lanceolate or narrowly ovate, gradually narrowed to tip, densely appressed-strigose. Ray florets yellow, in one row; corolla 9–13 mm long, 2- or 3-dentate. Disc florets yellow; corolla ca. 5 mm long, apex 5-dentate. Achenes 3–3.5 Χ 2–2.5 mm, cuneate at base, often 3-angled, coarsely strigose toward tip. Pappus bristles 2–2.5 mm long. Flowering all year rounded. 2n = 30.

Seasides. Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan [India, Indonesia, Japan, Malaysia, Philippines, Vietnam; Pacific islands].

1a. Ray florets 8–12, disc florets 20–35 per head .......................................................  4a. var. biflora

1b. Ray florets 14 or 15, disc florets 45–70 per head ..........................................  4b. var. ryukyuensis

4a. Wedelia biflora var. biflora

孪花蟛蜞菊(原变种)  luan hua peng qi ju (yuan bian zhong)

Verbesina biflora Linnaeus, Sp. Pl. ed. 2. 2: 1272. 1763.

Insert diagnosis.

Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan [India, Indonesia, Japan, Malaysia, Philippines, Vietnam; Pacific islands].

4b. Wedelia biflora var. ryukyuensis H. Koyama, Acta Phytotax. Geobot. 33: 245. 1982.

琉球蟛蜞菊  liu qiu peng qi ju

Leaves ovate, 6–12 Χ 3–8 cm, thick, petiole 1.5–3.6 cm long, base somewhat truncate. Heads 2.5–3 cm across; peduncle thick, 4–8 cm long; ray florets 14 or 15; disc florets 45–70 per head. 2n = 45.

Seasides. Taiwan [Japan].

Wedelia biflora var. ryukyuensis differs from the typical variety in having larger heads with more ray and disc florets and larger leaf blade with a longer petiole.

5. Wedelia wallichii Lessing, Linnaea 6: 162. 1831.

山蟛蜞菊  shan peng qi ju

Wedelia montana var. wallichii (Lessing) H. Koyama; W. urticaefolia Candolle.

Perennial herbs. Stems erect, stout, glabrous or scabrous. Leaves petiolate, petioles 1–2 cm, blade ovate or ovate-lanceolate, (3–)5–9 Χ (1–)3–4 cm, apex acuminate, base rounded or cuneate, margin crenulate, both surfaces scabrous. Heads ca. 15 mm in diam., solitary on elongate peduncle. Involucre campanulate, 7–10 mm long, 4–5 mm in diam. Phyllaries in two series; outer ones pappery, green, oblong, ca. 10 mm long, apex obtuse or shortly acute, dorsally scabrous; inner ones oblong to lanceolate, ca. 7 mm long, dorsally pubescent in upper part, apex acuminate. Paleae oblong, sparsely pubescent. Ray florets one series, yellow, ligules oblong, 4–6 mm long, ca. 2 mm wide, apex 2–3-toothed. Disc florets tubular, yellow, limbs 5-toothed. Achenes obovoid-triangonous, slightly compressed, ca. 5 mm long red brown. Pappus 2–3. Fl. Apr–Oct.

Streamside, roadside; 500–1200(–3000)m. Guangdong, Guangxi, Guizhou, Hainan, Yunnan [Bhutan, India, Myanmar, Nepal, Thailand].

17. Sclerocarpus Jacquin, Icon. Pl. Rar. 1: 17, plate 176. 1781.

硬果菊属  ying guo ju shu

Annuals, perennials or subshrubs. Stems sprawling to erect, branched. Leaves mostly cauline; opposite (proximal) or alternate; petiolate [sessile]; blades 3-nerved, ovate to rhombic or lanceolate, bases cuneate to rounded, margins coarsely toothed, faces glabrous or ± scabrous. Heads radiate, borne singly. Involucres rotate to hemispheric, 12–20 mm diam. Phyllaries persistent, 5–8(–21) in 1(–2) series. Receptacles low-conic, paleate; paleae each closely investing subtended floret, each forming a hard perigynium around a fruit and shed with it. Ray florets 5–8, neuter; corollas yellow to orange. Disc florets 10–50, bisexual, fertile; corollas usually yellow to orange, sometimes with purple, tubes much shorter than cylindric throats, lobes 5, lance-linear. Cypselae (within perigynia) weakly compressed, obliquely obovoid or arcuate (glabrous); pappi 0, or coroniform (of distinct or connate scales or bristles). x = 12.

About 12 species; mainly in United States, Mexico, Central America, one species in tropical Africa, tropical Asia. One species introduced in China.

1. Sclerocarpus africanus Jacquin, Icon. Pl. Rar. 1: 17, t. 176. 1781.

硬果菊  ying guo ju

Annual herbs. Stems 25–50 cm high, white hirsute. Leaves opposite; blade ovate, 3.5–7 Χ 1.5–2 cm, apex acute or obtuse, base decurrent to petioles, margins serrate, both surfaces scabrous. Head solitary, 8–12 mm diameter. Involucre campanulate; outer phyllaries ovate, ca. 1 cm long, apex acuminate; inner ones ovate-oblong, shorter than outer ones, abaxially densely tomentose. Ray florets in one series; ligules broad elliptical, apex 2-lobed; disc florets puberulous. Achenes oblong. Pappus coroniform of distinct or connate scales.

Naturalized in Xizang [native to tropical Africa and Asia].

18. Tithonia Desfontaines ex Jussieu, Gen. Pl. 189. 1789.

肿柄菊属  zhong bing ju shu

Annuals, perennials, subshrubs or shrubs. Stems erect, branched. Leaves all or mostly cauline; opposite (proximal) or mostly alternate; petiolate or sessile; blades often (1-), 3-, or 5-nerved, mostly deltate or pentagonal, sometimes 3- or 5-lobed, bases truncate or auriculate, ultimate margins serrate to crenate, faces glabrate, ± hirsute, pilose, soft-pubescent, or villous, often gland-dotted. Heads borne singly (peduncles usually distally dilated, fistulose). Involucres campanulate to hemispheric. Phyllaries persistent, in 2–5 series. Receptacles hemispheric to convex, paleate. Ray florets 8–30, neuter; corollas yellow or orange. Disc florets bisexual, fertile; corollas yellow, tubes shorter than throats, lobes 5, triangular. Cypselae compressed or flattened, often 3- or 4-angled or biconvex, cuneiform in silhouette, sometimes with basal elaiosomes; pappi 0, or coroniform, of connate scales, 1–2 scales sometimes subulate to aristate. x = 17.

About 11 species, SW United States, Mexico, Central America; one species introduced and naturalized in China.

1. Tithonia diversifolia (Hemsley) A. Gray, Proc. Amer. Acad. Arts. 19: 5. 1883.

肿柄菊  Zhong bing ju

Mirasolia diversifolia Hemsley, Biol. Cent.-Amer., Bot. 2 : 168, plate 47. 1881.

Perennials, subshrubs, or shrubs, to 250(–500) cm. Leaves: petioles 2–6 cm; blades ± deltate to pentagonal, 7–33 Χ 7–22 cm, sometimes 3- or 5-lobed, abaxial faces glabrous to hispid-pilose. Peduncles 7–24 cm. Phyllaries 16–28 in (3–)4 series, oblong to ovate; outer 6–10 Χ 4–7 mm, apices rounded to acute, abaxial faces usually glabrous; inner 10–20 Χ 3–10 mm, apices rounded to acute, abaxial faces glabrous. Paleae 10–13 Χ 2–3 mm, mucros 1.5–2.5 mm. Ray florets 7–14; corollas yellow, laminae linear, 48–69 Χ 9–16 mm. Disc florets 80–120+. Cypselae 4–6 mm. Fl. Sep–Jan. 2n = 34.

Introduced and naturalized in Guangdong, Taiwan, Yunnan [Mexico].

19. Helianthus Linnaeus, Sp. Pl. 904. 1753.

向日葵属  xiang ri kui shu

Annuals or perennials. Stems erect or ascending to decumbent or procumbent, usually branched distally. Leaves basal and/or cauline; opposite, or opposite (proximal) and alternate, or alternate; petiolate or sessile; blades mostly deltate, lance-linear, lanceolate, lance-ovate, linear, or ovate, bases cordate to narrowly cuneate, margins usually entire or serrate, rarely lobed, faces glabrous or hairy, often gland-dotted. Heads usually radiate or sometimes discoid, borne singly or in corymbiform, paniculiform, or spiciform arrays. Involucres usually hemispheric, sometimes campanulate or cylindric. Phyllaries persistent, in 2–3+ series, subequal to unequal. Receptacles flat to slightly convex or conic, paleate; paleae conduplicate, usually rectangular-oblong, usually 3-toothed, sometimes entire, apices sometimes reddish or purplish. Ray florets usually 5–30, rarely 0, neuter; corollas usually yellow. Disc florets numerous, bisexual, fertile; corollas yellow or reddish, tubes shorter than campanulate throats, lobes 5, triangular. Achenes usually purplish black, sometimes mottled, ± obpyramidal, ± compressed; pappi 0 or readily falling, of 2(–3) usually lanceolate, aristate, or erose scales plus 0–8 usually shorter scales, 0.2–2 mm. x = 17.

Species ca. 52. North America, Mexico; two species introduced in China.

1a.       Annual herbs without tuber; heads large, ca. 10–30 cm in diam.; disc florets brown or purple  1. H. annuus

1b.       Perennial herbs with tuber; heads ca. 2–5 cm in diam.; disc florets yellow
                                                                                                                                             2. H. tuberosus

1. Helianthus annuus Linnaeus, Sp. Pl. 904. 1753.

向日葵  xiang ri kui

Helianthus annuus subsp. jaegeri (Heiser) Heiser; H. annuus subsp. lenticularis (Douglas ex Lindley) Cockerell; H. annuus var. lenticularis (Douglas ex Lindley) Steyermark; H. annuus var. macrocarpus (Candolle) Cockerell; H. annuus subsp. texanus Heiser; H. aridus Rydberg; H. jaegeri Heiser; H. lenticularis Douglas ex Lindley; H. macrocarpus Candolle.

Annuals, 100–300 cm. Stems erect, usually hispid. Leaves mostly cauline; mostly alternate; petioles 2–20 cm; blades lance-ovate to ovate, 10–40 Χ 5–40 cm, bases cuneate to subcordate or cordate, margins serrate, abaxial faces usually ± hispid, sometimes gland-dotted. Heads 1–9. Peduncles 2–20 cm. Involucres hemispheric or broader, 15–40(–200) mm diam. Phyllaries 20–30(–100), ovate to lance-ovate, 13–25 Χ (3–)5–8 mm, (margins usually ciliate) apices abruptly narrowed, long-acuminate, abaxial faces usually hirsute to hispid, rarely glabrate or glabrous, usually gland-dotted. Paleae 9–11 mm, 3-toothed (middle teeth long-acuminate, glabrous or hispid). Ray florets (13–)17–30(–100+); laminae 25–50 mm. Disc florets 150–1000; corollas 5–8 mm, throats bulbous at bases, lobes usually reddish, sometimes yellow; anthers brownish to black, appendages yellow or dark. Achenes (3–)4–5(–15) mm, glabrate. Pappus of 2 lanceolate scales 2–3.5 mm plus 0–4 obtuse scales 0.5–1 mm. Fl. Jul–Sep, fr. Aug–Oct. 2n = 34.

Open areas; widely cultivated in China [native to North America].

Helianthus annuus is a major agronomic crop, intoduced nearly worldwide.

2. Helianthus tuberosus Linnaeus, Sp. Pl. 905. 1753.

菊芋  ju yu

Helianthus tomentosus Michaux; H. tuberosus var. subcanescens A. Gray.

Perennials, 50–200 cm, rhizomatous, producing tubers late in growing season. Stems erect, scabro-hispid to hirsute, sometimes glaucous. Leaves mostly cauline; opposite or alternate proximally, usually alternate distally; petioles 2–8 cm (often ± winged); blades (3-nerved from near bases) lanceolate to ovate, 10–23 Χ 7–15 cm, bases broadly to narrowly cuneate, margins entire or serrate, abaxial faces puberulent or hirsutulous to tomentulose and gland-dotted or ± scabrous. Heads 3–15. Peduncles 1–15 cm. Involucres hemispheric, (10–25 Χ) 8–12 mm. Phyllaries often dark green, drying nearly black, 22–35, subequal, bases appressed, apices ± spreading, sometimes reflexed in fruit, lanceolate, 8.5–15 Χ 2–4 mm, margins ciliate, apices acuminate, abaxial faces hispidulous or puberulent, gland-dotted. Paleae 8–9 mm, 3-toothed, apices hairy. Ray florets 10–20; laminae 25–40 mm. Disc florets 60+; corollas 6–7 mm, lobes yellow; anthers dark brown or black, appendages dark or yellowish. Cypselae 5–7 mm, glabrous or distally hairy. Pappus of 2 aristate scales 1.9–3 mm plus 0–1 deltate scales 0.5–0.8 mm. Fl. Aug–Sep. 2n = 102.

Roadsides, fields, waste areas; widely cultivated in China [native to North America].

Helianthus tuberosus is a vegetable crop and widely naturalized.

20. Acmella Richard in C. H. Persoon, Syn. Pl. 2: 472. 1807.

金钮扣属  jin niu kou shu

Spilanthes Jacquin sect. Acmella (L. C. Richard) Candolle.

Annual or perennial herbs. Leaves opposite and/or basally rosulate. Heads solitary or in cymes, radiate, disciform, or discoid. Involucres ± hemispheric to ovoid; phyllaries in one to three rows, subequal or with outer row spreading and longer, entire or irregularly dentate. Receptacles conic, paleate; paleae falling with fruit, ± navicular, membranous to scarious, each about equaling subtended floret. Ray florets, when present, 2- or 3-lobed, variously colored; disc florets 4- or 5-lobed,corolla yellow or orange, 4–5-lobed. Achenes ciliate, glabrous or sometimes with corky margins; ray achenes broadly ovate or elliptic, 3-angled; disc achenes ellipsoid, strongly compressed. Pappus absent or of up to 10 awnlike bristles. x = 13.

About 30 species: pantropical; three species in China.

1a.       Perennial herbs; leaf margin peaked serrate .......................................................  1. A. calva

1b.       Annual herbs; leaf margin dentate.

2a.       Leaves broadly ovate to deltate, 5–10 cm long; heads 10.5–23.5 mm tall
.................................................................................................................  2. A. oleracea

2b.       Leaves ovate-lanceolate to ovate, 2–4 cm long; heads 8.4–12.5 mm tall
..............................................................................................................  3. A. paniculata

1. Acmella calva (Candolle) R. K. Jansen, Syst. Bot. Monogr. 8: 41. 1985.

美形金钮扣  Mei xing jin niu kou

Spilanthes calva Candolle in Wight, Contr. Bot. Ind. 19. 1834; S. acmella var. calva (Candolle) Clarke ex J. D. Hooker; S. callimorpha A. H. Moore.

Perennial herbs. Stems creeping or prostrate, 20–60 cm long, glabrous, rooting at nodes, sparsely pilose. Leaves lanceolate, 3–7 Χ 1–3 cm, apex acuminate or caudate, base cuneate, margin peaked serrate, adaxilly pubescent, abxailly nearly glabrous or only pubescent along veins; petioles 5–8(–24) mm long, pubescent. Heads ovoid-conoid, 9–11(–14) mm long, 6–8 mm in diam.; peduncles 3–14 cm. Phyllaries ca. 8, in two series, subequal, green, ovate-oblong, 3–3.5 mm long, apex acute or obtuse, margin ciliate. Receptacle columned-conoid, 4–8 mm long, with oblong, navicular, membranous paleae. Flower yellow; ray florets female, ca. 4 mm long, ligules short, obovate, apex shallowly 3-lobed; disc florets bisexual florets tubular, ca. 2 mm long, 4–5-toothed. Achenes oblong, 1.5–2 mm long, brown, pubescent or glabrous. Pappus 2, arista-like. Fl. May–Dec.

Streamside, marshes, moist forest margin, fields; 1000–1900 m. Yunnan [India, Indonesia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand].

2. Acmella oleracea (Linnaeus) R. K. Jansen, Syst. Bot. Monogr. 8: 65. 1985.

桂圆菊  gui yuan ju

Spilanthes oleracea Linnaeus, Syst. Nat. 2: 534. 1767.

Plants annual. Stems decumbent to usually erect, not rooting at nodes, green to red, glabrous. Petiole 2–6.4 cm long, glabrous to very sparsely pilose, narrowly winged. Leaf blade 5–10 Χ 4–8 cm, broadly ovate to deltate, base truncate to short attenuate, apex short acuminate to usually acute, usually glabrous on both surfaces, margins dentate. Heads discoid, 10.5–23.5 mm tall, 11–17 mm in diam.; peduncles 3.5–12.5 cm long, glabrous to very sparsely pilose. Receptacle 8.3–21.5 mm tall, 3.5–8.5 mm in diam. Involucral bracts 15–18, triseriate, herbaceous, entire to sinuate, sparsely ciliate; bracts of outer series 5 or 6, 5.8–7.3 Χ 2.1–2.8 mm, usually narrowly ovate to lanceolate or sometimes ovate, acute. Disc florets 400–600; corolla 2.7–3.3 mm long, yellow, 5-merous; tube 0.5–0.7 mm long, 0.2–0.4 in diam.; lobes 0.3–0.6 Χ 0.2–0.4 mm; stamens 1.4–1.7 mm long; achenes 2–2.5 Χ 0.9–1.1 mm, moderately to densely ciliate with straight-tipped hairs. Pappus of 2 subequal bristles, longer one 0.5–1.5 mm long, shorter one 0.3–1.3 mm long.

Cultivated in Southern China and Taiwan [Known only from cultivation and escaped in weedy habitats].

Acmella oleracea is cultivated for medicinal, insecticidal and horticultural purposes.

3. Acmella paniculata (Wallich ex Candolle) R. K. Jansen, Syst. Bot. Monogr. 8: 67. 1985.

金钮扣  Jin niu kou

Spilanthes paniculata Wallich ex Candolle, Prodr. 5: 125. 1836; S. acmella var. paniculata (Wallich ex Candolle) J. D. Hooker; S. acmella auct. non (Linneaus) J. A. Murray, Li, Fl. Taiwan 4: 945. 1978.

Annual herbs. Stems branched, erect or ascending, to 30 cm or more tall, rarely rooting at nodes. Leaves petiolate, ovate to ovate-lanceolate, 2–4 cm long, 1–2.5 cm broad, apex acute, base cuneate, 3-nerved, entire, coarsely or crenately serrate; petiole 1–2 cm long. Heads discoid, solitary, terminal or axillary, 8.4–12.5 mm tall, 6.9–10 mm in diam.; peduncles 2.5–16 cm long, sparsely pilose. Receptacle 5–8 mm tall, 1.1–3 mm in diam., apex acuminate. Involucral bracts 9–12, in 2-series, ovate-lanceolate, ca. 6 mm long, herbaceous, glabrous. Disc florets 90–200; tubular, minute, 4- or 5-lobed. Achenes obovoid, 3-angled, ca. 3 mm long, apex slightly depressed, margins scabrid. Pappus of 2 subequal bristles, longer one 0.5–1.1 mm long, shorter one 0.4–0.9 mm long. Fl. Apr–Nov. 2n=26.

Fields, waste places, roadside, forest margin; 800–1900 m. Guangdong, Guangxi, Yunnan, Taiwan [India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].

21. Synedrella Gaertner, Fruct. Sem. Pl. 2: 456, plate 171, fig. 7. 1791.

金腰箭属  jin yao jian shu

Annual herbs. Stems erect or ascending. Leaves opposite, petiolate. Heads radiate, sessile or subsessile, in axillary glomerules or borne singly. Involucres cylindric to campanulate. Phyllaries persistent, 2–5+ in 1(–2) series. Receptacles convex, paleate. Ray florets in 1–2 series, pistillate, fertile; corollas yellowish. Disc florets bisexual, fertile; corollas yellowish, tubes about equaling slightly ampliate, cylindric throats, lobes 4, rounded-deltate. Cypselae dimorphic, strongly obcompressed or flattened, narrowly oval and winged or linear-cuneate and not winged; pappi persistent, of 2 triangular scales (ray) or 2–3 subulate scales or awns (disc).

Two species, Mexico, West Indies, Central America, South America, introduced in Asia, Africa, Pacific Islands, Australia; one species in China.

1. Synedrella nodiflora (Linnaeus) Gaertner, Fruct. Sem. Pl. 2: 456. 1791.

金腰箭  Jin yao jian

Verbesina nodiflora Linnaeus, Cent. Pl. I, 28. 1755.

Annuals, 10–80 cm high. Stems erect or ascending, branched from bases or ± throughout. Leaves cauline; opposite; petiolate; blades ovate to elliptic, 3–10 Χ 2–5 cm. usually 3-nerved, bases cuneate to rounded, margins toothed, faces ± scabrous. Heads radiate, sessile or subsessile in axillary glomerules or borne singly. Involucres cylindric to campanulate, 3–6 mm diam. Phyllaries persistent, 2–5+ in 1(–2) series, lanceolate, herbaceous to chartaceous. Receptacles convex, paleate; paleae lance-linear, scarious, flat or weakly cupped at bases. Ray florets 2–9, in 1–2 series, pistillate, fertile; corollas yellowish, ca. 2 mm long; ligules ovate to linear, 2–4 mm long. Disc florets 4–12+, bisexual, fertile; corollas yellowish, tubes ca. 4 mm long, about equaling slightly ampliate, cylindric throats, lobes 4, rounded-deltate. Flowering year round. 2n = 40.

Fields, waste places, roadsides. Guangdong, Taiwan, Yunnan [pantropical weed of American origin].

22. Calyptocarpus Lessing, Syn. Gen. Compos. 221. 1832.

金腰箭舅属  jin yao jian jiu shu

Small annual or perennial herbs, usually procumbent or prostrate. Leaves opposite, petiolate, margin crenate-serrate. Heads solitary or in small dense clusters, radiate. Involucreal bracts ca. 5. Receptacle with flat or concave paleae. Ray florets 5–8, pistillate, yellow; disc florets 4–5-lobed, yellow. Achenes oblanceolate-obconical, doriventrally compressed, tubuerculate or smooth. Pappus of 2 stout awns.

Three species in South America to North America; one species naturalized in China.

1. Calyptocarpus vialis Lessing, Syn. Gen. Compos. 221. 1832.

金腰箭舅  jin yao jian jiu

Perennial herbs. Stems prostrate, branched, rooting at nodes, densely appressed strigillose. Leaves ovate to broadly ovate, to 35 Χ 25 mm, apex acute, apiculate, base attenuate, margin crenate-serrate, both surfaces densely appressed strigillose; petiol 3–8 mm long, narrowly winged toward blade, margin ciliate. Heads axillary, solitary, subsessile, peduncle to 15 mm long. Involucre narrowly oblong-oblanceoloid, 6–7 Χ 2.5–3.5 mm. Phyllaries 4, in 2 series, concave, lanceolate, apex acuminate and apiculate, 6–7 Χ 3–3.5 mm. Paleae hyaline, narrowly elliptic, 3.5–4.8 Χ 0.8–1 mm. Ray florets 3–8, yellow; corolla 4.5–6.2 mm long, 1.4–1.8 mm diameter, 3-toothed; achenes oblanceolate, 3.5–4 mm long, 1.7–2 mm wide, pappus of 2 ascending divergent awns, 1.2–2.4 mm long. Disc florets 3–8; corolla 2.6–3.6 mm long, apex 2–3-toothed or entire, densely papillose within; achenes similar to those of ray florets but slightly narrower and thicker, sometimes 3-angled, 1.1–1.7 mm across, evidently muricate. 2n = 24.

Naturalized weed of roadsides. Taiwan [native to USA, Mexico and Cuba].

23. Coreopsis Linnaeus, Sp. Pl. 907. 1753

金鸡菊属  jin ji ju shu

Annuals, perennials, subshrubs, or shrubs, often rhizomatous or with cormiform bases. Stems usually 1, erect, branched distally or ± throughout. Leaves basal, basal and cauline, or cauline; opposite or alternate or both; petiolate or sessile; blades simple and usually entire, or pinnately or pedately lobed, faces glabrous or hairy. Heads radiate, borne singly or in open, corymbiform arrays. Calyculi of (3–)8+, distinct, ± herbaceous bractlets. Involucres ± globose to cylindric, 4–25+ mm diam. Phyllaries usually ca. 8 in ca. 2 series. Receptacles flat to convex, paleate; paleae falling, ovate to linear or subulate, flat, scarious. Ray florets mostly (5–)8(–12+), neuter, or styliferous and sterile, or pistillate and fertile; corollas usually yellow, sometimes red-brown to purple proximally, sometimes wholly purple or pink to white. Disc florets 8–150+, bisexual, fertile; corollas usually yellow, sometimes red-brown to purple at tips or throughout, tubes equaling or shorter than narrowly funnelform throats, lobes 4 or 5. Cypselae obcompressed, ± orbiculate to ovate, oblong, or linear, usually thin-margined or winged, wings membranous to chartaceous or corky, entire or lobed to toothed, sometimes ciliolate; faces smooth or ± papillate to tuberculate. Pappus 0, or persistent, of 2 bristly cusps or scales, sometimes pappi 0 and shoulders of cypsela wings bristly, pappus-like. x = 14.

About 35 species: mostly temperate North America, also tropical New World and Old World; three species introduced in China.

1a.       Ray florets above yellow, base reddish brown; disc florets reddish brown; achenes wingless  1. C. tinctoria

1b.       Ray florets and disc florets yellow; achenes winged.

2a.       Achenes broadly elliptic or nearly rounded; lower pinnately dissected
............................................................................................................  2. C. grandiflora

2b.       Achenes rounded; lower leaves entire ....................................................  3. C. lanceolata

1. Coreopsis tinctoria Nuttall, J. Acad. Nat. Sci. Philadelphia. 2: 114. 1821.

两色金鸡菊  liang se jin ji ju

Annuals, (10–)30–70(–150) cm. Leaves: proximal blades usually 1(–3)-pinnate, terminal lobes lance-ovate to oblanceolate, 10–60 Χ 5–25 mm; cauline blades usually 1–3-pinnate, rarely simple, simple blades or terminal lobes lance-linear to linear or filiform, 10–45 Χ 0.5–5 mm. Peduncles 1–15 cm. Calyculi of deltate-lanceolate bractlets 1–3+ mm. Phyllaries ± lance-oblong to lance-ovate, 4–9 mm. Ray laminae usually yellow with red-brown blotch, sometimes red-brown in proximal 1/3–9/10 and distally yellow, rarely yellow throughout, 12–18+ mm. Disc corollas 2.5–3+ mm. Cypselae 1.5–4 mm, wings 0 or 0.1–0.7 mm wide; pappi 0, or of 1–2 cusps or subulate scales 0.1–1 mm. Fl. Jun–Aug. 2n = 24.

Moist, sandy or clay soils, sometimes alkaline flats, prairies, ditches, disturbed places [native to North America].

Coreopsis tinctoria is widely grown in public and residential gardens, and commercially (for cut flowers), and has become widely established in the flora area.

2. Coreopsis grandiflora Hogg ex Sweet, Brit. Fl. Gard. 2: plate 175. 1826.

大花金鸡菊  da hua jin ji ju

Perennials, 40–60 cm. Aerial nodes proximal to first peduncle usually 6–10+, distalmost 1–3 internodes 4–7+ cm. Leaves: mostly cauline on proximal 2/3–7/8 of plant heights; petioles 0 or 10–35+ mm; blades usually 1(–2)-irregularly pinnately or ± pedately lobed with (3–)5–9+ lobes, rarely simple, simple blades or terminal lobes narrowly lanceolate to linear or filiform, 15–45(–90+) Χ (0.5–)2–8(–12+) mm. Peduncles 8–15(–25+) cm. Calyculi of lanceolate to linear bractlets 3.5–9+ mm. Phyllaries lance-ovate, 7–9(–12) mm. Ray laminae yellow, 12–25+ mm. Disc corollas 3.3–4.8 mm, apices yellow. Cypselae 2–3+ mm, wings spreading, ± chartaceous, entire or irregularly toothed to pectinate. Flowering May–Aug. 2n = 26.

Sandy soils, ditches and roadsides, other disturbed sites, granite and sandstone outcrops [native to North America].

3. Coreopsis lanceolata Linnaeus, Sp. Pl. 908. 1753.

剑叶金鸡菊  jian ye jin ji ju

Perennials, 10–30(–60+) cm. Aerial nodes proximal to first peduncle usually 1–3(–5+), distalmost 1–3 internodes 1–2(–8+) cm. Leaves: basal and cauline on proximal 1/4–1/3(–1/2) of plant heights; petioles 1–5(–8+) cm; blades simple or with 1–2+ lateral lobes, simple blades or terminal lobes lance-ovate or lanceolate to oblanceolate or lance-linear, 5–12 cm Χ 8–15(–18+) mm. Peduncles (8–)12–20(–35+) cm. Calyculi of lance-ovate to lance-linear or linear bractlets 4–8(–12) mm. Phyllaries deltate to lance-deltate, 8–12+ mm. Ray laminae yellow, 15–30+ mm. Disc corollas 6–7.5 mm, apices yellow. Achenes 2.6–4 mm long, wings ± spreading, ± chartaceous, entire. Flowering May–Jul. 2n = 26.

Sandy soils, ditches and roadsides, other disturbed sites [native to North America].

24. Dahlia Cavanilles, Icon. 1: 56 (t. 80). 1791

大丽花属  da li hua shu

Perennial herbs with rhizomes, developing elongated tubers. Leaves opposite, pinnatisect with toothed (or incised and parted) segments; sometimes simple leaves are also present on the same plant stems; as also the entire plant, glabrous. Heads large, usually drooping, long pedunculate. Involucre, consisting of an external row of recurvate herbaceous few bracts, basally, more or less, united, membranous at apex, occasionally arranged almost in two rows. Receptacle flat with large membranous bracts. Marginal ligulate florets pistillate or sterile; internal ones tubular, bisexual; forms with double-flowers are not rare. Achenes flattened, cuneate, without pappus or with its traces in the form of two denticules.

About 15 species: South America, Central America, Mexico; one species cultivated in China.

1. Dahlia pinnata Cavanilles, Icon. 1: 57. t. 80.1791.

大丽花  da li hua

Perennial. Stems 1.5–2.0 m high, erect, more or less, woody at base, hollow or unbranched. Leaves pinnate (sometimes bipinnate), ovate, oblong-ovate, often tapered at apex, margin serrate-toothed; bracts usually 5 in nember. Heads large, long-pedunculate, drooping. Ligulate florets arranged in one or a few rows, or all florets ligulate (in double-forms), variously colored, sometimes fancifully variable (convolute into tube or trulliform etc.). Achenes ??ate, 9–12 Χ 3–4 mm, or obascidiform, flattened. Fl. Jun–Dec, fr. Sep–Oct.

Widely cultivated in China [native to Mexico].

Dahlia pinnata is an important ornamental plant, and widely cultivated worldwide.

25. Cosmos Cavanilles, Icon. 1: 9, plate 14. 1791.

秋英属  qiu ying shu

Annuals, perennials or subshrubs. Stems usually 1, erect or ascending, branched distally or ± throughout. Leaves mostly cauline; opposite; petiolate or sessile; blades usually 1–3-pinnately lobed or undivided, ultimate margins usually entire, faces usually glabrous, sometimes glabrate, hispid, puberulent, or scabridulous. Heads radiate, borne singly or in corymbiform arrays. Calycles of (5–)8 basally connate, ± linear to subulate, herbaceous bractlets. Involucres hemispheric or subhemispheric, 3–15 mm diam. Phyllaries persistent, (5–)8 in ± 2 series, distinct, lanceolate, lance-oblong, lance-ovate, or oblong, ± equal, membranous or herbaceous, margins ± scarious. Receptacles flat, paleate; paleae falling, linear, flat or slightly concave-convex, scarious or entire. Ray florets neuter; corollas white to pink or purple, or yellow to red-orange. Disc florets bisexual, fertile; corollas yellow or orange, tubes shorter than funnelform throats, lobes 5, ± deltate. Achenes relatively slender, quadrangular-cylindric or -fusiform, sometimes slightly arcuate, attenuate-beaked, not winged [winged], faces glabrous or hispid to scabridulous or ± setose, sometimes papillate, usually with 1 groove; pappi of 2–4(–8) retrorsely barbed awns, sometimes 0. x = 12.

About 26 species: tropical and subtropical America, especially Mexico, widely introduced elsewhere; two species introduced in China.

1a.       Ray laminae pink, purple, purplish, rose-pink, violet, or white ...................  1. C. bipinnatus

1b.       Ray laminae yellow to red-orange ............................................................  2. C. sulphureus

1. Cosmos bipinnatus Cavanilles, Icon. 1: 10, plate 14. 1791.

秋英  qiu ying

Plants 30–200 cm high, glabrous or sparsely puberulent, sometimes scabridulous. Leaves sessile or with up to 1 cm long petioles; blades 6–11 cm long, ultimate lobes to 1.5 mm wide, margins entire, apices acute. Head solitary, 3–6 cm in diam.; peduncles 10–20 cm long. Calycles of spreading, linear to lanceolate bractlets 6–13 mm, apices acuminate. Involucres 7–15 mm diameter. Phyllaries erect, lanceolate to ovate-lanceolate, 7–13 mm, apices round or obtuse. Ray corollas white, pink, or purplish, laminae obovate to oblanceolate, 15–50 mm, apices ± truncate, dentate. Disc corollas 5–7 mm. Achenes 7–16 mm long, glabrous, papillose. Pappus 0, or of 2–3 ascending to erect awns 1–3 mm. Fl. Jun–Aug. 2n = 24.

Widely introduced in China [native to Mexico and southwestern United States].

A favorite garden plant, it has escaped and naturalized widely in warm climates almost worldwide.

2. Cosmos sulphureus Cavanilles, Icon. 1: 56, plate 79. 1791.

硫磺菊  liu huang ju

Plants 30–200 cm high, glabrous or sparsely pilose to hispid. Leaves petiolate; petioles 1–7 cm; blades 5–12(–25) cm, ultimate lobes 2–5 mm wide, margins sparsely spinulose-ciliate, apices apiculate. Peduncles 10–20 cm. Calycles of spreading-ascending, linear-subulate bractlets 5–7(–10) mm, apices acute. Involucres 6–10 mm diam. Phyllaries erect, oblong-lanceolate, 9–13(–18) mm, apices acute to rounded-obtuse. Ray corollas intensely yellow to red-orange, laminae obovate, 18–30 mm, apices ± truncate, denticulate. Disc corollas 6–7 mm. Achenes 15–30 mm long, usually hispidulous, rarely glabrous. Pappus 0, or of 2–3 widely divergent awns 1–7 mm. Fl. Jun–Sep. 2n = 24, 48.

Introduced in Beijing, Guangdong, Yunnan [native to Mexico].

26. Bidens Linnaeus, Sp. Pl. 831. 1753

鬼针草属  gui zhen cao

Annuals or perennials. Stems usually erect, branched distally or throughout. Leaves usually cauline; usually opposite, rarely whorled, distal sometimes alternate; petiolate or sessile; blades simple, compound (leaflets petiolulate), or 1–3-pinnatisect or -pinnately lobed, ultimate margins entire, dentate, laciniate, serrate, or toothed, faces usually glabrous, sometimes hirtellous, hispidulous, pilosulous, puberulent, scabrellous, or strigillose. Heads usually radiate or discoid, sometimes disciform, usually in corymbiform arrays, sometimes in 2’s or 3’s or borne singly. Calycles erect to spreading or reflexed, herbaceous bractlets or bracts. Involucres mostly hemispheric or campanulate to cylindric. Phyllaries persistent, mostly in 2 series, usually distinct, sometimes connate 0.05–0.1 Χ their lengths, mostly oblong or ovate to lance-oblong, chartaceous to membranous or scarious (usually striate with brownish nerves, margins usually hyaline). Receptacles flat or slightly convex, paleate; paleae usually falling, (usually stramineous, sometimes yellow to orange, with darker striae) flat to slightly navicular. Ray florets usually in one series, usually neuter, sometimes styliferous and sterile; corollas usually yellow, sometimes white or pinkish. Disc florets bisexual, fertile; corollas usually yellow to orange, sometimes whitish or purplish, tubes shorter than throats, lobes (3–)5, deltate (staminal filaments glabrous; style-branch appendages deltate or lanceolate to subulate). Achenes usually obcompressed to flat, unequally 3–4-angled, and cuneate to oblanceolate or obovate, sometimes (all or inner) equally 4-angled and linear-fusiform, rarely subterete, margins usually retrorsely, sometimes patently or antrorsely, barbed or ciliate, apices sometimes attenuate, not beaked, faces smooth, striate, or tuberculate, glabrous or hairy, each sometimes with 2 grooves; pappi 0, or persistent, of (1–)2–4(–8) usually retrorsely, sometimes antrorsely, barbellate or ciliate, rarely smooth, awns. x = 12.

150–250 species: widespread, especially in subtropical, tropical, and warm-temperate North America and South America; ten species in China.

1a.       Achenes broad, cuneate or obovate-cuneate, apex truncate.

2a.       Achenes 4-angled, barbed awns usually 4; disc floret corolla 5-dentate; ray florets present  1. B. cernua

2b.       Achenes compressed, barbed awns usually 2; disc floret corolla 4–5-dentate; ray florets absent.

3a.       Median cauline leaves of pinnate compound leaves .................................  2. B. frondosa

3b.       Median cauline leaves of dissected simple leaves, rarely entire leaves.

4a.       Leaves deeply dissected.

5a.       Heads subequal in length and wide; outer phyllaries 5–9; achenes 6–11 mm
                                                                                                           3. B. tripartita

5b.       Heads unequal in length and wide; outer phyllaries 9–14; achenes 3–4.5 mm  4. B. maximowicziana

4b.       Leaves not dissected or 3-lobed.

6a.       Outer phyllaries 9–12(–14), oblong-linear or lanceolate-linear, 8–20 mm long; corolla 4-toothed  5. B. radiata

6b.       Outer phyllaries usually 4, elliptic, oblong or linear, 1.5–3.8 cm long; corolla 5-toothed  6. B. leptophylla

1b.       Achenes linear, apex gradually narrower.

7a.       Achenes with 2 barbed awns; disc corolla 4-toothed ...............................  7. B. parviflora

7b.       Achenes with 3–4 barbed awns; disc corolla 5-toothed.

8a.       Involucral bracts spatulate, dilated toward apex; achenes numerous, ca. 50–70
                                                                                                                     8. B. pilosa

8b.       Involucral bracts linear, not dilated toward apex; achenes fewer than 50.

9a.       Leaves 2–3 pinnatisect, leaflet lanceolate, terminal segment narrow, few-toothed  9. B. bipinnata

9b.       Leaves 2–3 pinnatisect, leaflet lanceolate, terminal segment narrow, few-toothed  10. B. biternata

1. Bidens cernua Linnaeus, Sp. Pl. 832. 1753.

柳叶鬼针草  liu ye gui zhen cao

Bidens cernua var. elliptica Wiegand; B. cernua var. integra Wiegand; B. cernua var. minima (Hudson) Pursh; B. cernua var. oligodonta Fernald & H. St. John; B. cernua var. radiata Candolle; B. filamentosa Rydberg; B. glaucescens Greene; B. gracilenta Greene; B. minima Hudson; B. prionophylla Greene.

Annuals, 10–100 cm high. Leaves sessile; blades lance-ovate or oblanceolate to lanceolate or linear, 40–100(–200) Χ (2–)5–25(–45) mm, bases cuneate to rounded, margins usually coarsely dentate to serrate, sometimes entire, sometimes ciliate, apices acute to acuminate, faces glabrous. Heads borne singly or in open, corymbiform arrays. Peduncles 10–40(–100+) mm. Calycles of (3–)5–8(–10) spreading to reflexed, oblong to lance-linear, often ± foliaceous bractlets or bracts (3–)8–12(–25) mm, margins usually ciliate, abaxial faces usually glabrous, bases sometimes hispidulous. Involucres hemispheric or broader, (3–)6–10 Χ (8–)12–20+ mm. Phyllaries 6–8+, ovate or lance-ovate to lanceolate, 2–10 mm. Ray florets usually 6–8, sometimes 0; laminae orange-yellow, 2–15(–18) mm. Disc florets (10–)40–100(–150+); corollas orange-yellow, 3–4 mm. Achenes blackish or brown, usually flattened, sometimes 4-angled, cuneate, outer (3–)5–6 mm, inner 4–8 mm, margins thickened or winged, retrorsely ciliate, apices truncate to convex, faces ± striate, glabrous or tuberculo-strigillose; pappi of (2–)4, retrorsely barbed awns (1–)2–4 mm. Fl. Aug–Oct. 2n = 24, 48.

Swamps, marshes, peat and sedge bogs, flood plains; 0–2300 m. Sichuan, Xizang,Yunnan [Asia, Europe; North America].

2. Bidens frondosa Linnaeus, Sp. Pl. 832. 1753.

大狼杷草  da lang ba cao

Bidens frondosa var. anomala Porter ex Fernald; B. frondosa var. caudata Sherff; B. frondosa var. pallida (Wiegand) Wiegand; B. frondosa var. stenodonta Fernald & H. St. John; B. melanocarpa Wiegand.

Annuals, 20–120 cm high. Leaves petiolate; petioles 10–40(–60) mm; blades deltate to lance-ovate overall, 30–80(–150) Χ 20–60(–100) mm, 3(–5)-foliolate, leaflets petiolulate, lanceolate to lance-ovate, (15–)35–60(–120) Χ (5–)10–20(–30) mm, bases cuneate, margins dentate to serrate, sometimes ciliate, apices acuminate to attenuate, faces glabrous or hirtellous. Heads usually borne singly, sometimes in 2s or 3s or in open, corymbiform arrays. Peduncles 10–40(–80) mm. Calycles of (5–)8(–10) ascending to spreading, spatulate or oblanceolate to linear, sometimes ± foliaceous bractlets or bracts 5–20(–60) mm, margins usually ciliate, abaxial faces glabrous or hirtellous. Involucres campanulate to hemispheric or broader, 6–9 Χ 7–12 mm. Phyllaries 6–12, oblong or ovate to lance-ovate, 5–9 mm. Ray florets 0 or 1–3+; laminae golden yellow, 2–3.5 mm. Disc florets 20–60(–120+); corollas ± orange, 2.5–3+ mm. Cypselae blackish to brown or stramineous, ± obcompressed, obovate to cuneate, outer 5–7 mm, inner 7–10 mm, margins antrorsely or retrorsely barbed, apices ± truncate to concave, faces usually 1-nerved, sometimes tuberculate, glabrous or sparsely hirtellous. Pappus of 2 ± erect to spreading, antrorsely or retrorsely barbed awns 2–5 mm. Fl. Aug–Sep. 2n = 24, 48, 72.

Moist woods, meadows, thickets, fields, roadsides, railroads, borders of streams, ponds, sloughs, swamps, ditches. Introduced in Shanghai [native to North America].

3. Bidens tripartita Linnaeus, Sp. Pl. 831. 1753.

狼杷草  lang bao cao

Bidens repens D. Don; B. tripartita Linnaeus var. quinqueloba Z. X. An, syn. nov.; B. tripartita Linnaeus var. repens (D. Don) Sherff; B. tripartita Linnaeus var. shimadai (Hayata) Yamamoto; B. shimadai Hayata.

Annuals, 10–150 cm high. Leaves sessile or petioles 5–15(–35) mm (± winged); blades elliptic to ovate or lanceolate, 40–80(–150) Χ 15–40(–60) mm, sometimes laciniately 1-pinnatisect with 1–4+ lobes near bases, bases cuneate, margins entire or dentate to serrate, usually ciliate, apices acute to acuminate, faces glabrous or hirtellous. Heads borne singly or in 2s or 3s. Peduncles 10–40(–80) mm. Calyculi of (2–)6–7(–10) spreading, oblanceolate or lanceolate to linear, ± foliaceous bractlets or bracts 7–35(–60) mm, margins (entire or serrate) sometimes sparsely ciliate, abaxial faces hispidulous near bases, distally glabrous. Involucres campanulate to hemispheric or broader, (4–)5–7(–12) Χ (3–)6–12(–15) mm. Phyllaries (6–)7–8(–13), elliptic-ovate to lance-ovate, (4–)6–9(–12) mm. Ray florets usually 0, sometimes 1–5; laminae orange yellowish, 4–8 mm. Disc florets (5–)20–60(–150); corollas pale yellow to orange, (2–)3–4 mm, apex 4–5-toothed. Achenes blackish to purplish or brown, ± flattened, sometimes weakly 3(–4)-angled, usually cuneate to linear, outer (3–)6–7(–10) mm, inner (4–)6–9(–11) mm, margins proximally antrorsely to patently, distally retrorsely, barbed, apices ± truncate to concave, faces ± 1-nerved, usually smooth, seldom notably tuberculate, glabrous or sparsely strigillose; pappi 0, or of 1–3(–4) erect to spreading, retrorsely barbed awns (0.2–)2–3(–6) mm. Fl. Jul–Oct. 2n = 48.

Marshes and other wet sites, waste fields, roadside. Anhui, Fujian, Gansu, Guizhou, Hebei, Henan, Heilongjiang, Hubei, Hunan, Jilin, Jiangsu, Jiangxi, Liaoning, Nei Mongol, Ningxia, Qinghai, Shandong, Shaanxi, Sichuan, Taiwan, Xizang, Xinjiang, Yunnan, Zhejiang [India, Indonesia, Japan, Korea, Malaysia, Mongolia, Nepal, Philippines, Russia; Australia; Europe; North America; N Africa].

A worldwide weed.

4. Bidens maximowicziana Oettingen, Act. Hort. Jurjev. 6: 219. 1906.

羽叶鬼针草  yu ye gui zhen cao

Annual. Stems 30–50(80) cm high, erect, simple or branched with obliquely upright branches, glabrous, or, sparsely and finely hairy mainly in upper part. Leavees petiolate, pinnati sect (rarely ternate); segments narrow and long, oblong to linear, laterals (1)2–3 in number, 2–3 to 7–8 cm long, apical segments usually larger than laterals, serrate-toothed, margin fine with, appressed prickles. Heads solitary or in groups of 2–3 at the apex of stem and its branches, flat (wider than long and measures about 15 mm). External involucral bracts green, oblong, distinctly exceeding the head; internal ones shorter, lanceolate, brown-green, as long as achenes (including awns). Ligulate florets absent, all florets tubular, yellow. Achenes cuneate, 4 mm long, laterally compressed, tuberculate at margin, and slightly hairy with relatively long and plentiful bristly hairs pointing downwards, awns two in number. Fl. Jul–Aug.

River banks, wet padings on meadows, humus sandy soil. Heilongjiang, Jilin, Liaoning, Nei Mongol [Japan, Korea, Russia].

5. Bidens radiata Thuillier, Fl. Par. ed. 2: 432. 1799.

大羽叶鬼针草  da yu ye gui zhen cao

Bidens radiata Thuillier var. microcephala C. X. An, syn. nov.

Annuals. Stems (5)15–60(80) cm high, erect, branched, in upper part simple, glabrous or indistinctly hairy. Leaves yellowish-green, 3–5-parted or dissected, lateral parts lanceolate or ovate-rhombical, serrate-toothed, but the apical part much larger than laterals, glabrous or subglabrous, on rather long petioles, sometimes entire, large-toothed to parted (f. pseudocernuus Ganessch.): Heads erect, 12–15(20) mm wide, width more than length (sometimes, almost twice). Outer phyllaries (9)10–12(14) in number, oblong-linear or lanceolate-linear, 8–20 mm long, foliaceous, margins beset with fine prickles, considerably (sometimes, almost three times) exceeding the head; internal ones shorter, oval; bracts narrow-linear, exceeding in length or equal to the achenes (including awns). Florets tubular. Achenes cuneate, 3–4 mm long, glabrous, compressed, with two awns, equal to or almost shorter than achenes; sometimes awns four, but then, two of them longer than the other two. Fl. Jun–Sep.

Banks of rivers, lakes, marshes, wet meadows, wet ditches. Heilongjiang, Jilin, Nei Mongol, Xinjiang [Japan, Korea, Mongolia, Russia; Europe].

6. Bidens leptophylla Z. X. An, Fl. Xinjiang. 5: 476. pl. 27, 1–2. 1999.

薄叶鬼针草  bao ye gui zhen cao

Annuals, up tp 25 cm high. Leaves opposite, blade ovate-elliptic, ovate-rhombic or oblong, 1.8–5.8 Χ 0.5–2 cm together with petioles, apex shortly acute, base decurrent to petioles, margin 1–3-toothed, sometimes with a pair of lobes from base; lobes oblong-linear. Heads solitary, terminal or axillary. Involucre 8–10 mm long, 5–8 mm in diam. Outer phyllaries usually 4, foliaceous, elliptic, oblong or linear, 1.5–3.8 cm long, apex acute, margin entire or 1–3-toothed; inner phyllaries coriaceous, brown. Ray florets absent. Disc florets yellow, corolla 3–3.5 mm long, apex 5-dentate. Achenes cuneate, ca. 6 mm long, ca. 2 mm wide, compressed, two-ribbed. Pappus of two barbed awns ca. 2.5 mm long.

* Farmland. Xinjiang (Yiwu).

7. Bidens parviflora Willdenow, Enum. Hort. Berol. 840. 1809.

小花鬼针草  xiao hua gui zhen cao

Annuals. Stems (10–)20–50(–80) cm long, erect, branched, glabrous or bearing rarely indistinct hairs. Leaves many-fold (bi, tri) pinnatisect with narrow lanceolate or linear-lanceolate to linear segments, which, in their turn, bear small teeth and incisions on petioles, usually sparsely fine hairy to subglabrous. Heads solitary (or 2–3) at apices of the stem and its branches, on long peduncles, narrow, subcylindrical. External involucral bracts narrow-linear, green, herbaceous, finely hairly, shorter than internal ones, lanceolate-linear, greenish-brown, bearing rarely short bristles only on margins; bracts linear, shorter than achenes. Florets all tubular, yellow, not numerous. Achenes linear-tetragonate, more or less, compressed, beset with upright lucid bristles along ribs, otherwise, glabrous or finely hairy, unequal in length; internal ones 15–20(25) mm long, external ones shorter, two upright awns, 4–5 mm long, at the top of achenes. Fl. Jul–Sep.

Marshes, fields, streamsides. Anhui, Gansu, Guizhou, Hebei, Henan, Heilongjiang, Jilin, Jiangsu, Liaoning, Nei Mongol, Ningxia, Qinghai, Shandong, Shanxi, Shaanxi, Shandong, Sichuan [Japan, Korea, Mongolia, Russia].

8. Bidens pilosa Linnaeus, Sp. Pl. 832. 1753.

鬼针草  gui zhen cao

Bidens chilensis Willdenow; B. pilosa Linnaeus var. minor (Blume) Sherff; B. pilosa Linnaeus var. radiata Schultz-Bipontinus; B. pilosa Linnaeus f. rubiflorum S. S. Ying; B. sundaicus Blume var. minor Blume.

Annuals. Stems 30–180 cm high, glabrous or very sparsely pubescent in upper part. Leaves: petioles 10–30(–70) mm; blades either ovate to lanceolate, 30–70(–120) Χ 12–18(–45) mm, or 1-pinnately lobed, primary lobes 3–7, ovate to lanceolate, (10–)25–80 Χ (5–)10–40 mm, bases truncate to cuneate, ultimate margins serrate or entire, usually ciliate, apices acute to attenuate, faces pilosulous to sparsely hirtellous or glabrate. Heads usually borne singly, sometimes in open, ± corymbiform arrays. Peduncles 10–20(–90) mm. Calycles (6–)7–9(–13), appressed, spatulate to linear bractlets (3–)4–5 mm, margins ciliate, abaxial faces usually hispidulous to puberulent. Involucres turbinate to campanulate, 5–6 Χ 6–8 mm. Phyllaries (7–)8–9(–13), lanceolate to oblanceolate, 4–6 mm. Ray florets 0 or (3–)5–8; ligules whitish to pinkish, 5–15 mm long. Disc florets 20–40(–80); corollas yellowish, (2–)3–5 mm. Achenes: outer red-brown, ± flat, linear to narrowly cuneate, (3–)4–5 mm, margins antrorsely hispidulous, apices truncate or somewhat attenuate, faces obscurely 2-grooved, sometimes tuberculate-hispidulous; inner blackish, ± equally 4-angled, linear-fusiform, 7–16 mm, margins antrorsely hispidulous, apices attenuate, faces 2-grooved, tuberculate-hispidulous to sparsely strigillose; pappi 0, or of 2–3(–5), erect to divergent, retrorsely barbed awns (0.5–)2–4 mm. Flowering year round. 2n = 24, 36, 48, 72.

Roadside, fields, village; up to 2500 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Hebei, Henan, Hunan, Hubei, Jiangxi, Liaoning, Shandong, Shanxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Japan; tropical and subtropical regions].

9. Bidens bipinnata Linnaeus, Sp. Pl. 832. 1753.

婆婆针  po po zhen

Bidens pilosa var. bipinnata (Linnaeus) Hooker.

Annuals, (15–)30–100(–150+) cm. Leaves: petioles 20–50 mm; blades rounded-deltate to ovate or lanceolate overall, (20–)30–70+ Χ (20–)30–60+ mm, (1–)2(–3)-pinnatisect, ultimate lobes obovate or lanceolate, 15–45+ Χ 10–25+ mm, bases truncate to cuneate, ultimate margins entire, sometimes ciliolate, apices rounded to acute or attenuate, faces usually glabrous, sometimes hirtellous. Heads usually borne singly, sometimes in ± corymbiform arrays. Peduncles (10–)20–50(–100) mm. Calyculi of (7–)8(–10) linear bractlets 3–5 mm, ± appressed, margins ciliate, abaxial faces usually glabrous. Involucres ± campanulate, 5–7 Χ 3–4(–5) mm. Phyllaries 8–12, lanceolate to linear, 4–6 mm. Ray florets 0 or 3–5+; laminae yellowish or whitish, 1–2(–3) mm. Disc florets 10–20(–30+); corollas yellowish to whitish, 2–3 mm. Cypselae red-brown, outer weakly obcompressed, 7–15 mm, inner ± 4-angled, linear to linear-fusiform, 12–18 mm, margins not ciliate, apices ± attenuate, faces 2-grooved, often tuberculate-hispidulous. Pappus of (2–)3–4, erect to divergent, retrorsely barbed awns 2–4 mm. Fl. Aug–Oct. 2n = 24, 72.

Fields, forests, disturbed, wettish sites; up to 1800(–3000) m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Hebei, Jilin, Jiangsu, Jiangxi, Liaoning, Nei Mongol, Shandong, Shanxi, Shaanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Korea; North America; South America; Europe; Pacific Islands].

10. Bidens biternata (Loureiro) Merrill & Sherff, Bot. Gaz. 88: 293. 1829.

金盏银盘  jin zhan yin pan

Coreopsis biternata Loureiro, Fl. Cochinch. 508. 1790; Bidens chinensis Willdenow; B. robertianifolia H. Lιveillι & Vaniot.

Annuals. Stems erect, 30–150 cm high, subtetragonous, loosely crisp pilose, branched in upper portion. Leaves cauline; median leaves opposite, petiole 3–5 cm long, blade 9–15 cm long, prominently soft pubescent on both surfaces, once or twice divided into pinnate leaflets, terminal segment ovate, short acuminate, rather prominently toothed, lateral segments ovate, sometimes lower one pinnatifid; upper leaves gradually smaller, opposite or alternate, bi-pinnately parted. Heads radiate or discoid, 7–10 mm across; peduncle 1.5–5.5 cm long; phyllaries 8–10, in one series, herbaceous, linear, 3–65. mm long, acute; outer chaff 5–6 mm long, hyaline margined. Ray florets 0–5, sterile, ligule yellow, ca. 5.5 Χ 2.5–3 mm, tube 1.2–1.5 mm long; disc florets bisexual, fertile, corolla 4–5.5 mm long, tube 1.2–2 mm long. Achenes linear, 9–19 mm long, ca. 1 mm wide, compressed, 4-angled, short strigose; awns 3–4, 3–4 mm long. Fl. Sep–Nov.

Roadside, waste fields; up to 1300 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hainan, Hubei, Hunan, Jiangxi, Liaoning, Shaanxi, Shanxi, Shandong, Taiwan, Yunnan, Zhejiang [Indonesia, Japan, Korea, Malaysia, Philippines, Vietnam; Oceania; Africa].

27. Glossocardia Cassini, Bull. Soc. Philom. Paris 138. 1817.

鹿角草属  lu jiao cao shu

Glabrous perennials with a thick caudex; stems erect, branched; leaves few. Radical leaves crowded, long petiolate, pinnately parted; cauline leaves alternate. Heads small, solitary or few in corymbs, heterogamous. Involucral bracts in 1–4 series, outer bracts and palea oblong, slightly connate, somewhat inflated at base, inner palea narrowed, flat. Ray florets in 1 row, pistillate, fertile, corollas spreading, 3-toothed. Disc florets tubular, bisexual, corollas 4-lobed. Anthers obtuse at base. Achenes glabrous, flattened dorsally, linear, truncate. Pappus of 2 retrorsely scabrid awns.

Eleven species in northern Africa, tropical Asia to southern Australia and the Pacific islands; one species in China.

1. Glossocardia bidens (Retzius) Veldkamp, Blumea 35(2): 468. 1991.

鹿角草  lu jiao cao

Zinnia bidens Retzius, Observ. Bot. 5: 28. 1788; Bidens meyeniana Walpers; B. tenuifolia Labillardiθre; Glossogyne bidens (Retzius) Alston; G. oluanpiensis S. S. Ying; G. tenuifolia Cassini.

Perennial herbs; stems woody at base, 20–30 cm long, somewhat tufted. Radical leaves persistent, glabrous, long 4.5–9 cm long, sometimes undivided and linear, usually pinnately parted, segments 2 or 3 pairs, remote, spreading, linear, 1.8–2 mm wide, obtuse, lower segments 8–20 mm long, petiole 2.7–6 cm long; median leaves few, remote, petiolate, 3–4 cm long, pinnately parted or linear; upper leaves smaller, linear. Heads ca. 7 mm across, solitary, basal bracteoles ca. 1.5 mm long. Involucral bracts and palea ca. 7, oblong, ca. 3 mm long. Ray florets few, in 1 series, fertile, corolla ca. 3.5 mm long, lip 3-dentate. Disc florets: corolla ca. 2.5 mm long, apex 4-dentate. pappus awns erect, 1.5–2 cm long. 2n = 24.

Exposed coastal areas, sometimes on raised coral reefs; lower elevations. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Xizang [Bengal, India, Indonesia, Malaysia, Philippines; Oceania; New Caledonia].

28. Galinsoga Ruiz & Pavσn, Fl. Peruv. Prodr. 110, plate 24. 1794.

牛膝菊属  niu qi ju

Annuals, 2–62 cm. Stems erect. Leaves cauline; opposite; petiolate; blades (3-nerved) lanceolate to broadly ovate, margins entire or serrulate to serrate, faces glabrate to densely pilose. Heads radiate or discoid, in ± cymiform arrays. Involucres hemispheric to campanulate, 2.5–6 mm diam. Phyllaries persistent or falling, 6–9[–16] in 2[–3] series (elliptic, lance-ovate, oblong, or ovate, outer shorter, herbaceous or scarious, margins entire or minutely laciniate). Receptacles conic, paleate (paleae persistent or falling, scarious, proximal broadly elliptic to obovate, often connate at bases or nearly to apices, united in groups of 2–3 to adjacent proximal phyllary, each complex often enclosing and shed with a ray cypsela; distal persistent or falling, lanceolate to obovate, entire or 2- or 3-lobed, convex to conduplicate). Ray florets [0–](3–)5(–8)[–15], pistillate, fertile; corollas white or dull white to pinkish [purplish] (tubes pilose, laminae quadrate-obovate to oblong, lobes 0–3). Disc florets 5–50[–150], bisexual, fertile; corollas yellow, tubes (pilose) shorter than cylindric throats, lobes 5, deltate (anthers yellow; style-branch apices acute). Cypselae obconic to obpyramidal, glabrous or strigose (rays often shed with subtending phyllary plus 2–3 adjacent paleae); pappi 0, or persistent, of 5–20, white or gray, fimbriate, sometimes aristate scales. x = 8.

Species 15–33: North America, Mexico, West Indies, Bermuda, Central America, South America. Two species introduced in China.

1a.       Phyllaries persistent; inner paleae 3-lobed, lobes 1/3+ total lengths, acute; disc pappi 0, or of 15–20 often gray, sometimes white, linear, fimbriate, obtuse scales 0.5–2 mm ...............  1. G. parviflora

1b.       Phyllaries falling; inner paleae entire or 2- or 3-lobed, lobes to 1.3 total lengths, blunt; disc pappi 0, or of 1–5 or 14–20 white, lanceolate to oblanceolate, fimbriate, sometimes aristate, scales  2. G. quadriradiata

1. Galinsoga parviflora Cavanilles, Icon. 3: 41, plate 281. 1795.

牛膝菊  niu qi ju

Plants 4–60 cm. Leaf blades 7–110 Χ 3–70(–80) mm. Peduncles 1–40 mm. Involucres campanulate, 2.5–5 mm diam. Phyllaries persistent. Paleae: outer persistent with distal phyllaries or falling, elliptic to obovate; inner usually persistent, lanceolate to ovate or obovate, 2–3.5 mm, 3-lobed, lobes to 1/3+ total lengths, acute. Ray florets (3–)5(–8); corollas usually dull white (pink), laminae 0.5–1.8 Χ 0.7–1.5 mm. Disc florets 15–50. Cypselae: rays 1.5–2.5 mm; discs 1.3–2.5 mm, glabrous or strigose; pappi: rays 0 or of 5–10, laciniate scales 0.5–1 mm; discs 0, or of 15–20 white or gray, linear, fimbriate, obtuse (acute) scales 0.5–2 mm. Fl. Jul–Oct.

In fields, streamsides and sparse forests. A common weeds widely distributed in China [native to America].

2. Galinsoga quadriradiata Ruiz & Pavσn, Syst. Veg. Fl. Peruv. Chil. 1: 198. 1798.

粗毛牛膝菊  cu mao ni qi ju

Plants 8–62 cm. Leaf blades 20–60 Χ 15–45 mm. Peduncles 5–20 mm. Involucres hemispheric to campanulate, 3–6 mm diam. Phyllaries falling. Paleae: outer falling, broadly elliptic to obovate, 2–3 mm; inner falling, linear to lanceolate, 2–3 mm, entire or 2- or 3-lobed, lobes to 1/3 total lengths, blunt. Ray florets (4–)5(–8); corollas usually white, sometimes pink, laminae 0.9–2.5 Χ 0.9–2 mm. Disc florets 15–35. Cypselae: rays 1.5–2 mm; discs 1.3–1.8 mm; pappus: rays of 6–15 fimbriate scales 0.5–1 mm; discs 0, or of usually 14–20, rarely 1–5, white, lanceolate to oblanceolate, fimbriate, sometimes aristate, scales 0.2–1.7 mm. Fl. Jul–Oct. 2n = 32, 48, 64.

In forest, roadsides. Jiangxi [native to America].

29. Tridax Linnaeus, Sp. Pl. 900. 1753; Gen. Pl. ed. 5, 382. 1754.

羽芒菊属  yu mang ju shu

Perennials. Leaves opposite; petiolate or sessile; blades deltate, lanceolate, lance-ovate, or ovate, often pinnately or palmately lobed, ultimate margins coarsely toothed to subentire, faces glabrate, hirsute, scabrellous, or strigillose. Heads radiate or discoid, borne singly or several in cymiform arrays. Involucres cylindric to hemispheric, 4–8 mm diam. Phyllaries persistent, in 2–3+ series; outer usually shorter, more herbaceous, the inner often scarious. Receptacles convex to conic; paleae persistent, lance-linear, scarious, weakly conduplicate, often apically toothed. Ray florets pistillate, fertile; corollas pale yellow, white or purplish. Disc florets bisexual, fertile; corollas yellowish, whitish or purplish, tubes shorter than cylindric or funnelform throats, lobes 5, usually deltate, sometimes rays 0 and corollas of peripheral florets bilabiate. Cypselae obconic to obpyramidal, 3-, 4-, or 5-angled, densely piloso-sericeous, glabrous or villous. Pappus persistent, of plumose or ciliate, setiform scales. x = 10.

About 26 species: tropical America and Asia; one species naturalized in China.

1. Tridax procumbens Linnaeus, Sp. Pl. 900. 1753.

羽芒菊  yu mang ju

Perennial herbs, caulescent, decumbent. Stems procumbent, branched at base, branches slender, spreading or ascending, 20–50 cm long, hirsute. Leaves short petiolate, ovate to ovate-lanceolate, few, 2.5–5 cm long, apex acute or acuminate, margins deeply irregularly serrate, pinnatisect, segments few, narrow, base cuneate. Heads solitary, 1–1.5 cm across, peduncle 10–30 cm long. Involucre subcampanulate, 6–7 mm long. Involucral bracts hispid, in few series; outer bracts densely grayish white, elliptic, ca. 5 mm long, long hirsute; inner bracts tinged purplish, narrower, ca. 6 mm long. Ray florets 4, white, ligulate or 2-lipped. Disc florets yellow, tube ca. 5 mm long, limb 5-lobed, lobes reflexed, hairy. Achenes brown, oblong, ca. 2 mm long, densely silky pubescent. Pappus 5–6 mm long, of shiny feathery bristles. Fl. Nov–Mar. 2n = 36.

A noxious weed, along exposed, dry roadsides, on waste grounds and in lawns. Fujian, Hainan, Taiwan [India, Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam; tropical America].

Addendum

Gaillardia is transferred from the Helenieae, where it was treated in FRPS 75 (1979). Because the two species are only cultivated in China, presumably for ornament rather than as large-scale crops, these treatments should be deleted. However, the two species should be mentioned among other purely cultivated Heliantheae in China in a comment at the start of the tribe.

30. Gaillardia Fougeroux, Observ. Phys. 29: 55. 1786.

天人菊属  tian ren ju shu

Annuals, perennials, or subshrubs. Stems usually erect, usually branched. Leaves basal, basal and cauline, or mostly cauline; alternate; petiolate or sessile; blades mostly elliptic, lanceolate, linear, oblong, obovate, ovate, or spatulate, often pinnately lobed, ultimate margins toothed or entire, faces usually scabrellous to villous, rarely glabrous, usually notably gland-dotted. Heads usually radiate or discoid, borne singly. Involucres hemispheric to nearly rotate, mostly 9–20+ mm diam. Phyllaries persistent, in 2–3+ series, reflexed in fruit, usually distinct, oblanceolate, lanceolate, narrowly triangular, or linear, subequal to unequal, usually chartaceous proximally, distally herbaceous, apices often attenuate. Receptacles convex to hemispheric, smooth or pitted, epaleate. Ray florets 0, or 5–15+, usually neuter, rarely pistillate and fertile or styliferous and sterile; corollas yellow to white or orange to red, purplish, or brown, often bicolored (sometimes "ray" corollas funnelform, 5-lobed, heads radiant). Disc florets bisexual, fertile; corollas yellow or orange to red, purplish, or brown, tubes much shorter than ampliate, campanulate to cylindric or urceolate throats, lobes 5, broadly deltate or ovate. Cypselae obpyramidal to clavate, ca. 4-angled, sparsely to densely hairy. Pappus usually persistent, of 6–10+ medially thickened, laterally scarious scales in 1–2 series, rarely absent.

About 20 species: North America, South America; two species introduced in China.

1a.       Perennials (sometimes flowering first year); leaves basal and cauline; receptacular setae (2–)3–6+ mm     1. G. aristata

1b.       Annuals (seldom persisting); leaves all or mostly cauline; receptacular setae mostly 0.3–3 mm  2. G. pulchella

1. Gaillardia aristata Pursh, Fl. Amer. Sept. 2: 573. 1813.

宿根天人菊  su geng tian ren ju

Perennials, 20–80 cm high. Leaves basal and cauline or cauline; petiolar bases 5–15 cm; blades oblanceolate to lanceolate, 5–15 cm Χ 5–30(–40) mm, margins raggedly pinnately lobed to toothed or entire, faces scabrellous and/or sparsely to densely villous (hairs jointed). Peduncles (5–)20–35+ cm. Phyllaries 24–40+ ovate to lance-attenuate, 10–15+ mm, ciliate with jointed hairs (also strigose and gland-dotted). Receptacular setae 2–6 mm. Ray florets (6–)12–18+; corollas yellow or yellow/purple, rarely tubular and 5-lobed, usually distally laminate and 3-lobed, 15–35+ mm. Disc florets 60–120+; corollas usually purple or purple-tipped, sometimes yellow, tubes 0.5–1.5 mm, throats cylindric to urceolate, 4.5–5.5 mm, lobes lance-ovate to triangular-attenuate, 1–2 mm, jointed hairs ca. 0.3+ mm. Cypselae clavate (outer) to obpyramidal (inner), 2.5–6 mm, hairs 1.5–2.5 mm, inserted at bases. Pappus of 8 ovate to lanceolate, aristate scales 5–6 mm. Fl. May–Sep. 2n = 34, 68.

Widely cultivated in China [native to North America].

2. Gaillardia pulchella Fougeroux, Hist. Acad. Roy. Sci. Mιm. Math. Phys. (Paris, 4to). 1786: 5. 1788.

天人菊  tian ren ju

Gaillardia pulchelalla f. flaviflora S. S. Ying.

Annuals, 5–35(–60) cm high. Leaves cauline; petiolar bases 0–3+ cm; blades linear, oblong, or spatulate, 1–5(–12) cm Χ 4–12(–35) mm, margins usually entire, sometimes toothed or lobed, faces closely strigillose or hirtellous to ± villous. Peduncles 3–10(–20) cm. Phyllaries 18–28+, narrowly triangular- to linear-attenuate, 6–14+ mm long, usually ciliate with jointed hairs. Receptacular setae 1.5–3 mm. Ray florets usually 8–14, rarely 0; corollas usually reddish to purplish proximally, yellow to orange distally, rarely yellow, reddish, or purplish throughout, 13–30+ mm. Disc florets 40–100+; corollas yellowish to purple or brown, often bicolored, tubes 0.8–1.2 mm, throats campanulate to urceolate, 3–4 mm, lobes deltate to ovate, often attenuate, 1–3+ mm, jointed hairs ca. 0.3+ mm. Cypselae obpyramidal, 2–2.5 mm, hairs 1.5–2 mm, inserted at bases and on angles. Pappus of 7–8 deltate to lanceolate, aristate scales 4–7 mm. Fl. May–Aug. 2n = 34.

Widely cultivated in China [native to North America].



[1] Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

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