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4. Tribe CARDUEAE 1. SAUSSUREA Group Published online on 25 October 2011. Shi, Z., Raab-Straube, E. von, Greuter, W. & Martins, L. 2011. Cardueae. Pp. 42–194 in: Wu, Z. Y., Raven, P. H. & Hong, D. Y., eds., Flora of China Volume 20–21 (Asteraceae). Science Press (Beijing) & Missouri Botanical Garden Press (St. Louis). 4. Tribe CARDUEAE 飞廉族 fei lian zu Shi Zhu (石铸 Shih Chu); Eckhard von Raab-Straube, Werner Greuter, Ludwig Martins Herbs, annual to perennial. Leaves alternate, sometimes all basal. Capitula homogamous or heterogamous, bisexual, sometimes unisexual in dioecious plants, persistent or deciduous, solitary or clustered at end of stem and branches, sometimes surrounded by ± leaflike spiny bracts. Phyllaries imbricate, undivided. Receptacle densely covered with bristles, rarely with short scales, or naked. Florets few to numerous. Corolla usually actinomorphic or slightly zygomorphic, equally or unequally 5-lobed or sometimes strongly zygomorphic in radiant florets of heterogamous capitula. Stamen filaments glabrous, papillose, or hairy; anther sagittate, base caudate; connective with a sterile apical appendage. Style branches distinct and divergent or coherent medially for most of their length and only seceding along stigmatic margins; sweeping hairs usually confined to a ringlike sometimes swollen zone at base of style branches or longest there and distally abruptly shorter. Achene usually glabrous, rarely ± hairy, apex with a peripheral rim sur- rounding an apical plate on which pappus is inserted; rim sometimes salient as an entire or dentate crown, sometimes inconspicuous. About 75 genera and ca. 2,400 species: E and N Africa, Asia, Australia, Europe, North and South America; 37 genera (four endemic, three introduced) and 440 species (237 endemic, five introduced) in China. 1a. Achene attachment scar lateral or subbasal and oblique; capitula homogamous or heterogamous and mostly with radiant outer florets .................................................................................................................. 5. Subtribe Centaureinae 1b. Achene attachment scar ± basal; capitula always homogamous. 2a. Leaf margin unarmed, entire or with blunt to mucronulate non-pungent teeth, apex of phyllaries sometimes recurved but never hooked ........................................................................................................... 1. Saussurea group 2b. Leaf margin spiny or spinulose, or if ± entire then apex of phyllaries hooked. 3a. Pappus of distinct scabrid bristles falling off singly or (rarely) pappus absent; style branches distinct and divergent ................................................................................................................................... 2. Arctium group 3b. Pappus of scabrid to plumose bristles (rarely scales) basally connate into a ring and falling off together; style branches coherent for most of their length. 4a. Achene with apical rim either inconspicuous or forming a conspicuous and crenulate (rarely entire) crown, apical plate lacking an elaiosome ............................................................... 3. Onopordum group 4b. Achene with apical rim forming a smooth-margined upright crown, apical plate with a prominent oleaginous central umbo (elaiosome) ....................................................................... 4. Subtribe Carduinae 1. SAUSSUREA group 风毛菊群 feng mao ju qun Shi Zhu (石铸 Shih Chu); Eckhard von Raab-Straube Cauliferous herbs, stemless rosette herbs, or subshrubs, biennial, perennial, or rarely annual. Leaves without spines; leaf blade undivided to pinnatisect, margin entire or with blunt to mucronate non-pungent teeth. Capitula homogamous, bisexual. Phyllaries in several rows, without spines, apically sometimes with a green or purplish membranous or herbaceous appendage or mucronate. Receptacle densely covered with long subulate bristles, short scales, apiculate papillae, or naked and alveolate. Florets bisexual, all fertile. Corolla ± actinomorphic. Filaments distinct, glabrous. Style branches usually distinct and divergent, rarely partly coherent. Achene usually glabrous, sometimes glandular, rarely pubescent or papillose, apical rim inconspicuous or forming a ± developed crenulate or denticulate crown. Pappus homomorphic or heteromorphic, in 1 to several distinct or connate rows of scabrid to plumose bristles or rarely of small scales; inner pappus long, usually overtopping involucre. About 15 genera and ca. 700 species: NW Africa, Asia, Europe, North America; nine genera (two endemic, one introduced) and 323 species (205 endemic, one introduced) in China. 1a. Plants annual; outer pappus an incomplete row of small scales and inner pappus a ring of plumose bristles .......... 21. Hemisteptia 1b. Plants perennial or biennial; pappus of only bristles. 2a. Pappus bristles in several rows, of different length, usually scabrid or barbellate, rarely plumose; receptacle with short scales or naked and alveolate, rarely with bristles. 3a. Pappus bristles silvery, dirty white, or white, inserted on a conic cupule, 0.5–1.5 cm; receptacle with short scales or bristles .................................................................................................................................................... 14. Jurinea 3b. Pappus bristles dirty white, yellowish, brown, or reddish, not inserted on a conic cupule, 1.5–4 cm; receptacle either alveolate or with long bristles. 4a. Capitula several and clustered or solitary in center of leaf rosette; phyllaries not densely lanate; receptacle alveolate, naked or with very short apiculate papillae, without bristles .................................................. 17. Dolomiaea 4b. Capitulum solitary on a subscapose stem; phyllaries densely lanate; receptacle with long bristles ................................................................................................................................................... 18. Bolocephalus 42 CARDUEAE 43 2b. Pappus bristles in 1 or 2 rows, with bristles of inner row of same length and always plumose; receptacle usually with ± long, subulate bristles, sometimes naked and alveolate, or with short apiculate papillae. 5a. Pappus in 2 rows; achene smooth, ribbed, or sometimes wrinkled, apical rim inconspicuous or rarely with a short crown. 6a. Outer pappus a row of scabrid or rarely plumose caducous bristles (rarely lacking), inner pappus a ring of plumose bristles ........................................................................................................................... 22. Saussurea 6b. Outer and inner pappus bristles both plumose and connate with each other into a ring ..................... 15. Diplazoptilon 5b. Pappus in 1 row; achene wrinkled or muricate, apical rim always forming a distinct crown. 7a. Corolla limb with sessile glands; achene blackish, 2–5 mm, with a dentate crown .............................. 16. Himalaiella 7b. Corolla limb glabrous; achene straw-colored or brown with black spots, 5–8 mm, with a crenulate crown. 8a. Capitulum solitary, long pedunculate; petiole of basal leaves unwinged ............................................. 19. Frolovia 8b. Capitula 2–7 and clustered, sessile or subsessile; petiole of basal leaves with a broad lobed wing .......................................................................................................................................... 20. Aucklandia 14. JURINEA Cassini, Bull. Sci. Soc. Philom. Paris 1821: 140. 1821. 苓菊属 ling ju shu Shi Zhu (石铸 Shih Chu); Eckhard von Raab-Straube Pilostemon Iljin. Herbs, perennial [or shrublets]. Leaves undivided to pinnatisect. Capitula solitary or several in a corymbiform synflorescence. Involucre bowl-shaped, ovoid, campanulate, hemispheric, or cylindric. Phyllaries herbaceous or leathery, appressed or reflexed. Receptacle with broad rigid shortly acuminate scales or bristles. Corolla red to purple, with sessile glands. Anther tails lacerate- caudate. Style branches short, divergent. Achene isodiametric, narrowly obovoid, narrowly ellipsoid, or obconic, 3- or 4-angled, gla- brous or glandular, sometimes spinulose-tuberculate, apical rim forming a crown. Pappus bristles in several rows, white, scabrid, bar- bellate, or plumose, unequal in length, sometimes 2–5 inner bristles much longer than others; bristles inserted on a conspicuous coni- cal cupule, persistent or caducous as a whole. About 250 species: NW Africa, C and SW Asia, Europe; ten species (one endemic) in China. The genus Pilostemon differs from Jurinea only in the pilose anther connectives. All other diagnostic characters given in the protologue of the genus, especially concerning the receptacle, achene, and pappus structure, can be found at least in some sections of Jurinea as well. Pilostemon is therefore not accepted in this treatment. 1a. Achene not spinulose-tuberculate. 2a. Leaf blade linear-filiform, 1–2 mm wide; involucre obconic; pappus plumose, 1–1.5 cm ...................................... 3. J. filifolia 2b. Leaf blade broadly linear to elliptic, more than 3 mm wide; involucre cylindric or bowl-shaped; pappus scabrid or barbellate, 0.8–1.2 cm. 3a. Leaf blade broadly linear, abaxially grayish white, adaxially green, undivided; involucre cylindric; phyllaries scarious ........................................................................................................................................... 1. J. multiflora 3b. Leaf blade elliptic, obovate, or ovate, green to grayish green, almost concolorous, pinnately dentate, pinnately lobed, or pinnately divided; involucre
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