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187m.8. Asteraceae Martinov (tribe Heliantheae) subtribe Ecliptinae Lessing, Linnaea. 6: 153. 1831.
Enceliinae Panero; Engelmanniinae Stuessy; Spilanthinae Panero; Verbesininae Bentham & Hooker f.; Zinniinae Bentham & Hooker f.
Annuals, perennials, subshrubs, or shrubs, (3–)10–200(–400+) [2500+] cm. Leaves usually cauline, sometimes mostly basal, or basal and cauline; opposite, alternate, or both; petiolate or sessile; blades cordate, deltate, elliptic, lanceolate, linear, rhombic, orbiculate, or ovate (and intermediate shapes), often 1(–2+)-palmately or -pinnately lobed, ultimate margins entire or toothed, faces glabrous or hairy (often hispid or scabrous), often gland-dotted or stipitate-glandular. Heads usually radiate, sometimes discoid, borne singly or in cymiform, corymbiform, paniculiform, racemiform, or spiciform arrays. Calyculi 0. Involucres cylindric, or campanulate to hemispheric or broader, or rotate. Phyllaries persistent or falling, usually 8–40+ in 2–4+ series, usually distinct, usually lanceolate, linear, orbiculate, or ovate, subequal or unequal (outer usually shorter, rarely longer, than inner). Receptacles convex, conic, or flat, paleate (paleae usually falling, oblong to linear, herbaceous to chartaceous or scarious, usually conduplicate). Ray florets usually (1–)3–40+, sometimes 0, usually pistillate and fertile, sometimes styliferous and sterile, or neuter; corollas usually yellow to orange, sometimes pink to purple, red, brown, or white (sometimes sessile, persistent, and becoming papery, e.g., in Heliopsis, Sanvitalia, and Zinnia). Disc florets 4–200+, bisexual and fertile, or functionally staminate; corollas usually yellow to orange, sometimes brown, greenish, maroon, pink, purple, or red, rarely white or whitish, tubes shorter to longer than campanulate, cylindric, or funnelform throats, lobes (4–)5, deltate to lanceolate (usually equal); anther thecae usually dark (collars not continuous around filaments); stigmatic papillae usually continuous, sometimes none, rarely in 2 lines. Cypselae obpyramidal or prismatic (3–4-angled), or terete to ovoid, or strongly compressed or flattened and ± orbiculate to obovate or cuneate (lengths seldom more than 2 times diams.), sometimes ribbed, sometimes winged, glabrous or hairy; pappi usually persistent, sometimes fragile or readily falling, usually of scales, sometimes of bristles or bristlelike awns or coroniform, rarely of awns or 0.
Genera 75, species 600 (31 genera, 124 species, including 4 hybrids, in the flora): mostly subtropical and warm-temperate North America, especially arid areas, also in the Old World.
The circumscription of Ecliptinae adopted by H. Robinson (1981) and followed here differs from the treatment by P. O. Karis and O. Ryding (1994), who placed Eclipta as unassigned to a subtribe and placed Echinacea in Rudbeckiinae, Acmella, Heliopsis, Sanvitalia, and Zinnia in Zinniinae, Berlandiera, Chrysogonum, Engelmannia, Lindheimera, and Silphium in Engelmanniinae, and Balsamorhiza (including Agnorhiza), Borrichia, Calyptocarpus, Encelia, Enceliopsis, Flourensia, Geraea, Helianthella, Jefea, Lasianthaea, Melanthera, Pascalia, Phoebanthus, Sphagneticola, Synedrella, Verbesina, Wedelia, and Wyethia (including Scabrethia) in Verbesininae. Based on studies of chloroplast DNA sequences, J. L. Panero (2005) placed Encelia, Enceliopsis, Flourensia, Geraea, and Helianthella together in Enceliinae and Acmella in Spilanthinae, indicating a sister relationship between Spilanthinae and traditionally circumscribed Zinniinae.
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| 1 |
Ray florets usually 5–21 (more in "double" cultivars), sometimes 0 (in Zinnia anomala, subshrubs or shrublets to 12 cm, leaves linear, Texas), corollas usually yellow to orange, sometimes purple, red, or whitish (usually persistent, sessile, becoming papery) |
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(2) |
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Ray florets usually (2–)5–35, sometimes 0, corollas usually yellow to orange, sometimes white (seldom sessile, laminae usually borne on tubes, never persistent and becoming papery) |
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(4) |
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| 2 (1) |
Leaf margins serrate to coarsely toothed |
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264 Heliopsis, p. 67 |
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Leaf margins entire |
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(3) |
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| 3 (2) |
Leaves petiolate or sessile; 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) |
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265 Sanvitalia, p. 70 |
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Leaves sessile; cypselae 3-angled or flattened (none with uncinate hairs, none winged) |
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266 Zinnia, p. 71 |
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| 4 (1) |
Disc florets functionally staminate (only ray florets produce cypselae) |
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(5) |
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Disc florets bisexual, fertile |
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(9) |
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| 5 (4) |
Phyllaries 8–10 in 2 series |
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(6) |
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Phyllaries 12–45+ in (2–)3–4 series |
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(7) |
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| 6 (5) |
Perennials; leaves opposite, petiolate, margins crenate |
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267 Chrysogonum, p. 74 |
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Annuals; leaves mostly alternate (distal sometimes opposite), ± sessile, margins coarsely toothed (mostly distal 1/2) |
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268 Lindheimera, p. 75 |
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| 7 (5) |
Ray florets 8–35 (in 1–3 series); cypselae (shed alone without accessory structures) |
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269 Silphium, p. 77 |
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Ray florets usually (2–)8(–13); cypselae (each shed together with subtending phyllary and 2–4 adjacent paleae and disc florets) |
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(8) |
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| 8 (7) |
Phyllaries usually 14–22 in 2–3 series (broadly obovate to orbiculate, without linear appendages); ray corollas pale yellow to orange-yellow (abaxially red to maroon or with greenish or red to maroon veins) |
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270 Berlandiera, p. 83 |
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Phyllaries 18–24+ in ± 3 series (at least outer with relatively short, expanded, indurate bases and longer, linear, herbaceous tips); ray corollasyellow (without notably colored veins) |
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271 Engelmannia, p. 87 |
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| 9 (4) |
Leaves mostly basal, or basal and cauline, or cauline, mostly alternate |
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(10) |
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Leaves mostly cauline, mostly opposite |
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(21) |
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| 10 (9) |
Cypselae prismatic, or nearly so, 3–4-angled |
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(11) |
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Cypselae compressed to flattened |
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(15) |
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| 11 (10) |
Ray florets usually 8–21 (corollas mostly pink, red, purple, or white, yellow in E. paradoxa var. paradoxa, laminae often drooping or reflexed);receptacles mostly conic (heights often 2+ times diams.) |
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272 Echinacea, p. 88 |
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Ray florets (1–)5–25+ (corollas yellow or orange, laminae not drooping or reflexed), sometimes 0; receptacles flat to convex |
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(12) |
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| 12 (11) |
Leaves mostly basal (cauline usually notably smaller than basal); pappi 0 |
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273 Balsamorhiza, p. 93 |
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Leaves basal and cauline, or mostly cauline; pappi usually coroniform, sometimes of 1–4+ scales or 0 |
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(13) |
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| 13 (12) |
Leaves mostly cauline (blades narrowly oblong to linear, 5–25 mmwide) |
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274 Scabrethia, p. 99 |
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Leaves basal and cauline, or mostly cauline (blades mostly 30–120 mm wide) |
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(14) |
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| 14 (13) |
Leaves mostly elliptic, lanceolate, or oblong (basal and cauline, basal usually notably larger than cauline, cauline mostly sessile) |
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275 Wyethia, p. 100 |
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Leaves mostly orbiculate, ovate, or rounded-deltate (mostly cauline, mostly petiolate, proximal and distal usually ± similar) |
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276 Agnorhiza, p. 104 |
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| 15 (10) |
Cypselae winged; pappi persistent, of 2(–3) scales (scales often aristate or subulate, without additional scales) |
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277 Verbesina (in part), p. 106 |
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Cypselae sometimes thin-edged (margins sometimes ciliate or corky-thickened, never truly winged); pappi usually of (1–)2, subulate scales or bristlelike awns plus 2–4+ shorter scales, sometimes 0 (rarely of 2–3 aristate scales without additional scales) |
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(16) |
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| 16 (15) |
Perennials (scapiform); leaves all or mostly basal; involucres 20–30+ mm diam |
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278 Enceliopsis, p. 112 |
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Perennials (rarely scapiform), subshrubs, or shrubs; leaves usually cauline, sometimes basal and cauline; involucres 4–30 mm diam |
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(17) |
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| 17 (16) |
Perennials (rhizomatous); leaves linear to filiform |
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279 Phoebanthus, p. 113 |
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Perennials or shrubs (not rhizomatous); leaves mostly deltate, elliptic, lanceolate, or ovate (and most intermediate shapes, not linear to filiform) |
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(18) |
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| 18 (17) |
Ray florets 8–21, pistillate and fertile |
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280 Helianthella (in part), p. 114 |
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Ray florets 0 or 8–40, neuter, or styliferous and sterile |
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(19) |
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| 19 (18) |
Subshrubs or shrubs (glabrous or ± scabrellous, usually vernicose); phyllaries 12–40 in 2–4+ series (subequal or unequal, outer longer) |
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281 Flourensia, p. 117 |
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Annuals, perennials, or shrubs (glabrous or canescent, hirtellous, scabrellous, strigose, or tomentose, often gland-dotted or glandular-puberulent to stipitate-glandular, seldom vernicose); phyllaries 18–30(–50+) in 2–3+ series (subequal or unequal, outer shorter) |
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(20) |
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| 20 (19) |
Perennials (E. nutans), subshrubs, or shrubs; pappi usually 0, sometimes fragile, of 2 weak, villous scales |
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282 Encelia, p. 118 |
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Annuals or perennials; pappi usually persistent, of 2 subulate scales |
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283 Geraea, p. 122 |
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| 21 (9) |
Heads discoid (corollas white or whitish); pappi 0, or readily falling, of 2–12 barbellate bristles or awns |
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284 Melanthera, p. 123 |
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Heads radiate or discoid (corollas seldom white or whitish); pappi 0, or usually persistent and coroniform or cyathiform (each an erose, fimbriate, or lacerate cup, with or without additional awns or bristles, borne on rostrums), or of 2–4+ awns, bristles, and/or scales |
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(22) |
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| 22 (21) |
Pappi usually coroniform or cyathiform (cypselae often rostrate, each with apical boss or neck), sometimes 0 (prostrate perennials) |
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(23) |
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Pappi usually of 2–4+ awns, bristles, and/or scales (not cyathiform, cypselae not rostrate), sometimes coroniform or 0 (not prostrate perennials) |
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(24) |
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| 23 (22) |
Subshrubs or shrubs (erect); cypselae (some or all) strongly compressed, nota-bly winged |
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285 Wedelia, p. 125 |
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Perennials (prostrate); cypselae strongly biconvex to plumply 3–4-angled (not com-pressed, not winged, epidermes usually corky, often tuberculate) |
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286 Sphagneticola, p. 126 |
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| 24 (22) |
Some or all cypselae winged (each bordered by wing of membranous or corky tissue different from that of body of cypsela) |
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(25) |
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Cypselae sometimes sharp-edged (not winged) |
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(27) |
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| 25 (24) |
Heads in glomerules or borne singly (sessile or subsessile in axils); cypselaewinged (rays, not discs, wings lacerate) |
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287 Synedrella, p. 127 |
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Heads borne singly or in corymbiform, dichasiiform, or paniculiform arrays (not sessile); cypselae winged (rays and discs, wings not lacerate) |
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(26) |
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| 26 (25) |
Phyllaries 9–30 in 1–4 series (outer 2–5 similar to others, unlike foliage); pappi of 2(–3) persistent (often aristate or subulate) scales without additional scales |
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277 Verbesina (in part), p. 106 |
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Phyllaries 26–38+ in 3–4+ series (outer 2–6+ similar to foliage in shape, texture, and indument); pappi of 2–3 fragile or persistent awns or subulate scales plus 2–8+, distinct or basally connate, erose or lacerate scales (often each cypsela with additional seta on inner shoulder) |
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288 Jefea, p. 127 |
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| 27 (24) |
Cypselae 3–4-angled (weakly or not at all compressed or obcompressed, epidermes usually thick, corky) |
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(28) |
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Cypselae (all or at least disc) strongly compressed or obcompressed or flattened (epidermes seldom thick and corky) |
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(30) |
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| 28 (27) |
Corollas white or whitish (paleae linear-filiform, not conduplicate) |
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289 Eclipta, p. 128 |
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Corollas yellow to orange (paleae lanceolate to ovate, conduplicate) |
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(29) |
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| 29 (28) |
Leaves elliptic, linear, oblanceolate, obovate, or ovate, glabrous or puberulent to villous and/or sericeous (outer phyllaries elliptic, oblanceolate, or ovate) |
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290 Borrichia, p. 129 |
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Leaves lanceolate to lance-linear, sparsely scabrous (outer phyllaries lance-lin-ear to linear) |
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291 Pascalia, p. 131 |
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| 30 (27) |
Annuals or perennials (mostly 5–30+ cm; larger leaves mostly 1–5+ cm); involucres 3–8 mm diam.; phyllaries 5 in 1(–2) series, or 8–15+ in 1–3 series |
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(31) |
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Perennials (coarse, 10–150 cm; larger leaves mostly 5–25 cm); involucres 10–30 mm diam.; phyllaries 12–35 in 2–5 series |
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(32) |
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| 31 (30) |
Disc florets 25–100(–200+); receptacles conic; cypselae ellipsoid to obovoid; pappi 0, or fragile, of 1–3 bristlelike awns |
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292 Acmella, p. 132 |
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Disc florets 10–20; receptacles convex; cypselae cuneate; pappi of 2(–5+) stout awns |
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293 Calyptocarpus, p. 133 |
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| 32 (30) |
Leaf blades oblanceolate to lanceolate or lance-linear (longer usually 8–25 cm), margins entire |
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280 Helianthella (in part), p. 114 |
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Leaf blades rounded-deltate to ovate or lance-ovate (longer usually 5–8 cm),margins coarsely serrate |
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294 Lasianthaea, p. 133 |
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List of lower taxa
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