treatments/Asteraceae Cichorieae

From Botanical Knowledge
Jump to: navigation, search


Asteraceae   Martinov tribe   Cichorieae Lamarck & de Candolle Syn. Pl. Fl. Gall.,. 255. 1806.



Annuals, biennials, perennials, subshrubs, or shrubs [trees, vines] (sap usually milky). Leaves basal and/or cauline; alternate (proximal opposite in Shinnersoseris) [opposite]; petiolate or sessile; margins usually dentate or pinnately lobed (then frequently runcinate), sometimes prickly, sometimes entire, rarely much divided (bases often clasping). Heads homogamous (liguliflorous), usually in corymbiform or paniculiform arrays, sometimes borne singly (on scapiform peduncles), sometimes subsessile in axillary clusters on stems or among leaves of basal rosettes [aggregated in second-order heads]. Calyculi 0 or of 1–15+ bractlets in 1–3+ series. Phyllaries usually persistent, usually in 3–5+ series, distinct, and unequal, sometimes in 1–2 series, distinct or connate, and subequal to equal, margins (seldom prickly) and/or apices sometimes notably scarious (phyllaries sometimes enfold and fall with subtended florets or cypselae, e.g., in Rhagadiolus). Receptacles flat to convex, epaleate (sometimes bristly-setose) or paleate (paleae enfold and fall with subtended cypselae in Scolymus). Florets ligulate (bisexual, fertile); corollas usually yellow to orange, sometimes cyanic or white (zygomorphic, 5-toothed); anther bases usually tailed, apical appendages ovate to lanceolate or hardly developed (pollen sometimes brightly colored); styles abaxially usually papillate to hirsute (mostly distally), branches filiform to stout, adaxially continuously stigmatic from bases almost to apices, apices rounded to acute, appendages essentially 0. Cypselae usually monomorphic within heads, ± clavate, columnar, ellipsoid, fusiform, or prismatic, often compressed, obcompressed, or flattened, often beaked or apically tapered, bodies smooth, muricate, rugose, or tuberculate, often ribbed, sometimes winged (glabrous or hairy); pappi (rarely 0) persistent or readily falling, usually of fine to coarse, often barbellate, sometimes plumose bristles, sometimes of awns or scales, sometimes combinations of bristles, awns, and/or scales (scales often aristate).


Genera ca. 100, species ca. 1600 (49 genera, 229 species in the flora):nearly worldwide; mostly in Old World; mostly at temperate latitudes; some species widely introduced.;


SELECTED REFERENCES
Jansen, R. K. et al. 1991b. Systematic implications of chloroplast DNA variation in the subtribe Microseridinae (Asteraceae: Lactuceae). Amer. J. Bot. 78: 1015–1027.Kim, S. C., D. J. Crawford, and R. K. Jansen. 1996. Phylogenetic relationships among the genera of the subtribe Sonchinae (Asteraceae): Evidence from ITS sequences. Syst. Bot. 21: 417–432.Lee, J. and B. G. Baldwin. 2004. Subtribes of principally North American genera of Cichorieae (Compositae). Novon 14: 309–313.Lee, J., B. G. Baldwin, and L. D. Gottlieb. 2002. Phylogeny of Stephanomeria and related genera (Compositae–Lactuceae) based on 18S-26S nuclear rDNA ITS and ETS sequences. Amer. J. Bot. 89: 160–168.Lee, J., B. G. Baldwin, and L. D. Gottlieb. 2003. Phylogenetic relationships among the primarily North American genera of Cichorieae (Compositae) based on analysis of 18S–26S rDNA ITS and ETS sequences. Syst. Bot. 28: 616–626.Price, H. J. and K. Bachmann. 1975. DNA content and evolution in the Microseridinae. Amer. J. Bot. 62: 262–267.Stebbins, G. L. 1953. A new classification of the tribe Cichorieae, family Compositae. Madroño 12: 33–64.Tomb, A. S. 1974. Chromosome numbers and generic relationships in subtribe Stephanomeriinae (Compositae: Cichorieae). Brittonia 26: 203–216.Whitton, J., R. S. Wallace, and R. K. Jansen. 1995. Phylogenetic relationships and patterns of character change in the tribe Lactuceae (Asteraceae) based on chloroplast DNA restriction site variation. Canad. J. Bot. 73: 1058–1073.


1. Cypselae beaked (outer sometimes beakless in Hypochaeris ). (2.)
2. Stems scapiform. (3.)
3. Heads usually in spiciform, paniculiform, cymiform, or corymbiform arrays (sometimes borne singly). (4.)
4. Leaf margins spiny; calyculi 0; phyllaries in 3–5+ series, unequal; pappi of crisped (frizzy), outer and straight, coarse, inner bristles 47. Launaea (in part), p. 272
4. Leaf margins not spiny; calyculi of 5–12 bractlets; phyllaries in 1(–2) series, equal; pappi of uniform bristles 36. Crepis (in part), p. 222
3. Heads borne singly. (5.)
5. Pappi of outer scales and inner bristles, or of aristate scales. (6.)
6. Cypsela beaks relatively long or 0; pappi of outer scales and inner, plumosebristles; annuals or perennials; leaves oblanceolate to oblong 52. Leontodon (in part), p. 294
6. Cypsela beaks relatively short; pappi of lanceolate, apically notched, aristate scales, aristae smooth; annuals; leaves linear to narrowly lanceolate, grasslike 65. Uropappus (in part), p. 322
5. Pappi of bristles. (7.)
7. Receptacles paleate; corollas pink, purple, lavender, or nearly white; perennials 81. Pinaropappus, p. 374
7. Receptacles usually epaleate (if paleate, corollas yellow; some Agoseris ); corollas usually yellow to orange, rarely ochroleucous, pink, pinkish, purplish, or white; annuals, biennials, or perennials. (8.)
8. Calyculi of (6–)8–18(–20) bractlets in 1–3 series; phyllaries equal; perennials 37. Taraxacum, p. 239
8. Calyculi 0; phyllaries equal or unequal; annuals, biennials, or perennials. (9.)
9. Leaves entire, toothed, or pinnately lobed (not spiny); heads borne singly; corollas yellow, orange, pinkish, purplish, or white; pappi of ± barbellate bristles; annuals or perennials 66. Agoseris (in part), p. 323
9. Leaves often pinnately lobed, ultimate margins dentate (spiny); heads borne singly or in spiciform or paniculiform arrays; corollas yellow to ochroleucous; pappi of crisped (frizzy), outer and straight, coarse, inner bristles; annuals or biennials 47. Launaea (in part), p. 272
2. Stems leafy. (10.)
10. Pappi of plumose bristles or subulate scales; annuals or biennials. (11.)
11. Corollas white, sometimes abaxially rose- or purple-veined; calyculi of spreading to reflexed, unequal bractlets; annuals 70. Rafinesquia, p. 348
11. Corollas yellow, orange, purple, or pinkish to purplish; calyculi 0; annuals or biennials. (12.)
12. Leaves linear to lance-linear or lance-attenuate, entire; peduncles inflated distally; involucres campanulate; pappi of subulate scales; biennials (winter annuals); heads borne singly; corollas yellow or purple 59. Tragopogon, p. 303
12. Leaves mostly obovate to oblong-obovate, the distal ovate to linear, usually pinnately lobed, sometimes dentate or entire; peduncles little, if at all, inflated distally; involucres urceolate; pappi of bristles; annuals; heads borne singly or in corymbiform arrays; corollas yellow, sometimes abaxially striped with red 53. Urospermum, p. 296
10. Pappi of smooth or barbellate bristles, or of subulate scales or awns, or of aristate scales; annuals, biennials, or perennials. (13.)
13. Perennials. (14.)
14. Involucres narrowly cylindric, 1–2+ mm diam.; phyllaries (4–)5; florets 5; cypselae obovoid to lanceoloid 43. Mycelis (in part), p. 257
14. Involucres cylindric to campanulate, 2–5 mm diam.; phyllaries 5–21; florets 6–150+; cypselae cylindric to fusiform or lanceoloid. (15.)
15. Corollas usually bluish; cypselae lanceoloid 44. Mulgedium (in part), p. 258
15. Corollas yellow to white; cypselae cylindric to fusiform. (16.)
16. Phyllaries 8–21+; pappi: outer crowns of spreading, white hairs plus 2–3+ inner series of rufous to stramineous bristles 82. Pyrrhopappus, p. 376
16. Phyllaries 5–10; pappi of white bristles. (17.)
17. Calyculi of 3–4, minute bractlets; involucres cylindric; cypselae cylindric, 5+-ribbed (without rings of scales); pappi of 40–50+ bristles 38. Chondrilla, p. 252
17. Calyculi of 3–10+, deltate to lanceolate bractlets; involucres campanulate to cylindric; cypselae fusiform, 10-ribbed or -winged (with rings of scales at bases of beaks); pappi of 20–30 bristles 39. Ixeris, p. 254
13. Annuals or biennials. (18.)
18. Peduncles inflated distally (fistulose); pappi of aristate scales 65. Uropappus (in part), p. 322
18. Peduncles not inflated distally; pappi of bristles or subulate scales (not aristate). (19.)
19. Receptacles paleate; outer cypselae beakless; all or inner pappus bristles plumose 54. Hypochaeris, p. 297
19. Receptacles epaleate; all cypselae beaked; pappus bristles usually barbellulate or smooth (sometimes plumose in Helminthotheca ). (20.)
20. Calyculi 0; leaves spiny 47. Launaea (in part), p. 272
20. Calyculi of 2–16 bractlets; leaves not spiny. (21.)
21. Cypselae dimorphic, outer gibbous, inner ellipsoid to fusiform, not ribbed (rugulose); pappi of subulate to setiform scales 55. Helminthotheca, p. 300
21. Cypselae monomorphic, ellipsoid, oblong, obovoid, or lanceoloid, or cylindric to fusiform, ribbed; pappi of bristles (sometimes with minute outer crowns as well). (22.)
22. Cypselae ellipsoid to oblong or obovoid to lanceoloid; pappi of minute outer crowns plus inner bristles (sometimes 2–3+ series of bristles in Lactuca ). (23.)
23. Involucres narrowly cylindric, 1–2+ mm diam.; calyculi of 2–4 bractlets in 1 series 43. Mycelis (in part), p. 257
23. Involucres campanulate to cylindric, 2–5 mm diam.; calyculi of 3– 10+ bractlets in 2–3 series 45. Lactuca, p. 259
22. Cypselae fusiform, not or little compressed; pappi of bristles. (24.)
24. Stems dotted with tack-glands; pappus bristles basally connate (fall- ing in rings), white 61. Calycoseris, p. 307
24. Stems eglandular or glandular (without tack-glands); pappus bristles distinct or inner basally connate, white to tawny. (25.)
25. Heads usually in cymiform, corymbiform, or paniculiform arrays, sometimes borne singly; corollas usually yellow or orange, sometimes white, pink, or reddish; cypselae 10–20- ribbed 36. Crepis (in part), p. 222
25. Heads borne singly; corollas white to pale yellow; cypselae 4– 5-angled or -ribbed (angles roughened) 75. Glyptopleura, p. 361
1. Cypselae beakless. (26.)
26. Stems scapiform. (27.)
27. Pappi 0 or coroniform. (28.)
28. Perennials; leaves (± fleshy) linear to oblanceolate, margins entire; calyculi 0 80. Phalacroseris, p. 374
28. Annuals or biennials; leaves obovate or oblanceolate to spatulate, margins ± dentate or lobed; calyculi of 1–10+ bractlets or 0. (29.)
29. Leaf margins pinnately lobed; cypselae oblong, 10–13-ribbed 40. Lapsanastrum, p. 254
29. Leaf margins ± dentate to toothed; cypselae obovoid or subcylindric to weakly clavate, 4–5- or 8–10-ribbed. (30.)
30. Leaves oblanceolate to spatulate; heads borne singly or 2–3; peduncles naked, inflated distally; involucres broadly campanulate to urceolate; phyllaries basally connate; cypselae obovoid, 8–10-ribbed 49. Arnoseris, p. 276
30. Leaves obovate; heads in corymbiform arrays; peduncles sometimes bracteate, not inflated distally; involucres cylindro-campanulate; phyllaries distinct; cypselae subcylindric or weakly clavate, 4–5-ribbed (ribs corky) 62. Atrichoseris, p. 309
27. Pappi of bristles and/or scales, or of aristate scales. (31.)
31. Heads usually in corymbiform, paniculiform, or cymiform arrays; calyculi of 3–12 bractlets. (32.)
32. Leaves entire or dentate to pinnatifid (often lyrate or runcinate); cypselae 10– 20-ribbed; pappi of usually distinct, white to tawny bristles 36. Crepis (in part), p. 222
32. Leaves pinnately lobed (lyrate); cypselae 11–13-ribbed (ribs spiculate); pappi of basally connate, white bristles 41. Youngia, p. 255
31. Heads usually borne singly (sometimes in spiciform or paniculiform arrays in Launaea ); calyculi 0. (33.)
33. Pappi of subulate or aristate scales. (34.)
34. Heads erect; cypselae narrowed distally; pappi of lustrous, white subulate or aristate scales (bodies narrowly lanceolate to subulate) 67. Nothocalaïs (in part), p. 335
34. Heads nodding (at least in bud); cypsela apices truncate; pappus scales silvery to yellowish, brownish, or blackish (rarely white), plumose, barbellate, or barbellulate. (35.)
35. Involucres fusiform to ovoid or globose; pappi of deltate, lanceolate, linear, oblong, orbiculate, or ovate aristate scales, aristae plumose, barbellate, or barbellulate 68. Microseris (in part), p. 338
35. Involucres campanulate; pappi of narrowly lanceolate aristate scales, aristae barbellulate 69. Stebbinsoseris (in part), p. 346
33. Pappi wholly or partly of bristles (in Leontodon , pappi of outer cypselae of scales, of inner cypselae of bristles; sometimes scales + bristles in Krigia ). (36.)
36. Pappi of outer cypselae of scales, of inner cypselae of plumose bristles 52. Leontodon (in part), p. 294
36. Pappi of bristles or of outer scales plus inner bristles. (37.)
37. Phyllary margins papery (wider than midribs); receptacles paleate; pappus bristles plumose 63. Anisocoma, p. 309
37. Phyllary margins not papery; receptacles usually epaleate (sometimes paleate in Agoseris ); pappus bristles mostly smooth, barbellulate, or ± barbellate. (38.)
38. Leaves spiny; corollas yellow to ochroleucous; pappi of crisped (frizzy), outer and coarse, straight, inner bristles 47. Launaea (in part), p. 272
38. Leaves not spiny; corollas usually yellow to orange, sometimes pink, purple, or red; pappi of ± barbellate bristles. (39.)
39. Phyllaries in 1–2 series; cypselae 10–20-nerved or -ribbed; pappi of 5 outer scales plus 5–45 inner bristles 76. Krigia (in part), p. 362
39. Phyllaries in 2–5(–7) series; cypselae 8–10(–15)-ribbed; pappi of bristles in 1–6 series. (40.)
40. Phyllaries hairy or glabrous; cypselae usually beaked (6– 31 mm) 66. Agoseris (in part), p. 323
40. Phyllaries usually glabrous; cypselae not beaked (5–10 mm) (41.)
41. Plants taprooted; pappi of 30–80, whitish bristles 67. Nothocalaïs (in part), p. 335
41. Plants rhizomatous (usually in bogs); pappi of 24–48, brownish bristles 68. Microseris (in part), p. 338
26. Stems leafy. (42.)
42. Pappi 0, or of scales or aristate scales, or of awns plus bristles, or coroniform. (43.)
43. Pappi 0. (44.)
44. Stems winged, wings spiny; receptacles paleate 34. Scolymus (in part), p. 220
44. Stems not winged; receptacles epaleate. (45.)
45. Heads borne singly; florets 5–35; cypselae ± monomorphic (1.3–1.7 mm) 76. Krigia (in part), p. 362
45. Heads in corymbiform or thyrsiform arrays; florets 5–15; cypselae heteromorphic (3–25 mm, outer longer than inner). (46.)
46. Leaves ovate to suborbiculate, coarsely dentate; heads in corymbiform to thyrsiform arrays; cypselae subcylindric (curved), terete to slightly compressed, ± 20-ribbed 42. Lapsana, p. 257
46. Leaves ovate-lanceolate to linear, entire or dentate to pinnately lobed; heads in ± corymbiform arrays; cypselae ± terete (outer straight or arcuate, inner straight to coiled), not ribbed 56. Rhagadiolus, p. 300
43. Pappi of scales or aristate scales, or of awns or subulate scales plus bristles, or coroniform. (47.)
47. Corollas usually blue, sometimes pink or white; cypselae prismatic, 3–5-angled; pappi coroniform (erose scales) 35. Cichorium, p. 221
47. Corollas orange or yellow to white or pink to lavender; cypselae columnar or cylindric to fusiform or obconic, 6–15-ribbed; pappi usually of scales (scales sometimes aristate), sometimes of awns or subulate scales plus bristles, or coroniform ( Krigia wrightii , annuals, cypselae 1.3–1.6 mm). (48.)
48. Phyllaries enfolding outer cypselae. (49.)
49. Peduncles inflated distally; calyculi of 3–10+, deltate to lanceolate bractlets; phyllaries 5–13+ in 1 series; florets 8–30+; cypselae cylindric to fusiform, 12–15-ribbed; pappi of outer cypselae coroniform (short scales), of inner cypselae of lance-aristate to subulate-aristate scales (0–5+ outer and 5+ inner) 57. Hedypnois, p. 302
49. Peduncles not inflated distally; calyculi of 8–13, linear to filiform bractlets; phyllaries 20–25+ in 2+ series; florets 30–100+; cypselae columnar, 6–8(–10)-ribbed; pappi of setiform scales plus smooth or barbellate bristles 50. Tolpis, p. 277
48. Phyllaries not enfolding outer cypselae. (50.)
50. Heads ± nodding (at least in bud); pappi of aristate scales . (51.)
51. Perennials; involucres campanulate, fusiform, ovoid, or globose; pappus scales deltate, lanceolate, linear, oblong, orbiculate, or ovate, aristae plumose, barbellate, or barbellulate 68. Microseris (in part), p. 338
51. Annuals; involucres campanulate; pappus scales narrowly lanceolate, aristae barbellulate 69. Stebbinsoseris (in part), p. 346
50. Heads erect; pappi of setiform, subulate, or aristate scales or of awns or subulate scales plus bristles. (52.)
52. Perennials; leaves linear to linear-lanceolate, entire; corollas pale lavender to white; cypselae 5-angled or -ridged; pappi ofawns or subulate scales plus bristles 77. Chaetadelpha, p. 368
52. Annuals, biennials, or perennials; leaves ovate-lanceolate or oblong to lanceolate or linear, entire or pinnately lobed; corollas white, yellow, or purplish; cypselae sometimes 10-nerved; pappi of setiform, subulate, or aristate scales. (53.)
53. Leaves oblong or lanceolate to oblanceolate or linear, spiny; involucres campanulate to urceolate; receptacles paleate; corollas yellow; cypselae obovoid, ribs 0; pappi (0 or) of 2–4 setiform to aristate scales plus 0–4 muticous to lanceolate scales 34. Scolymus (in part), p. 220
53. Leaves ovate-lanceolate to lanceolate or linear, not spiny; involucres ovoid to cylindric; receptacles epaleate; corollas whitish to yellow or purplish; cypselae columnar to obclavate or fusiform, not compressed, usually 10-nerved; pappi of 28–50+ plumose to barbellate, subulate to setiform scales 60. Scorzonera, p. 306
42. Pappi wholly or mostly of bristles (sometimes with minute outer scales). (54.)
54. Shrubs; California 71. Munzothamnus, p. 349
54. Annuals, biennials, or perennials. (55.)
55. Corollas yellow, orange, ochroleucous, pinkish to reddish, or white (sometimes abaxially pinkish, reddish, or purplish). (56.)
56. Leaves spiny (on margins, bases often auriculate); phyllaries in 3–5+ series. (57.)
57. Involucres 3–5 mm diam.; phyllaries 18–25; cypselae weakly compressed, cylindric to prismatic or fusiform, 4–5-ribbed (or grooved), ribs muricate; pappi of crisped (frizzy, often basally coherent or connate), outer plus straight, coarse, inner bristles 47. Launaea (in part), p. 272
57. Involucres 5–15+ mm diam.; phyllaries 27–50; cypselae compressed, oblong or oblanceoloid to elliptic, 2–4(–5)-ribbed, smooth, rugose, or tuberculate; pappi of distinct outer bristles plus basally coherent or connate inner bristles 48. Sonchus, p. 273
56. Leaves not spiny (bases not auriculate); phyllaries in 1–2+, or 2–3, or 4–6 series. (58.)
58. Calyculi 0. (59.)
59. Heads borne singly; corollas yellow to orange; pappi (usually 2-seriate): outer of 5+ scales, inner of 5–45 bristles 76. Krigia (in part), p. 362
59. Heads in corymbiform to paniculiform arrays; corollas yellow to white; pappi (1 or 2-seriate): outer pappi 0, or of minute teeth plus 0–6 bristles, inner (or only) pappi of 15–35, basally coherent bristles 64. Malacothrix, p. 310
58. Calyculi usually of 3–16+ bractlets (sometimes intergrading with phyllaries). (60.)
60. Perennials; phyllaries in 2+ series; cypselae columnar or prismatic to ± urceolate 51. Hieracium, p. 278
60. Annuals, biennials, or perennials; phyllaries in 1–2 series; cypselae subcylindric to fusiform. (61.)
61. Corollas yellow or orange (white, pink, reddish), not abaxially reddish; cypselae 10–20-ribbed, sometimes spiculate; pappi persistent, of distinct bristles 36. Crepis (in part), p. 222
61. Corollas yellow, often abaxially reddish; cypselae 5–10-ribbed, transversely rugulose; pappi falling, of basally connate bristles 58. Picris, p. 302
55. Corollas usually purple, lavender, pink, or blue, sometimes white, rarely yellow. (62.)
62. Annuals. (63.)
63. Pappus bristles distinct, plumose (at least distally) 72. Stephanomeria (in part), p. 350
63. Pappus bristles basally connate, smooth or barbellulate. (64.)
64. Stems glandular-puberulent to glabrescent; leaves basal and cauline, blades spatulate to oblanceolate or minute, margins irregularly dentate to runcinate, often spinulose; peduncles minutely bracteate; cypselae columnar (tapering slightly proximally), 5-ribbed 73. Prenanthella, p. 359
64. Stems glabrous; leaves cauline (opposite proximally), blades linear, margins entire; peduncles ebracteate; cypselae subcylindric (apices abruptly constricted), 10-ribbed 78. Shinnersoseris, p. 368
62. Perennials. (65.)
65. Heads (usually nodding) in racemiform, thyrsiform, corymbiform, or paniculiform arrays; corollas usually white to pink or lavender, rarely yellow 46. Prenanthes, p. 264
65. Heads (erect) usually borne singly (sometimes in ± corymbiform to paniculiform arrays); corollas usually bluish, pink to lavender, or rose, rarely white. (66.)
66. Leaves oblong, elliptic, or ovate to lanceolate or linear; phyllaries 8–13+ in 1–2 series; florets (1–)15–50+; cypselae compressed, lanceoloid 44. Mulgedium (in part), p. 258
66. Leaves linear to oblong (distal sometimes bractlike or subulate distally); phyllaries 3–10 in ± 1 series; florets 3–16; cypselae not compressed, columnar. (67.)
67. Stems rigid, spine-tipped (thorny); leaves cauline, entire; phyllaries 3–5; florets 3–5 74. Pleiacanthus, p. 361
67. Stems not spine-tipped; leaves basal and cauline, entire, dentate, or pinnately lobed; phyllaries 4–10; florets (4–)5–16. (68.)
68. Cypselae subcylindric, subterete, angled, or sulcate; pappus bristles barbellate 79. Lygodesmia, p. 369
68. Cypselae columnar, 5-angled; pappus bristles (wholly or partly) plumose 72. Stephanomeria (in part), p. 350