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187. Asteraceae Martinov

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Key to Genera of Group 14 Heads eradiate; receptacles epaleate; pappi wholly, or partially, of awns or scales

1 Florets 1(–2) or (1–)2–4(–5+) per head and heads in second-order heads or in headlike clusters   (2)
+ Florets (2–)10–100(–1000) per head and heads borne singly or in usually corymbiform, paniculiform, racemiform, or spiciform arrays (heads ± crowded in Orochaenactis, annuals with pappi of 11–17 basally connate, oblanceolate scales, and in Eriophyllum mohavense, annuals with pappi of 12–14 distinct, linear to spatulate scales)   (7)
2 (1) Subshrubs or shrubs   (3)
+ Annuals or perennials   (4)
3 (2) Leaves alternate, blades narrowly lance-linear, margins serrate (± thickened, spiny near bases); corollas purplish   1 Hecastocleis (in part), v. 19, p. 71
+ Leaves opposite, blades lance-ovate to ovate, margins ± serrate (not notably thick- ened or spiny); corollas yellow   295 Lagascea (in part), v. 21, p. 136
4 (2) Annuals, 1–5 cm; leaves opposite (basal and cauline, crowded, seemingly whorled)   307 Dimeresia, v. 21, p. 182
+ Perennials, mostly 10–200 cm; leaves alternate (basal or basal and cauline, basal usually crowded)   (5)
5 (4) Plants mostly 100–200 cm (thistlelike); leaf blades 1–3-pinnately lobed, ultimate margins toothed (prickly or spiny)   9 Echinops (in part), v. 19, p. 85
+ Plants mostly 10–120 cm (not thistlelike); leaf blades not lobed, margins usually toothed, rarely entire (not prickly or spiny)   (6)
6 (5) Heads (1–)10–40 per cluster, each cluster subtended by (2–)3, ± deltate bracts; pappi of 5(–6), 1-aristate scales (look closely for squamiform, gradually to abruptly tapering base of each arista), no scales tipped with plicate aristae   29 Elephantopus, v. 19, p. 202
+ Heads 1–5+ per cluster, each cluster subtended by 1–2, lanceolate to spatulate or linear bracts; pappi of 6–10, ± laciniate to aristate scales, 2(–3+) of the aristate scales each with its awnlike arista plicate (2-folded) distally   30 Pseudelephantopus, v. 19, p. 204
7 (1) Leaves opposite (at least proximal, cauline sometimes mostly alternate) or whorled   (8)
+ Leaves alternate   (27)
8 (7) Corollas mostly yellow (sometimes with purple, red, or red-brown)   (9)
+ Corollas mostly white to cream or blue, lavender, pink, or purple   (17)
9 (8) Leaves and/or phyllaries dotted or streaked with pellucid (schizogenous) glands containing strong-scented oils   321 Tagetes (in part), v. 21, p. 235
+ Leaves and/or phyllaries rarely dotted or streaked (never with pellucid, schizogenous glands containing strong-scented oils, plants sometimes with sessile or stipitate, surface glands and sometimes otherwise strong-scented)   (10)
10 (9) Phyllaries often ± conduplicate and navicular; disc corollas usually 4-lobed; cypselae strongly flattened or weakly 3–4-angled, usually callous-margined, often ciliate   (11)
+ Phyllaries usually flat to weakly navicular (none in Carlquistia in which receptacular paleae function as an involucre); disc corollas (4–)5-lobed; cypselae usually obpyramidal to obconic, sometimes columnar, seldom cla-vate or cylindric, often 4–5-angled (seldom strongly compressed or flattened, callous-margined, and ciliate)   (12)
11 (10) Leaf blades usually lobed (mostly 5–30 mm); phyllaries 8–16 in 2–3 series, distinct   352 Perityle, v. 21, p. 317
+ Leaf blades usually triangular-hastate to narrowly deltate, seldom notably lobed (30–120 mm); phyllaries 15–21 in 1(–2) series, wholly or partially connate   353 Pericome, v. 21, p. 334
12 (10) Pappus scales (9–17) subulate, plumose   349 Carlquistia, v. 21, p. 302
+ Pappus scales not both subulate and plumose   (13)
13 (12) Leaves usually sessile, sometimes obscurely petiolate (rarely truly petiolate); pappus scales not notably medially thickened   (14)
+ Leaves usually petiolate; pappus scales usually notably medially thickened   (15)
14 (13) Leaves glabrous (often granular-glandular, not woolly)   355 Amblyopappus, v. 21, p. 348
+ Leaves ± woolly or tomentose (usually stems and/or phyllaries as well)   359 Eriophyllum (in part), v. 21, p. 353
15 (13) Annuals; leaves all or mostly opposite; pappus scales 8+   367 Schkuhria (in part), v. 21, p. 381
+ Annuals or biennials; leaves proximally opposite, mostly alternate; pappus scales 11–18   (16)
16 (15) Leaves lobed (lobes linear, 1–2 mm wide); disc corolla lobes lance-linear, lance-oblong, or linear (lengths mostly 2+ times widths); pappus scales distinct, narrowly lanceolate to subulate, some or all ± aristate   371 Hymenothrix (in part), v. 21, p. 387
+ Leaves lance-oblong to linear (not lobed); disc corolla lobes mostly deltate, lance-deltate, lanceolate, or ovate (lengths mostly 1–2 times widths); pappus scales basally connate (falling together), oblanceolate, obtuse   379 Orochaenactis, v. 21, p. 414
17 (8) Style-branch appendages essentially 0 or ± penicillate or deltate to lanceolate, linear, or filiform (lengths mostly 0–3+ times lengths of stigmatic areas or lines)   (18)
+ Style-branch appendages usually terete to clavate (lengths usually 2–5+ times lengths of stigmatic lines)   (20)
18 (17) Phyllaries often ± conduplicate and navicular; disc corollas usually 4-lobed; cypselae strongly flattened or weakly 3–4-angled, usually callous-margined, often ciliate   352 Perityle (in part), v. 21, p. 317
+ Phyllaries usually flat to weakly navicular; disc corollas 5-lobed; cypselae obpyramidal, 4-angled (not callous-margined and ciliate)   (19)
19 (18) Leaf blades broadly lanceolate to linear (not lobed); corollas usually pinkish or purplish, sometimes whitish; cypselae densely to sparsely hairy (hairs straight)   372 Palafoxia (in part), v. 21, p. 388
20 (17) Involucres narrowly cylindric, (1–)2–3 mm diam.; phyllaries 5(–6) in ± 1– 2 series; florets 5(–6)   397 Stevia (in part), v. 21, p. 483
+ Involucres campanulate, cylindric, ellipsoid, hemispheric, or obconic, (2–)3–7(–25) mm diam.; phyllaries (5–)8–45(–65+) in (1–)2–8+ series; florets (3–)10–125(–200+)   (21)
21 (20) Cypselae 8–10-ribbed (pappi of 1–5+ muticous, erose, lacerate, or lanceolate to subulate scales plus 9–12+ aristate scales)   398 Carphochaete, v. 21, p. 486
+ Cypselae (3–)4–5(–8)-ribbed   (22)
22 (21) Pappi of 8–13 plumose, setiform scales (coherent or ± connate, falling together or in groups)   409 Carminatia (in part), v. 21, p. 511
+ Pappi usually of 2–6(–12) muticous or aristate to subulate scales plus 0–6(–12) setiform scales, rarely coroniform (Ageratum)   (23)
23 (22) Phyllaries unequal; receptacles flat to convex (not warty)   (24)
+ Phyllaries ± equal; receptacles convex to conic or hemispheric (sometimes warty)   (25)
24 (23) Leaves mostly sessile (or nearly so), blades linear; cypselae ± fusiform   407 Malperia (in part), v. 21, p. 509
+ Leaves petiolate, blades ovate, deltate, or rhombic to lanceolate; cypselae prismatic   408 Pleurocoronis, v. 21, p. 510
25 (23) Leaves whorled (4 or 6 per node), blades linear; heads borne singly (plants ± aquatic)   401 Sclerolepis, v. 21, p. 488
+ Leaves mostly opposite (distal sometimes alternate), blades elliptic, lanceolate, or oblong; heads usually in cymiform to corymbiform arrays, sometimes borne singly   (26)
26 (25) Leaves petiolate; involucres 3–6 mm diam.; phyllaries usually 2-nerved; pappi usually of 5–6 aristate scales, rarely coroniform   396 Ageratum (in part), v. 21, p. 481
+ Leaves sessile; involucres 3–4(–5) mm diam.; phylla-ries obscurely 3–4-nerved; pappi of 2–6 setiform scales.   399 Trichocoronis, v. 21, p. 487
27 (7) Leaf margins prickly to spiny (plants ± thistlelike)   (28)
+ Leaf margins not prickly or spiny (tips sometimes spinose or apiculate; plants not thistlelike)   (34)
28 (27) Stems winged   (29)
+ Stems not or rarely winged (some Cirsium spp.)   (30)
29 (28) Cypsela attachments basal; pappus scales basally connate   10 Onopordum (in part), v. 19, p. 87
+ Cypsela attachments slightly lateral; pappus scales usually distinct, sometimes basally connate   12 Carduus (in part), v. 19, p. 91
30 (28) Leaves variegated (with white veins or mottlings; stamen filaments connate).   14 Silybum, v. 19, p. 164
+ Leaves not variegated (stamen filaments distinct)   (31)
31 (30) Corollas white or purplish to red; cypsela attachments basal; pappi of basally connate, plumose, setiform scales ("flattened bristles")   (32)
+ Corollas yellow to orange, red, or ± purple; cypsela attachments lateral; pappi 0 or of distinct, barbellulate, setiform scales ("flattened bristles") or subulate scales   (33)
32 (31) Involucres 35–100+ mm diam. (larger leaves 60–150 cm; receptacles becoming fleshy)   11 Cynara (in part), v. 19, p. 89
+ Involucres 10–50 mm diam. (larger leaves usually 20–50, sometimes to 110 cm; receptacles usually not notably fleshy)   13 Cirsium (in part), v. 19, p. 95
33 (31) Heads discoid (all florets bisexual, fertile); cypselae 4-angled   23 Carthamus (in part), v. 19, p. 178
+ Heads disciform (peripheral florets neuter); cypselae terete, 20-ribbed   24 Centaurea (in part), v. 19, p. 181
34 (27) Corollas mostly white or blue, lavender, pink, or purple   (35)
+ Corollas mostly yellow to orange (sometimes with purple, red, or red-brown)   (53)
35 (34) Anthers tailed (styles swollen or with rings of hairs proximal to branches)   (36)
+ Anthers not tailed (sometimes sagittate; styles not swollen or with rings of hairs proximal to branches)   (44)
36 (35) Heads discoid (all florets bisexual and fertile)   (37)
+ Heads disciform or radiant (peripheral florets usually neuter)   (39)
37 (36) Cypsela attachments basal (receptacles becoming fleshy; florets 100–250+; pappus scales basally connate)   11 Cynara (in part), v. 19, p. 89
+ Cypsela attachments ± lateral   (38)
38 (37) Phyllary appendages dentate or fringed (often spiny); pappi of persis- tent, nonplumose scales   24 Centaurea (in part), v. 19, p. 181
+ Phyllary appendages entire or lacerate, not fringed; pappi of readily falling, distally plumose bristles   17 Acroptilon, v. 19, p. 171
39 (36) Heads disciform   (40)
+ Heads radiant   (41)
40 (39) Phyllary appendages dentate or fringed, often spiny   24 Centaurea (in part), v. 19, p. 181
+ Phyllary appendages none (apices acute, entire)   22 Crupina, v. 19, p. 177
41 (39) Phyllary appendages none (apices spiny)   (42)
+ Phyllary appendages present   (43)
42 (41) Biennials or perennials; spines on phyllary apices soon falling; cypsela apices not coronate   19 Mantisalca (in part), v. 19, p. 173
+ Annuals; spines on phyllary apices persistent; cypsela apices coronate   20 Volutaria, v. 19, p. 174
43 (41) Cypselae compressed (oblong; attachment scars rimmed, rims whitish, swollen), apices denticulate   18 Amberboa, v. 19, p. 172
+ Cypselae ± terete (barrel-shaped; attachment scars not rimmed), apices entire   24 Centaurea (in part), v. 19, p. 181
44 (35) Phyllaries 18–70+ in 3–8+ series   (45)
+ Phyllaries 5–15(–21) in 1–2(–3) series   (48)
45 (44) Heads pseudo-radiant or -radiate (corollas of peripheral, bisexual florets enlarged, zygomorphic); phyllary margins (at least of the outer), pectinately spinose-toothed   28 Stokesia, v. 19, p. 201
+ Heads discoid; phyllary margins not pectinately spinose-toothed   (46)
46 (45) Annuals (perhaps persisting); cypselae not ribbed   31 Cyanthillium, v. 19, p. 204
+ Perennials (sometimes functionally annuals); cypselae 8–10-ribbed   (47)
47 (46) Heads each subtended by 3–8+, ± foliaceous bracts; pappi readily falling   32 Centratherum, v. 19, p. 206
+ Heads not each subtended by foliaceous bracts; pappi persistent   33 Vernonia, v. 19, p. 206
48 (44) Style-branch appendages usually terete to clavate (lengths usually 2–5+ times lengths of stigmatic lines)   (49)
+ Style-branch appendages mostly deltate to lanceolate, linear, or filiform, sometimes essentially none (lengths mostly 0–3+ times lengths of stigmatic areas or lines)   (50)
49 (48) Phyllaries 5(–6) in 1 series; florets 5(–6)   397 Stevia (in part), v. 21, p. 483
+ Phyllaries 12–15 in 2–3 series; florets 7–10   413 Hartwrightia (in part), v. 21, p. 540
50 (48) Heads in (usually secund) spiciform arrays; florets 3   153 Thurovia, v. 20, p. 86
+ Heads borne singly or in corymbiform or cymiform arrays; florets 8–70   (51)
51 (50) Cypselae stoutly obconic to obpyramidal, usually 4-angled and 12–16-ribbed (lengths usually 1–2, rarely to 3.5 times diams.; phyllary margins usually notably membranous to scarious)   351 Hymenopappus (in part), v. 21, p. 309
+ Cypselae narrowly clavate to cylindric or compressed, obscurely 8–20-angled, or ± quadrangular with 8–12 obscure nerves (lengths usually 3+ times diams., if stouter, usually ± compressed; phyllary margins not notably membranous or scarious)   (52)
52 (51) Leaves mostly basal; heads borne singly; florets 10–30+ (per-ennials 2–9 cm, 10–20+ cm across; leaf blades 1- or 3-nerved, cordate, elliptic, ovate, or rounded, margins entire or distally ± crenate, revolute to ± plane, faces ± strigose and gland-dotted, adaxial sometimes glabrescent)   376 Chamaechaenactis, v. 21, p. 395
+ Leaves basal and cauline or mostly cauline; heads borne singly or in ± cymiform arrays; florets 8–70+   378 Chaenactis (in part), v. 21, p. 400
53 (34) Shrubs; pappi of 12–30 scales (sometimes with bristles as well)   (54)
+ Annuals, biennials, perennials, subshrubs, or shrubs; pappi mostly of 1–12(–20+) scales (scales sometimes subtending 20–100+ bristles; if shrubs, pappus scales 1 or 3–5)   (58)
54 (53) Phyllaries 4–6, or 8–18, in 2 series, ± equal   (55)
+ Phyllaries 7–60+ in 3–5 series, mostly unequal   (56)
55 (54) Florets 4–8 (cypselae copiously pilose, hairs white, 4–14 mm, sometimes obscuring pappi)   237 Tetradymia (in part), v . 20, p. 629
+ Florets 12–21 (cypselae hirsute, hairs tawny to reddish, 0.5–1 mm; pappi often of both scales and bristles)   363 Peucephyllum (in part), v. 21, p. 378
56 (54) Phyllaries 40–60+ in 3–5 series (often in vertical ranks)   175 Chrysothamnus (in part), v. 20, p. 187
+ Phyllaries 7–25 in 3 series   (57)
57 (56) Involures campanulate, globose, or hemispheric, 4–13 × 7–16 mm; florets 13–45   173 Acamptopappus (in part), v. 20, p. 184
+ Involures turbino-cylindric, 4–5.5 × 2–3 mm; florets 3–7 (functionally staminate)   174 Amphipappus (in part), v. 20, p. 185
58 (53) Shrubs (stems ± tomentose; leaf blades obscurely pinnate; heads borne singly or in loose, corymbiform arrays; pappi of 1 or 3–5 scales)   125 Pentzia (in part), v. 19, p. 543
+ Annuals, biennials, perennials, or subshrubs   (59)
59 (58) Leaves usually 1–3-pinnate (usually aromatic when crushed); phyllary margins usually notably scarious; style-branch appendages essentially none; pappus scales usually rudimentary (3–5, subulate in Sphaeromeria)   (60)
+ Leaves usually not lobed, sometimes 1–4-pinnate (seldom notably aromatic when crushed); phyllary margins seldom notably scarious; style-branch appendages mostly deltate to lanceolate or linear; pappus scales usually conspicuous (sometimes associated with bristles)   (62)
60 (59) Annuals; heads discoid   124 Matricaria (in part), v. 19, p. 540
+ Perennials; heads disciform (peripheral florets pistillate)   (61)
61 (60) Heads usually in lax to dense, corymbiform arrays, rarely borne singly; involucres (3–)5–10 mm diam.; florets 60–300+   112 Tanacetum (in part), v. 19, p. 489
+ Heads usually (8–20+) in tight, capitate arrays; involucres (2–)3–4 mm diam.; florets 30–50+   118 Sphaeromeria (in part), v. 19, p. 499
62 (59) Phyllaries 25–125 in 3–9 series; pappi of scales only or of 5–20 scales plus 20–100+ bristles   (63)
+ Phyllaries 6–25(–60) in 2–3 series; pappi mostly wholly of scales (usually 5, 10, 15, or 20, subulate-aristate scales in Pentachaeta)   (65)
63 (62) Phyllary apices often looped, hooked, patent, recurved, straight, or incurved; pappi fragile or readily falling, of 8–15 subulate to setiform scales   202 Grindelia (in part), v. 20, p. 424
+ Phyllary apices usually erect and straight, sometimes spreading or reflexed; pappi persistent or readily falling, usually of scales plus bristles   (64)
64 (63) Heads usually in corymbiform, ± paniculiform, or subumbelliform arrays, sometimes borne singly; phyllaries in 3–5 series, unequal, usually thickened or keeled, 1-nerved (nerves not golden-resinous); cypselae 3-angled or 4–12-ribbed   185 Heterotheca (in part), v. 20, p. 230
+ Heads borne singly or in 2s or 3s; phyllaries in 2–3 series, equal or subequal, flat, usually 1–3-nerved (nerves golden-resinous); cypselae 2-nerved   186 Erigeron (in part), v. 20, p. 256
65 (62) Cypselae narrowly clavate to oblanceoloid, or columnar to obconic or obpyramidal (lengths usually 3+ times diams., if stouter, usually ± compressed)   (66)
+ Cypselae stoutly obconic to obpyramidal (lengths usually 1–2, rarely to 3.5, times diams.)   (68)
66 (65) Leaves usually sessile, sometimes obscurely petiolate (rarely truly petiolate); pappus scales often aristate, not medially thickened   145 Pentachaeta (in part), v. 20, p. 46
+ Leaves usually petiolate (± sessile in some spp. of Chaenactis); pappus scales sometimes aristate or lacerate, often notably medially thickened   (67)
67 (66) Pappus scales 4–20, not lacerate (usually erose, sometimes uniaristate)   378 Chaenactis (in part), v. 21, p. 400
+ Pappus scales 5, lacerate (divisions bristlelike)   381 Trichoptilium, v. 21, p. 418
68 (65) Phyllaries usually strongly reflexed in fruit; receptacles mostly globose (sometimes with setiform enations); disc corollas often brown-purple to red-brown or tipped with brown-purple to red-brown (tubes much shorter than abruptly much-dilated, urceolate to campanulate throats, lobes often shaggily hairy, hairs ± moniliform)   (69)
+ Phyllaries mostly erect to spreading in fruit; receptacles flat, conic, domed, hemispheric, or ovoid (smooth or pitted, without setiform enations); disc corollas usually uniformly yellow to cream, sometimes purplish to reddish (tubes much shorter than to about equaling slightly dilated, funnelform to cylindric throats, lobes not shaggily hairy with moniliform hairs)   (70)
69 (68) Stems not winged (receptacles usually with setiform enations; style-branch apices ± attenuate)   384 Gaillardia (in part), v. 21, p. 421
+ Stems often winged (by decurrent leaf bases; receptacles rarely with setiform enations; style-branch apices penicillate or truncate)   385 Helenium (in part), v. 21, p. 426
70 (68) Leaf blades simple or 1–2-pinnately lobed (lobes mostly filiform, linear, or oblong), ultimate margins entire or toothed, faces glabrous or hairy, usually ± gland-dotted (often in pits); phyllaries persistent (or inner falling), usually (6–)6–30(–40) in 2 series and unequal (outer usually connate), sometimes 28–50 in 2–3 series and subequal (usually spreading to erect in fruit)   386 Hymenoxys (in part), v. 21, p. 435
+ Leaf blades mostly oblanceolate to linear or filiform, sometimes lobed, ultimate margins usually entire, sometimes toothed, faces glabrous or ± hairy, eglandular or ± gland-dotted; phyllaries usually persistent, 11–60+ in 2–3 series (mostly spreading to erect in fruit, distinct, herbaceous; outer with or without scarious margins, abaxial faces ± hairy; mid usually same number as, alternating with, and similar to outer, almost always with ± scarious margins; inner narrower than others, margins scarious)   389 Tetraneuris (in part), v. 21, p. 447

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