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beggar-ticks, beggars-ticks, bident, bur-marigold, fourchette, sticktight

Habit Annuals, perennials, subshrubs, or shrubs, 10–400 cm (sometimes rhizomatous or with cormiform bases, stoloniferous in Coreopsis auriculata). Annuals or perennials [shrubs, vines], 5–400 cm.
Stems

usually 1, usually erect, (terete or 4-angled, often striate or sulcate) branched distally or ± throughout.

Leaves

usually cauline (sometimes mostly basal); usually mostly opposite (distal sometimes alternate), rarely whorled;

petiolate or sessile;

blades (often pinnately or palmately lobed, sometimes compound), or lobes or leaflets, mostly deltate, elliptic, filiform, lanceolate, linear, oblanceolate, or ovate, ultimate margins entire or toothed, faces usually glabrous or glabrate, sometimes hairy (rarely, if ever, gland-dotted).

usually cauline; usually opposite, rarely whorled, distal sometimes alternate;

petiolate or sessile;

blades simple, compound (leaflets petiolulate), or 1–3+-pinnatisect or -pinnately lobed (submerged leaves multifid in B. beckii, an aquatic), ultimate margins entire, dentate, laciniate, serrate, or toothed, faces usually glabrous, sometimes hirtellous, hispidulous, pilosulous, puberulent, scabrellous, or strigillose.

Involucres

mostly campanulate, cylindric, turbinate, or hemispheric or broader.

mostly hemispheric or campanulate to cylindric, (1–)4–12(–25+) mm diam.

Receptacles

mostly flat to convex, paleate (paleae often stramineous to white with red-brown to purplish striae, orbiculate to oblong or linear, ± flat to slightly cupped).

flat or slightly convex, paleate;

paleae usually falling, (usually stramineous, sometimes yellow to orange, with darker striae) ± flat to slightly navicular.

Ray florets

0 or 1–21+ (more in cultivars), pistillate and fertile, or neuter, or styliferous and sterile;

corollas usually yellow to orange (sometimes bicolored: yellow to orange with brown, red-brown, or purple), sometimes cyanic or white.

usually 1–21+ (often 3, 5, 8, or 13), sometimes 0, usually neuter, sometimes styliferous and sterile;

corollas usually yellow, sometimes white or pinkish.

Disc florets

3–150+, usually bisexual, fertile (functionally staminate in Dicranocarpus);

corollas usually yellow to orange, sometimes brown, red-brown, or purple, tubes usually shorter than, sometimes equaling funnelform throats (longer than throats in some Thelesperma spp.), lobes (3–)5, ± deltate to lance-ovate; (staminal filaments hairy in Cosmos) anther thecae pale or dark;

stigmatic papillae in 2 lines.

(5–)12–60(–150+), bisexual, fertile;

corollas usually yellow to orange, sometimes whitish [purplish], tubes shorter than throats, lobes (3–)5, ± deltate (staminal filaments glabrous; style-branch appendages deltate or lanceolate to subulate).

Phyllaries

persistent, 3–34+ in ± 2 series (usually distinct, notably connate in Thelesperma), usually ± membranous, margins usually ± scarious.

persistent, mostly (4–)8–21(–30+) in ± 2 series, usually distinct, sometimes connate 0.05–0.1 their lengths, mostly oblong or ovate to lance-oblong, chartaceous to membranous or scarious (usually striate with brownish nerves, margins usually hyaline).

Calyculi

usually of 3–8(–21+) bractlets or bracts (usually ± herbaceous, usually shorter than phyllaries and/or reflexed, sometimes ± foliaceous and surpassing phyllaries).

of (3–)5–13(–21+) erect to spreading or reflexed, ± herbaceous (sometimes foliaceous) bractlets or bracts (sometimes surpassing phyllaries).

Heads

radiate or discoid, borne singly or in ± corymbiform, cymiform, or paniculiform arrays.

usually radiate or discoid, sometimes ± disciform, usually in corymbiform arrays, sometimes in 2s or 3s or borne singly.

Cypselae

usually either obcompressed to obflattened, usually cuneate, linear, oblong, orbiculate, or ovate (bodies unequally 3–4-angled, margins often winged), or (all or inner) ± equally 4-angled and linear-fusiform, sometimes ± beaked, faces glabrous or hairy (often striate, tuberculate, or papillate);

pappi 0, or persistent, of (1–)2–4(–8), usually ± barbellate awns or scales, rarely coroniform or of 1–2, smooth to ciliate or barbed awns or scales.

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 (± corky-winged in B. aristosa, B. cernua, and B. polylepis) usually retrorsely, sometimes patently or antrorsely, barbed or ciliate, apices sometimes attenuate, not beaked [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

= 13.

Asteraceae (tribe Heliantheae) subtribe Coreopsidinae

Bidens

Distribution
Mostly subtropical and warm-temperate New World and Old World
map from USDA
Widespread; especially in subtropical; tropical; and warm-temperate North America and South America
[BONAP county map]
Discussion

Genera 20, species 374 (7 genera, 69 species in the flora).

The circumscription of Coreopsidinae followed here (H. Robinson 1981) is a bit narrower than the traditional one. The subtribe is remarkable among Heliantheae for having a relatively high number of species native in subtropical and tropical Old World, especially Africa. Distinctions among some genera (e.g., Bidens, Coreopsis, and Cosmos) are often subtle.

In Coreopsidinae, each involucre is subtended by a calyculus of more or less herbaceous (sometimes leaflike) bractlets or bracts (sometimes surpassing the phyllaries). In keys and descriptions here, shapes, heights, and diameters given for involucres are based on the phyllaries collectively (exclusive of calyculi) at flowering; the involucres are sometimes notably larger in fruit.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species 150–250+ (25 in the flora).

Within species of Bidens, leaves may be simple, 1-pinnately compound, or 1–3-pinnatisect or -pinnately or -ternately lobed. For compound leaves, leaflets are described; for leaves mostly 1-pinnately lobed, primary lobes are described; for leaves 2–3-pinnatisect or -pinnately lobed, the ultimate lobes are described. Involucres in Bidens species are subtended by more or less herbaceous (sometimes foliaceous) bractlets or bracts; collectively, they constitute calyculi and they often well surpass the phyllaries. Together, phyllaries constitute involucres. In keys and descriptions here, shapes, heights, and diameters given for involucres are for involucres at flowering; involucres are sometimes notably larger in fruit. In some Bidens species, the outer and inner cypselae are obcompressed (unequally 3–4-angled) to flat, are mostly cuneate to obovate or nearly linear (broadest at or near apices), and are more or less similar except for sizes (outer smaller); in other species, the outer cypselae are obcompressed to flat and obovate to cuneate or linear and the inner are more or less equally 4-angled (in cross section) and linear-fusiform (thickest near their middles and more or less attenuate toward their tips).

Etymology: Latin bis, two, and dens, tooth, alluding to 2-awned pappi of the original species

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Parent taxa Asteraceae > tribe Heliantheae Asteraceae > tribe Heliantheae > subtribe Coreopsidinae
Subordinate taxa
Bidens, Coreocarpus, Coreopsis, Cosmos, Dicranocarpus, Heterosperma, Thelesperma
B. amplissima, B. aristosa, B. aurea, B. beckii, B. bidentoides, B. bigelovii, B. bipinnata, B. cernua, B. connata, B. discoidea, B. eatonii, B. frondosa, B. heterodoxa, B. heterosperma, B. hyperborea, B. laevis, B. lemmonii, B. leptocephala, B. mitis, B. pilosa, B. polylepis, B. tenuisecta, B. trichosperma, B. tripartita, B. vulgata
Key
1. Phyllaries (excluding calyculi) 3–6 in 1(–2) series; disc florets 3–4+ (functionally staminate)
Dicranocarpus
1. Phyllaries (3–)8–34+ in ± 2 series; disc florets 3–10, 10–20, or (5–)12–150+ (bisexual, fertile)
→ 2
2. Phyllaries connate 1/5–7/8+ their lengths
Thelesperma
2. Phyllaries usually distinct, rarely connate ± 1/10 their lengths
→ 3
3. Cypselae (at least inner) ± 4-angled, ± linear-fusiform, often apically attenuate or beaked (none winged)
→ 4
3. Cypselae all ± obcompressed (sometimes winged)
→ 5
4. Disc florets 10–20 (staminal filaments hairy near anthers); cypselae usually with 1 groove on each face
Cosmos
4. Disc florets (5–)12–150+ (staminal filaments not hairy); cypselae with 0 or 2grooves on each face
Bidens
5. Annuals; ray florets 1–3 (laminae 1–2+ mm); cypselae mostly ellipsoid orobovoid (inner obovoid to obscurely urceolate, ± beaked)
Heterosperma
5. Annuals, perennials, subshrubs, or shrubs; ray florets usually 1, 2, 3, 5, 8, 13, or 21+, sometimes 0 (laminae mostly 4–30+ mm); cypselae mostly cuneate, linear, oblanceolate, oblong, ± orbiculate, or ovate (not beaked).
→ 6
6. Cypselae rarely winged (margins sometimes thickened, winged in B. aristosa and B. polylepis); pappi usually of barbellate (rarely smooth) awns, sometimes 0
Bidens
6. Cypselae (some or all) usually thin-margined or ± winged; pappi usually 0, sometimes coroniform, or of 2 bristly cusps or scales (in Coreopsis), or of 1–2 retrorsely barbellate awns (in Coreocarpus)
→ 7
7. Ray florets usually neuter or styliferous and sterile; wings of cypselae membranous, chartaceous, or corky, entire or lobed to toothed, sometimes ciliate
Coreopsis
7. Ray florets usually pistillate and fertile; wings of cypselae ± corky, ± pectinately toothed
Coreocarpus
1. Leaves (some or most) multifid (aquatics, multifid leaves usually submersed, ultimate lobes mostly 0.1–0.3 mm diam.); pappus awns 13–25(–40) mm
B. beckii
1. Leaves simple, 1-pinnately compound, or 1–3-pinnatisect or -pinnately lobed (plants sometimes rooted in mud, submersed leaves usually 0; leaflets or lobes mostly 0.5–40+ mm wide); pappus awns 0 or (0.1–)2–4(–8.5+) mm
→ 2
2. All or inner cypselae ± equally 4-angled, mostly linear-fusiform (usually thickest in middle 1/2 and distally ± attenuate proximally and distally, each of 4 faces usually 2-grooved)
→ 3
2. All cypselae ± flattened (if 3–4-angled, 2 angles ± acute and 1–2 ± obtuse), usually cuneate, linear, oblanceolate, or obovate, seldom ± linear-fusiform (usually broadest in distal 1/3)
→ 10
3. Involucres ± campanulate to cylindric, 1–3(–4) mm diam.; disc florets (3–)5–9(–13)
→ 4
3. Involucres campanulate to turbinate or hemispheric or broader, (3–)4–8 mm diam.; disc florets (5–)20–80+
→ 7
4. Leaves: primary lobes lance-rhombic or ovate to lanceolate, 15–30(–50+) × 5–15(–30+) mm; pappi (1–)2–4 mm
B. bigelovii
4. Leaves: primary or ultimate lobes ovate, oblanceolate, lanceolate, or spatulate to linear or filiform, 2–25+ × 0.5–5(–15) mm; pappi 0 or 1–3 mm
→ 5
5. Calyculus bractlets spatulate to linear (or 3–5+ lobed, some ± foliaceous), 3–10(–25+) mm (usually surpassing phyllaries)
B. lemmonii
5. Calyculus bractlets linear to subulate or filiform (not foliaceous), 1–4 mm (seldom surpassing phyllaries)
→ 6
6. Leaves: ultimate lobes 0.5–1(–2) mm wide; cypselae usually glabrous
B. heterosperma
6. Leaves: ultimate lobes (1–)2–3(–15) mm wide; cypselae ± hispidulous (at least distally)
B. leptocephala
7. Leaves: ultimate lobes ± linear, (2–)5–15+ × (0.5–)2–3+ mm
B. tenuisecta
7. Leaves: blades simple, or primary or ultimate lobes lance-rhombic, lance-deltate, ovate, or lanceolate, (10–)15–30(–90+) × (5–)10–15(–40+) mm
→ 8
8. Leaf blades either ovate to lanceolate and 30–70(–120) × 12–18(–45) mm, or 1-pinnately lobed, primary lobes 3–7, ovate to lanceolate, (10–)25–80+ × (5–)10–40+ mm; disc florets 20–40(–80+)
B. pilosa
8. Leaf blades rounded-deltate to ovate or lanceolate overall, (20–)25–90 × 15–35(–60+) mm, usually 2–3-pinnatisect, ultimate lobes lanceolate, lance-rhombic, obovate, or ovate, 15–30(–50+) × 5–15(–30+) mm; disc florets 10–20(–30+)
→ 9
9. Outer cypselae 6–7 mm, inner cypselae 10–14 mm; pappi of 2(–3) awns
B. bigelovii
9. Outer cypselae 7–15 mm, inner cypselae 12–18 mm; pappi of (2–)3–4 awns
B. bipinnata
10. Blades of all or most leaves either 1-pinnately compound (leaflets 3–5+, petiolulate) or 1–2-pinnately lobed (ultimate lobes lanceolate or oblanceolate to lance-linear or linear, usually 1–12 mm, sometimes to 25 mm wide); calyculus bractlets seldom foliaceous
→ 11
10. Blades of all or most leaves simple, margins ± dentate to serrate or margins coarsely incised (lobes 3–7+); calyculus bractlets or bracts often foliaceous.
→ 18
11. Ray florets 0, or 1–3+, or 3–5, laminae 2–3.5 mm
→ 12
11. Ray florets (5–)8–13, laminae 10–30 mm
→ 14
12. Calyculus bractlets (3–)4(–5+), seldom ciliate; disc florets (10–)15–20+
B. discoidea
12. Calyculus bractlets 5–21, usually ciliate; disc florets 20–150
→ 13
13. Leaves usually 3(–5)-foliolate (leaflets lanceolate to lance-ovate); calyculus bractlets (5–)8(–10)
B. frondosa
13. Leaves usually laciniately 1-pinnatisect or 2–5-foliolate (leaflets or lobes ± lanceolate), rarely 2–3-pinnatisect; calyculus bractlets 10–16(–21)
B. vulgata
14. Phyllaries ovate to lanceolate (margins usually yellow); ray florets 5–6; margins of cypselae not barbed or ciliolate
B. aurea
14. Phyllaries lance-linear, lanceolate, lance-ovate or oblong (margins not notably yellow); ray florets (5–)8–13; margins of cypselae usually barbed or ciliate (except B. mitis)
→ 15
15. Cypselae narrowly cuneate (lengths mostly 2.5–4 times widths)
B. trichosperma
15. Cypselae broadly cuneate, oblanceolate, or obovate (lengths mostly 1.5–2.5 times widths)
→ 16
16. Cypselae 2.5–5 mm, margins (not winged) not barbed or ciliate, faces glabrous or hirtellous; pappi 0, or of 2 antrorsely barbed awns or± deltate scales 0.5(–1) mm
B. mitis
16. Cypselae (4–)5–8 mm, margins (often ± corky-winged) usually barbed or ciliate, faces glabrate, glabrous, or strigillose; pappi 0, or of 2(–4) antrorsely or retrorsely barbed awns or scales 0.1–6 mm
→ 17
17. Calyculus bractlets 8–12(–16), (4–)5–7(–12) mm
B. aristosa
17. Calyculus bractlets 12–21+, (6–)8–12(–20+) mm
B. polylepis
18. Leaves sessile
→ 19
18. Leaves petiolate, petioles (5–)10–25(–60) mm (sometimes ± winged)
→ 24
19. Calyculus bractlets (3–)8–12(–25+) mm, or (6–)10–12(–20+) mm, usually spreading to reflexed
→ 20
19. Calyculus bractlets (6–)10–30(–75+) mm, usually erect, sometimes spreading
→ 21
20. Rays usually 6–8, sometimes 0, laminae 2–15(–18) mm; outer cypselae (3–)5–6+ mm, inner 4–8 mm (margins ± thickened or winged); pappi of (2–)4 awns (1–)2–4 mm (paleae stramineous to yellowish at tips)
B. cernua
20. Rays usually 7–8, rarely 0, laminae (10–)15–25(–30) mm; outer cypselae 6–8 mm, inner cypselae 8–10 mm (margins not notably thickened or winged); pappi of 2–4 awns 3–5 mm (paleae usually orange at tips)
B. laevis
21. Involucres (4–)7–10 mm diam.; calyculus bractlets erect; disc florets 15–30(–60);cypselae narrowly cuneate (faces ± 1-nerved and ± striate)
B. hyperborea
21. Involucres (3–)6–12(–15+) or 18–25 mm diam.; calyculus bractlets ± erect to spreading or reflexed; disc florets (5–)25–60(–150+); cypselae cuneate to oblanceolate or linear (faces usually smooth or weakly striate, sometimes tuberculate)
→ 22
22. Involucres 6–10 × 18–25 mm; ray florets usually 6–11
B. amplissima
22. Involucres (4–)5–7(–12) × (3–)6–12(–15+) mm; ray florets usually 0, sometimes 1–5
→ 23
23. Cypselae ± flattened, sometimes weakly 3(–4)-angled, faces usually smooth, seldom notably tuberculate (disc corollas gradually ampliate, anthers usually pale)
B. tripartita
23. Cypselae (at least inner) usually ± 4-angled, faces usually tuberculate (disc corollas abruptly ampliate, anthers usually dark, blackish)
B. connata
24. Ray florets 8–13 (laminae 12–25+ mm); cypselae 2.5–4.5 mm (margins not barbed or ciliate)
B. mitis
24. Ray florets 0 or 1–11 (laminae 2–12 mm); cypselae (3–)6–13 mm (margins sometimes barbed or ciliate)
→ 25
25. Involucres ± hemispheric or broader, 6–10 × 18–25+ mm; ray florets 6–11 (laminae 4–8 mm)
B. amplissima
25. Involucres cylindric, campanulate, or hemispheric or broader, 4–10(–16) × (3–)5–8(–15) mm; ray florets 0 or 1–7 (laminae 2–12 mm)
→ 26
26. Involucres campanulate to hemispheric or broader; disc florets (5–)20–60(–150+); cypsela faces usually smooth or tuberculate (not notably striate)
→ 27
26. Involucres usually campanulate to cylindric, sometimes ± hemispheric; disc florets (6–)10–25(–60); cypsela faces usually ± striate, sometimes tuberculate
→ 28
27. Cypselae ± flattened, sometimes weakly 3(–4)-angled, faces usually smooth, seldom notably tuberculate (disc corollas gradually ampliate, anthersusually pale)
B. tripartita
27. Cypselae (at least inner) usually ± 4-angled, faces usually tuberculate (disc corollas abruptly ampliate, anthers usually dark, blackish)
B. connata
28. Petioles ± winged; involucres usually ± campanulate to cylindric, sometimes ± hemispheric, 5–7(–9) × (4–)7–10 mm; disc florets 15–30(–60)
B. hyperborea
28. Petioles not notably winged; involucres mostly campanulate to cylindric, (4–)8–10(–16) × (4–)7–9(–12) mm; disc florets 6–20(–35+)
→ 29
29. Cypselae narrowly cuneate to linear, margins evenly antrorsely strigillose, faces ± evenly antrorsely strigillose
B. bidentoides
29. Cypselae broadly to narrowly cuneate or oblanceolate to linear, margins usually barbed or ciliate (proximally antrorsely, distally retrorsely), faces glabrous or sparsely setulose
→ 30
30. Involucres (4–)8–10(–12) × (4–)7–9 mm; cypselae: outer 5.5–10 mm, inner 6.5–11 mm
B. eatonii
30. Involucres (4–)6–7 × (4–)5–7+ mm; cypselae: outer 4–5(–6) mm, inner (5–)6–7(–8+) mm
B. heterodoxa
Synonyms subtribe Coreopsideae, subtribe Petrobiinae Megalodonta
Name authority Lessing: Linnaea 5: 153. (1830) Linnaeus: Sp. Pl. 2: 831. (1753): Gen. Pl. ed. 5, 362. (1754)
Source FNA vol. 21, p. 183. FNA vol. 21, p. 205. Treatment authors: John L. Strother, Ronald R. Weedon.
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