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FNA | Family List | FNA Vol. 21 | Asteraceae

386. Hymenoxys Cassini in F. Cuvier, Dict. Sci. Nat. ed. 2. 55: 278. 1828.

Bitterweed, rubberweed [Greek hymen, membrane, and oxys, sharp, alluding to aristate pappus scales]

Mark W. Bierner

Dugaldia Cassini; Plummera A. Gray

Annuals, biennials, or perennials, 5–150 cm (sometimes with ± branched, woody caudices or stout rhizomes). Stems 1–30+, erect, unbranched or branched, green throughout to purple-red-tinted proximally or distally to purple-red-tinted throughout, glabrous or ± hairy. Leaves alternate; blades simple or 1–2-pinnately lobed, ultimate margins entire or toothed, faces glabrous or hairy, usually ± gland-dotted (often in pits). Heads radiate [discoid], borne singly or in paniculiform to corymbiform arrays. Involucres subhemispheric, hemispheric, globoid, campanulate, or urceolate, (2.5–)6–32 mm diam. Phyllaries persistent (or inner falling), usually (6–)16–30(–40) in 2 series and unequal, sometimes 28–50 in 2–3 series and subequal (usually spreading to erect in fruit). Receptacles usually hemispheric, globoid, ovoid, or conic (flat in H. ambigens), smooth or pitted, epaleate. Ray florets usually (3–)8–13(–16), sometimes 14–34 [0], pistillate, fertile; corollas (usually withering after flowering, falling early or tardily) yellow or yellow-orange to orange (laminae fan-shaped, lobes 3–5). Disc florets usually 25–150(–400+), usually bisexual and fertile (6–15, functionally staminate in H. ambigens); corollas yellow to yellow-brown proximally, yellow distally, tubes shorter than cylindric to cylindric-campanulate throats, lobes 5, ± deltate. Cypselae obconic or obpyramidal, glabrous or hairy; pappi 0, or persistent, of 2–11(–15) usually aristate scales. x = 15.

Species 25 (17 in the flora): North America, Mexico, Central America, South America. Subgenus Hymenoxys, which occurs only in South America, contains four species, three of which are discoid.

Subgenus Phileozera (Buckley) Cockerell includes Hymenoxys odorata and H. chrysanthemoides (Kunth) de Candolle (Mexico) (M. W. Bierner 2001). Subgenus Plummera (A. Gray) Bierner comprises only Hymenoxys ambigens (Bierner 1994, 2001). Subgenus Dugaldia (Cassini) Bierner includes Hymenoxys hoopesii, H. integrifolia (Kunth) Bierner (Mexico and Guatemala), and H. pinetorum (Standley) Bierner (Mexico; Bierner 1994, 2001). Subgenus Rydbergia (Greene) Bierner comprises Hymenoxys brandegeei, H. grandiflora, and H. insignis (Mexico) (Bierner 2001, 2005). Hymenoxys bigelovii is the sole member of subg. Macdougalia (A. Heller) Bierner (Bierner 2001, 2004). Hymenoxys richardsonii, H. subintegra, H. cooperi, H. lemmonii, H. rusbyi, H. jamesii, and H. brachyactis belong to subg. Picradenia (Hooker) Cockerell (Bierner 2001). J. L. Anderson et al. (1996) and M. W. Bierner and R. K. Jansen (1998) provided evidence that Hymenoxys helenioides is a hybrid between H. hoopesii and H. richardsonii var. floribunda. Bierner (2001) recognized H. helenioides as a species because he was unable to determine whether all of the plants were F1 hybrids or at least some of them had given rise to breeding populations. Plants identified as Hymenoxys anthemoides, a discoid, South American annual that resembles H. odorata, were reported as "adventive on ballast," Mobile County, Alabama, by C. T. Mohr (1901).


Bierner, M. W. 1974. A systematic study of Dugaldia (Compositae). Brittonia 26: 385–392. Bierner, M. W. 1994. Submersion of Dugaldia and Plummera in Hymenoxys (Asteraceae: Heliantheae: Gaillardiinae). Sida 16: 1–8. Bierner, M. W. 2001. Taxonomy of Hymenoxys subgenus Picradenia and a conspectus of the subgenera of Hymenoxys (Asteraceae, Helenieae, Tetraneurinae). Lundellia 4: 37–63. Bierner, M. W. 2004. Taxonomy of Hymenoxys subgenus Macdougalia (Asteraceae, Helenieae, Tetraneurinae). Sida 21: 657–663. Bierner, M. W. 2005. Taxonomy of Hymenoxys subgenus Rydbergia (Asteraceae, Helenieae, Tetraneurinae). Lundellia 8: 28–37. Bierner, M. W. and R. K. Jansen. 1998. Systematic implications of DNA restriction site variation in Hymenoxys and Tetraneuris (Asteraceae, Helenieae, Gaillardiinae). Lundellia 1: 17–26. Spring, O., B. Zitterell-Haid, M. W. Bierner, and T. J. Mabry. 1994. Chemistry of glandular trichomes in Hymenoxys and related genera. Biochem. Syst. & Ecol. 22: 171–195. Wagner, W. L., R. Fletcher, and R. K. Shannon. 1999. Another look at Hymenoxys subgenus Plummera (Asteraceae: Heliantheae: Gaillardiinae) from Arizona and New Mexico. Brittonia 51: 79–86.

1 Annuals   (2)
+ Biennials or perennials   (3)
2 (1) Plants 5–10(–15) cm (delicate); ray corollas 2–3 × 0.7–1 mm (not surpassing phyllaries; Texas)   1 Hymenoxys texana
+ Plants 10–80 cm (robust); ray corollas 8.5–11 × 3–5.5 mm (well surpassing phyllaries; widespread)   2 Hymenoxys odorata
3 (1) Disc florets 6–15 (functionally staminate; receptacles flat)   3 Hymenoxys ambigens
+ Disc florets 25–400+ (bisexual; receptacles hemispheric to globoid, ovoid or conic)   (4)
4 (3) Phyllaries 22–50 in 2–3 series, subequal   (5)
+ Phyllaries (11–)16–30(–40) in 2 series, unequal   (7)
5 (4) Plants 30–100 cm; leaf blades simple; phyllaries 36–50 in 2 series; ray corollas usually yellow-orange to orange   4 Hymenoxys hoopesii
+ Plants 8–30 cm; leaf blades simple and/or lobed; phyllaries 22–48+ in 2–3 series; ray corollas yellow   (6)
6 (5) Basal leaves simple or (some, not all) lobed (lobes 3–7); involucres 13–16 × 19–23 mm; Arizona, Colorado, New Mexico   5 Hymenoxys brandegeei
+ Basal leaves lobed (lobes 3–15); involucres 15–25 × 18–30 mm; Colorado, Idaho, Montana, Utah, Wyoming   6 Hymenoxys grandiflora
7 (4) Outer phyllaries basally connate to 1/5 their lengths; inner phyllaries narrowly lanceolate to oblanceolate, 8.5–12.6 mm, apices aristate   7 Hymenoxys bigelovii
+ Outer phyllaries basally connate 1/4–2/3 their lengths; inner phyllaries obovate to oblanceolate, 2.8–8 mm, apices usually mucronate, sometimes acuminate   (8)
8 (7) Stems 1–20(–30+; plants usually with highly branched, woody caudices; basal leaf bases densely long-villous-woolly)   8 Hymenoxys richardsonii
+ Stems 1–10(–20; plants often with sparingly or moderately branched, woody caudices; basal leaf bases sparsely, if at all, long-villous-woolly)   (9)
9 (8) Involucres 4–8 mm diam   (10)
+ Involucres (8–)10–18 mm diam   (12)
10 (9) Perennials (polycarpic); mid blades simple or lobed (lobes 3, terminal lobes 2–4 mm wide)   15 Hymenoxys rusbyi
+ Biennials or perennials (monocarpic); mid blades lobed (lobes 3–7, terminal lobes 0.8–2.2 mm wide)   (11)
11 (10) Stems usually purple-red-tinted proximally; disc corollas 2.4–3 mm; pappi 0.8–2.1 mm; Mogollon Plateau area, c Arizona   16 Hymenoxys jamesii
+ Stems green throughout; disc corollas 3.1–4.2 mm; pappi 2.5–3 mm; c New Mexico   17 Hymenoxys brachyactis
12 (9) Stems (and leaves) usually densely sericeous (mainly Kaibab Plateau in Arizona, Utah)   9 Hymenoxys subintegra
+ Stems (and leaves) sometimes densely hairy (only H. cooperi and H. richardsonii var. floribunda on Kaibab Plateau in Arizona, Utah).   (13)
13 (12) Outer phyllaries 5(–8); s Arizona (mostly Huachuca Mountains)   10 Hymenoxys quinquesquamata
+ Outer phyllaries 7–15; not s Arizona   (14)
14 (13) Leaf blades lobed (lobes 3–23); outer phyllaries basally connate 1/2–2/3 their lengths   11 Hymenoxys vaseyi
+ Leaf blades simple or lobed (lobes 3–13); outer phyllaries basally connate 1/4–1/2 their lengths   (15)
15 (14) Mid blades usually lobed (lobes 3, terminal lobes 2–5.5 mm wide), sometimes not lobed; outer phyllaries (weakly keeled) basally connate 1/4–1/3 their lengths; raycorollas yellow to yellow-orange   12 Hymenoxys helenioides
+ Mid blades lobed (lobes 3–7, terminal lobes 0.7–2.5 mm wide); outer phyllaries (weakly to strongly keeled) basally connate 1/4–1/2 their lengths; ray corollas yellow   (16)
16 (15) Biennials or perennials, (10–)20–80(–100) cm (monocarpic); leaves ± hairy   13 Hymenoxys cooperi
+ Perennials, 30–50 cm (polycarpic); leaves glabrous or sparsely hairy   14 Hymenoxys lemmonii

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