Asteraceae tribe Heliantheae subtribe Madiinae |
Holozonia |
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Holozonia |
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Habit | Annuals, perennials, subshrubs, or shrubs, (0.5–)1–250 cm. | Perennials, 30–150 cm (rhizomatous). | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Stems | (aerial) ± erect. |
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Leaves | mostly basal, or basal and cauline, or mostly cauline; opposite and/or alternate; usually sessile, rarely petiolate; blades usually spatulate to oblanceolate or linear to linear-elliptic, sometimes pinnately lobed, ultimate margins entire or toothed, faces usually hairy and glandular (glands sometimes elaborate, apices sometimes gland- or spine-tipped). |
mostly cauline; proximal opposite (basally connate), distal alternate; sessile; blades lanceolate to linear, margins entire, faces hirsute and (distal leaves) glandular-hirtellous (glands cup-shaped). |
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Involucres | campanulate, cylindric, ellipsoid, globose, fusiform, hemispheric, obconic, or urceolate. |
± obconic or turbinate, 2–4+ mm diam. |
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Receptacles | flat or convex to conic, ± paleate (paleae either in 1 series interior to ray florets, then usually connate and persistent, or 1 palea subtending each disc floret, then usually distinct and soon falling, or, in discoid heads, paleae functioning as phyllaries in 1 series and together constituting an “involucre”). |
flat to convex, glabrous or setulose, paleate (paleae falling, in 1 series between rays and discs, connate). |
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Ray florets | 0, or 1–75+, pistillate, fertile; corollas usually yellow or whitish, sometimes proximally yellow and distally whitish, often marked with red or purple or turning red on drying, rarely orange or red-orange (apices usually ± deeply divided, lobes usually 3, sometimes 2). |
4–10, pistillate, fertile; corollas whitish (abaxially purplish-veined). |
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Disc florets | 1–200+, bisexual and fertile, or functionally staminate; corollas usually yellow, sometimes brown, orange, reddish, or whitish, tubes much shorter than to about equaling cylindric to funnelform throats, lobes 5, deltate; anther thecae usually dark (reddish to purple, often described as “black”), sometimes pale (yellow to brown); stigmatic papillae in 2 lines. |
9–28, functionally staminate; corollas white (pubescent), tubes shorter than funnelform throats, lobes 5, deltate (anthers ± dark purple; styles glabrous proximal to branches). |
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Phyllaries | persistent or falling, 1–75+ in 1(–2) series or 0 (the “involucre” then interpreted as constituted of 1 series of receptacular paleae; phyllaries distinct or weakly connate, narrowly ovate to lanceolate or linear, equal or subequal, herbaceous, usually conduplicate, often each partly or wholly investing a subtended floret, apices often attenuate, faces usually glandular). |
4–10 in 1 series (each mostly or wholly enveloping a ray ovary, ± lance-linear, herbaceous, abaxially hirsute, sometimes glandular-hirtellous, glands cup-shaped). |
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Calyculi | usually 0, sometimes 2–12+ bractlets (Centromadia, some members of Hemizonia and Lagophylla). |
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Heads | usually radiate, rarely discoid, usually in corymbiform, glomerulate, paniculiform, spiciform, or ± umbelliform arrays, sometimes borne singly. |
radiate, borne singly or in loose, corymbiform arrays (peduncles filiform). |
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Cypselae | obcompressed, compressed, or terete, often obpyramidal, clavate, or fusiform, glabrous or hairy (ray and disc often different); pappi 0, or persistent or falling, of awns, bristles, or scales (sometimes in combination) in 1–2 series (the elements often ciliate or plumose). |
(black) obcompressed, ± clavate (basal attachments centered, apices beakless, areolae broadly cupulate, faces glabrous); pappi (rays) 0 or coroniform (0.1–0.3 mm), or (discs) 0 or (readily falling) of 1–5 subulate scales. |
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Peduncular | bracts: pit-glands, tack-glands, and/or spines 0. |
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x | = 14. |
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Asteraceae tribe Heliantheae subtribe Madiinae |
Holozonia |
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Distribution |
HI; South America; Mostly w North America |
CA |
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Discussion | Genera 24, species 121 (20 genera, 83 species in the flora). Madiinae has long included the “tarweeds,” which are centered in or restricted to the California Floristic Province, the silversword alliance of the Hawaiian Islands (Argyroxiphium, Dubautia, and Wilkesia), and Raillardella (in a broad sense), which was formerly included in Senecioneae (Carlquist et al. 2003). Such a circumscription was used by H. Robinson (1981) and is followed here. In papers by B. G. Baldwin and B. Wessa (2000) and Baldwin et al. (2002), Madiinae were linked to genera traditionally included in Heliantheae (e.g., Eriophyllum, Hulsea, Lasthenia, Venegasia) and in Senecioneae (e.g., Arnica) and Madiinae plus Arnicinae, Baeriinae, Hulseinae, and Venegasiinae were included in tribe Madieae. Key to genera of Madiinae here is based on key by B. G. Baldwin (in S. Carlquist et al. 2003). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Species 1. Holozonia has been treated as congeneric with Hemizonia and with Lagophylla. Like Lagophylla, Holozonia has functionally staminate disc florets, cup-shaped glands, and obcompressed cypselae, each completely or mostly invested by a phyllary. Additional morphologic considerations and biosystematic studies led W. C. Thompson (1983) to reject the hypothesis of a close relationship between Holozonia and Lagophylla. Molecular phylogenetic data are in keeping with a closer relationship of Holozonia to other continental tarweeds with white corollas and x = 14 (i.e., Blepharizonia and Hemizonia), than to Lagophylla (S. Carlquist et al. 2003). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
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Key |
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Source | FNA vol. 21, p. 254. | FNA vol. 21, p. 294. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Name authority | Bentham & Hooker f.: Gen. Pl. 2: 198. (1873) | Greene: Bull Torrey Bot. Club 9: 122. (1882) | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Web links |