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Chylismia claviformis

brown-eyed primrose, browneyes, clavate fruit primrose

Habit Herbs usually annual, sometimes perennial, rarely biennial.Leaves basal and cauline, usually with well-developed basal rosette; blade usually pinnately or bipinnately lobed, sometimes with scattered, irregular lobes, sometimes lateral lobes greatly reduced or absent, terminal lobe elliptic, narrowly to broadly ovate to oblong, lanceolate, oblanceolate, cordate, or subcordate. Herbs annual, glabrous, strigillose, glandular puberulent, or, sometimes, villous.
Stems

branched mostly from base, 3–70 cm.

Leaves

primarily in basal rosette, cauline reduced or absent, 1.5–20 × 0.3–3.5 cm;

petiole 0.7–12 cm;

blade usually pinnately lobed, sometimes lateral lobes poorly developed or absent, terminal lobe usually narrowly ovate to lanceolate, sometimes cordate or subcordate, 0.8–9 × 0.2–4.5 cm, margins dentate, sinuate-dentate, or serrate, brown oil cells conspicuously lining veins abaxially.

Racemes

nodding, elongating after anthesis.

Flowers

usually opening at sunrise, rarely at sunset;

floral tube 0.4–9 mm;

petals usually bright yellow, rarely white or cream, usually with red dots basally, or lavender to purple with white or yellow basally, sometimes with darker flecks near base, fading yellow, orange, reddish, or lavender;

pollen shed singly.

opening at sunset or sunrise;

buds with or without subapical or apical free tips;

floral tube 2–6.5 mm, villous inside proximally;

sepals 2–8 mm;

petals pale to bright yellow or white, sometimes red- or purple-dotted near base, fading purple, sometimes red or orange, or not changing color, 1.5–8 mm;

stamens subequal, filaments 1.5–5.5 mm, anthers 1.5–6 mm, ciliate;

style 5–16 mm, stigma exserted beyond anthers at anthesis.

Capsules

ascending to spreading, clavate, 8–40 mm;

pedicel 4–40 mm.

Seeds

0.6–1.5 mm.

Chylismia sect. Chylismia

Chylismia claviformis

Distribution
w United States; nw Mexico
w United States; nw Mexico
[WildflowerSearch map]
[BONAP county map]
Discussion

Species 14 (14 in the flora).

Section Chylismia consists of 10 diploid (2n = 14) species, and two that are partly polyploid (2n = 14, 28) [C. scapoidea subsp. scapoidea and C. walkeri subsp. walkeri (only one tetraploid population)]; no chromosome counts are available for the remaining two species, C. atwoodii and C. confertiflora (P. H. Raven 1962, 1969). Species of sect. Chylismia usually occur on sandy desert slopes, flats, and washes, often in sagebrush shrubland in the northern part of its range, or on rock slides or cliffs, mainly in the Mojave and northwestern Sonoran deserts, the Great Basin, and the lower elevations of the surrounding Sierra Nevada and Rocky Mountains. The limits of this range stretch from southeastern Oregon, central and southern Idaho, and central Wyoming, south through Nevada and Utah to eastern and southeastern California, northern Baja California and northwestern Sonora, Mexico, Arizona, northwestern New Mexico, and western Colorado. Several species are rare; C. confertiflora and C. specicola are known only from the Grand Canyon in northwestern Arizona, C. megalantha only from the vicinity of the type locality in Nye County, Nevada, and C. atwoodii also only from a narrow area around its type locality in Kane County, Utah. Others are widespread, especially the very diverse C. claviformis (11 subspp.), C. scapoidea (4 subspp.), C. walkeri (2 subspp.), and C. brevipes (3 subspp.). Chylismia scapoidea is the only species in the genus to occur east of the continental divide, both in Colorado on the upper Arkansas River in Fremont and Pueblo counties, and much more widely in Wyoming. Chylismia does not occur west of the Cascade-Sierra Nevada axis. Because R. A. Levin et al. (2004) included only C. claviformis in their analysis, they did not test the monophyly of sect. Chylismia; however, this section is both geographically distinct and morphologically set apart by the characteristic pinnate leaves (modified in some species, which have retained the entire apical lobe but do not have the smaller lateral lobes). Most species have bright yellow petals with red dots proximally and ultraviolet reflectance distally; some subspecies of C. claviformis have white petals; three species (C. atwoodii, C. heterochroma, and C. megalantha) have lavender or purple petals, often with lavender or purple flecks toward base, and white or yellow at the base and no reflectance, clearly a derived condition within the section (Raven 1962, 1969).

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

Subspecies 11 (10 in the flora).

P. H. Raven (1962) subdivided this species into 12 subspecies and, subsequently (1969), he combined two of them. The latter approach is used here. Only subsp. wigginsii P. H. Raven does not occur in the United States; its narrow range is restricted to northern Baja California. Raven (1962, 1969) determined this species to be self-incompatible.

Chylismia claviformis is the most complex and, along with C. scapoidea, the most widely distributed species of the genus. The central part of its geographical range is occupied by five closely related white-petaled subspecies (aurantiaca, claviformis, funerea, integrior, and peeblesii) that are very similar morphologically. South of this area four additional subspecies occur, all yellow-petaled (peirsonii, rubescens, wigginsii, and yumae). These four subspecies have sepals and petal color similar to those of C. brevipes, and P. H. Raven (1962, 1969) thought it likely that they were derived following hybridization between that species and one of the white-petaled populations of C. claviformis. North of the range of the white-petaled subspecies are found two additional yellow-petaled subspecies (cruciformis and lancifolia). Most populations of subsp. cruciformis consist of plants in which the flowers open in the early morning; in all other subspecies the flowers open in the late afternoon (Raven 1962, 1969). The following key will separate them, but there are many intergrades among the subspecies so that not all specimens will be easily identified.

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

Key
1. Stamens usually 4, rarely 8, then antipetalous anthers abortive, smaller.
C. exilis
1. Stamens 8, anthers all fertile.
→ 2
2. Petals lavender or purple, often with purple or lavender flecks near base.
→ 3
3. Stigma surrounded by anthers at anthesis, plants autogamous; petals 2–6 mm.
C. heterochroma
3. Stigma exserted beyond anthers at anthesis, plants outcrossing; petals 7–14 mm.
→ 4
4. Capsules 8–14 mm, erect or ascending; floral tubes 4–9 mm.
C. megalantha
4. Capsules 11–25 mm, spreading to reflexed; floral tubes 0.6–1 mm.
C. atwoodii
2. Petals yellow or white, often with red dots at base.
→ 5
5. Capsules distinctly clavate.
→ 6
6. Stigma surrounded by anthers at anthesis; petals 1.5–5.5(–8) mm.
C. scapoidea
6. Stigma exserted beyond anthers at anthesis; petals 2–10 mm.
→ 7
7. Mature capsules sharply reflexed; petals bright yellow.
C. munzii
7. Mature capsules ascending, spreading, erect, or slightly reflexed; petals yellow or white.
→ 8
8. Raceme branches intricate and filiform; leaves in poorly defined basal rosette; capsules 4–10 mm.
C. parryi
8. Raceme branches not filiform; leaves primarily in well-defined basal rosette; capsules 8–40 mm.
→ 9
9. Stamens subequal; leaves usually pinnately lobed, with well-developed lateral lobes, sometimes these reduced or absent, margins dentate, sinuate-dentate, or serrate.
C. claviformis
9. Stamens unequal, differentiated into 2 sets; leaves usually unlobed, rarely pinnately lobed with reduced lateral lobes, margins entire or sparsely denticulate.
C. eastwoodiae
5. Capsules oblong-cylindrical.
→ 10
10. Stigma surrounded by anthers at anthesis; petals 1–6 mm; styles 1.5–6(–7) mm; racemes erect in bud, mature buds individually reflexed.
→ 11
11. Plants perennial, base sometimes woody; leaf blades glabrous or sparsely villous; anthers glabrous.
C. specicola
11. Plants annual or short-lived perennial, base not woody; leaf blades moderately to densely villous; anthers glabrous or sparsely ciliate.
C. walkeri
10. Stigma exserted beyond anthers at anthesis; petals 3–18 mm; styles 7–18 mm; racemes erect or nodding in bud, mature buds individually reflexed or not.
→ 12
12. Racemes erect to nodding, elongating in bud, mature buds individually reflexed; plants virgate; floral tubes 1–3 mm.
C. multijuga
12. Racemes nodding, mostly elongating after flowers open; plants not virgate; floral tubes 3–8 mm.
→ 13
13. Buds without free tips or with subapical free tips 1–2 mm, or with minute apical free tips less than 1 mm, and then bud pubescent but not glandular puberulent; plants branched or unbranched distally; stamens subequal.
C. brevipes
13. Buds with subapical free tips 1–2 mm, glandular puberulent; plants well branched; stamens differentiated into 2 sets.
C. confertiflora
1. Herbs villous proximally; buds with subapical free tips; petals usually yellow, rarely white.
→ 2
2. Petals not changing color in fading, 4.5–7 mm; lateral lobes of leaf blades well developed.
subsp. peirsonii
2. Petals fading brick red, 3–5 mm; lateral lobes of leaf blades poorly developed, small or absent.
subsp. rubescens
1. Herbs strigillose, glandular puberulent, or glabrous proximally; buds with or without subapical or apical free tips; petals yellow or white.
→ 3
3. Petals yellow.
→ 4
4. Herbs strigillose, sometimes also glandular puberulent distally.
subsp. yumae
4. Herbs glabrous or glandular puberulent distally.
→ 5
5. Leaf blades with narrowly ovate to subcordate terminal lobes, at least some lateral lobes developed; flowers usually opening at dawn.
subsp. cruciformis
5. Leaf blades with lanceolate terminal lobes, lateral lobes usually greatly reduced or absent; flowers usually opening at dusk.
subsp. lancifolia
3. Petals usually white, rarely pale yellow (in subsp. claviformis).
→ 6
6. Herbs usually glabrous distally, rarely sparsely strigillose or glandular puberulent; lateral lobes of leaf blades usually well developed.
subsp. claviformis
6. Herbs usually strigillose and/or glandular puberulent, rarely glabrate; lateral lobes of leaf blades reduced, absent, or well developed.
→ 7
7. Lateral lobes of leaf blades well developed.
→ 8
8. Herbs strigillose, sometimes glabrate distally.
subsp. aurantiaca
8. Herbs glandular puberulent and strigillose.
subsp. peeblesii
7. Lateral lobes of leaf blades poorly developed, small, or absent.
→ 9
9. Leaf blades usually with at least some poorly developed lateral lobes; plants strigillose.
subsp. funerea
9. Leaf blades often with only terminal lobe developed; plants strigillose proximally, strigil-lose and glandular puberulent or glabrate distally.
subsp. integrior
Source FNA vol. 10. FNA vol. 10.
Parent taxa Onagraceae > subfam. Onagroideae > tribe Onagreae > Chylismia Onagraceae > subfam. Onagroideae > tribe Onagreae > Chylismia > sect. Chylismia
Sibling taxa
C. arenaria, C. atwoodii, C. brevipes, C. cardiophylla, C. confertiflora, C. eastwoodiae, C. exilis, C. heterochroma, C. megalantha, C. multijuga, C. munzii, C. parryi, C. scapoidea, C. specicola, C. walkeri
Subordinate taxa
C. atwoodii, C. brevipes, C. claviformis, C. confertiflora, C. eastwoodiae, C. exilis, C. heterochroma, C. megalantha, C. multijuga, C. munzii, C. parryi, C. scapoidea, C. specicola, C. walkeri
C. claviformis subsp. aurantiaca, C. claviformis subsp. claviformis, C. claviformis subsp. cruciformis, C. claviformis subsp. funerea, C. claviformis subsp. integrior, C. claviformis subsp. lancifolia, C. claviformis subsp. peeblesii, C. claviformis subsp. peirsonii, C. claviformis subsp. rubescens, C. claviformis subsp. yumae
Synonyms Camissonia section tetranthera, Oenothera section tetranthera Oenothera claviformis, Camissonia claviformis, C. scapoidea var. claviformis, O. scapoidea var. claviformis
Name authority unknown (Torrey & Frémont) A. Heller: Muhlenbergia 2: 105. (1906) — (as Chylisma clavaeformis)
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