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evening primrose

California sun cup, southern suncup

Habit Herbs,usually annual, rarely short-lived perennial, caulescent. Herbs annual, rarely short-lived perennial, usually villous, sometimes strigillose.
Stems

prostrate to ascending or erect, often with reddish brown or white exfoliating epidermis.

1–several from base, ascending or decumbent, to 80 cm.

Leaves

cauline and often in a basal rosette, alternate;

stipules absent;

sessile or petiolate;

blade margins dentate, denticulate, or serrulate.

1.2–12 × 0.2–1.5 cm;

petiole 0–4 cm, distal ones 0–0.3 cm;

blade (basal) narrowly elliptic or (cauline) usually narrowly lanceolate or lanceolate, rarely linear, base (basal) narrowly cuneate, (cauline) cuneate or subcordate, margins usually sparsely and inconspicuously denticulate, apex acute.

Inflorescences

spikes, erect or nodding at anthesis.

Flowers

bisexual, actinomorphic, buds erect;

floral tube deciduous (with sepals, petals, and stamens) after anthesis, with basal nectary;

sepals 4, usually reflexed in pairs, sometimes separately;

petals 4, yellow, fading red, with 1+ red dots basally;

stamens 8, in 2 unequal series, anthers versatile, pollen shed singly;

ovary 4-locular, without apical projection, style glabrous or pubescent distally, stigma entire, subcapitate to subglobose, surface unknown, probably wet and non-papillate.

opening near sunrise;

floral tube 2–5(–7.5) mm;

sepals (2.3–)5–8(–11) mm;

petals yellow, each usually with 1 bright red dot, rarely 2, near base, (4.2–)7–15 mm; episepalous filaments (1–)1.5–3.5 mm, epipetalous filaments (0.5–)1–2.5 mm, anthers (0.5–)1.3–2(–2.5) mm, less than 5% of pollen grains 4- or 5-pored;

style (5.5–)7–12 mm, stigma exserted beyond anthers at anthesis.

Fruit

a capsule, contorted or curled 1 to 5 times, or straight, narrowly cylindrical and thickened proximally, 4-angled (at least when dry), regularly but tardily loculicidally dehiscent, not swollen by seeds;

sessile.

Capsules

straight or somewhat contorted, weakly 4-angled, 12–40 × 1.5–2.5 mm.

Seeds

numerous, in 1 row per locule, flattened, narrowly obovoid, dull.

0.9–1 mm.

xI> = 7.

2n

= 14.

Camissoniopsis

Camissoniopsis bistorta

Phenology Flowering Mar–Jun.
Habitat Sandy or clayey soils, coastal strands, grasslands, coastal sage scrub, chaparral, oak woodlands, margins of Sonoran and Mojave deserts, rarely higher elevation meadows.
Elevation 0–1600(–2600) m. (0–5200(–8500) ft.)
Distribution
w United States; nw Mexico
[BONAP county map]
from FNA
CA; Mexico (Baja California)
[WildflowerSearch map]
[BONAP county map]
Discussion

Species 14 (13 in the flora).

Camissoniopsis proavita (P. H. Raven) W. L. Wagner & Hoch is known from northern Baja California, Mexico. It is a diploid, closely related to C. micrantha but differing in having numerous flowers in the basal rosette, which is densely leafy.

All species of Camissoniopsis occur near coasts or on dry slopes or desert flats inland from 0–2500 m. R. A. Levin et al. (2004) found strong molecular support for Camissoniopsis in a clade with Neoholmgrenia and Tetrapteron. Camissoniopsis was segregated from Camissonia as delimited by P. H. Raven (1969). Camissoniopsis is distinguished by having 4-angled fruits, at least when dry, and not swollen by seeds, dull seeds usually smaller than 1 mm, and by flowering from both basal and distal nodes (Raven). Relationships within Camissoniopsis are complex and reticulate. Several diploids (especially C. hirtella) appear to have contributed to the formation of the tetraploids and, in turn, the hexaploids (Raven), and, as a result, are very similar morphologically to each other. Identification of the polyploid species of Camissoniopsis is aided by their pollen having a high proportion of grains with higher number of pores than typical Onagraceae 3-pored pollen, usually 4- or 5-pored. This can be observed under low magnification (for example, 10\×) since the 3-pored pollen is triangular while the 4-pored is quadrangular and 5-pored is pentangular. Raven proposed Camissonia sect. Holostigma as a new combination based on Spach’s generic name. He was unaware that Holostigma Spach, like Agassizia Spach, is a later homonym and thus illegitimate; however, he satisfied all requirements for valid publication of a new sectional name in Camissonia. Reproductive features include: self-incompatible (C. cheiranthifolia and C. bistorta) or self-compatible; flowers diurnal; outcrossing and pollinated by bees (E. G. Linsley et al. 1963, 1964, 1973) or autogamous (Raven).

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

Camissoniopsis bistorta occurs in California from Ventura County south and east through the counties of southern Los Angeles, southwestern San Bernardino, Orange, western Riverside, and the western two-thirds of San Diego, reaching the margins of the desert in San Bernardino and San Diego counties, and southward in cismontane Baja California to Ojos Negros and San Vicente. The species occurs at exceptionally high elevations in the Santa Ana drainage of the San Bernardino Mountains. P. H. Raven (1969) indicated that there were occasional apparent hybrids between C. cheiranthifolia subsp. suffruticosa and C. bistorta occurring in intermediate habitats in areas where the two species co-occur. He determined that C. bistorta is self-incompatible.

Camissoniopsis bistorta was apparently introduced with stream gravel in 1959 in Goleta Marsh, Santa Barbara, California, and on ballast heaps at Nanaimo, Vancouver Island, British Columbia, in 1893. It has apparently not persisted at either site.

Oenothera heterophylla Nuttall ex Hooker & Arnott (1839), not Spach (1836), is an illegitimate name that pertains to Camissoniopsis bistorta.

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

Key
1. Herbs perennial; coastal habitats.
C. cheiranthifolia
1. Herbs usually annual, rarely short-lived perennial (in C. bistorta); primarily inland habitats.
→ 2
2. Stigma exserted beyond anthers at anthesis; sepals (2.3–)5–8(–11) mm; petals (4.2–)7–15 mm.
C. bistorta
2. Stigma surrounded by all anthers, or at least those of longer filaments, at anthesis; sepals 1–6(–8.5) mm; petals 1.5–10.5(–13) mm.
→ 3
3. Capsules 2.8–3.5 mm diam. near base, straight or slightly curved outward, deeply grooved along lines of dehiscence.
C. guadalupensis
3. Capsules 0.7–2.2 mm diam. near base, straight or curved into 1+-coiled spirals, not deeply grooved.
→ 4
4. Pollen with 25–100% of grains 4- or 5-pored.
→ 5
5. Inflorescences exclusively villous; 25–60% of pollen grains 4- or 5-pored.
C. luciae
5. Inflorescences villous and glandular puberulent; 70–100 % of pollen grains 4- or 5-pored.
→ 6
6. Capsules 1.3–1.6 mm diam., subterete in living material (obscurely 4-angled when dry); southernmost Monterey County to central San Luis Obispo County, California.
C. hardhamiae
6. Capsules 1.5–2 mm diam., 4-angled in living material; San Diego County, California, adjacent Baja California, and offshore islands.
C. robusta
4. Pollen with less than 5% of grains 4-pored (rarely more in C. intermedia).
→ 7
7. Capsules 1.8–2.2 mm diam., conspicuously 4-angled in living material.
C. lewisii
7. Capsules 0.7–1.2(–1.8) mm diam., terete, subterete, or obscurely 4-angled, at least in living material.
→ 8
8. Distal leaves petiolate, blade base attenuate; capsules usually much contorted, irregularly to 5-coiled; herbs moderately to sparsely strigillose, sometimes also sparsely villous.
C. ignota
8. Distal leaves usually subsessile, blade base rounded, cuneate, or truncate; capsules straight to 1–2-coiled; herbs strigillose to villous.
→ 9
9. Herbs conspicuously grayish in appearance, densely strigillose; lateral stems usually decumbent; plants of the deserts.
C. pallida
9. Herbs not conspicuously gray in appearance, mostly villous; lateral stems erect to decumbent; plants not of deserts or only at desert margins (except C. confusa in central Arizona).
→ 10
10. Capsules 0.7–0.9 mm diam.; distal leaf blades elliptic-ovate or ovate; stems ascending to erect.
C. hirtella
10. Capsules 0.9–1.2(–1.8) mm diam.; distal leaf blades narrowly lanceolate to narrowly ovate; stems decumbent or erect.
→ 11
11. Stems decumbent; inflorescences usually densely villous, rarely also glandular puberulent.
C. micrantha
11. Stems erect; inflorescences usually moderately to densely villous, also glandular puberulent.
→ 12
12. Floral tube (1.8–)2–3.8 mm; petals (2.5–)5–10.5 mm; styles (2.5–)4.5–7.5 mm; herbs densely villous, often also stigillose.
C. confusa
12. Floral tube 1.2–2 mm; petals 1.5–3.5(–4.5) mm; styles 2–3.5 mm; herbs moderately villous.
C. intermedia
Source FNA vol. 10. Author: Warren L. Wagner. FNA vol. 10.
Parent taxa Onagraceae > subfam. Onagroideae > tribe Onagreae Onagraceae > subfam. Onagroideae > tribe Onagreae > Camissoniopsis
Sibling taxa
C. cheiranthifolia, C. confusa, C. guadalupensis, C. hardhamiae, C. hirtella, C. ignota, C. intermedia, C. lewisii, C. luciae, C. micrantha, C. pallida, C. robusta
Subordinate taxa
C. bistorta, C. cheiranthifolia, C. confusa, C. guadalupensis, C. hardhamiae, C. hirtella, C. ignota, C. intermedia, C. lewisii, C. luciae, C. micrantha, C. pallida, C. robusta
Synonyms Agassizia, Camissonia section holostigma, Holostigma Oenothera bistorta, Camissonia bistorta, O. bistorta var. veitchiana, Sphaerostigma bistortum, S. bistortum var. veitchianum, S. veitchianum
Name authority W. L. Wagner & Hoch: Syst. Bot. Monogr. 83: 123. (2007) (Nuttall ex Torrey & A. Gray) W. L. Wagner & Hoch: Syst. Bot. Monogr. 83: 204. (2007)
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