The green links below add additional plants to the comparison table. Blue links lead to other Web sites.
enable glossary links

evening-primrose family

evening primrose, sun cup, suncups

Habit Herbs, annual or perennial, shrubs, or subshrubs, [lianas or trees], terrestrial, amphibious, or aquatic, unarmed, not clonal; often with epidermal oil cells, usually with internal phloem, abundant raphides in vegetative cells. Herbs, annual, caulescent; with a taproot.
Stems

erect to decumbent or prostrate.

erect, decumbent, or ascending, usually branched from base and distally, epidermis white or reddish brown, often exfoliating.

Leaves

usually deciduous, usually alternate or opposite, sometimes whorled, simple, usually cauline, sometimes basal and forming rosettes;

stipules present, intrapetiolar, usually caducous, relatively small, or absent (tribes Epilobieae and Onagreae);

sessile or subsessile to petiolate;

blade margins usually entire, toothed, or pinnately lobed, rarely bipinnately lobed.

cauline, proximalmost often clustered near base, alternate;

stipules absent;

subsessile;

blade margins entire, serrulate, or serrate.

Inflorescences

axillary, flowers solitary, leafy spikes, racemes, or panicles.

usually leafy spikes, sometimes racemes, nodding at anthesis, erect in fruit.

Flowers

usually bisexual, (protandrous in Chamaenerion, Clarkia, Epilobium, [and most species of Lopezia]; protogynous in Circaea and Fuchsia), sometimes unisexual (gynodioecious or dioecious, [subdioecious]), usually actinomorphic, sometimes zygomorphic, (2–)4(–7)-merous;

perianth and androecium epigynous;

sepals persistent after anthesis (in Ludwigia), or all flower parts deciduous after anthesis;

floral tube present or absent in Chamaenerion, Ludwigia, [and most species of Lopezia];

sepals usually green or red, rarely pink or purple, valvate;

petals present, rarely absent, often fading darker with age, imbricate or convolute, sometimes clawed;

nectary present;

stamens 2 times as many as sepals and in 2 series, antisepalous set usually longer, rarely all equal (Chamaenerion), or as many as sepals, [in Lopezia reduced to 2 or 1 plus 1 sterile staminode];

filaments distinct;

anthers usually versatile, sometimes basifixed, dithecal, polysporangiate, with tapetal septa, sometimes also with parenchymatous septa, opening by longitudinal slits, pollen grains united by viscin threads, (2 or)3(–5)-aperturate, shed singly or in tetrads or polyads;

ovary inferior, usually with as many carpels and locules as sepals, rarely 1 or 2 (Circaea and Gayophytum), septa sometimes thin or absent at maturity;

placentation axile or parietal;

style 1, stigma 1, with as many lobes as sepals or clavate to globose, papillate or not, and wet with free-running secretions to dry without the secretions;

ovules 1 to numerous per locule, in 1 or several rows or clustered, anatropous, bitegmic.

bisexual, actinomorphic, buds erect;

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

sepals 4, reflexed separately or in pairs;

petals 4, yellow, fading red, often with red dots basally;

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

ovary 4-locular, without apical projection, stigma subentire, subcapitate to subglobose, surface unknown, probably wet and non-papillate.

Fruit

a loculicidal capsule or indehiscent berry or nutlike.

a capsule, straight to flexuous, cylindrical, subterete, regularly, sometimes tardily, loculicidally dehiscent; usually sessile, sometimes pedicellate.

Seeds

smooth or sculptured, sometimes with a coma or wings, with straight, oily embryo, 4-nucleate embryo sac, endosperm absent.

numerous, in 1 row per locule, narrowly obovoid to narrowly oblanceoloid, triangular in cross-section, appearing smooth, glossy.

xI> = 7, 8, 10, 11, 15, 18.

xI> = 7.

Onagraceae

Camissonia

Distribution
North America; Mexico; Central America; South America; West Indies; Bermuda; Eurasia; Africa; Atlantic Islands; Indian Ocean Islands; Pacific Islands; Australasia; nearly worldwide; primarily New World
[BONAP county map]
from USDA
w North America; nw Mexico; w South America
[BONAP county map]
Discussion

Genera 22, species 664 (17 genera, 277 species in the flora).

Members of the Onagraceae are especially richly represented in North America. The family comprises annual and perennial herbs, with some shrubs and a few small to medium-sized trees. Most species occur in open habitats, ranging from dry to wet, with a few species of Ludwigia aquatic, from the tropics to the deserts of western North America, temperate forests, and arctic tundra; some species of Epilobium, Ludwigia, and Oenothera can be weeds in disturbed habitats. Members of the family are characterized by 4-merous flowers (sometimes 2-, 5-, or 7-merous), an inferior ovary, a floral tube in most species, stamens usually two times as many as sepals, and pollen connected by viscin threads. Flowers are usually bisexual, sometimes unisexual, and plants are gynodioecious, matinal, diurnal, or vespertine, self-compatible or self-incompatible, often outcrossing and then pollinated by a wide variety of insects or birds, or autogamous (P. H. Raven 1979; W. L. Wagner et al. 2007).

Onagraceae are known in considerable systematic detail, and information is available on comparative breeding systems and pollination biology, on chromosome numbers and cytogenetic relations, often involving translocations, and on vegetative, floral, and seed anatomy, palynology, and embryology. The phylogeny of the family is known in reasonably good detail, with most parts of the trees generally well-supported.

The suprageneric and generic classification presented by W. L. Wagner et al. (2007) differs in a number of ways from the previous classification (P. H. Raven 1979, 1988). Onagraceae are divided into two subfamilies based on a fundamental basal split recognized in all phylogenetic studies (R. H. Eyde 1981; P. C. Hoch et al. 1993; R. A. Levin et al. 2003, 2004; V. S. Ford and L. D. Gottlieb 2007), with Ludwigia on one branch (as Ludwigioideae), and the rest of the family on a second branch (as Onagroideae). Onagroideae are subdivided into six tribes: Circaeeae (including Fuchsieae), Epilobieae, Gongylocarpeae, Hauyeae, Lopezieae, and Onagreae. The Epilobieae and Onagreae are diverse; together they constitute fully two-thirds of the species in the family and include 15 of the 22 genera. The classification following Wagner et al. can be viewed on the Onagraceae web site by Wagner and Hoch at http://botany.si.edu/Onagraceae.

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

Species 12 (11 in the flora).

Species of Camissonia occur in desert scrub, grasslands, or pinyon-juniper woodlands, on brushy or open slopes and flats, washes, and, sometimes, on serpentine barrens, at elevations 0 to 2300 meters. Camissonia campestris and C. kernensis are self-incompatible diploids; C. pusilla and C. sierrae are self-compatible diploids; C. contorta is an autogamous hexaploid; the other species are autogamous tetraploids. Identification of the polyploid species of Camissonia is aided by their pollen having a high proportion of grains with more pores, usually 4 or 5, than typical 3-pored pollen in Onagraceae. This can be observed under low magnification (for example, 10\×) since 3-pored pollen is triangular, while 4-pored is quadrangular, and 5-pored is pentangular.

P. H. Raven (1969) delineated a group of four closely related species (Camissonia kernensis, C. parvula, C. pubens, and C. pusilla), marked by having sepals reflexed separately (rather than in pairs), which occur mainly in the Great Basin. Of the remaining species of Camissonia in the flora area, several (C. benitensis, C. campestris, C. integrifolia, C. lacustris, and C. sierrae) have more or less restricted ranges within California, or (C. strigulosa) extend also to Baja California, Mexico, or (C. contorta) to Washington, Idaho, and disjunctly to Vancouver Island, British Columbia. Camissonia dentata (Cavanilles) Reiche is the twelfth species in the genus, disjunct in western South America from southern Peru southward into Chile and Argentina.

W. L. Wagner et al. (2007) departed significantly from the most recent monograph by P. H. Raven (1969) in the delimitation of Camissonia based on the molecular analysis (R. A. Levin et al. 2004); they recognized eight genera in addition to the much reduced Camissonia: Camissoniopsis, Chylismia, Chylismiella, Eremothera, Eulobus, Neoholmgrenia, Taraxia, and Tetrapteron. Raven noted that Camissonia was the most heterogeneous genus in Onagreae, consisting of sharply distinct sections. He further noted that the capitate or subglobose stigma found in Camissonia, by which he distinguished the genus from the broadly circumscribed Oenothera of P. A. Munz (1965), was also found in Gayophytum, Gongylocarpus Schlechtendal & Chamisso, and Xylonagra Donnell Smith & Rose, thus the primary defining character state for the genus at that time is a plesiomorphy (P. C. Hoch et al. 1993). The redefined Camissonia is morphologically delimited by having subterete capsules that are more or less swollen by seeds, linear to narrowly elliptic leaves, and glossy seeds that are triangular in cross-section and mostly smaller than 1 mm in length; and flowering only at the distal, not basal, nodes; and is without ultraviolet reflectance pattern on petals (Raven). Reproductive features include: self-incompatible (C. campestris, C. kernensis) or self-compatible; flowers diurnal; outcrossing and pollinated by bees (E. G. Linsley et al. 1963, 1963b, 1964, 1973), or autogamous, rarely cleistogamous (Raven).

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

Parent taxa Onagraceae > subfam. Onagroideae > tribe Onagreae
Subordinate taxa
Camissonia, Camissoniopsis, Chamaenerion, Chylismia, Chylismiella, Circaea, Clarkia, Epilobium, Eremothera, Eulobus, Fuchsia, Gayophytum, Ludwigia, Neoholmgrenia, Oenothera, Taraxia, Tetrapteron
C. benitensis, C. campestris, C. contorta, C. integrifolia, C. kernensis, C. lacustris, C. parvula, C. pubens, C. pusilla, C. sierrae, C. strigulosa
Key
1. Sepals persistent or tardily caducous after anthesis; flowers (3 or)4 or 5(–7)-merous; floral tube absent; petals yellow or white [a. Onagraceae subfam. Ludwigioideae, p. 70].
Ludwigia
1. Sepals deciduous after anthesis (along with other flower parts); flowers (2–)4-merous; floral tube usually present, often elongate, if absent then petals usually rose purple or pink, rarely white [b. Onagraceae subfam. Onagroideae, p. 101].
→ 2
2. Stipules present and soon deciduous; fruit indehiscent (berry or burlike capsule with hooked hairs) [b1. Onagraceae subfam. Onagroideae tribe Circaeeae, p. 101].
→ 3
3. Fruit a berry; seeds few to ca. 500; flowers 4-merous.
Fuchsia
3. Fruit a capsule, burlike, with stiff, hooked hairs; seeds 1 or 2; flowers 2-merous.
Circaea
2. Stipules absent; fruit usually a capsule, sometimes indehiscent.
→ 4
4. Seeds usually comose, coma rarely secondarily lost; sepals erect or spreading; stigmas with dry multicellular papillae, entire or 4-lobed, lobes commissural; x = 18 [b2. Onagraceae subfam. Onagroideae tribe Epilobieae, p. 107].
→ 5
5. Floral tube absent; stamens subequal; style deflexed at anthesis, later erect, stamens initially erect, later deflexed; leaves usually alternate, rarely subopposite or subverticillate proximally.
Chamaenerion
5. Floral tube present; stamens in 2 unequal whorls; style and stamens erect; leaves opposite, at least near base of stem.
Epilobium
4. Seeds not comose; sepals reflexed; stigmas usually wet, non-papillate, and entire or (3 or)4-lobed (non-commissural), sometimes (Clarkia) lobes commissural and then with dry unicellular papillae; x = 7 [b3. Onagraceae subfam. Onagroideae tribe Onagreae, p. 159].
→ 6
6. Stigmas with commissural lobes and dry, unicellular papillae; flowers usually protandrous.
Clarkia
6. Stigmas hemispherical to subglobose or subcapitate, peltate, or 4-lobed, not commissural, surface wet, non-papillate; flowers not protandrous.
→ 7
7. Ovaries 2-locular; stems usually delicate.
Gayophytum
7. Ovaries (3 or)4-locular; stems usually not especially delicate.
→ 8
8. Seeds with concave and convex sides, concave side with a thick wing, convex side covered with glasslike, clavate hairs; petals white with yellow basal area.
Chylismiella
8. Seeds not concave/convex and not with a wing and clavate hairs; petals yellow, purple, red, or white, if white, mostly without yellow base.
→ 9
9. Ovaries with a slender, sterile apical projection; plants usually acaulescent.
→ 10
10. Herbs perennial; sterile projection of ovary persistent with fertile part in fruit, projection without visible abscission lines at its junctures with floral tube or fertile part of ovary.
Taraxia
10. Herbs annual; sterile projection of ovary with visible abscission lines at its junctures with both short floral tube and fertile part of ovary.
Tetrapteron
9. Ovaries without an apical projection; plants usually caulescent, sometimes acaulescent (in Oenothera).
→ 11
11. Styles with peltate indusium at base of stigma, at least at younger stages prior to anthesis; stigmas (3 or)4-lobed, receptive all around (or peltate to discoid or nearly square in sect. Calylophus).
Oenothera
11. Styles without indusium; stigmas usually subglobose to globose, subcapitate, capitate, or cylindrical (Eulobus), rarely conical-peltate and ± 4-lobed.
→ 12
12. Seeds in 2 rows per locule; capsules pedicellate; leaves mostly basal, blades often pinnately lobed, rarely bipinnately, sometimes unlobed, or lateral lobes greatly reduced or absent, terminal lobe usually large, abaxial surface of leaves or leaf margins with conspicuous, usually brown, oil cells.
Chylismia
12. Seeds in 1 row per locule; capsules usually sessile, rarely very shortly pedicellate; leaves not predominately basal, blades not lobed or pinnatifid, leaves without oil cells.
→ 13
13. Petals usually white, rarely red or tinged red; flowers vespertine.
Eremothera
13. Petals yellow, often with red flecks or spots; flowers diurnal.
→ 14
14. Flowering stems virgate; leaf blades pinnatifid to lobed; petals yellow with red flecks near base; seeds usually with purple spots; floral tube with a lobed disc.
Eulobus
14. Flowering stems not virgate; leaf blades not pinnatifid, margins entire or toothed; petals yellow, sometimes with 1+ red spots at base; seeds without purple spots; floral tube without a lobed disc.
→ 15
15. Stems densely leafy distally, nearly leafless proximally, with many slender, ascending branches from base; capsules strongly flattened, straight.
Neoholmgrenia
15. Stems usually leafy throughout, branched throughout or with a few basal branches; capsules not flattened, subterete or 4-angled, often flexuous or curled, sometimes straight.
→ 16
16. Capsules subterete; flowers only from distal nodes; seeds appearing smooth, glossy, triangular in cross section.
Camissonia
16. Capsules 4-angled, at least when dry; flowers from basalmost to distal nodes; seeds dull, flattened.
Camissoniopsis
1. Sepals reflexed separately.
→ 2
2. Petals 8–15(–18) mm; sepals 5–9(–11) mm; stigma exserted beyond anthers at anthesis.
C. kernensis
2. Petals 1.5–4 mm; sepals 1.2–3.8 mm; stigma surrounded by anthers at anthesis.
→ 3
3. Leaves cauline, none clustered near base, blade margins entire or subentire; plants usually glabrous or densely strigillose, rarely villous proximally, sometimes sparsely glandular puberulent distally.
C. parvula
3. Leaves cauline with some clustered near base, blade margins serrulate or undulate-serrulate; plants moderately to densely villous, usually also glandular puberulent, especially distally.
→ 4
4. Leaf blades 0.04–0.2 cm wide; floral tubes 0.8–1.6 mm; sepals 1.2–2 mm; capsules 18–32 mm.
C. pusilla
4. Leaf blades 0.2–0.6 cm wide; floral tubes 1.3–3 mm; sepals 2.2–3.8 mm; capsules (18–)26–50 mm.
C. pubens
1. Sepals reflexed in pairs.
→ 5
5. Stigma exserted beyond anthers at anthesis; sepals 3.5–8(–12) mm; petals (3.5–)5–15.5 mm.
C. campestris
5. Stigma surrounded by anthers at anthesis (except sometimes slightly exserted beyond anthers in C. sierrae); sepals (1.2–)1.6–4(–5.5) mm; petals 2–7 mm.
→ 6
6. Leaf blade margins usually entire, rarely with 1 or 2 small teeth; plants densely strigillose distally.
C. integrifolia
6. Leaf blade margins usually sparsely serrulate (C. sierrae sometimes with 1–several small teeth); plants glandular puberulent, usually also villous (except also strigillose in C. strigulosa) distally.
→ 7
7. Leaf blades usually lanceolate to narrowly ovate, sometimes elliptic, base rounded.
C. sierrae
7. Leaf blades linear to narrowly elliptic, base cuneate or attenuate.
→ 8
8. Plants usually densely strigillose, often also glandular puberulent, especially distally, sometimes glandular puberulent only and glabrate, sometimes also villous near base; sepals 1.6–4 mm; less than 10% of pollen grains 4-pored.
C. strigulosa
8. Plants villous, often also glandular puberulent distally, rarely strigillose and glandular puberulent, if so, more than 30% of pollen grains 4 or 5-pored; sepals 1.6–5.5 mm; less than 10% or more than 30% of pollen grains 4 or 5-pored.
→ 9
9. Sepals (3–)3.8–5.5 mm; petals (4–)4.5–7 mm.
C. lacustris
9. Sepals 1.6–4 mm; petals 2.5–4(–5) mm.
→ 10
10. Anthers with less than 10% of pollen grains 4-pored; leaves green or slightly bluish green; floral tubes 1.2 mm; plants villous and also glandular puberulent distally; San Benito County and adjacent Fresno and Monterey counties, California.
C. benitensis
10. Anthers usually with more than 30% of pollen grains 4 or 5-pored; leaves usually bluish green; floral tubes 1.6–2.7 mm; plants usually villous throughout, often also glandular puberulent distally, rarely entirely strigillose and glandular puberulent; British Columbia, California, Idaho, Nevada, Oregon, Washington.
C. contorta
Synonyms C. section sphaerostigma, Oenothera section sphaerostigma, Oenothera unraked sphaerostigma, Oenothera subg. sphaerostigma, Sphaerostigma
Name authority Jussieu Link: Jahrb. Gewächsk. 1(1): 186. (1818)
Source FNA vol. 10. Treatment authors: Warren L. Wagner, Peter C. Hoch. FNA vol. 10. Treatment author: Warren L. Wagner.
Web links