clarkia, farewell to spring, godetia
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Herbs, annual, caulescent. |
Herbs (annual or perennial), [shrubs]. |
slender to stout, erect to prostrate or decumbent, unbranched to sparsely branched. |
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cauline, alternate; stipules absent; sessile or petiolate, petiole usually shorter than blade; blade margins entire or denticulate. |
alternate or basal; stipules absent. |
usually racemes or spikes, rarely panicles; buds erect or pendent. |
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bisexual, usually actinomorphic, often protandrous; floral tube deciduous (with sepals, petals, and stamens) after anthesis, usually with basal nectary and a ring of hairs inside; sepals 4, often pink to purplish red, usually connate to tip in bud, reflexed singly, in pairs, or all together to 1 side at anthesis; petals 4, usually lavender or pink to dark reddish purple, pale yellow or white, [rarely blue (C. tenella)], often spotted, flecked or streaked with red, purple, or white; stamens 8, in 2 equal or unequal series, or 4 in 1 series, filaments filiform or expanded distally, sometimes subtended by hairy scales, anthers basifixed, often with short acute sterile tip, pollen cream, yellow, blue-gray, lavender, or red, shed singly; ovary 4-locular, stigma 4-lobed, commissural, lobes receptive only on dry, unicellular-papillose inner surfaces. |
usually actinomorphic, rarely slightly zygomorphic (in Oenothera), (3 or)4-merous; stamens 2 times as many, or rarely as many, as sepals; pollen usually shed in monads, rarely tetrads (Chylismia sect. Lignothera). |
a capsule, elongate, straight or curved, cylindrical, fusiform, or subclavate, often 4-angled (shallowly to deeply 4- or 8-grooved) or terete, loculicidal, often tardily dehiscent, sometimes with sterile beak, rarely short, indehiscent and nutlike (C. heterandra); sessile or pedicellate. |
a dry capsule, usually dehiscent, sometimes indehiscent. |
usually numerous, rarely (C. heterandra) 1 or 2, usually angled, cubic, or elongate to spindle-shaped, often with crest of elongated cells, scaly or minutely tuberculate. |
few to numerous, without hairs or wings, [very rarely with asymmetrical dry wing (Xylonagra)], or with dry (Oenothera), erose or smooth wing, or with thick, papillate wings (Chylismiella). |
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w North America; w South America |
North America; Mexico; Central America; South America; West Indies |
Species 42 (41 in the flora). Molecular phylogenetic analysis places a strongly monophyletic Clarkia as sister to Gayophytum and Chylismiella (R. A. Levin et al. 2004). All but one species of Clarkia are endemic to western North America; the only exception is C. tenella (Cavanilles) H. Lewis & M. E. Lewis, which occurs in Mediterranean-climate regions of Argentina and Chile. Several species of Clarkia are grown ornamentally, especially C. amoena, the so-called Godetia of horticulture, and C. unguiculata, the common garden clarkia. Both species and cultivars developed from them are used in annual border plantings or in hanging baskets. Clarkia pulchella is also commonly cultivated, especially in Europe. Clarkia has been the subject of detailed systematic and evolutionary studies for more than half a century, and there are many reports of these studies in the literature (H. Lewis and M. E. Lewis 1955; V. S. Ford and L. D. Gottlieb 2003; V. M. Eckhart et al 2004). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Genera 13, species 265 (12 genera, 199 species in the flora). Onagreae account for more than half the total genera in Onagraceae and diversified from a center in southwestern North America (L. Katinas et al. 2004). Delimitation of the tribe by W. L. Wagner et al. (2007) differs from previous ones by the exclusion of Gongylocarpus, now in its own tribe, by the segregation of eight genera (Camissoniopsis, Chylismia, Chylismiella, Eremothera, Eulobus, Neoholmgrenia, Taraxia, and Tetrapteron) from Camissonia, and by the inclusion of three previously separate genera (Calylophus, Gaura, and Stenosiphon) in Oenothera. Within the branch of the family that lacks stipules (Gongylocarpeae, Epilobieae, and Onagreae), the last two tribes form a clade that has very strong molecular support (R. A. Levin et al. 2003, 2004), but no obvious morphological synapomorphy. The clade may be defined by a cytogenetic change from the base chromosome number of x = 11 found in Circaeeae, Gongylocarpeae, and Lopezieae, to x = 18 in Epilobieae, and x = 7 in Onagreae; however, these changes could also have occurred independently. Other than the new chromosome number x = 7, the only apparent morphological synapomorphy for Onagreae alone is pollen with prominent apertural protrusions (J. Praglowski et al. 1987, 1989), a character state also found in Circaeeae (Praglowski et al. 1994). The monophyly of Onagreae has moderate (Levin et al. 2004) to strong support (V. S. Ford and L. D. Gottlieb 2007). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
1. Capsules indehiscent, nutlike, 2–3 mm, 1 or 2 seeded [6h.7. sect. Phaeostoma subsect. Heterogaura]. | C. heterandra |
1. Capsules loculicidal, 10–70 mm, many-seeded. | → 2 |
2. Stamens 4; floral tube slender, 13–35 mm; petals conspicuously 3-lobed [6a. sect. Eucharidium]. | → 3 |
3. Petal length 2 times width, lobes ± equal or middle lobe wider. | C. concinna |
3. Petal length equal to width, middle lobe longer and much narrower. | C. breweri |
2. Stamens 8; floral tube obconic to campanulate or funnelform, 1–10(–15) mm; petals lobed or not. | → 4 |
4. Inner anthers sterile; petals with 3 subequal lobes [6b. sect. Clarkia]. | C. pulchella |
4. Inner anthers fertile, anthers similar or inner much smaller, paler; petals not 3-lobed. | → 5 |
5. Inflorescence axis recurved at tip in bud; buds pendent. | → 6 |
6. Petal claw broad with pair of lateral basal lobes; stamens subtended by ciliate scales [6d. sect. Myxocarpa]. | → 7 |
7. Stigma not or rarely exserted beyond anthers; petals 6–12(–14) mm. | → 8 |
8. Petals not spotted; pollen yellow. | C. stellata |
8. Petals spotted or mottled or with darker flecks; pollen blue-gray. | → 9 |
9. Inflorescence axis recurved only at tip in bud, straight 4+ nodes distal to open flowers. | C. virgata |
9. Inflorescence axis in bud recurved 1–3 nodes distal to open flowers. | C. rhomboidea |
7. Stigma exserted beyond anthers (rarely so in C. virgata); petals (7–)12–20 mm. | → 10 |
10. Inflorescence axis recurved in bud, straight 1–3 nodes distal to open flowers; petal length 1.4–1.6 times width. | C. mildrediae |
10. Inflorescence axis recurved only at tip in bud, straight 4+ nodes distal to open flowers; petal length 1.5–3 times width. | → 11 |
11. Flower buds fusiform, tip acute. | C. borealis |
11. Flower buds narrowly obovoid, tip obtuse. | → 12 |
12. Petal length 1.5–2 times width. | C. mosquinii |
12. Petal length 1.9–3 times width. | → 13 |
13. Leaf blades elliptic to ovate. | C. virgata |
13. Leaf blades linear to lanceolate. | C. australis |
6. Petals not clawed or claw not greater than 2 mm, without lateral lobes; stamens not subtended by scales. | → 14 |
14. Petals shallowly to deeply 2-lobed [6h.1. sect. Phaeostoma subsect. Lautiflorae]. | C. biloba |
14. Petals not 2-lobed, sometimes emarginate. | → 15 |
15. Stamens subequal, anthers of similar size and color. | → 16 |
16. Ovary 4-grooved; capsules usually wider distally [6c.3. sect. Rhodanthos subsect. Jugales]. | C. gracilis |
16. Ovary conspicuously 8-grooved; capsules not wider distally [6c.2. sect. Rhodanthos subsect. Flexicaules]. | → 17 |
17. Stigma not exserted beyond anthers; pedicel in fruit 0–3 mm. | C. lassenensis |
17. Stigma exserted beyond anthers; pedicel in fruit 5–15 mm. | C. arcuata |
15. Inner stamens shorter, inner anthers much smaller, paler. | → 18 |
18. Stigma not exserted beyond anthers; petals 5–12 mm. | → 19 |
19. Petals white to pale cream, not flecked, fading pink [6h.3. sect. Phaeostoma subsect. Micranthae]. | C. epilobioides |
19. Petals pale to dark pink, usually darker flecked. | → 20 |
20. Petals pink, usually darker flecked [6h.1. sect. Phaeostoma subsect. Lautiflorae]. | C. modesta |
20. Petals pale pink, shading nearly white near base, purple- flecked [6h.2. sect. Phaeostoma subsect. Prognatae]. | C. similis |
18. Stigma exserted beyond anthers; petals 10–35 mm. | → 21 |
21. Corollas rotate; petals oblanceolate [6h.1. sect. Phaeostoma subsect. Lautiflorae]. | C. lingulata |
21. Corollas bowl-shaped; petals fan-shaped. | → 22 |
22. Ovary 8-grooved [6h.1. sect. Phaeostoma subsect. Lautiflorae] . | C. dudleyana |
22. Ovary 4-grooved [6h.5. sect. Phaeostoma subsect. Sympherica]. | → 23 |
23. Width of outer filaments about 2 times inner; floral tube 2–7 mm ring of hairs in floral tube below rim. | C. cylindrica |
23. Width of all filaments about equal or inner slightly thinner; floral tube 1.5–4 mm ring of hairs in floral tube at rim. | → 24 |
24. Capsule beak 0–3 mm . | C. lewisii |
24. Capsule beak 7–15 mm. | C. rostrata |
5. Inflorescence axis straight or erect; buds erect or pendent. | → 25 |
25. Buds pendent; corollas rotate or bowl-shaped; inner stamens shorter, inner anthers smaller, paler. | → 26 |
26. Petals 2-lobed, with a slender central tooth [6g. sect. Fibula]. | C. xantiana |
| → 27 |
27. Corollas bowl-shaped; petals not clawed [6g. sect. Fibula]. | → 28 |
| C. bottae |
| C. jolonensis |
27. Corollas rotate; petals clawed. | → 29 |
29. Petal claw shorter than blade [6h.4. sect. Phaeostoma subsect. Connubium]. | C. delicata |
29. Petal claw equal to or longer than blade [6h.6. sect. Phaeostoma subsect. Phaeostoma]. | → 30 |
30. Sepals and ovary puberulent, mixed with longer, straight spreading hairs to 3 mm. | C. unguiculata |
30. Sepals and ovary sparsely to densely puberulent, without longer, straight spreading hairs. | → 31 |
31. Leaf blades not glaucous, bright green. | C. exilis |
31. Leaf blades glaucous, gray-green or reddish green. | → 32 |
32. Sepals usually dark red-purple; stigma exserted beyond anthers. | C. springvillensis |
32. Sepals green, red-tinged or not; stigma exserted or not beyond anthers. | C. tembloriensis |
25. Buds erect; corollas bowl-shaped; stamens subequal, anthers of similar size and color. | → 33 |
33. Sepals remaining connate along edges and reflexed together to 1 side. | → 34 |
| → 35 |
35. Petals 30–60 mm; ovary fusiform [6c.1. sect. Rhodanthos subsect. Primigenia]. | C. amoena |
35. Petals 5–15 mm; ovary cylindrical [6f. sect. Biortis]. | C. affinis |
34. Ovary 4-grooved [6c.1. sect. Rhodanthos subsect. Primigenia]. | → 36 |
36. Petals with distinct red spot or mark near middle. | C. amoena |
36. Petals red or reddish purple at base. | → 37 |
37. Petals 10–30 mm; stigma exserted beyond anthers. | C. rubicunda |
37. Petals 5–13 mm; stigma not exserted beyond anthers. | C. franciscana |
33. Sepals reflexed individually or in pairs [6e. sect. Godetia]. | → 38 |
38. Stigma not exserted beyond anthers; petals 5–15 mm. | → 39 |
39. Stems erect; leaf blades usually linear to lanceolate, apex acute. | C. purpurea |
39. Stems prostrate to decumbent; leaf blades elliptic to oblanceolate, apex usually obtuse. | → 40 |
40. Petals 5–11 mm, without spot. | C. davyi |
40. Petals 10–15 mm, with red spot above base. | C. prostrata |
38. Stigma exserted beyond anthers; petals 10–30 mm. | → 41 |
41. Inflorescences dense; ovary at anthesis longer than adjacent internode. | → 42 |
42. Floral tube conspicuously veined; petals with large, wedge-shaped purplish red spot near apex. | C. imbricata |
42. Floral tube not conspicuously veined; petals with red spot near or proximal to middle. | C. speciosa |
41. Inflorescences open; ovary at anthesis shorter than adjacent internode. | → 43 |
43. Buds mucronate, sepal tips distinct in bud. | C. williamsonii |
43. Buds not mucronate, sepal tips connate to tip. | → 44 |
44. Petals without red or red-purple spot. | C. speciosa |
44. Petals with conspicuous red or purple spot. | → 45 |
45. Petal spot at or proximal to middle. | C. speciosa |
45. Petal spot distal to middle. | C. purpurea |
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FNA vol. 10. Authors: Harlan Lewis†, Peter C. Hoch. |
FNA vol. 10. |
Onagraceae > subfam. Onagroideae > tribe Onagreae |
Onagraceae > subfam. Onagroideae |
C. affinis, C. amoena, C. arcuata, C. australis, C. biloba, C. borealis, C. bottae, C. breweri, C. concinna, C. cylindrica, C. davyi, C. delicata, C. dudleyana, C. epilobioides, C. exilis, C. franciscana, C. gracilis, C. heterandra, C. imbricata, C. jolonensis, C. lassenensis, C. lewisii, C. lingulata, C. mildrediae, C. modesta, C. mosquinii, C. prostrata, C. pulchella, C. purpurea, C. rhomboidea, C. rostrata, C. rubicunda, C. similis, C. speciosa, C. springvillensis, C. stellata, C. tembloriensis, C. unguiculata, C. virgata, C. williamsonii, C. xantiana |
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Pursh: Fl. Amer. Sept. 1: 256, 260 [ — as Clarckia], plate 11. 1813 |
Dumortier: Fl. Belg., 89. (1827) |
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