Oenothera sect. Calylophus |
Onagraceae |
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evening-primrose family |
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Habit | Herbs perennial, rarely annual, sometimes suffrutescent, caulescent; from a stout to slender taproot. | 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. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Stems | decumbent to ascending or erect, branched or unbranched, epidermis gray to brown, ± exfoliating. |
erect to decumbent or prostrate. |
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Leaves | cauline, 0.3–9 cm, fascicles of small leaves often present in larger leaf axils; blade margins entire or subentire, serrate, or serrulate, sometimes spinulose. |
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. |
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Inflorescences | solitary flowers in axils of distal leaves. |
axillary, flowers solitary, leafy spikes, racemes, or panicles. |
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Flowers | opening near sunset or sunrise, sometimes in afternoon, usually with a sweet scent; buds erect, terete or quadrangular, with free tips; floral tube 2–60(–70) mm; sepals flat or with keeled midribs, reflexed individually; petals yellow, usually fading dark yellow, orange, pale pink, or pale purple, suborbiculate to rhombic or obcordate; stigma usually yellow to yellow-green, blue-black in O. capillifolia subsp. capillifolia, peltate, discoid to quadrangular, sometimes shallowly 4-lobed. |
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. |
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Fruit | a loculicidal capsule or indehiscent berry or nutlike. |
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Capsules | woody and hard to thin and ± papery, straight, cylindrical to obtusely 4-angled, often tapering at each end, dehiscent 1/2 to throughout their length; sessile. |
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Seeds | usually numerous, in 2 rows per locule, usually obovoid and somewhat angled, rarely oblanceoloid, surface smooth. |
smooth or sculptured, sometimes with a coma or wings, with straight, oily embryo, 4-nucleate embryo sac, endosperm absent. |
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2n | = 14 (28). |
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Oenothera sect. Calylophus |
Onagraceae |
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Distribution | w North America; Mexico; c North America |
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 |
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Discussion | Species 7 (7 in the flora). Section Calylophus consists of 7 species (13 taxa) classified in two subsections distributed throughout the Great Plains to Arizona and south to central Mexico with a center of diversity in Texas (H. F. Towner 1977). P. H. Raven (1964) separated Calylophus from Oenothera as treated by P. A. Munz (1965) based on its peltate stigma and unusual sporogenous tissue (Towner). The peltate stigma can now be more properly interpreted as a variation from the typical Oenothera stigma, with the indusium enlarged and the lobes reduced (W. L. Wagner et al. 2007). Molecular studies (R. A. Levin et al. 2004) strongly support both the inclusion of Calylophus within Oenothera and the monophyly of the section by inclusion of a species from each of the subsections. Subsequent detailed analyses (B. Cooper, unpubl.) indicate that subsect. Salpingia is not monophyletic with subsect. Calylophus nested within it, indicating that subsections may not be justifiable as currently defined. All species, except O. serrulata, are self-incompatible and outcrossing; flowers diurnal to vespertine, opening in the early morning or from midafternoon to near sunset, wilting in one and one-half to two days; those species with diurnal flowers are pollinated by bees [especially Halictidae (halictids) and Anthophoridae (anthophorids), often oligolectic species], those with larger vespertine flowers are pollinated by hawkmoths (Towner). The self-compatible O. serrulata is autogamous and a PTH species. Recent work (M. Moore, pers. comm.) suggests that there is additional edaphic endemism within sect. Calylophus, which is being investigated using morphology and molecular analyses (B. Cooper, unpubl.), and will likely result in detection of previously unknown species in the section. Meriolix Rafinesque ex Endlicher 1840 is a superfluous name that pertains here. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
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.) |
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Key |
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Source | FNA vol. 10. | FNA vol. 10. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Synonyms | Calylophus, O. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Name authority | (Spach) W. L. Wagner & Hoch: Syst. Bot. Monogr. 83: 147. (2007) | Jussieu | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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