Onagraceae |
Circaea |
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| evening-primrose family |
circaea, enchanter's nightshade, nightshade |
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| 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, perennial, caulescent, colonial; stolons numerous. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| Stems | erect to decumbent or prostrate. |
erect, unbranched or sparsely branched. |
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| 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, opposite; stipules present, soon deciduous; petiolate; blade margins dentate to prominently dentate. |
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| Inflorescences | axillary, flowers solitary, leafy spikes, racemes, or panicles. |
simple or branched racemes, terminal on main stem or also at apex of branches, erect. |
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| 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, zygomorphic, buds erect; floral tube inconspicuous, deciduous (with sepals, petals, and stamens) after anthesis, with a nectary wholly within and filling proximal portion of floral tube or elongated and projecting above opening of floral tube as a fleshy, cylindrical or ringlike disc; sepals 2, reflexed to spreading; petals 2, alternate sepals, white or pink, without spots, clawed, apex notched; stamens 2, anthers basifixed, pollen shed singly; ovary 1- or 2-locular, stigma bilobed or obpyramidal, surface wet, minutely papillate. |
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| Fruit | a loculicidal capsule or indehiscent berry or nutlike. |
a capsule, spreading or slightly reflexed, globose to clavoid or obovoid, indehiscent, surface smooth or with prominent longitudinal grooves (sulci) and rounded ridges, burlike, with stiff, hooked hairs; pedicellate, deciduous at maturity. |
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| Seeds | smooth or sculptured, sometimes with a coma or wings, with straight, oily embryo, 4-nucleate embryo sac, endosperm absent. |
1 or 2, ellipsoid, glabrous, without appendages. |
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Onagraceae |
Circaea |
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| 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 |
North America; Europe; Asia; n Africa |
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| 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 8 (3, including 1 hybrid, in the flora). Circaea occurs throughout the temperate and boreal northern hemisphere, but is most diverse in eastern Asia, where all but one species occur. Reproductive features include: self-compatible; flowers diurnal, outcrossing, and pollinated by syrphid flies and small bees, or, sometimes, autogamous. It is found in rich, moist soils in deciduous forests and thickets, forest margins, and in moss or soil in mixed, coniferous-broadleaved deciduous, boreal forests. Circaea alpina subsp. alpina and C. canadensis subsp. canadensis often grow in close proximity and hybridize in eastern North America to produce C. ×sterilis. The unilocular C. alpina, with petals less than 2 mm, is self-pollinating under adverse weather conditions, but outcrosses on warm, sunny days. Because of its shorter style and much smaller pollen grains, it is probably the pollen recipient during hybridization events. Artificial hybridization experiments in England using C. alpina as the pollen donor and C. lutetiana as the pollen recipient failed to result in offspring, although hybrids were easily produced in the other direction (P. M. Benoit 1966). Recent molecular phylogenetic analysis supported the separation of the C. canadensis complex into two species; C. alpina subsp. pacifica was found to be sister to the remainder of the genus rather than being nested with other members of C. alpina (Xie L. et al. 2009). Thus, despite the strong morphological similarities of taxa within the C. canadensis and C. alpina complexes, these North American taxa may be better treated as separate species. Further detailed molecular studies are underway to examine this in more detail (Xie et al., unpubl.). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
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| Name authority | Jussieu | Linnaeus: Sp. Pl. 1: 8. (1753): Gen. Pl. ed. 5, 10. (1754) | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| Source | FNA vol. 10. | FNA vol. 10. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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