Eremothera |
Onagraceae |
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evening primrose, mooncup |
evening-primrose family |
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Habit | Herbs, annual, caulescent; with a 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 | usually erect, sometimes ascending, usually well-branched from base, sometimes also distally, with white or reddish green exfoliating epidermis. |
erect to decumbent or prostrate. |
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Leaves | cauline, proximal ones often clustered near base, alternate; stipules absent; petiolate, often subsessile distally; blade margins denticulate, crenate-dentate, serrulate, sinuate-toothed, or entire. |
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 | spikes, erect or nodding at anthesis, or flowers also in proximal leaf axils in some taxa. |
axillary, flowers solitary, leafy spikes, racemes, or panicles. |
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Flowers | bisexual, actinomorphic, buds erect; floral tube deciduous (with sepals, petals, and stamens) after anthesis, with basal nectary; sepals 4, reflexed singly or in pairs; petals 4, usually white, rarely red or tinged red, without spots, fading pink or red; stamens 8 in 2 unequal series, episepalous ones rarely abortive (E. minor), anthers versatile, pollen shed singly; ovary 4-locular, without apical projection, style villous near base, strigillose, or glabrous, stigma entire, subglobose, surface unknown, probably wet and non-papillate. |
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 capsule, straight or much contorted, narrowly cylindrical throughout or thickened proximally, terete or 4-angled, regularly but tardily loculicidal; sessile. |
a loculicidal capsule or indehiscent berry or nutlike. |
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Seeds | numerous, in 1 row per locule, usually monomorphic and narrowly obovoid to oblanceoloid, sometimes dimorphic, with seeds near base of capsule sharply angular and truncate-ellipsoid, finely reticulate, or seeds near base of capsule coarsely papillose. |
smooth or sculptured, sometimes with a coma or wings, with straight, oily embryo, 4-nucleate embryo sac, endosperm absent. |
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Eremothera |
Onagraceae |
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Distribution | w United States; sc United States; nw Mexico |
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). Species of Eremothera are found mainly in the interior deserts and bordering areas of the western United States. R. A. Levin et al. (2004) found strong molecular support for paraphyly in the broadly delimited Camissonia of P. H. Raven (1969). There was some support for a clade of Camissonia and Eremothera and another clade of Camissoniopsis, Neoholmgrenia, and Tetrapteron (Levin et al.), but without morphological features linking the members of these two clades. The monophyletic subclades of these two clades were recognized as genera by W. L. Wagner et al. (2007) whereas they were all treated by Raven as clearly distinguishable sections. Raven recognized four distinct groups within Eremothera (as Camissonia sect. Eremothera): E. refracta and its autogamous derivative, E. chamaenerioides; the very diverse E. boothii (with six subspecies) and two rare autogamous derivatives, E. gouldii and E. pygmaea; the local clay endemic E. nevadensis; and the widespread autogamous and often cleistogamous E. minor. Levin et al. included one species from each of these four groups in their molecular analyses and found strong support for Eremothera as circumscribed by Raven and maintained by Wagner et al. Eremothera is well defined by white petals that open in the evening and an entire, subglobose stigma; some species are visited by moths at anthesis and by bees the following morning (Raven). Reproductive features include: self-incompatible (E. boothii, E. refracta, and, possibly, E. nevadensis) or self-compatible; flowers vespertine; outcrossing and pollinated in the evening by small moths and the following morning by bees, in E. boothii subsp. decorticans by large oligolectic andrenid bees (E. G. Linsley et al. 1963, 1964, 1973), or autogamous, rarely cleistogamous (Raven). (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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Source | FNA vol. 10. | FNA vol. 10. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Synonyms | Camissonia section eremothera, Oenothera section eremothera | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Name authority | (P. H. Raven) W. L. Wagner & Hoch: Syst. Bot. Monogr. 83: 125. (2007) | Jussieu | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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