Epilobium hirsutum |
Epilobium |
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codlins-and-cream, fiddle grass, fiddle grass willowherb, great or great hairy willowherb, great willowherb, hairy willlowherb, hairy willow-herb, épilobe hirsute |
spike-primrose, willow-herb, willow-weed |
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Habit | Herbs usually robust and rank, sometimes woody near base, with thick, ropelike stolons to 1 m with scattered cataphylls and, often, terminal leafy rosette. | Herbs, annual or perennial, sometimes suffrutescent, caulescent, often with basal rosettes, fleshy decussate turions, soboles, stolons, which may be tipped with turions, or rarely buds (gemmae) in leaf axils; with woody base or caudex, or with taproots. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Stems | erect to ascending, often clumped, terete, 25–120(–250) cm, unusually thick, 3–9 mm diam., well branched mainly in distal 1/2, densely long-villous throughout, usually mixed glandular puberulent distally, rarely sparsely villous or densely white-tomentose. |
erect to ascending or decumbent, simple to well-branched distally or sometimes from base, in some species proximal epidermis exfoliating, strigillose, glandular-puberulent, villous, often mixed, or glabrous, often with raised hairy lines decurrent from leaf axils. |
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Leaves | opposite proximal to inflorescence, alternate distally, sessile and ± clasping stem; blade elliptic-lanceolate to narrowly obovate or elliptic, 4–12(–23) × 0.3–4(–5) cm, base cuneate to attenuate, margins serrulate, 15–50 teeth per side, veins 6–9 per side, apex acute to acuminate or obtuse proximally, surfaces ± densely villous; bracts moderately reduced. |
cauline, sometimes also basal, opposite and decussate proximal to inflorescence or only in proximal pairs, or rarely throughout, alternate or, sometimes, fasciculate distally; stipules absent; subsessile to petiolate; blade margins entire or toothed; bracts usually reduced distally. |
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Inflorescences | erect racemes or panicles, usually densely villous and glandular puberulent, rarely tomentose. |
terminal, usually racemes or spikes, rarely panicles, flowers solitary in leaf axils, erect or ascending, sometimes also flowering from proximal nodes; bracteoles absent. |
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Flowers | erect; buds 5–9 × 1.8–4.5 mm, sometimes beaked; pedicel 3–11 mm; floral tube 1.3–2.9 × 2.2–4 mm, conspicuous ring of spreading hairs near mouth inside; sepals oblong-linear, often keeled, 6–12 × 1–3 mm, abaxial surface densely pubescent; petals bright pink to rose-purple, rarely white, broadly obcordate, 9–20 × 7–15 mm, apical notch 1–3 mm; filaments white or pink, those of longer stamens 5–10 mm, those of shorter ones 2.5–6 mm; anthers cream, 1.5–3 × 0.6–1.2 mm; ovary 15–34 mm, densely villous and glandular puberulent; style white or pink, 5–12 mm, usually glabrous, stigma deeply 4-lobed, 1.8–2.2 × 3–5.5 mm, lobes recurved or spreading, exserted beyond anthers. |
bisexual, usually actinomorphic, rarely zygomorphic, buds often erect, sometimes recurved; floral tube deciduous (with sepals, petals, and stamens) after anthesis, obconic or cylindrical, sometimes with bulbous base, with scales or ring of hairs or raised ring of tissue near mouth inside, nectary at base of tube; sepals 4, spreading individually, usually green or flushed with red or cream, rarely same color as petals, lanceolate; petals 4, usually rose-purple to white, rarely cream-yellow or orange-red, usually obcordate or broadly obovate, emarginate; stamens 8, in 2 unequal series with episepalous longer or rarely subequal, erect, anthers versatile, on smallest flowers appearing basifixed, pollen usually shed in tetrads, rarely singly; ovary 4-locular, style erect, stigma entire and clavate to capitate, or deeply 4-lobed, commissural, receptive only on inner surfaces, surface dry with multicellular papillae. |
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Fruit | a capsule, straight or slightly curved, narrowly cylindrical to fusiform or rarely narrowly ellipsoidal, usually terete, rarely ± 4-angled, loculicidally dehiscent, usually splitting to base with intact central column, rarely splitting only on distal 1/3 with central column disintegrating; pedicellate or sessile. |
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Capsules | often flushed purple, 25–90 mm, surfaces usually densely villous and glandular puberulent, rarely glabrescent; pedicel 5–20 mm. |
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Seeds | narrowly obovoid, 0.8–1.2 × 0.3–0.6 mm, chalazal collar inconspicuous, dark brown, surface coarsely papillose; coma easily detached, tawny or dull white, 7–10 mm. |
usually numerous (1–100+ per locule), usually in 1, rarely 2, rows per locule, very rarely 1 row per capsule by dissolution of septa and central column, surface papillose to finely reticulate or longitudinally ridged, sometimes constricted near micropylar end, rarely with inflated rim around body of seed on adaxial side, with persistent coma at chalazal end or coma absent. |
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x |
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2n | = 36. |
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Epilobium hirsutum |
Epilobium |
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Phenology | Flowering Jun–Sep. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Habitat | Low wet areas along streams, rivers, ponds, and lakes, roadside ditches, along railroad tracks, marshes and swampy areas. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Elevation | 0–150[–3000] m. (0–500[–9800] ft.) | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Distribution |
CO; CT; IL; IN; KY; MA; MD; ME; MI; NH; NJ; NY; OH; OR; PA; RI; UT; VT; WA; WI; WV; BC; NB; NS; ON; PE; QC; Eurasia; Africa [Introduced in North America]
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North America; Mexico; Central America; South America; West Indies; Eurasia; Africa; Pacific Islands (New Zealand); Australasia |
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Discussion | Epilobium hirsutum is very widespread in cool temperate Eurasia and montane regions, occurring throughout Europe except in the far north (P. H. Raven 1968), through the Caucasus and central Asia (E. I. Steinberg 1949) to Nepal (Raven 1962), China (Chen C. J. et al. 1992), and Japan (A. W. Lievens and P. C. Hoch 1999). It occurs as well along the Mediterranean coast of Africa, through East Africa to southern Africa, and in the Canary and Cape Verde Islands (Raven 1967). Epilobium hirsutum exceeds almost all other species of the genus in stature, so its size, very large flowers, and densely villous aspect make it easy to identify. R. L. Stuckey (1970) provided a detailed account of the introduction and spread of E. hirsutum in North America, noting the earliest known collection (July 1829) was from Newport, Rhode Island. Most early collections appeared in waste areas, particularly near harbor ballast piles, although some may have been grown in gardens. By the 1890s this species was well established along the Atlantic coastal region from New Jersey and Philadelphia through New England, and around Niagara Falls in the Great Lakes region. During the twentieth century, E. hirsutum spread extensively in southern Ontario and Quebec, south along the Atlantic coast to Maryland, and to all of the states along the southern shores of the Great Lakes, most recently including Wisconsin (1970), and Indiana (1972). It occurs in much the same habitat as that of another, more widely publicized invader, Lythrum salicaria, and sometimes is recorded as a companion species. The earliest known collection in western North America was made in 1933 in Bingen (Klickitat County), Washington. Whether from that introduction or others, E. hirsutum is now naturalized and widespread in the Pacific Northwest. It also was reported recently from the Denver region in Colorado and near Midway in Utah. Epilobium grandiflorum F. H. Wiggers and E. grandiflorum Allioni are illegitimate names that pertain here. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Species ca. 165 (41 in the flora). Epilobium is the largest genus in Onagraceae, distributed mainly in cool temperate, montane, boreal, and arctic areas on all continents except Antarctica. P. H. Raven (1976) proposed many elements of the current classification of Epilobieae, recognizing six sections in Epilobium, including for the first time the genus Zauschneria as a separate section, based on patterns of variation in chromosome number (H. Lewis and Raven 1961; M. Kurabayashi et al. 1962), wood anatomy (S. Carlquist 1975), seed morphology (S. R. Seavey et al. 1977), and pollen wall (J. J. Skvarla et al. 1976) and viscin thread morphology (Skvarla et al. 1978). Raven excluded Boisduvalia from Epilobium, though it later was included (P. C. Hoch and Raven 1992) as two distinct sections, and Raven included Chamaenerion as a section, which was later excluded (Hoch and Raven 1999; W. L. Wagner et al. 2007). Both of these changes from the 1976 classification were based in large part on relationships supported by molecular data (D. A. Baum et al. 1994). The monophyly of this re-defined Epilobium was very strongly supported in analyses of ITS alone (Baum et al.) and ITS plus trnL-F (R. A. Levin et al. 2004), as was the segregation of Chamaenerion from Epilobium. In addition, both analyses strongly supported a clade that includes the former segregated genera Boisduvalia and Zauschneria as well as the sections of Epilobium with chromosome numbers other than n = 18. As treated here, Epilobium is divided into eight sections, one of which has two subsections (Wagner et al.). The phylogeny of Epilobium as elucidated by D. A. Baum et al. (1994) and later R. A. Levin et al. (2004) suggests complex chromosomal evolution in the genus. Since the outgroup (Chamaenerion) and one major clade (sect. Epilobium) share the base number x = 18, the derivation of the diverse other gametic numbers (n = 9, 10, 12, 13, 15, 16, and 19) appears to have involved aneuploid reduction from x = 18. One explanation of the relationships proposed by the molecular data suggests the early evolution of a lineage with n = 15 (sects. Cordylophorum, Epilobiopsis, and Zauschneria), from which arose a branch with n = 12 (sect. Xerolobium), a separate branch with n = 13 and n = 16 (sect. Crossostigma), and a third branch with n = 9, 10, and 19 (sect. Pachydium). This hypothesis requires the fewest aneuploidy changes, but confirming it will require additional molecular and cytological analysis. Almost all Epilobium species tested are self-compatible, but at least E. obcordatum is apparently self-incompatible (S. R. Seavey and S. K. Carter 1994, 1996). All species have hermaphroditic, diurnal flowers that usually remain open for more than one day. Many species are modally autogamous, a few primarily cleistogamous, but others, including 15 species in North America, are partly or wholly outcrossing. In the latter group, flowers are often markedly protandrous, with a 4-lobed stigma exserted beyond anthers. Primary pollinators include bees, flower flies, butterflies, moths, and sometimes hummingbirds (sect. Zauschneria). In general, flowering commences in the first year of growth. Only the nine species (eight in the flora area) of sects. Pachydium, Crossostigma, Epilobiopsis, and Xerolobium are annuals; all other taxa of Epilobium (sects. Cordylophorum, Epilobium, Macrocarpa, and Zauschneria) are perennials. Annual species have taproots and proximal stems usually with peeling epidermis. Perennial species of Epilobium, which have more fibrous root systems and rarely peeling epidermis, vary greatly in the habit and stature, degree of clumping, degree of branching, and mode of perennation. The various types of perennating structures in Epilobium (R. C. Keating et al. 1982; Chen C. J. et al. 1992) are extremely useful for identification of many taxa; unfortunately, many plants are collected without the structures. Unless care is taken to dig up the bases of the plants, some of the most useful diagnostic features such as turions are lost, making identification more difficult. Perennating structures are less useful for phylogenetic analysis due to the difficulty in determining whether the common possession of a particular type represents an analogous or homologous pattern. However, one type of structure (soboles) is found in at least some taxa of all perennial sections and may be the most generalized type in the genus. Soboles are here defined as simple, scaly, underground shoots with somewhat elongated internodes and arising from the caudex. Soboles generally give rise to more or less clumped plants with ascending stems. In certain species, including Epilobium rigidum (sect. Macrocarpa), all taxa of sects. Cordylophorum and Zauschneria, and some in sect. Epilobium, soboles arise from more or less woody caudices, a character state that may be plesiomorphic in the genus; all other types of perennating structures are found only in sect. Epilobium and are discussed there. In addition to habit and perennating structures, certain other morphological features particularly useful for diagnosing plants include: leaf size, shape, attachment, and arrangement on stem; seed surface (papillose, reticulate, or ridged), size, and shape (S. R. Seavey et al. 1977); pubescence type (mainly strigillose: short incurved hairs; villous: long, thin, spreading hairs; and/or glandular-puberulent) and pattern on stem (all around or restricted to raised lines decurrent from base of petioles) and leaves; and some floral features including size, shape, and color. In fact, floral features are diagnostic for only certain sections or species; most species have very similar flowers. (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. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Parent taxa | Onagraceae > subfam. Onagroideae > tribe Epilobieae > Epilobium > sect. Epilobium | Onagraceae > subfam. Onagroideae > tribe Epilobieae | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Sibling taxa | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Synonyms | Chamaenerion hirsutum, E. amplexicaule, E. aquaticum, E. hirsutum var. villosum, E. villosum | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Name authority | Linnaeus: Sp. Pl. 1: 347. (1753) | Linnaeus: Sp. Pl. 1: 347. (1753): Gen. Pl. ed. 5, 164. (1754) | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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