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FOC | Family List | FOC Vol. 13 | Onagraceae

4. Epilobium Linnaeus, Sp. Pl. 1: 347. 1753.

柳叶菜属 liu ye cai shu

Authors: Jiarui Chen, Peter C. Hoch & Peter H. Raven

Boisduvalia Spach; Chamaenerion Séguier; Zauschneria Presl.

Herbs perennial [or annual, sometimes suffrutescent], with leafy rosettes, stolons, soboles (shoots), or turions (subterranean globose buds with fleshy scales). Stems glabrous to pubescent, often with lines of hairs decurrent from margins of petioles. Leaves opposite, becoming alternate and bractlike in inflorescence; petiolate or sessile; stipules absent; bracteoles absent. Inflorescences simple or branched racemes, panicles, spikes, or corymbs. Flowers 4-merous, often protandrous, with floral tube, producing nectar at base of style. Petals pink to rose-purple or white [or rarely cream-colored or orange-red], obcordate or obtrullate, notched at apex. Stamens 8, in two unequal whorls; pollen yellow, shed in tetrads. Style erect; stigma entire or 4-lobed. Fruit an elongate, slender capsule, 4-loculed, loculidical. Seeds many [or rarely only 4], generally with terminal coma of silky hairs [or coma rarely lacking]. 2n = [18, 20, 24, 26, 28, 30, 32,] 36, [38, 60].

About 165 species: montane, boreal, and/or arctic regions of Africa, Asia, Australasia, Europe, and North and South America, from sea level to 5000 m, often in moist, disturbed places: 33 species (nine endemic) in China.

The genus is divided into seven sections, all present in North America but only one, Epilobium sect. Epilobium, in China, where the species occur in most temperate to montane habitats except extreme deserts or warm, subtropical forests.

Careful gathering of ripe seeds and perennating structures, usually at or just below the ground surface, facilitate identification. The pattern of vestiture on stems, also extremely valuable for identification, is sometimes obscure on specimens collected very late in the growing season.

1 Stigma deeply 4-lobed (rarely shallowly so in E. blinii)   (2)
+ Stigma entire or shallowly emarginate   (5)
2 (1) Stems (18-)30-120(-250) cm tall, with long spreading and short erect glandular hairs; leaves 3-12 cm, subacute to acuminate, with (15-)20-60 teeth per side; seeds 0.8-1.2 mm, coarsely papillose   (3)
+ Stems 3-25(-45) cm tall, strigillose; leaves 0.8-3 cm, obtuse, with 3-7 indistinct teeth per side; seeds 1.2-1.8 mm, finely papillose or reticulate   (4)
3 (2) Leaves clasping; petals (7-)10-20 mm; stigma exserted beyond anthers at anthesis.   1 E. hirsutum
+ Leaves subsessile but not clasping; petals 5-8 mm; stigma surrounded by anthers of longer stamens at anthesis.   2 E. parviflorum
4 (2) Loosely matted herb, stems 3-18 cm tall, branched; leaf petioles 1-3 mm; petals 1.6-3.1 cm; fruiting pedicels 4-7 mm; seeds reticulate; Taiwan.   4 E. nankotaizanense
+ Ascending or erect herb, stems 10-25(-45) cm tall, usually simple; leaves subsessile; petals 1-1.5 cm; fruiting pedicels 15-35 mm; seeds finely papillose; SW China.   3 E. blinii
5 (1) Stems pubescent throughout, either lacking raised pubescent lines decurrent from margins of petioles, or lines inconspicuous   (6)
+ Stems subglabrous below inflorescence except for 2 or 4 raised strigillose lines decurrent from margins of petiole   (17)
6 (5) Plants forming short basal shoots (soboles), often clumped; stems occasionally with persistent basal scales   (7)
+ Plants with basal rosettes, turions, or stolons, not basal shoots, rarely clumped; stems often with persistent basal scales   (13)
7 (6) Inflorescence variously pubescent, but without glandular hairs   (8)
+ Inflorescence variously pubescent, but always with glandular hairs, especially on inflorescence   (10)
8 (7) Ovary white canescent; seeds with conspicuous chalazal collar; leaves oblong-lanceolate to narrowly ovate, 2-7 cm.   29 E. minutiflorum
+ Ovary strigillose, not white canescent; seeds with inconspicuous chalazal collar; leaves sublinear to elliptic or lanceolate, 1-4.5 cm   (9)
9 (8) Plants erect, loosely clumped or not clumped; stems mostly well-branched throughout; leaves 1-4.5 cm, linear to narrowly lanceolate.   10 E. platystigmatosum
+ Plants ascending, forming distinct clumps or mats; stems simple or branched only from base; leaves 1-2 cm, elliptic to lanceolate.   11 E. hohuanense
10 (7) Mid-cauline and upper leaves with distinct petioles 2-7 mm, and with narrowly cuneate bases.   13 E. royleanum
+ Mid-cauline and upper leaves subsessile or with petioles to 2 mm, and with subcordate, rounded, or broadly cuneate bases   (11)
11 (10) Leaves subentire, lanceolate-elliptic, seed coma tawny.   28 E. fastigiatoramosum
+ Leaves denticulate, ovate to lanceolate or oblong-lanceolate; seed coma white or dingy   (12)
12 (11) Petals 3-4.3(-5) mm; ovary white canescent; seeds with conspicuous chalazal collar.   29 E. minutiflorum
+ Petals 7-11 mm; ovary pubescent but not white canescent; seeds with inconspicuous chalazal collar.   16 E. brevifolium
13 (6) Plants throughout densely appressed tomentose; plants with leafy rosettes; petals 8-16 mm.   15 E. pannosum
+ Plants strigillose, villous, and/or glandular pubescent, but not tomentose; plants forming stolons or turions, not leafy rosettes; petals 4-8 mm (except to 15 mm in E. kermodei)   (14)
14 (13) Plants forming fleshy turions, leaving dense leathery basal scales; fruiting pedicels 5-10 mm; plants 7-25 cm tall; Taiwan.   12 E. taiwanianum
+ Plants forming stolons, usually with few or no basal scales; fruiting pedicels 7-50 mm; plants 25-120 cm tall (except E. palustre, (5-)15-70 cm tall); widespread, but not in Taiwan   (15)
15 (14) Plants forming thick, ropelike stolons with fleshy terminal buds; capsules 7-11 cm; seeds 0.8-1.2 mm.   14 E. kermodei
+ Plants forming filiform, threadlike stolons, sometimes with terminal buds; capsules 3-9 cm; seeds 1.3-2.2 mm   (16)
16 (15) Leaves sublinear to narrowly lanceolate, subentire to obscurely denticulate; stolons with fleshy terminal turions; fruiting pedicels 1-5 cm; seed comas dingy white.   27 E. palustre
+ Leaves ovate or broadly oblong to lanceolate, sharply serrulate; stolons without terminal turions; fruiting pedicels 0.7-1.5 cm; seed comas reddish.   17 E. pyrricholophum
17 (5) Inflorescence glabrous or rarely with scattered hairs on ovary and sepals   (18)
+ Inflorescence moderately to densely pubescent   (19)
18 (17) Plants low, mat-forming; stems 3-20 cm tall; leaves 0.8-2.5 cm; petals 2.5-6.5 mm; capsules 1.7-3.6 cm.   33 E. anagallidifolium
+ Plants erect, clumped; stems (5-)10-25(-60) cm tall; leaves (0.8-)1.5-7.5 cm; petals 7-14 mm; capsules 5-9 cm.   22 E. sikkimense
19 (17) Inflorescence strigillose and/or villous, but lacking short, erect glandular hairs   (20)
+ Inflorescence variously pubescent, always with some short, erect glandular hairs   (26)
20 (19) Leaves subsessile (lower ones often with petioles to 2 mm)   (21)
+ Leaves with distinct petioles 2-11 mm   (23)
21 (20) Stems 10-50 cm tall, erect; leaves 3.4-6 cm; seeds papillose.   5 E. roseum
+ Stems 7-25 cm tall, ascending; leaves 1-2.5 cm; seeds reticulate   (22)
22 (21) Leaves 1.1-1.6 cm; capsules 3.5-4 cm, pedicels 6-7 mm; seeds 0.8-0.9 mm; SW China.   31 E. clarkeanum
+ Leaves 1.5-2.5 cm; capsules 4.5-5 cm, pedicels 15-20 mm; seeds 1-1.1 cm; Taiwan.   32 E. pengii
23 (20) Leaves narrowly spatulate to rarely broader, with prominent pale abaxial midvein, often crowded and spirally arranged on mid-stem; petioles 2-11 mm; fruiting pedicels 1.3-4 cm.   9 E. sinense
+ Leaves sublinear to narrowly ovate, without pale midvein, always opposite below inflorescence; petioles 2-7(-10) mm; fruiting pedicels 0.5-3 cm   (24)
24 (23) Leaves narrowly lanceolate to sublinear; petioles 3-7(-10) mm; seeds 0.8-1 mm, papillose.   7 E. cylindricum
+ Leaves lanceolate to narrowly ovate; petioles 2-5 mm; seeds 1-1.3 mm, reticulate or papillose   (25)
25 (24) Plants well-branched, not much clumped; petals 5-8 mm; stigma capitate to broadly clavate; seeds reticulate; Himalayas and SW China.   8 E. tibetanum
+ Plants mostly simple, forming clumps; petals 5.5-6.5 mm; stigma clavate or rarely subcapitate; seeds coarsely papillose; W China (Tian Shan).   6 E. tianschanicum
26 (19) Plants forming fleshy turions at or below ground level; stem bases with thick, brown, leathery scales   (27)
+ Plants forming soboles, stolons, or rosettes, but not fleshy turions; stem bases with rather loose, ± herbaceous scales, or scales absent   (31)
27 (26) Leaves elliptic or oblong to elliptic-lanceolate, with mostly cuneate bases and distinct petioles (1-)2-15 mm   (28)
+ Leaves ovate to lanceolate, with ± rounded bases and obscure petioles to 3 mm (except in E. laxum with petioles 2-8 mm on lower leaves)   (29)
28 (27) Petioles 3-10(-15) mm; stigma broadly clavate to subcapitate; seeds 1-1.2 × 0.45-0.55 mm, obovoid; Xinjiang.   5 E. roseum
+ Petioles 1-4(-6) mm; stigma capitate; seeds 1.1-1.4 × 0.3-0.45 mm, narrowly obovoid; SW China.   26 E. fangii
29 (27) Petals 4.5-5.5 mm; stigma clavate to subcapitate; seeds 0.9-1 mm.   25 E. gouldii
+ Petals 5.5-16 mm; stigma capitate or nearly so; seeds 1.1-1.5 mm   (30)
30 (29) Leaves 2-7 × 1.2-2.6 cm, ovate to narrowly ovate, crowded, usually longer than internodes; petals (7-)10-16 mm; fruiting pedicels 1-7(-10) mm; W Himalayas and Tian Shan   21 E. laxum
+ Leaves 1.5-5.5 × 0.5-1.5 cm, narrowly ovate to lanceolate, not crowded, usually shorter than internodes; petals 5.5-11 mm; fruiting pedicels 4-12 mm; SW China.   24 E. subcoriaceum
31 (26) Plants forming leafy rosettes; seed surface with conspicuous longitudinal ridges of flattened fused papillae; NE China.   30 E. ciliatum
+ Plants forming soboles or stolons; seed surface papillose or reticulate, without longitudinal ridges   (32)
32 (31) Stems ascending, forming clumps or mats; leaves subentire to scarcely denticulate; capsules 1.7-3.6 cm; pedicels 1-3.5(-5) cm; seeds reticulate.   33 E. anagallidifolium
+ Stems mostly erect, loosely or not clumped; leaves denticulate or serrulate; capsules (1.5-)3.5-11 cm; pedicels 0.3-2.5 cm; seeds papillose   (33)
33 (32) Plants forming thick ropelike stolons 1-12 cm with fleshy terminal bud; robust stems 40-120(-200) cm tall; capsules 7-11 cm.   14 E. kermodei
+ Plants forming soboles, not stolons; stems 4-80(-150) cm, mostly less than 50 cm; capsules 1.5-7.5 cm (rarely to 9 cm in E. sikkimense)   (34)
34 (33) Stem bases with crowded brown scales; stems 5-25(-60) cm tall   (35)
+ Stem bases without scales; stems (10-)20-150 cm tall, mostly more than 25 cm tall (except E. kingdonii, with stems 8-25 cm tall)   (36)
35 (34) Stems mostly erect; leaves (0.8-)1.5-7.5 cm; petals 7-14 mm; capsules 5-9 cm; pedicels 0.6-2(-2.5) cm.   22 E. sikkimense
+ Stems mostly ascending; leaves 0.7-2.2 cm; petals 5-6.5 mm; capsules 3.5-5(-6) cm; pedicels 0.4-1 cm.   23 E. williamsii
36 (34) Plants 8-25 cm tall, ascending; floral tube with sparse, even pubescence; seeds 1.4-1.6 mm, very finely papillose.   19 E. kingdonii
+ Plants (10-)20-150 cm tall, erect; floral tube usually with tufts of hairs on costae and at insertion of sepals at mouth of floral tube; seeds 0.8-1 mm, coarsely papillose   (37)
37 (36) Stems 15-80 cm tall, somewhat 4-angled, with (2 or)4 raised strigillose lines on internodes; leaves oblong, oblong-ovate, or elliptic, apex obtuse or rarely acute; petals 5.5-13 mm; fruiting pedicels 1-2.5 cm.   20 E. wallichianum
+ Stems (10-)20-150 cm tall, subterete, with 2 raised strigillose lines on internodes; leaves ovate to oblong-lanceolate, apex acute or acuminate; petals 4.5-8 mm; fruiting pedicels 0.3-1.3 cm.   18 E. amurense

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