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milk-flower willowherb, white-flower willow-herb, épilobe à fleurs blanches

Habit Herbs with short, leafy epigeal soboles. Herbs, annual or perennial, sometimes suffrutescent.
Stems

ascending to suberect, often clumped, terete, 15–50 cm, usually simple, rarely branched proximally, subglabrous proximal to inflorescence except for raised densely strigillose lines decurrent from margins of petioles, usually mixed strigillose and glandular puberulent distally.

Leaves

opposite proximal to inflorescence or just proximal 1/3, alternate distally, petiole 3–12 mm, ± winged;

blade broadly spatulate to ovate proximally, narrowly ovate to narrowly lanceolate distally, 2–5.5 × 0.8–2.4 cm, base attenuate to cuneate, margins subentire proximally to denticulate distally with 7–16 teeth per side, more marked distally, lateral veins inconspicuous, 4–8 per side, apex obtuse proximally to subacute distally, surfaces glabrous except for strigillose margins;

bracts reduced and narrower.

opposite at least near base or throughout, alternate distally, or sometimes alternate throughout;

stipules absent.

Inflorescences

nodding in bud, later erect, ± open racemes, mixed strigillose and glandular puberulent.

Flowers

suberect;

buds 2–5 × 1.5–3.5 mm;

pedicel 5–15 mm;

floral tube 1–2.2 × 1–3 mm, inner surface glabrous without ring;

sepals often flushed purplish red, frequently keeled, (2–)3–5.5 × 0.9–1.8 mm, abaxial surface sparsely glandular puberulent, sometimes mixed strigillose;

petals white, rarely with red veins or flushed light pink, 3–8.5 × 1.6–4.5 mm, apical notch 0.7–1.4 mm;

filaments white to cream, those of longer stamens 1.4–4 mm, those of shorter ones 1.1–3 mm;

anthers light yellow, 0.4–0.9 × 0.3–0.6 mm;

ovary 20–40 mm, glandular puberulent;

style cream or white, 1.4–4.6 mm, stigma clavate or rarely subcapitate and indented apically, entire, 1.2–2.5 × 0.4–1.6 mm, surrounded by anthers.

actinomorphic or slightly zygomorphic, 4-merous;

sepals erect or spreading;

stamens 2 times as many as sepals;

pollen shed in tetrads or monads.

Fruit

a slender, cylindrical, loculicidal capsule.

Capsules

slender, sometimes flushed reddish green, ± ascending, 50–100 mm, surfaces sparsely glandular puberulent;

pedicel 15–45 mm.

Seeds

narrowly obovoid, 1.1–1.7 × 0.4–0.6 mm, chalazal collar 0.05–0.1 mm, blond or light brown, surface reticulate or sometimes barely rugose;

coma easily detached, white, 7–14 mm.

(1 or) many per locule, with tuft of hairs (coma) at chalazal end, sometimes without coma.

2n

= 36.

Epilobium lactiflorum

Onagraceae tribe Epilobieae

Phenology Flowering Jun–Sep.
Habitat Montane stream banks, moist crevices and ledges, gravelly roadsides, burned-over woodlands, sandy moraines, subalpine forests, alpine meadows.
Elevation 50–3800 m. (200–12500 ft.)
Distribution
from FNA
AK; AZ; CA; CO; ID; ME; MT; NH; NM; NV; OR; UT; VT; WA; WY; AB; BC; NB; NL; NS; NT; ON; QC; YT; Greenland; Eurasia
[WildflowerSearch map]
[BONAP county map]
North America; Mexico; Central America; South America; West Indies; Eurasia; Africa; Atlantic Islands; Australasia [Introduced in Pacific Islands]
Discussion

Epilobium lactiflorum has a nearly circum-subarctic distribution in North America (including coastal Greenland) and Eurasia, extending south into alpine and cool montane habitats along mountain axes. This distribution is similar to that of E. anagallidifolium and E. hornemannii (all with CC chromosomal arrangement), and these species commonly grow in similar habitats as well.

Petal color can be variable in many Epilobium species, but E. lactiflorum (white flowers) differs quite consistently from E. hornemannii (rose-purple to light pink or rarely white) in that feature. Mature fruits and pedicels are also fairly longer in E. lactiflorum. Despite their morphological similarities and broadly overlapping ranges and habitats, E. lactiflorum and E. hornemannii subsp. hornemannii do not appear to hybridize with much frequency, although intermediates, with only moderately reduced seed fertility, might be difficult to detect.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera 2, species 173 (2 genera, 43 species in the flora).

Epilobium and its close relatives have been recognized historically either as part of Onagreae (A. P. de Candolle 1828b), sometimes as a subtribe (É. Spach 1834–1848, vol. 4; J. Torrey and A. Gray 1838–1843, vol. 1), or as the distinct Epilobieae (R. Raimann 1893; P. A. Munz 1965; P. H. Raven 1976). The synapomorphies for Epilobieae as currently delimited include its highly condensed, heteropycnotic chromosomes with a base chromosome number of x = 18, sepals held erect or spreading (not reflexed) throughout anthesis, and the presence of a coma of hairs on the seeds (secondarily lost in some species). Molecular support for the tribe is strong (97–100% BOOTSTRAP support; D. A. Baum et al. 1994; R. A. Levin et al. 2004).

Endlicher established Epilobieae and included Oenothera within it; it is unclear how or whether his concept differed from Onagreae of A. P. de Candolle (1828b). É. Spach (1834–1848, vol. 4) recognized these genera as Onagreae and differentiated so-called sect. Oenotherinae from sect. Epilobieae, placing in the latter not only Epilobium and related groups, but also Clarkia and its segregates. J. Torrey and A. Gray (1840) excluded Clarkia from their Epilobiinae and also excluded Boisduvalia, a delimitation also followed by R. Raimann (1893) for his Epilobieae. Epilobieae did not assume its current delimitation, including only Epilobium and its close relatives, until the works of P. A. Munz (1941) and P. H. Raven (1964).

G. L. Stebbins (1971) and P. H. Raven (1976) considered the diverse chromosome numbers in Epilobieae and proposed that the species of Boisduvalia (now a section of Epilobium) with n = 9 or 10 represented the original base chromosome number for the tribe, and that these numbers were derived from x = 11 which is found in Circaeeae, Gongylocarpeae, and Lopezieae (W. L. Wagner et al. 2007). They proposed a series of aneuploid reductions from n = 9 or 10 to n = 6, followed by polyploidy to produce the array of numbers in Epilobium (n = 12, 13, 15, 16, 18, 30) and Boisduvalia (n = 9, 10, 15, 19). D. A. Baum et al. (1994) demonstrated that molecular data did not support that hypothesis and found that a monophyletic Chamaenerion (n = 18, 36, 54) forms a strongly distinct sister branch to Epilobium, within which sect. Epilobium (n = 18) is monophyletic and sister to the rest of Epilobium, including the former segregates Boisduvalia and Zauschneria. The data from Baum et al. suggested that Epilobieae are primitively polyploid, with numbers based on x = 18, which is unique in the family. Using comparable sampling and some additional genes, R. A. Levin et al. (2004) found strong support for the phylogeny proposed by Baum et al.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Source FNA vol. 10. FNA vol. 10.
Parent taxa Onagraceae > subfam. Onagroideae > tribe Epilobieae > Epilobium > sect. Epilobium Onagraceae > subfam. Onagroideae
Sibling taxa
E. anagallidifolium, E. arcticum, E. brachycarpum, E. campestre, E. canum, E. ciliatum, E. clavatum, E. cleistogamum, E. coloratum, E. davuricum, E. densiflorum, E. densum, E. foliosum, E. glaberrimum, E. hallianum, E. hirsutum, E. hornemannii, E. howellii, E. leptocarpum, E. leptophyllum, E. luteum, E. minutum, E. mirabile, E. montanum, E. nevadense, E. nivium, E. obcordatum, E. obscurum, E. oreganum, E. oregonense, E. pallidum, E. palustre, E. parviflorum, E. rigidum, E. saximontanum, E. septentrionale, E. siskiyouense, E. smithii, E. suffruticosum, E. torreyi
Subordinate taxa
Synonyms E. alpinum var. lactiflorum, E. canadense, E. canadense var. albescens, E. hornemannii var. lactiflorum Epilobioideae alph., Boisduvaliinae raimann, Epilobiinae torrey
Name authority Haussknecht: Oesterr. Bot. Z. 29: 89. (1879) Endlicher: Fl. Poson., 366. (1830)
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