FNA: Epilobium hallianum vs. Onagraceae tribe Epilobieae

	Epilobium hallianum	Onagraceae tribe Epilobieae
	glandular willowherb, Hall's willowherb, Sierra willowherb
Habit	Herbs with small, 3–6 mm, round or oblong, compact turions 1–5 cm below ground.	Herbs, annual or perennial, sometimes suffrutescent.
Stems	strict, erect, terete, 2–50(–60) cm, rarely branched only in larger plants, subglabrous proximal to inflorescence except for raised strigillose lines decurrent from margins of petioles, or sometimes ± densely long-villous throughout with inconspicuous decurrent lines.
Leaves	opposite proximally, alternate on inflorescence, subsessile or proximally with petioles 1–1.5 mm; blade ovate proximally to lanceolate or narrowly elliptic distally, 0.5–4.7 × 0.2–1.4 cm, base rounded to cuneate, margins subentire proximally to denticulate distally, 8–20 teeth per side, veins inconspicuous, 3–6 per side, apex obtuse to subacute, surfaces mostly glabrous with strigillose margins; bracts much reduced.	opposite at least near base or throughout, alternate distally, or sometimes alternate throughout; stipules absent.
Inflorescences	usually nodding in bud, erect later, open racemes, sometimes congested, usually mixed strigillose and glandular puberulent, rarely also mixed villous, or rarely subglabrous.
Flowers	erect; buds 2–5 × 1–2 mm; pedicel 3–8 mm; floral tube 0.5–1.7 × 0.8–1.6 mm, with slightly raised ring of spreading hairs at mouth inside; sepals green, 1.2–2.8 × 0.5–1 mm, abaxial surface subglabrous or sparsely glandular puberulent; petals white, often fading pink, 1.6–5.5 × 1.2–3 mm, apical notch 0.3–1.2 mm; filaments white or cream, those of longer stamens 0.6–2.5 mm, those of shorter ones 0.4–1.5 mm; anthers cream, 0.2–0.9 × 0.2–0.5 mm; ovary 10–14 mm, strigillose and glandular puberulent or subglabrous; style cream, 0.8–5 mm, stigma clavate, entire, 0.4–1.2 × 0.3–0.7 mm, usually surrounded by, rarely exserted beyond, 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	very narrowly cylindrical, (15–)24–60 mm, surfaces usually subglabrous to mixed strigillose and glandular puberulent, rarely sparsely villous; pedicel 8–40 mm.
Seeds	narrowly fusiform to narrowly obovoid, 1.1–1.6 × 0.4–0.6 mm, chalazal collar ± conspicuous, 0.05–0.2 mm, light brown, surface papillose; coma easily detached, white, 3–6 mm.	(1 or) many per locule, with tuft of hairs (coma) at chalazal end, sometimes without coma.
2n	= 36.

	Epilobium hallianum	Onagraceae tribe Epilobieae
Phenology	Flowering Jun–Sep.
Habitat	Semi-shaded stream banks, wet grassy slopes or meadows, bogs, seasonally wet sites, vernal pools.
Elevation	100–3700 m. (300–12100 ft.)
Distribution	AZ; CA; CO; ID; MT; NM; NV; OR; SD; UT; WA; WY; AB; BC; SK [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 hallianum has condensed fleshy turions and generally strict habit, suggesting an affinity with E. ciliatum or E. saximontanum, both with the AA arrangement. However, E. hallianum has the BB arrangement (S. R. Seavey and P. H. Raven 1978) and apparently more distant relationship to those other species. In different parts of its geographical range Epilobium hallianum shows considerable morphological variability, especially in leaf shape and margins, and in type and pattern of stem vestiture, including plants in the southern part of its range with densely villous stems that have been treated as E. ursinum. In part because of this variability and in part because the very characteristic condensed turions are easily lost during collection and/or often overlooked, E. hallianum is frequently misidentified. (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. hirsutum, E. hornemannii, E. howellii, E. lactiflorum, 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. brevistylum var. pringleanum, E. brevistylum var. subfalcatum, E. brevistylum var. tenue, E. brevistylum var. ursinum, E. delicatum var. tenue, E. glandulosum var. tenue, E. pringleanum, E. pringleanum var. tenue, E. ursinum, E. ursinum var. subfalcatum	Epilobioideae alph., Boisduvaliinae raimann, Epilobiinae torrey
Name authority	Haussknecht: Monogr. Epilobium, 261. (1884) — (as halleanum)	Endlicher: Fl. Poson., 366. (1830)
Web links	Local floras: BC, CA, OR, WA Local Web sites: CalFlora, CalPhotos, Flora NW, PNW Herbaria WildflowerSearch iNaturalist (observations) USDA Plants Database LBJ Wildflower Center SEINet Plants of the World Online Encyclopedia of Life Wikipedia Google Image Search	Google search

Onagraceae tribe Epilobieae

glandular willowherb, Hall's willowherb, Sierra willowherb

Habit

Herbs with small, 3–6 mm, round or oblong, compact turions 1–5 cm below ground.

Herbs, annual or perennial, sometimes suffrutescent.

Stems

strict, erect, terete, 2–50(–60) cm, rarely branched only in larger plants, subglabrous proximal to inflorescence except for raised strigillose lines decurrent from margins of petioles, or sometimes ± densely long-villous throughout with inconspicuous decurrent lines.

Leaves

opposite proximally, alternate on inflorescence, subsessile or proximally with petioles 1–1.5 mm;

blade ovate proximally to lanceolate or narrowly elliptic distally, 0.5–4.7 × 0.2–1.4 cm, base rounded to cuneate, margins subentire proximally to denticulate distally, 8–20 teeth per side, veins inconspicuous, 3–6 per side, apex obtuse to subacute, surfaces mostly glabrous with strigillose margins;

bracts much reduced.

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

stipules absent.

Inflorescences

usually nodding in bud, erect later, open racemes, sometimes congested, usually mixed strigillose and glandular puberulent, rarely also mixed villous, or rarely subglabrous.

Flowers

erect;

buds 2–5 × 1–2 mm;

pedicel 3–8 mm;

floral tube 0.5–1.7 × 0.8–1.6 mm, with slightly raised ring of spreading hairs at mouth inside;

sepals green, 1.2–2.8 × 0.5–1 mm, abaxial surface subglabrous or sparsely glandular puberulent;

petals white, often fading pink, 1.6–5.5 × 1.2–3 mm, apical notch 0.3–1.2 mm;

filaments white or cream, those of longer stamens 0.6–2.5 mm, those of shorter ones 0.4–1.5 mm;

anthers cream, 0.2–0.9 × 0.2–0.5 mm;

ovary 10–14 mm, strigillose and glandular puberulent or subglabrous;

style cream, 0.8–5 mm, stigma clavate, entire, 0.4–1.2 × 0.3–0.7 mm, usually surrounded by, rarely exserted beyond, 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

very narrowly cylindrical, (15–)24–60 mm, surfaces usually subglabrous to mixed strigillose and glandular puberulent, rarely sparsely villous;

pedicel 8–40 mm.

Seeds

narrowly fusiform to narrowly obovoid, 1.1–1.6 × 0.4–0.6 mm, chalazal collar ± conspicuous, 0.05–0.2 mm, light brown, surface papillose;

coma easily detached, white, 3–6 mm.

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

= 36.

Epilobium hallianum

Onagraceae tribe Epilobieae

Phenology

Flowering Jun–Sep.

Habitat

Semi-shaded stream banks, wet grassy slopes or meadows, bogs, seasonally wet sites, vernal pools.

Elevation

100–3700 m. (300–12100 ft.)

Distribution

AZ; CA; CO; ID; MT; NM; NV; OR; SD; UT; WA; WY; AB; BC; SK

[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 hallianum has condensed fleshy turions and generally strict habit, suggesting an affinity with E. ciliatum or E. saximontanum, both with the AA arrangement. However, E. hallianum has the BB arrangement (S. R. Seavey and P. H. Raven 1978) and apparently more distant relationship to those other species.

In different parts of its geographical range Epilobium hallianum shows considerable morphological variability, especially in leaf shape and margins, and in type and pattern of stem vestiture, including plants in the southern part of its range with densely villous stems that have been treated as E. ursinum. In part because of this variability and in part because the very characteristic condensed turions are easily lost during collection and/or often overlooked, E. hallianum is frequently misidentified.

(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.

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. hirsutum, E. hornemannii, E. howellii, E. lactiflorum, 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. brevistylum var. pringleanum, E. brevistylum var. subfalcatum, E. brevistylum var. tenue, E. brevistylum var. ursinum, E. delicatum var. tenue, E. glandulosum var. tenue, E. pringleanum, E. pringleanum var. tenue, E. ursinum, E. ursinum var. subfalcatum

Epilobioideae alph., Boisduvaliinae raimann, Epilobiinae torrey

Name authority

Haussknecht: Monogr. Epilobium, 261. (1884) — (as halleanum)

Endlicher: Fl. Poson., 366. (1830)

Web links

Local floras: BC, CA, OR, WA
Local Web sites: CalFlora, CalPhotos, Flora NW, PNW Herbaria
WildflowerSearch
iNaturalist (observations)
USDA Plants Database
LBJ Wildflower Center
SEINet
Plants of the World Online
Encyclopedia of Life
Wikipedia
Google Image Search

Google search

Unless otherwise noted, all data comes from the online version of the Flora of North America, who have made most of their content available via a Creative Commons license (CC BY 4.0).

Choose a data source: Flora of North America, OregonFlora, UW Burke Herbarium

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Epilobium hallianum

Onagraceae tribe Epilobieae

Epilobium hallianum

Onagraceae tribe Epilobieae