Epilobium siskiyouense
Epilobium ciliatum
Siskiyou fireweed, Siskiyou rock-fringe, Siskiyou willow-herb
American willowherb, ciliate willowherb, fringe willow-herb, northern purple-leaf willowherb, purple-leaf willowherb, slender willow herb, Watson's willowherb, épilobe cilié
several to many, erect to ascending, loosely clumped, terete, 10–25 cm, rarely branched distal to base, usually short-villous and strigillose throughout, mixed sparsely glandular puberulent distally, rarely subglabrous proximal to inflorescence.
erect, green or tan to reddish green, terete, (3–)10–120(–190) cm, often thick, well branched or simple, subglabrous proximal to inflorescence with raised strigillose lines decurrent from margins of petioles, ± densely mixed strigillose and glandular puberulent distally, rarely densely strigillose or densely villous throughout.
opposite proximal to inflorescence, alternate and usually crowded distally, sessile;
blade gray-green, narrowly to broadly ovate, 1.3–2.6 × 0.8–2 cm, base rounded to subcordate, margins usually serrulate, 6–12 teeth per side, rarely subentire, veins inconspicuous, 3–5 per side, apex rounded proximally to acute distally, surfaces sparsely short-villous to subglabrous and glaucous;
bracts much reduced.
opposite proximal to inflorescence, alternate distally, petiole 0–5(–10) mm, often subsessile distally, sometimes clasping;
blade narrowly obovate, obovate, broadly elliptic, or spatulate proximally, to very narrowly lanceolate to ovateor broadly elliptic distally, (1–)3–12(–16) × (0.2–)0.6–5.5 cm, base rounded to cuneate or short-attenuate, margins serrulate, (8–)15–40 irregular teeth per side, veins prominent, 4–10 per side, apex obtuse to acute or subacuminate, surfaces usually subglabrous with strigillose margins, rarely densely strigillose or villous;
bracts scarcely reduced to very reduced and narrower.
erect, compact racemes, densely canescent and glandular puberulent, or subglabrous, only ovaries pubescent.
usually erect, rarely nodding, racemes or panicles, well branched and open, to simple and congested, ± densely strigillose and glandular puberulent.
erect;
buds often purplish green, 9–11 × 3.5–5 mm, blunt;
pedicel 6–12 mm;
floral tube 2.1–4 × 2.9–5 mm, prominent ring of tissue 0.3–0.6 mm wide, edged by spreading hairs, 0.9–1.8 mm from base of tube inside;
sepals purplish green, 5–10.5 × 2–3.5 mm, apex acute;
petals pink to rose-purple, obcordate, 10–22.5 × 9.5–15.5 mm, apical notch 2–6.5 mm;
filaments cream, those of longer stamens 6.5–14 mm, those of shorter ones 3.5–11 mm;
anthers cream 1.9–3.3 × 0.7–1.2 mm;
ovary 12–22 mm, ± densely canescent and glandular puberulent;
style white to light pink, 10.5–18 mm, sparsely villous just proximal to stigma, stigma broadly 4-lobed, 1–1.8 × 2.4–4.2 mm, exserted beyond anthers.
erect;
buds 1.5–7 × 1–3 mm;
pedicel 2–14(–20) mm;
floral tube 0.5–2.6 × 0.9–3.5 mm, ring of spreading hairs at mouth inside;
sepals often reddish green, lanceolate, sometimes keeled, 2–7.5 × 0.7–2.5 mm;
petals white or pink to rose-purple, obovate, 2–14 × 1.3–6.3 mm, apical notch 0.4–2.5 mm;
filaments white to dark pink, those of longer stamens 1.4–7 mm, those of shorter ones 0.6–5.2 mm;
anthers light yellow to cream, 0.5–1.8 × 0.3–0.9 mm;
ovary often reddish green, 8–40 mm, ± densely mixed strigillose and glandular pubescent;
style cream to light yellow, 1.1–8.5 mm, stigma cream to orange-yellow, narrowly to broadly clavate or subcapitate, 0.8–2.8 × 0.4–1.2 mm, rarely indented apically, usually surrounded by, rarely exserted beyond, anthers.
25–45 mm, surfaces canescent and glandular puberulent;
pedicel 6–25 mm.
erect, (15–)30–100 mm, surfaces usually strigillose and glandular puberulent, rarely glabrescent;
pedicel 2–15(–40) mm, rarely subsessile.
narrowly obovoid, 1.4–1.9 × 0.6–0.7 mm, with low, obscure chalazal collar, light brown, surface papillose;
coma easily detached, somewhat tawny, 4–8 mm.
narrowly obovoid or subfusiform, (0.6–)0.8–1.6(–1.9) × 0.3–0.6 mm, chalazal collar ± conspicuous, 0.1–0.3 × 0.2–0.4 mm, grayish tan to brown, surface with conspicuous parallel longitudinal ridges of laterally flattened papillae;
coma readily detached, white or dingy white, 2–8 mm, very rarely absent.
= 36.
= 36.
Epilobium siskiyouense
Epilobium ciliatum
Epilobium siskiyouense is endemic to the Klamath region in southwestern Oregon (Jackson County) and north-central California in the Salmon, Scott Bar, and Siskiyou mountains of Siskiyou and Trinity counties. As noted by Hoch and Raven, this geographical range and several morphological features appear to be intermediate between those of E. obcordatum and E. rigidum. All three species have unusually large flowers (12–26 mm) with four-lobed stigmas, and as a group are quite distinct from their congeners in the region. Despite these similarities, the three taxa differ substantially in details of floral structure, especially regarding the dimensions of the floral tube. Specifically, E. rigidum has mean petal length 18.2 mm, floral tubes 1–1.6 × 2.5–3.6 mm; E. siskiyouense mean petal length 17.1 mm, floral tubes 2.1–4 × 2.9–5 mm; and E. obcordatum mean petal length 18.6 mm, floral tubes 3.2–5.2 × 2.2–3.6 mm. Thus, in flowers that are similar in overall size and aspect, E. rigidum has a very short, broad floral tube, E. obcordatum has a relatively long, narrow tube, and E. siskiyouense has a tube intermediate in size and shape. In terms of the ratio of tube length to width, the three taxa do not overlap. Although these characters are difficult to include in a key (since they require floral dissection and/or precise measurements), they are diagnostic for these species.
Epilobium siskiyouense has an additional diagnostic floral character that is unique in the genus. Whereas most other species of Epilobium have a simple ring of spreading hairs, sometimes with a low ridge of tissue near the mouth of the floral tube, E. siskiyouense has a relatively broad ring of tissue (0.3–0.6 mm wide), shaped like a washer, from which spreading hairs arise; this feature may provide protection for the nectar.
Epilobium siskiyouense has two distinct patterns of vestiture on the stems. In some specimens, the lower stems are mixed canescent and glandular puberulent and the inflorescence only glandular puberulent. In other specimens, the stems are subglabrous below a sparsely canescent and glandular puberulent inflorescence. There is no obvious correlation of this difference with any other morphological, ecological, or geographical factors.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
Subspecies 3 (3 in the flora).
Epilobium ciliatum, which has the AA chromosomal arrangement, shows extraordinary variation in morphology. It has the largest geographical range among North American Epilobium species, and has spread invasively outside of its native range. Although almost certainly originating in North America, E. ciliatum is also considered native in South America (J. C. Solomon 1982) and East Asia (Chen C. J. et al. 1992), but adventive in Europe and western Russia (P. H. Raven 1968), Pacific Islands, especially New Zealand, and Australia (Raven and T. E. Raven 1976). Its chromosomal affinities and morphological similarities to a small group of species in western North America strongly suggest that that region is its center of origin.
Within the enormous variation displayed by Epilobium ciliatum, three broadly defined entities can be recognized: subsp. watsonii, characteristically with bracts scarcely reduced on an extended, crowded corymbose inflorescence, found only along the Pacific coast, usually within sight of the ocean; subsp. glandulosum, generally large, few-branched plants with condensed turions just below ground and crowded inflorescences of relatively large rose-purple flowers, found mainly in damp, cool, and relatively undisturbed habitats; and subsp. ciliatum, which range from small and simple to large and well-branched, usually with leafy basal rosettes and open inflorescences, relatively narrow leaves and small white flowers, found most often in disturbed damp to dry habitats throughout the entire range of the species. These subspecies often intergrade in regions where their ranges overlap, resulting in populations with diverse mixtures of intermediate characters, yet the subspecies consistently retain their main morphological characteristics in populations throughout most of their respective ranges. Each shows some degree of endogenous variability, most notably in the very widespread subsp. ciliatum.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Leaf blades very narrowly lanceolate to narrowly ovate or elliptic, proximally narrowly obovate to spatulate; bracts very reduced on open inflorescence; petals 2–6(–9) mm, white or sometimes pink; herbs usually with rosettes, rarely fleshy turions. | subsp. ciliatum |
1. Leaf blades narrowly ovate to ovate to broadly elliptic, sometimes lanceolate, proximally obovate to broadly elliptic; bracts little reduced on crowded inflorescence; petals 4.5–12(–15) mm, usually rose-purple to pink, rarely white; herbs usually with fleshy turions or rosettes, rarely fleshy shoots. | → 2 |
2. Herbs usually with large, condensed subsessile turions 1–10 cm below ground, leaving dark scales, rarely with rosettes of fleshy leaves; inflorescences simple or branched, not corymbose. | subsp. glandulosum |
2. Herbs with leafy basal rosettes, sometimes fleshy shoots from woody caudex; inflorescences ± simple, subcorymbose. | subsp. watsonii |
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