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coastal willow, dune willow, Hooker's willow

glaucous willow, gray willow, gray-leaf willow, grey-leaf willow

Habit Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or stem fragmentation). Plants 0.2–6 m, not clonal.
Stems

branches (highly brittle at base), yellow-brown, gray-brown, red-brown, or violet, not or weakly glaucous, glabrous, tomentose, woolly, or sparsely villous to glabrescent (nodes hairy);

branchlets gray-brown, red-brown, or yellow-brown (sometimes color obscured by hairs), glabrous, pilose, moderately densely villous, tomentose, or woolly, scale with inner membranaceous layer free, (not separating from outer layer).

erect or decumbent;

branches brownish, yellow-brown, gray-brown, or red-brown, villous or pilose to glabrescent;

branchlets yellow-brown or red-brown, sparsely to densely villous or tomentose to glabrescent.

Leaves

stipules rudimentary or absent on early ones, foliaceous (early deciduous) or rudimentary (sometimes obscured) on late ones, (2.5–7.8–18 mm), apex acuminate, acute, or rounded;

petiole convex to flat, or shallowly grooved adaxially, 4–29 mm, villous, woolly, pilose, or tomentose adaxially;

largest medial blade (sometimes hemiamphistomatous), narrowly to broadly elliptic, oblanceolate, or obovate to broadly obovate, 36–123 × 18–63 mm, 1.5–4.2 times as long as wide, base convex, rounded, subcordate, cordate, or cuneate, margins slightly revolute, crenate, serrate, shallowly serrulate, sinuate, or entire, apex acuminate, acute, or convex, abaxial surface glaucous, pilose, moderately densely tomentose, villous, or woolly, midrib hairy, hairs (white, sometimes also ferruginous), wavy or straight, adaxial highly or slightly glossy, glabrous, pilose, villous, or moderately densely tomentose, midrib and veins hairy (hairs white, sometimes also ferruginous);

proximal blade margins entire or shallowly serrulate;

juvenile blade yellowish, reddish green (sometimes obscured by hairs), pilose or sparsely to densely long-silky, tomentose, woolly, or villous abaxially, hairs white, sometimes also ferruginous, or yellowish.

stipules (marcescent or not), foliaceous or rudimentary on early and late ones;

petiole (usually deeply to shallowly grooved adaxially), 1–27 mm, (much longer than subtended bud);

largest medial blade usually hypostomatous, sometimes hemiamphistomatous or amphistomatous, usually narrowly elliptic, elliptic, usually oblanceolate or obovate, sometimes narrowly oblong or obovate, 27–82 × 6–39 mm, 1.4–4.8 times as long as wide, base usually cuneate or convex, sometimes rounded, rarely subcordate, margins slightly revolute or flat, usually entire, apex acute, acuminate, convex, or rounded, abaxial surface densely villous or villous-silky, tomentose, short- or long-silky, or pilose, hairs usually wavy or straight, sometimes curved, adaxial usually slightly glossy, sometimes dull, moderately densely villous, pilose, or long-silky to glabrescent;

proximal blade margins entire or serrulate;

juvenile blade sparsely or densely villous, tomentose, or long-silky abaxially.

Staminate flowers

adaxial nectary oblong, ovate, or narrowly oblong, 0.5–1.4 mm;

filaments distinct or slightly basally connate, glabrous or hairy on proximal 1/2 or basally;

anthers yellow, cylindrical or ellipsoid, (0.5–)0.7–1 mm.

abaxial nectary 0.1–1 mm, adaxial nectary narrowly oblong, oblong, square, ovate, or flask-shaped, 0.5–1.3 mm, nectaries distinct, or connate and cup-shaped;

filaments distinct or slightly or partly connate, glabrous, or hairy on proximal 1/2;

anthers 0.4–0.8 mm.

Pistillate flowers

adaxial nectary narrowly oblong, oblong, or square, 0.5–1.4 mm, shorter than stipe;

stipe 0.5–1.8(–2.8) mm;

ovary obclavate or pyriform, glabrous, tomentose, villous, or woolly (hairs wavy), beak sometimes abruptly tapering to styles;

ovules 12–20 per ovary;

styles 0.6–2.3 mm;

stigmas broadly to slenderly cylindrical, 0.3–0.8 mm.

abaxial nectary absent, adaxial nectary usually narrowly oblong, oblong, or ovate, sometimes flask-shaped, 0.4–1.8 mm, shorter to longer than stipe;

stipe 0.3–2.8 mm;

ovary pyriform or obclavate, densely villous, tomentose, short-silky, or pubescent, beak usually gradually tapering to styles, sometimes gradually tapering to or slightly bulged below styles;

ovules 6–22 per ovary;

styles connate to distinct 1/2 their lengths or more, 0.3–1.6 mm;

stigmas flat, abaxially non-papillate with rounded tip, or slenderly or broadly cylindrical, 0.2–0.8 mm.

Capsules

5–10 mm.

4.5–9 mm.

Catkins

flowering before or as leaves emerge; staminate slender or stout, 26–73 × 10–27 mm, flowering branchlet 0–10 mm; pistillate densely flowered, slender or stout, 36–92(–140 in fruit) × 10–25 mm, flowering branchlet 0–20 mm;

floral bract brown, black, or bicolor, 1.1–3.6 mm, apex convex, rounded, or acute, abaxially hairy, hairs straight or wavy.

staminate 14–53 × 5–17 mm, flowering branchlet 1–25 mm; pistillate densely to sometimes loosely flowered, slender, stout, subglobose, or globose, 15–83 × 7–21 mm, flowering branchlet 2–37 mm;

floral bract tawny, brown, bicolor, or greenish, 1–3.4 mm, apex convex or rounded, entire, abaxially hairy, hairs wavy, crinkled, or straight.

2n

= 114.

= 76, 95, 114, 152.

Salix hookeriana

Salix glauca

Phenology Flowering mid Apr-mid Jun.
Habitat Marine coastal beaches and sand dunes, interdunal depressions, coastal marshes, pine barrens, floodplains, ravines, wet sedge meadows, lakeshores, morainal flats, sandy or gravelly substrates
Elevation 0-1800 m (0-5900 ft)
Distribution
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
from FNA
AK; CO; MT; NM; OR; UT; WA; WY; AB; BC; MB; NS; NT; NU; ON; QC; SK; YT; Eurasia (China [Altay Shan], Chukotka, Mongolia, Novaya Zemlya, Russian Far East, Sakhalin, Scandinavia, arctic, e, w Siberia)
[WildflowerSearch map]
[BONAP county map]
Discussion

Salix hookeriana is primarily a coastal species occurring from northern California northward to Oregon, Washington, and southern Vancouver Island, with disjunct populations on Queen Charlotte Islands, British Columbia, and northward to Yakatut Bay, Turnagain Arm, and Kodiak, Alaska. It was treated by G. W. Argus (1973) and R. D. Dorn (2000) in a broad sense because of an absence of strong distinguishing characters and intergradation in characters that could be used to divide it. It is highly variable and three very similar taxa have been named: S. amplifolia, S. hookeriana (including vars. tomentosa and laurifolia), and S. piperi. Although extremes of these taxa sometimes are recognizable, the intergradation displayed is so great that even attempts to recognize them as varieties are thwarted. The amplifolia variant in Alaska is characterized by having only white leaf hairs, hairy ovaries, no stipules, and catkins often borne on distinct flowering branchlets, but variation can occur within the same population, and typical S. hookeriana on Vancouver Island sometimes displays the same characteristics. The piperi variant, an inland population in western Oregon and Washington, is usually recognized by local botanists as different from coastal populations. It is characterized by leaves and branchlets soon becoming glabrate and stipules prominent. These characteristics, however, sometimes appear in northern California coastal populations, and some inland populations in Oregon include very hairy individuals that are indistinguishable from coastal variants of S. hookeriana. In general, very hairy populations of S. hookeriana are probably an adaptation to marine coastal environments, but some variation may be due to hybridization and introgression with S. scouleriana. Inland populations suggest the influence of S. lasiolepis. Two hexaploid chromosome numbers reported for S. hookeriana from Vancouver Island (R. L. Taylor and S. Taylor 1977) and Queen Charlotte Islands (R. L. Taylor and G. A. Mulligan 1968), British Columbia, indicate that hybridization has played a role in the evolution of this complex. It is possible that each variant of S. hookeriana has had a different, possibly even recurrent, polyploid origin. Further cytological and genetic study is indicated.

The following comparisons may help to distinguish Salix hookeriana, S. lasiolepis, and S. scouleriana.

Vegetative specimens of Salix hookeriana can be distinguished from S. lasiolepis by having floral buds ellipsoid, beaks distinctly long-tapered, densely long-hairy (villous), red-brown, blades usually pilose, villous, or woolly on abaxial surfaces, usually 18–63 mm wide, and 1.5–4.2 times as long as wide; S. lasiolepis has floral buds ovoid, beaks inconspicuous and blunt, sparsely to moderately densely short-hairy (velvety), yellowish to red-brown, blades usually tomentose on abaxial surfaces, usually 6–32 mm wide, and 3.2–9.6 times as long as wide.

Salix hookeriana is distinguished from S. scouleriana by having branchlets with spreading hairs (woolly or tomentose to glabrate), petioles usually pilose to tomentose, blades typically narrowly elliptic but variable, stigmas 0.3–0.74, short in relation to styles (0.6–2.3 mm), and pistillate nectaries 0.5–1.4 mm, shorter or longer than stipes; S. scouleriana has branchlets usually with short, erect hairs (velutinous), sometimes spreading (villous or tomentose), petioles velvety or villous adaxially, blades typically oblanceolate but variable, stigmas 0.4–1.04 mm, long in relation to styles (0.2–0.6 mm), and pistillate nectaries 0.2–0.8 mm, shorter than stipes.

Hybrids:

Salix hookeriana forms natural hybrids with S. barclayi and S. scouleriana. Variation in some S. hookeriana populations suggests hybridization with S. lasiolepis but no positive identifications have been made. R. D. Dorn (2000) doubted that hybridization in California between these species with different chromosome numbers was possible, but species with different chromosome numbers do hybridize [for example, S. athabascensis (4x) × S. pedicellaris (2x)]; synthetic hybridization studies are indicated.

Salix hookeriana × S. scouleriana: Plants from southern British Columbia with leaves similar to S. hookeriana but with prominent stipules, catkins both erect and recurving, and relatively long stigmas were thought by J. K. Henry (1915) to be this hybrid.

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

Varieties 5 (4 in the flora).

Occurrence of Salix glauca is disjunct in western Siberia; var. glauca is known from Scandinavia.

Salix glauca is ubiquitous, highly polymorphic, and polyploid. It appears to have four major variations that are treated in different ways. European floras have treated it as an undivided polymorphic species(A. K. Skvortsov 1999), as several distinct species (K. H. Rechinger 1964b), or as comprising several subspecies (Rechinger 1993; G. W. Argus et al. 1999; B. Jonsell and T. Karlsson 2000+, vol. 1). In North America, Argus (1965) treated the four central tendencies as widely intergrading “phases” of a single species, later adopting varietal rank, whereas E. Hultén (1968) recognized them as subspecies. In view of the fact that the four elements are confluent over wide areas, there may be some merit in recognizing them as informal phases, but varietal rank is used here.

All specimens from Iceland named Salix glauca are S. arctica and those from Svalbard, Norway, are S. lanata Linnaeus.

The major reason for the high variability within Salix glauca seems to be high, and probably recurrent, polyploidy. Tetraploids, pentaploids, hexaploids, and octoploids are known in the species, and two of the subspecies include three ploidal levels. There also are intergrading geographical variations that are the basis for the recognition of infraspecific taxa.

Hybrids:

Salix glauca forms natural hybrids with S. arctica, S. arctophila, S. ballii, S. barclayi, S. boothii, S. brachycarpa, S. eastwoodiae, S. myricoides, S. niphoclada, S. pedicellaris, and S. planifolia.

The following key will help identify the varieties, but there are extensive areas of overlap among them and many intermediates.

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

Key
1. Stipules often marcescent, foliaceous on late leaves, usually prominent, linear, narrowly elliptic, or lanceolate, 2-17 mm; branches red-brown or gray-brown or brownish; branchlets densely villous to glabrescent; proximal blades: margins entire; floral bracts 1.2-2.5 mm; staminate flowering branchlets 1-14 mm; ovaries obclavate or pyriform; Alaska, Canada
→ 2
1. Stipules sometimes marcescent or rudimentary, if foliaceous, usually inconspicuous, oblong, elliptic, or ovate, 1-8(-10) mm; branches usually yellow-brown or red-brown; branchlets usually sparsely villous or tomentose; proximal blades: margins entire or serrulate; floral bracts 1-3.4 mm; staminate flowering branchlets 1.5-25 mm; ovaries pyriform; northern Canada east of Mackenzie River, Greenland, Rocky Mountains
→ 3
2. Shrubs, 0.3-1 m; branchlets densely villous; petioles 1-9 mm; largest medial blades: apex acute, sometimes acuminate, or convex, adaxial surface moderately densely villous, or long-silky to glabrescent; staminate catkins 14-26 mm; filaments distinct or partly connate, glabrous or hairy on proximal 1/2; pistillate catkins stout to subglobose, flowering branchlets 2-19 mm; stipes 0.4-1.8 mm; Alaska, Northwest Territories, w Nunavut, n Yukon.
var. stipulata
2. Shrubs, 0.3-6 m; branchlets soon becoming pilose or glabrescent; petioles 4-27 mm; largest medial blades: apex acuminate to convex, adaxial surface often sparsely long-silky or pilose; staminate catkins 19-45 mm; filaments distinct, glabrous; pistillate catkins slender to stout, flowering branchlets 3-37 mm; stipes 0.5-2.8 mm; c Alaska, British Columbia, Northwest Territories, Yukon.
var. acutifolia
3. Stipules 0.9-2.8-8(-10) mm, sometimes marcescent; floral bracts usually brownish; largest medial blades: petiole 3-14 mm; Alberta, British Columbia, Rocky Mountains, Saskatchewan to New Mexico.
var. villosa
3. Stipules 1-2.1-4 mm, usually deciduous (in autumn); floral bracts usually tawny; largest medial blades: petiole 2-9 mm; c, e Canada, Greenland, St. Pierre and Miquelon.
var. cordifolia
Source FNA vol. 7, p. 127. FNA vol. 7, p. 89.
Parent taxa Salicaceae > Salix > subg. Vetrix > sect. Cinerella Salicaceae > Salix > subg. Chamaetia > sect. Glaucae
Sibling taxa
S. alaxensis, S. alba, S. amygdaloides, S. arbusculoides, S. arctica, S. arctophila, S. argyrocarpa, S. arizonica, S. athabascensis, S. atrocinerea, S. aurita, S. babylonica, S. ballii, S. barclayi, S. barrattiana, S. bebbiana, S. bonplandiana, S. boothii, S. brachycarpa, S. breweri, S. calcicola, S. candida, S. caprea, S. caroliniana, S. cascadensis, S. chamissonis, S. chlorolepis, S. cinerea, S. columbiana, S. commutata, S. cordata, S. daphnoides, S. delnortensis, S. discolor, S. drummondiana, S. eastwoodiae, S. elaeagnos, S. eriocephala, S. euxina, S. exigua, S. famelica, S. farriae, S. floridana, S. fuscescens, S. geyeriana, S. glauca, S. gooddingii, S. hastata, S. herbacea, S. humboldtiana, S. humilis, S. interior, S. irrorata, S. jejuna, S. jepsonii, S. laevigata, S. lasiandra, S. lasiolepis, S. lemmonii, S. ligulifolia, S. lucida, S. lutea, S. maccalliana, S. melanopsis, S. monochroma, S. monticola, S. myricoides, S. myrsinifolia, S. myrtillifolia, S. nigra, S. niphoclada, S. nivalis, S. nummularia, S. orestera, S. ovalifolia, S. pedicellaris, S. pellita, S. pentandra, S. petiolaris, S. petrophila, S. phlebophylla, S. planifolia, S. polaris, S. prolixa, S. pseudomonticola, S. pseudomyrsinites, S. pulchra, S. purpurea, S. pyrifolia, S. raupii, S. reticulata, S. richardsonii, S. rotundifolia, S. scouleriana, S. sericea, S. serissima, S. sessilifolia, S. setchelliana, S. silicicola, S. sitchensis, S. sphenophylla, S. stolonifera, S. taxifolia, S. thurberi, S. tracyi, S. triandra, S. turnorii, S. tweedyi, S. tyrrellii, S. uva-ursi, S. vestita, S. viminalis, S. wolfii, S. ×fragilis, S. ×jesupii, S. ×pendulina, S. ×sepulcralis, S. ×smithiana
S. alaxensis, S. alba, S. amygdaloides, S. arbusculoides, S. arctica, S. arctophila, S. argyrocarpa, S. arizonica, S. athabascensis, S. atrocinerea, S. aurita, S. babylonica, S. ballii, S. barclayi, S. barrattiana, S. bebbiana, S. bonplandiana, S. boothii, S. brachycarpa, S. breweri, S. calcicola, S. candida, S. caprea, S. caroliniana, S. cascadensis, S. chamissonis, S. chlorolepis, S. cinerea, S. columbiana, S. commutata, S. cordata, S. daphnoides, S. delnortensis, S. discolor, S. drummondiana, S. eastwoodiae, S. elaeagnos, S. eriocephala, S. euxina, S. exigua, S. famelica, S. farriae, S. floridana, S. fuscescens, S. geyeriana, S. gooddingii, S. hastata, S. herbacea, S. hookeriana, S. humboldtiana, S. humilis, S. interior, S. irrorata, S. jejuna, S. jepsonii, S. laevigata, S. lasiandra, S. lasiolepis, S. lemmonii, S. ligulifolia, S. lucida, S. lutea, S. maccalliana, S. melanopsis, S. monochroma, S. monticola, S. myricoides, S. myrsinifolia, S. myrtillifolia, S. nigra, S. niphoclada, S. nivalis, S. nummularia, S. orestera, S. ovalifolia, S. pedicellaris, S. pellita, S. pentandra, S. petiolaris, S. petrophila, S. phlebophylla, S. planifolia, S. polaris, S. prolixa, S. pseudomonticola, S. pseudomyrsinites, S. pulchra, S. purpurea, S. pyrifolia, S. raupii, S. reticulata, S. richardsonii, S. rotundifolia, S. scouleriana, S. sericea, S. serissima, S. sessilifolia, S. setchelliana, S. silicicola, S. sitchensis, S. sphenophylla, S. stolonifera, S. taxifolia, S. thurberi, S. tracyi, S. triandra, S. turnorii, S. tweedyi, S. tyrrellii, S. uva-ursi, S. vestita, S. viminalis, S. wolfii, S. ×fragilis, S. ×jesupii, S. ×pendulina, S. ×sepulcralis, S. ×smithiana
Subordinate taxa
S. glauca var. acutifolia, S. glauca var. cordifolia, S. glauca var. stipulata, S. glauca var. villosa
Synonyms S. amplifolia, S. hookeriana var. laurifolia, S. hookeriana var. tomentosa, S. piperi
Name authority Barratt ex Hooker: Fl. Bor.-Amer. 2: 145, plate 180. (1838) Linnaeus: Sp. Pl. 2: 1019. (1753)
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