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coastal willow, dune willow, Hooker's willow

gray willow, large gray or gray willow, large gray willow

Habit Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or stem fragmentation). Shrubs, 3–7 m. Stems: branches brownish, not glaucous, pilose, villous, or tomentose to glabrescent, (peeled wood with striae to 62 mm); branchlets yellow-brown, pilose, velvety, or densely villous.
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).

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 rudimentary or foliaceous on early ones, foliaceous on late ones, apex acute or rounded;

petiole convex to flat adaxially, 4–15 mm, tomentose adaxially;

largest medial blade elliptic, broadly elliptic, oblanceolate, or obovate, 65–105 × 22–52 mm, 2–3 times as long as wide, base convex or cuneate, margins slightly revolute, entire, crenate, or sinuate, (glands submarginal), apex acuminate or convex, abaxial surface glaucous, tomentose, hairs erect or spreading, curly, adaxial dull or slightly glossy, pubescent or tomentose;

proximal blade margins entire;

juvenile blade yellowish green, sparsely to densely tomentose abaxially, hairs white.

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.

adaxial nectary oblong or ovate, 0.5–1 mm;

filaments distinct, glabrous or hairy basally;

anthers yellow or purple turning yellow, ellipsoid or shortly cylindrical, 0.7–1 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.

adaxial nectary oblong or square, 0.4–1 mm, shorter than stipe;

stipe 1.2–2.7 mm;

ovary pyriform, long-silky, beak slightly bulged below styles;

ovules 12 per ovary;

styles 0.2–0.5 mm;

stigmas slenderly or broadly cylindrical, 0.3–0.6 mm.

Capsules

5–10 mm.

5–5.6 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.

flowering before leaves emerge; staminate stout or subglobose, 26–39 × 12–26 mm, flowering branchlet 0–5 mm; pistillate densely flowered, stout or subglobose, 27–54(–75 in fruit) × 4–15 mm, flowering branchlet 1–5(–10) mm;

floral bract dark brown, black, or bicolor, 2–3 mm, apex acute or convex, abaxially hairy, hairs straight.

2n

= 114.

= 76.

Salix hookeriana

Salix cinerea

Phenology Flowering mid Apr-mid Jun. Flowering mid Mar-late May.
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 Stream shores, mesic woodlands, gravelly or sandy beaches, waste ground
Elevation 0-1800 m (0-5900 ft) 0-700 m (0-2300 ft)
Distribution
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
from FNA
AL; CT; DC; GA; IA; KY; LA; MA; MD; MI; MO; NC; NJ; NY; OH; PA; RI; SC; SD; TN; UT; VA; WI; WV; ON; Eurasia [Introduced in North America]
[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.)

The Ohio occurrence of Salix cinerea is based on information from T. Cooperrider (pers. comm.).

Salix cinerea and S. atrocinerea are very closely related. Their occurrence in the flora area, as naturalized introductions, is not well understood, probably because they usually are introduced under the name S. caprea, and that species often is not treated in North American floristic literature (e.g., C. K. Schneider 1921; M. L. Fernald 1950). They probably are introductions of long-standing brought to the New World for their value as ornamentals and bee-plants. Salix atrocinerea was first documented in the southeastern United States (G. W. Argus 1986) after plants with ferruginous hairs and prominently striate wood were found in North Carolina; since that time, it has been found in other states and provinces. In the northeastern states, S. atrocinerea and S. cinerea are thought to be invasive species. The species do reproduce by seed and hundreds of seedlings were observed in a drained reservoir (A. Zinovjev, pers. comm.) and on sandy pond shores (T. Rawinski, pers. comm.), where they are thought to compete with native species.

The presence of long, prominent, striae on the peeled wood of 4–5 year old branches is commonly used in European literature (K. H. Rechinger 1993; A. K. Skvortsov 1999) to separate Salix cinerea and S. atrocinerea from S. caprea etc., in which the wood is smooth or with fewer, shorter striae. In the flora area, long striae also occur in S. bebbiana, S. discolor, and S. humilis, but usually they are not as long as or as prominent in S. cinerea and S. atrocinerea. Some floras (e.g., F. Martini and P. Paiero 1988) use the relative prominence of striae to separate S. cinerea and S. atrocinerea, but their separation remains difficult. The presence of ferruginous hairs on the leaves of S. atrocinerea is the best diagnostic characteristic, but they are not always present or easily observed. For a comparison of these species, see the key to species under subg. Vetrix. For further discussion of morphologies, see Salix ×smithiana Willdenow [p. 132] and 76. S. discolor.

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

Source FNA vol. 7, p. 127. FNA vol. 7, p. 132.
Parent taxa Salicaceae > Salix > subg. Vetrix > sect. Cinerella Salicaceae > Salix > subg. Vetrix > sect. Cinerella
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. 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. 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
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: 1021. (1753)
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