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coastal willow, dune willow, Hooker's willow

Arizona willow

Habit Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or stem fragmentation). Plants 0.3–2.6 m. Stems: branches red-brown or yellow-brown, not or weakly glaucous, glabrous or pilose at nodes; branchlets yellow-green, red-brown, or brownish, pilose.
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 foliaceous, apex convex or rounded;

petiole convex to flat, or shallowly grooved adaxially, 2–7.5 mm, villous or pubescent to glabrescent adaxially;

largest medial blade (sometimes amphistomatous), elliptic or broadly elliptic, 20–50 × 10–31 mm, 1.6–2–2.8(–3.6) times as long as wide, base convex, rounded, or cordate, margins flat, serrulate or entire, apex acute, convex, or acuminate, abaxial surface not glaucous, pilose or glabrous, hairs wavy, adaxial slightly glossy, pilose or glabrous;

proximal blade margins entire, serrulate, or crenulate;

juvenile blade green, glabrous or pilose 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 narrowly oblong to oblong, 0.4–0.8 mm;

filaments distinct, glabrous;

anthers purple turning yellow, 0.4–0.6 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 narrowly oblong to oblong, 0.4–1 mm, shorter to longer than stipe;

stipe 0.2–1 mm;

ovary pyriform, glabrous, beak gradually tapering to or slightly bulged below styles;

ovules 8–12 per ovary;

styles 0.5–1.2 mm;

stigmas broadly cylindrical, 0.14–0.21–0.36 mm.

Capsules

5–10 mm.

3.2–4.5 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 as leaves emerge; staminate stout, subglobose, or globose, 7–17 × 6–10 mm, flowering branchlet 1–3 mm; pistillate densely or moderately densely flowered, stout or subglobose, 12–38 × 6–12 mm, flowering branchlet 1.5–10 mm;

floral bract brown, black, or bicolor, 1–2 mm, apex acute or convex, abaxially hairy, hairs wavy.

2n

= 114.

= 38.

Salix hookeriana

Salix arizonica

Phenology Flowering mid Apr-mid Jun. Flowering late May-late 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 Subalpine sedge meadows, along streams, wet drainageways, cienegas
Elevation 0-1800 m (0-5900 ft) 2600-3400 m (8500-11200 ft)
Distribution
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
from FNA
AZ; CO; NM; UT
[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.)

Of conservation concern.

Salix arizonica is very similar to S. boothii. They were separated by Dorn on the presence of five flavonoid compounds identified in S. boothii not found in S. arizonica. Some morphological differences were noted but the characters used to separate them are quite variable. The most important feature seems to be the usually broader leaves of S. arizonica. In addition, the diploid chromosome number for S. arizonica separates it from the tetraploid S. boothii. Although the two are distinct species, the overlap in their morphological characters suggests that positive identification needs to be based on chromosome number or chromatographic analysis.

Salix arizonica is distinguished from S. boothii by having stipule apices convex to rounded, petioles 3–7.5 mm, juvenile blades glabrous or hairy, hairs white, largest medial blades elliptic or broadly elliptic, 20–50 mm, 1.6–3.6 times as long as wide, abaxial surfaces with white hairs, staminate catkins 1–1.7 times as long as broad, pistillate catkins 1.2–2.8 times as long broad, floral bract apices acute to convex, nectaries narrowly oblong to oblong, staminate nectaries 0.4–0.8 mm, anthers 0.4–0.6 mm, filaments distinct, glabrous, pistillate nectaries shorter to longer than stipes, stipes 0.2–1 mm, stigmas broadly cylindrical, and capsules 3.2–4.5 mm; S. boothii has stipule apices acuminate or acute to rounded, petioles 3–17 mm, juvenile blades hairy, hairs white, sometimes also ferruginous, largest medial blades lorate to narrowly or broadly elliptic, 26–102 mm, 2–5.2 times as long as wide, abaxial surfaces with white, sometimes also ferruginous, hairs, staminate catkins 1.2–3.1 times as long as broad, pistillate catkins 1.4–4.1 times as long as broad, floral bract apices rounded or retuse, nectaries narrowly oblong to ovate or flask-shaped, staminate nectaries 0.6–1.5 mm, anthers 0.48–0.8 mm, filaments distinct to connate about half their lengths, glabrous or hairy, pistillate nectaries shorter than stipes, stipes 0.5–2.5 mm, stigmas flat, abaxially non-papillate with rounded tip, slenderly cylindrical or plump, and capsules 2.5–6 mm.

Salix arizonica, originally known from Mt. Baldy in east-central Arizona, was proposed for listing under the United States Endangered and Threatened Wildlife and Plants Act, but the proposal was withdrawn when additional populations were discovered in southern Utah and in New Mexico; it is now listed as a “sensitive species” (K. Decker, www.fs.us/r2/projects/scp/assessments/salixarizonica.pdf). Major conservation concerns are from browsing by cattle and elk (J. Maschinski 2001) and Melampsora infection (M. L. Fairweather 1993; R. A. Obedzinski et al. 2001). The Arizona populations receive minimal protection from cattle and wildlife browsing by exclosures and by introduction into new localities. The Arizona and Utah populations have a genetic similarity of ca. 37%, which has been attributed to a period of panmixis followed by a long period of isolation in regions with different environments (J. T. Thompson et al. 2003).

Salix arizonica is in the Center for Plant Conservation’s National Collection of Endangered Plants.

Hybrids:

Salix arizonica forms natural hybrids with S. brachycarpa, S. eastwoodiae, and S. lutea.

Salix arizonica × S. brachycarpa var. brachycarpa occurs in southern Utah. Its parentage is supported by chromatographic data (E. D. McArthur, pers. comm.).

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

Source FNA vol. 7, p. 127. FNA vol. 7, p. 112.
Parent taxa Salicaceae > Salix > subg. Vetrix > sect. Cinerella Salicaceae > Salix > subg. Vetrix > sect. Hastatae
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. 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. 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) Dorn: Canad. J. Bot. 53: 1499. (1975)
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