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coastal willow, dune willow, Hooker's willow

heart-leaf willow, Missouri or diamond or heart-leaf willow, Missouri River willow, Missouri willow

Habit Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or stem fragmentation). Shrubs, 0.2–6 m, (sometimes forming clones by stem fragmentation).
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).

branches (sometimes highly brittle at base), red-brown, not glaucous, glabrous or glabrescent;

branchlets yellow-brown to red-brown, pilose, moderately to densely velvety, pubescent, or villous, (inner membranaceous bud-scale layer free).

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, (4.5–13 mm), apex rounded or acute;

petiole shallowly grooved adaxially, 3–18 mm, tomentose adaxially;

largest medial blade narrowly oblong, very narrowly elliptic or obovate, 58–96–136 × 9–21–36 mm, 2.3–4.6–8 times as long as wide, base cordate, convex, rounded, subcordate, or, sometimes, cuneate, margins flat, serrate or serrulate, apex acute to acuminate, abaxial surface thickly glaucous, glabrous, puberulent, sparsely pubescent or short-silky, adaxial highly glossy, glabrous or sparsely villous (hairs white, sometimes also ferruginous);

proximal blade margins entire or shallowly serrulate;

juvenile blade reddish or yellowish green, glabrous, pilose, or villous 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, oblong, or ovate, 0.2–1 mm;

filaments distinct or connate less than 1/2 their lengths, glabrous;

anthers yellow or purple turning yellow (ellipsoid or shortly cylindrical), 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 oblong or flask-shaped, 0.3–0.8 mm, shorter than stipe;

stipe 1.2–2.8 mm;

ovary pyriform, glabrous, beak slightly bulged below styles;

ovules 12–16 per ovary;

styles 0.3–0.6 mm;

stigmas flat, abaxially non-papillate with rounded tip, or broadly cylindrical, or 2 plump lobes, 0.16–0.28 mm.

Capsules

5–10 mm.

3.5–7 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 flowering just before leaves emerge, pistillate as leaves emerge; staminate slender or stout, 19–44 × 7–14 mm, flowering branchlet 0.5–5 mm; pistillate densely or moderately densely flowered, slender or stout, 22–65 × 7–14 mm, flowering branchlet 2–10 mm;

floral bract dark brown or bicolor, 0.8–1.6 mm, apex rounded, abaxially hairy, hairs wavy.

2n

= 114.

= 38.

Salix hookeriana

Salix eriocephala

Phenology Flowering mid Apr-mid Jun. Flowering early 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 Gravelly or rocky stream banks, marshy fields, in mixed mesophytic woods on alluvium
Elevation 0-1800 m (0-5900 ft) 0-1200 m (0-3900 ft)
Distribution
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
from FNA
AL; AR; CT; DC; DE; FL; GA; IA; IL; IN; KS; KY; MA; MD; ME; MI; MN; MO; ND; NE; NH; NJ; NY; OH; PA; RI; SD; TN; VA; VT; WI; WV; NB; NF; NS; ON; PE; QC
[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.)

Salix eriocephala sometimes is very difficult to separate from S. myricoides. Some of the confusion may be due to hybridization but no definite hybrids have been seen.

Salix eriocephala can be distinguished from S. myricoides by having stipules on early leaves foliaceous, apices acute to rounded, largest medial blades 4.9–23.3 times as long as petiole, abaxial surface usually thickly glaucous (stomata can be seen through the grayish wax), margins serrulate or serrate, floral bracts 0.8–1.6 mm, moderately to very densely hairy, styles 0.3–0.6 mm, and stigmas 0.16–0.28 mm; S. myricoides has stipules on early leaves rudimentary or foliaceous, apices acuminate or acute, largest medial blades 4.7–13.4 times as long as petiole, abaxial surface usually with very thick wax (stomata cannot be seen through the wax), margins crenulate to serrulate, floral bracts 1.2–3 mm, sparsely to moderately densely hairy, styles 0.3–1.3 mm, and stigmas 0.28–0.56 mm.

Hybrids:

Salix eriocephala forms natural hybrids with S. candida, S. famelica, S. humilis, S. interior, S. lasiandra, S. petiolaris, and S. sericea. Hybrids with S. amygdaloides, S. bebbiana, S. myricoides, and S. pedicellaris have been reported (M. L. Fernald 1950) but no convincing specimens have been seen. Controlled pollinations made with S. discolor had low success and many seedlings were abnormal (A. Mosseler 1990). In controlled pollination using S. eriocephala as the maternal parent, seeds were rarely produced due to pollen-stigma incompatibility (Mosseler 1989).

Salix eriocephala × S. famelica: Hybrids and intergrades occur in the area of overlap (R. D. Dorn 1995). Specimens from a population in Douglas County, Nebraska, which included successive collections and cultivated specimens, have branches with yellow-mottled coloration of S. famelica and villous indumentum of S. eriocephala; they may be this hybrid.

Salix eriocephala × S. petiolaris: Controlled pollinations (A. Mosseler 1990) had low seed-set but a high percent of seed germination and seedling survival. Because reproductive barriers between these species are weak, it was suggested that their morphological variability may be due to interspecific gene flow (Mosseler). Natural hybrids are known from Illinois, Maine, Massachusetts, Michigan, Missouri, New York, Ontario, Quebec, and West Virginia.

Salix eriocephala × S. sericea: This hybrid is relatively common wherever the ranges of the parents overlap. It has been studied in the southeastern United States (G. W. Argus 1986) and in eastern Canada. The results of a molecular study (T. M. Hardig et al. 2000) have been discussed already under the genus. In general, the hybrids resemble S. eriocephala but have leaves that are sparsely to moderately densely short-silky on abaxial surfaces and ovaries hairy as in S. sericea. Foliaceous stipules are often present on late leaves and sometimes even on early leaves, as in S. eriocephala, but they are not as prominent. In S. sericea stipules usually are lacking or rudimentary, but on late leaves they may be foliaceous. Petioles and branchlets of hybrids are finely velvety as in S. sericea. This hybrid was described from Maine (O. W. Knight 1907), where it was noted that the catkins were usually abortive but sometimes produced one or two fertile seeds.

Salix eriocephala is distinguished from S. sericea in having stipules on early and late leaves foliaceous, 4–6.2–8.3 × 2.5–3.6–4.6 mm, 1.5–2 times as long as wide, ovaries glabrous, juvenile blades glabrous or sparsely hairy, hairs white, largest medial blade abaxial surfaces glabrous, puberulent, sparsely pubescent, or short-silky, stipes 1.2–2.8 mm, and capsules 3.5–7 mm; S. sericea has stipules on early leaves absent or rudimentary, on late leaves rudimentary to foliaceous, 1.1–1.6–2.1 × 0.4–0.6–0.8 mm, 2.3–3 times as long as wide, ovaries short-silky, juvenile blades very densely short-silky, hairs white, sometimes also ferruginous, largest medial blade abaxial surfaces densely short-silky, stipes 0.6–1.5 mm, and capsules 2.5–4 mm.

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

Source FNA vol. 7, p. 127. FNA vol. 7, p. 120.
Parent taxa Salicaceae > Salix > subg. Vetrix > sect. Cinerella Salicaceae > Salix > subg. Vetrix > sect. Cordatae
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. 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 S. angustata, S. cordata, S. cordata var. abrasa, S. missouriensis, S. rigida, S. rigida var. angustata, S. rigida var. vestita
Name authority Barratt ex Hooker: Fl. Bor.-Amer. 2: 145, plate 180. (1838) Michaux: Fl. Bor.-Amer. 2: 225. (1803)
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