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coastal willow, dune willow, Hooker's willow

Barclay's willow

Habit Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or stem fragmentation). Plants (0.3–)1–3(–5) m. Stems: branches usually red-brown, sometimes yellow-brown, not to strongly glaucous, glabrous or villous; branchlets yellow-green, yellow-brown, or red-brown, densely villous to pubescent, (buds caprea-type, inner membranaceous bud-scale layer free, separating from outer layer).
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, (2–14 mm), apex rounded or acuminate;

petiole shallowly grooved, or convex to flat adaxially, 3–14(–20) mm, villous or pilose adaxially;

largest medial blade oblong, narrowly elliptic, elliptic, oblanceolate, or obovate, 33–70(–100) × 12–35(–48) mm, 1.6–2.8(–4) times as long as wide, base usually rounded or convex, sometimes subcordate, margins slightly revolute or flat, serrulate, apex acute, acuminate, or convex, abaxial surface glaucous, glabrous or glabrescent, hairs straight, adaxial slightly glossy or dull, glabrous or pilose, midrib pilose;

proximal blade margins serrulate or entire;

juvenile blade sometimes reddish, densely villous or glabrous 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, 0.5–1 mm;

filaments distinct, glabrous;

anthers yellow, 0.6–1 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 ovate, 0.4–0.8 mm, shorter than or equal to stipe;

stipe 0.4–1.5 mm;

ovary obclavate or pyriform, glabrous, beak gradually tapering to styles;

ovules 18–24 per ovary;

styles 0.6–2.5 mm;

stigmas slenderly cylindrical, 0.28–0.48–0.72 mm.

Capsules

5–10 mm.

3–8 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, 13–60 × 10–25 mm, flowering branchlet 0–17 mm; pistillate moderately densely flowered, stout, subglobose, or slender, 26–80 × 9–18 mm, flowering branchlet 4–24 mm;

floral bract brown or black, 1.6–2.8 mm, apex acute or rounded, abaxially hairy, hairs straight, wavy, or curly.

2n

= 114.

= 76 (based on putative Salix barclayi × S. barrattiana, see below).

Salix hookeriana

Salix barclayi

Phenology Flowering mid Apr-mid Jun. Flowering late May-early Aug.
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 Lake and streamshores, fens, moist to mesic forest openings, subalpine and alpine slopes, glacial moraines
Elevation 0-1800 m (0-5900 ft) 0-2800 m (0-9200 ft)
Distribution
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
from FNA
AK; ID; MT; OR; WA; WY; AB; BC; NT; YT
[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.)

Vegetative specimens of Salix hastata often are misidentified as S. barclayi. The characters that indicate S. hastata, or sometimes possible hybrids, are the presence of ferruginous hairs on juvenile leaves or adaxial midribs of mature leaves, leaf margins often entire or with teeth present only distally, and buds with inner membranaceous layer not separating from outer layer.

Salix barclayi is distinguished from S. farriae and S. hastata by having bud-scales with inner membranaceous layer separating from outer layer, juvenile blades glabrous, pilose, or moderately densely villous, largest medial blades oblong, narrowly elliptic, elliptic, oblanceolate, or obovate, margins always toothed, pistillate flowering branchlets 4–24 mm, staminate flowering branchlets 0–17 mm, floral bracts brown to black, moderately densely hairy, anthers 0.6–1 mm, styles 0.6–2.5 mm, and stipes 0.4–1.5 mm; S. farriae has bud-scales with inner membranaceous layer separating from outer layer, juvenile blades glabrous or sparsely villous, largest medial blades narrowly elliptic to elliptic, margins usually entire, pistillate flowering branchlets 1.5–14 mm, staminate flowering branchlets 1–5 mm, floral bracts bicolor, brown, or black, sparsely hairy, anthers 0.3–0.6 mm, styles 0.3–1.2 mm, and stipes 0.5–1.2 mm; and S. hastata has bud-scales with inner membranaceous layer not separating from outer layer, juvenile blades sparsely pubescent, largest medial blades narrowly to broadly elliptic, narrowly ovate, or ovate, margins usually entire, pistillate flowering branchlets 1.5–9 mm, staminate flowering branchlets 1–7 mm, floral bracts brown or bicolor, sparsely hairy, anthers 0.4–0.6 mm, styles 0.2–0.48 mm, and stipes 0.4–1.2 mm.

Hybrids:

Salix barclayi is morphologically highly variable. While it is possible that much of this variability is inherent, hybridization and introgression have played an important role.

Salix barclayi forms natural hybrids with S. arctica, S. barrattiana, S. brachycarpa, S. cascadensis, S. commutata, S. farriae, S. hastata, S. hookeriana, S. richardsonii, and S. stolonifera.

Salix barclayi × S. barrattiana is characterized by oily buds that stain the pressing paper yellow, and leaves often entire, or nearly so, and closely gland-dotted, and not glaucous abaxially. Some hybrids resemble S. barclayi in having ovaries glabrous and leaves serrulate and slightly glaucous abaxially, while others resemble S. barrattiana in having stipules marcescent and leaves short-hairy on both surfaces and not glaucous abaxially, but it is the presence of the oily buds characteristic of S. barrattiana that indicates their hybridity. The hybrids usually were from populations that included both parents. An important ramification of the discovery of this hybrid is that the specimen on which the chromosome number for S. barclayi (R. D. Dorn 1976) is based is a plant with oily buds, which has leaves with a mixture of weakly developed teeth and closely gland-dotted, entire margins, and is putatively identified as this hybrid. Further chromosome counts for S. barclayi are needed. Hybrids are known in British Columbia from the mountains in the Peace and Liard river basins, and in Mt. Robson National Park. The hybrid should be suspected wherever the parents occur together.

Salix barclayi × S. cascadensis, collected in Washington in alpine mats growing with both parents, has ovaries with hairy beaks indicating hybridization.

Salix barclayi × S. commutata resembles S. barclayi in having leaves glaucous abaxially, and S. commutata in having leaves hairy on both surfaces (northern British Columbia, Unalaska Island, and southeastern Alaska).

Salix barclayi × S. farriae is represented by a series of specimens resembling S. barclayi but with ferruginous hairs on juvenile leaves or on adaxial midribs of mature leaves, found from the Rocky Mountains of Alberta to southern Yukon and adjacent Northwest Territories. Similar specimens from Alaska may be the hybrid S. barclayi × S. hastata. Both of these hybrids require further study. See 63. S. hastata and 64. S. farriae.

Salix barclayi × S. hookeriana hybrids or introgressants at Tanis Lake, Alaska, have subentire leaves and stipes glabrous or hairy.

Salix barclayi × S. richardsonii hybrids and intergrades are relatively common in Alaska, British Columbia, and Yukon. They resemble S. barclayi in having catkins borne on distinct flowering branchlets, small stipules, anthers yellow and longer, and S. richardsonii in having marcescent stipules, branchlets and branches coarsely villous with spreading hairs, pistillate nectaries longer than stipes, and catkins sometimes on relatively short flowering branchlets.

Salix barclayi × S. scouleriana has the general appearance of S. barclayi but the densely hairy ovaries and the relatively long stigmas (0.9–0.92 mm) of S. scouleriana. It was growing with S. barclayi in openings created in white spruce by bark beetles.

Salix barclayi × S. stolonifera is characterized by relatively small leaves with irregularly serrulate margins, some young leaves reddish green, and a prostrate habit (G. W. Argus 1973).

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

Source FNA vol. 7, p. 127. FNA vol. 7, p. 113.
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. arizonica, S. athabascensis, S. atrocinerea, S. aurita, S. babylonica, S. ballii, 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) Andersson: Öfvers. Kongl. Vetensk.-Akad. Förh. 15: 125. (1858)
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