Salix hookeriana
Salix candida
coastal willow, dune willow, Hooker's willow
hoary willow, sage willow, sage-leaf willow
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 dark gray-brown to yellow-brown, not glaucous, woolly in patches or floccose to glabrescent;
branchlets yellow-brown to red-brown or gray-brown, densely (white) woolly or tomentose, sometimes floccose.
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, late ones 2–3.6 mm, apex acute;
petiole shallowly to deeply grooved adaxially, 3–10 mm, tomentose or densely (white) woolly adaxially (obscured by hairs);
largest medial blade lorate, narrowly elliptic or oblanceolate, 47–103 × 5–20 mm, base convex or cuneate, margins strongly to slightly revolute, entire, or sinuate, apex acute or convex, abaxial surface glaucous (generally obscured by hairs), very densely to sparsely tomentose-woolly (cobwebby in age), hairs dull white, crinkled, adaxial dull or slightly glossy, moderately densely to sparsely tomentose, floccose, hairs dull white;
proximal blade margins entire;
juvenile blade yellowish green, very densely tomentose abaxially, hairs white.
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, 0.6–1 mm;
filaments distinct or connate less than 1/2 their lengths;
anthers purple turning yellow, ellipsoid, long-cylindrical, or globose, 0.4–0.6 mm.
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, 0.4–1 mm;
ovary pyriform, beak sometimes slightly bulged below styles;
ovules 12–18 per ovary;
styles 0.3–1.9 mm.
5–10 mm.
4–6 mm.
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 or subglobose, 17–39 × 8–16 mm, flowering branchlet 0.5–7 mm; pistillate densely to moderately densely flowered, stout or slender, 20–66 × 9–18 mm, flowering branchlet 1–24 mm;
floral bract tawny or brown, 1.2–1.8 mm, apex rounded or acute, abaxially hairy, hairs straight to wavy.
= 114.
= 38.
Salix hookeriana
Salix candida
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.)
Occurrence of Salix candida in Nunavut is on Akimiski Island in James Bay.
Salix candida is geographically wide-ranging but limited to calcareous habitats and, for that reason, it is quite local or even rare in some parts of its range.
Hybrids:
Salix candida forms natural hybrids with S. bebbiana, S. brachycarpa var. brachycarpa, S. calcicola, S. eriocephala, S. famelica, S. myrtillifolia, S. petiolaris, and S. planifolia. Hybrids with S. discolor, S. petiolaris, and S. sericea have been reported (the latter also by C. K. Schneider 1921; M. L. Fernald 1950) but no convincing specimens have been seen. Salix candida hybrids are recognized from their woolly indumentum that often is conspicuous on leaves, stems, and ovaries. In hybrids, these characters, especially woolly patches on ovaries, stand out as discordant variation.
Salix candida × S. eriocephala (S. ×rubella Bebb ex C. K. Schneider) was described by W. W. Rowlee and K. M. Wiegand (1896) as S. candida × S. cordata. In addition to woolly patches on the ovaries, they noted that buds of the hybrids usually are shorter, more divergent, and blunter than those in S. eriocephala, and are glabrous or hairy. Known from New York and Newfoundland; it should be expected throughout the sympatric range of the parental species.
Salix candida × S. famelica: The Saskatchewan specimen resembles S. famelica but has the leaf indumentum of S. candida.
Salix candida × S. myrtillifolia: Saskatchewan specimens combine characters of the two parents.
Salix candida × S. petiolaris: Intermediates between these species are known from Michigan and New York (W. W. Rowlee and K. M. Wiegand 1896) as well as Ontario and Saskatchewan, but can be expected wherever the two grow together. The invalid name “S. ×clarkei” is sometimes used for this hybrid.
The glabrescent form of Salix candida, forma denudata (Andersson) Rouleau, may be of hybrid origin.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
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