Flora of North America species comparison

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.)

Variety monica applies to the diminutive alpine form that sometimes is recognized in the southern Rocky Mountains (S. J. Brunsfeld and F. D. Johnson 1985); it occurs at higher elevations (2200–4000 m) and is characterized by low growth form (0.14–1 m) and smaller, slightly broader leaves. Although it can be distinctive, it is morphologically confluent with the typical species. B. G. O. Floderus (1939) may be correct in characterizing it as an alpine ecotype.

Salix planifolia and S. pulchra are closely related. Their ranges overlap in northwestern Canada, from northern British Columbia across the southern quarter of the Yukon and northeastward into the Great Bear Lake area. Specimens identified as S. pulchra occur as far northeastward as Coppermine and northeast of Bathurst Inlet; S. planifolia has been recognized in the Mackenzie Delta and Eskimo Lake regions, Northwest Territories. Outlying records should be treated with caution because identification of individual specimens out of context may not be definitive. G. W. Argus (1969, 1973) treated these taxa as subspecies based on their intergradation in northwestern British Columbia, their tetraploid chromosome number, and their similar leaf flavonoid chromatographic patterns, but this taxonomy needs reconsideration.

The primary differences between the two species are stipule size, shape, and persistence and the pubescence on juvenile leaves. Stipules of Salix planifolia are oblong to narrowly elliptic or obovate, 0.8–3 mm (or –4.5 mm at Back River, Northwest Territories), distinctly shorter than petioles, and rarely marcescent for more than one year; stipules of S. pulchra are linear to narrowly oblong, 3–32 mm, usually longer than petioles, and usually marcescent for two or more years. Juvenile leaves of S. planifolia are usually more densely hairy, but vary from glabrescent to sparsely or very densely pubescent or long-silky, whereas juvenile leaves of S. pulchra are usually glabrous or, sometimes, sparsely hairy. The occurrence of rhombic mature leaf blades in S. pulchra sometimes is distinctive, but overlap in leaf shape between the two taxa is very great.

The area of geographic overlap in Yukon and western Northwest Territories is large, but evidence suggests that there the two species may be separated by elevation. In the vicinity of Whitehorse, Yukon, Salix pulchra occurs at higher elevations (1400–1900 m) than S. planifolia (ca. 1000 m); no mixed populations were seen. In Nahanni National Park, Northwest Territories, where S. planifolia is more common than S. pulchra, the latter occurs only in alpine and subalpine habitats (1200–1400 m). Evidence from both localities indicates an elevational separation of the two taxa. Within the area of overlap there is little evidence of intergradation except that S. planifolia has stipules that tend to be more marcescent (40% are marcescent) and sometimes longer (2–3.5 mm) than is usual outside the area of overlap. Nevertheless, specimens from the area of overlap can be easily assigned to one taxon or the other with only a few apparent intermediates. The problem in recognizing intermediacy is that there are only a few, variable characters that separate the two. In contrast, in 1973, G. W. Argus described evidence of hybridization and introgression along the Haines Road in northwestern British Columbia. This was based on variation in stipule size, presence, and persistence in what appeared to be a hybrid swarm. Further data are needed to answer questions about actual hybridization. Are the species separated by habitat or elevation, and are there reproductive barriers? Answers could be gained by population studies and controlled hybridization. Until that is done it is best to treat these taxa as species.

See 76. Salix discolor and 95a. S. alaxensis var. alaxensis for comparative descriptions.

Hybrids:

Salix planifolia forms natural hybrids with S. alaxensis var. alaxensis, S. argyrocarpa, S. brachycarpa var. brachycarpa, S. candida, S. drummondiana, S. humilis, S. pellita, S. pulchra, and S. scouleriana. Hybrids with S. glauca var. cordifolia have been reported (C. K. Schneider 1921) but no convincing specimens have been seen.

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