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sandbar willow

coastal willow, dune willow, Hooker's willow

Habit Shrubs or trees, 4–9 m. Stems: branches gray-brown to red-brown, glabrous or villous; branchlets yellow brown to red-brown, densely tomentose or villous to glabrescent. Shrubs or trees, (0.6–)2–8 m, (sometimes forming clones by layering or 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).

Leaves

stipules absent or rudimentary on early ones, rudimentary or foliaceous on late ones;

petiole 1–5(–9) mm, glabrous or sparsely villous adaxially;

largest medial blade linear to lorate, 60–160 × 4–11 mm, (6.5–)11–19(–31) times as long as wide, base cuneate, margins flat, remotely spinulose-serrulate (teeth 2–5 per cm), apex acute or subacuminate, abaxial surface thinly glaucous, densely villous or long-silky to glabrescent, adaxial slightly glossy, pilose or densely villous to glabrescent;

proximal blade margins entire;

juvenile blade reddish or yellowish green, moderately densely to sparsely long-silky abaxially.

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.

Staminate flowers

abaxial nectary 0.5–1.1 mm, adaxial nectary ovate, narrowly oblong, or flask-shaped, 0.6–1.4 mm, nectaries distinct;

filaments hairy;

anthers 0.4–0.9 mm.

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.

Pistillate flowers

adaxial nectary narrowly oblong, 0.4–1.1 mm, shorter to longer than stipe;

stipe 0.4–0.8 mm;

ovary obclavate to pyriform, glabrous, glabrescent, or long-silky, beak abruptly tapering to styles;

ovules 16–36 per ovary;

styles 0–0.2 mm;

stigmas flat, abaxially non-papillate with pointed tip, or broadly cylindrical, 0.3–0.7 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.

Capsules

(4–)5–8(–10) mm.

5–10 mm.

Catkins

(flowering throughout season); staminate 20–61 × 4–10 mm, flowering branchlet 3–20 mm; pistillate loosely flowered, slender or stout, 20–67 × 5–9 mm, flowering branchlet 3–19 mm;

floral bract (sometimes greenish), 1.5–3.5 mm, apex acute, acuminate, or rounded, entire, erose, or toothed, abaxially hairy either proximally or distally, hairs wavy.

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.

2n

= 38.

= 114.

Salix interior

Salix hookeriana

Phenology Flowering early Apr-early Jul. Flowering mid Apr-mid Jun.
Habitat Sandy to silty flood plains, margins of lakes, ponds, and prairie sloughs, dry prairie sand hills, marshes, disturbed areas Marine coastal beaches and sand dunes, interdunal depressions, coastal marshes, pine barrens, floodplains, ravines, wet sedge meadows, lakeshores, morainal flats, sandy or gravelly substrates
Elevation 10-1800 m (0-5900 ft) 0-1800 m (0-5900 ft)
Distribution
from FNA
AK; AR; CO; CT; DC; DE; IA; IL; IN; KS; KY; LA; MD; ME; MI; MN; MO; MS; MT; ND; NE; NJ; NY; OH; OK; PA; SD; TN; TX; VA; WI; WV; WY; AB; BC; MB; NB; NT; ON; QC; SK; YT; Mexico (Tamaulipas, Veracruz)
[WildflowerSearch map]
[BONAP county map]
from FNA
AK; CA; OR; WA; BC
[WildflowerSearch map]
[BONAP county map]
Discussion

Sometimes Salix interior is treated as a subspecies of S. exigua (R. D. Dorn 1998). Salix exigua and S. interior hybridize and apparently intergrade in the western Great Plains; because the area of overlap is relatively small and distinctiveness of the two taxa is not compromised by hybridization and introgression, it is best to treat them as separate species.

Leaves on sylleptic shoots are usually very densely silky. Salix interior sometimes has shoots that arise from buds on either side of the normal axillary bud. They do not seem to be directly related to the stipules because they are enclosed by the petiole. Catkins with both staminate and pistillate flowers are rare in S. interior, but a Quebec specimen had some catkins predominantly pistillate and others staminate; most were a mixture. The flowers were not teratological, but a mature capsule contained aborted ovules.

Hybrids:

Salix interior forms natural hybrids with S. exigua var. exigua. Controlled pollinations using S. interior (as S. exigua) from southern Ontario (A. Mosseler 1990) successfully produced F1 hybrids with S. bebbiana, S. discolor, S. eriocephala, and S. petiolaris. Seed production was usually relatively low, except in crosses with S. discolor. In general, F1 viability was relatively low in crosses with these members of subg. Vetrix. No seeds were produced in crosses with members of subgenera Protitea or Salix. Morphology of the hybrids usually was intermediate between the two parents, but when S. petiolaris was used as the maternal parent, the F1s more closely resembled that species. J. Salick and E. Pfeffer (1999) extended these findings to show that, although crosses between S. interior (as S. exigua) and S. eriocephala are partially sterile, their clonal growth parameters (sprouting, shoot length, and biomass production) are strong and thus permit these partially sterile hybrids to exist as successful individuals and perhaps to “... make a contribution to interspecific gene flow over time.” Of particular taxonomic interest is that, in this cross, the staminate parent has a significant influence on leaf shape, whereas in the cross S. eriocephala × S. petiolaris it is the pistillate parent that is significant for leaf shape. Relatively few hybrids resembling those produced by Mosseler have been recognized in nature, but it is possible that the unusually broadly leaved plants named S. interior var. exterior and var. wheeleri, from northern Maine, Nebraska, New York, and West Virginia, and probably elsewhere, may be hybrids. Phenological isolation may be strong enough to prevent crosses in nature (A. Mosseler and C. S. Papadopol 1989) with the earlier flowering S. eriocephala and S. petiolaris, a barrier that even an occasional period of overlap cannot breach.

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

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.)

Source FNA vol. 7, p. 56. FNA vol. 7, p. 127.
Parent taxa Salicaceae > Salix > subg. Longifoliae Salicaceae > Salix > subg. Vetrix > sect. Cinerella
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. hookeriana, S. humboldtiana, S. humilis, 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. 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
Synonyms S. rubra, S. exigua var. exterior, S. exigua subsp. interior, S. exigua var. pedicellata, S. exigua var. sericans, S. fluviatilis var. sericans, S. interior var. exterior, S. interior var. pedicellata, S. interior var. wheeleri, S. linearifolia, S. longifolia var. interior, S. longifolia var. pedicellata, S. longifolia var. sericans, S. longifolia var. wheeleri, S. wheeleri S. amplifolia, S. hookeriana var. laurifolia, S. hookeriana var. tomentosa, S. piperi
Name authority Rowlee: Bull. Torrey Bot. Club 27: 253. (1900) Barratt ex Hooker: Fl. Bor.-Amer. 2: 145, plate 180. (1838)
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