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Pacific willow (var. lasiandra), shining willow, whiplash willow (ssp. caudata)

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

Habit Shrubs or trees, 4–6 m. Stems: branches flexible to highly brittle at base, yellow-brown, gray-brown, or red-brown, slightly to highly glossy, glabrous or villous to glabrescent; branchlets yellow-brown, gray-brown, or red-brown, glabrous, pilose, densely villous, or velvety, hairs spreading, straight, wavy, or crinkled. Shrubs, 0.2–6 m, (sometimes forming clones by stem fragmentation).
Stems

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 foliaceous, apex convex to rounded;

petiole shallowly to deeply grooved adaxially, 5–13 mm, with clusters of spherical or foliaceous glands distally, glabrous, pilose, or densely villous adaxially;

largest medial blade usually hypostomatous or hemiamphistomatous, rarely amphistomatous, lorate, very narrowly elliptic, narrowly elliptic, or lanceolate, (24–)55–133 × 11–43 mm, 2.5–6.2 times as long as wide, base convex or cuneate, margins flat, serrulate, apex acuminate to caudate, abaxial surface usually not glaucous (rarely so), glabrous, pilose, or moderately densely villous or long-silky, hairs appressed or spreading, white and/or ferruginous, straight or wavy, (coarse, caducous), adaxial (secondary veins flat or protruding), slightly or highly glossy, glabrous, pilose, or long-silky, hairs white and/or ferruginous;

proximal blade margins entire and glandular-dotted, or serrulate or crenulate;

juvenile blade reddish or yellowish green, glabrous or densely villous or long-silky abaxially, hairs white and ferruginous.

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

abaxial nectary 0.5–1.1 mm, adaxial nectary square or ovate, 0.3–0.9 mm, nectaries connate and cup-shaped;

stamens 3–6;

filaments distinct, hairy on proximal 1/2 or basally;

anthers ellipsoid, shortly cylindrical, obovoid, or globose, 0.6–0.8 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 square or ovate, (swollen), 0.2–0.5 mm, shorter than stipe;

stipe 0.5–2 mm;

ovary pyriform, beak slightly bulged below or gradually tapering to styles;

ovules 18–24 per ovary;

styles connate or distinct 1/2 their lengths, 0.5–0.8 mm;

stigmas flat, abaxially non-papillate with rounded tip, broadly cylindrical, or 2 plump lobes, 0.24–0.31–0.42 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

5–7 mm.

3.5–7 mm.

Catkins

staminate 19–69 × 4–14 mm, flowering branchlet 5–23 mm; pistillate (fruiting in summer), moderately densely to loosely flowered, slender to stout, 23–56(–70 in fruit) × 8–12 mm, flowering branchlet 8–25 mm;

floral bract 1.5–3 mm, apex convex or rounded, entire or toothed, abaxially sparsely hairy throughout or proximally, hairs 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

= 76.

= 38.

Salix lucida

Salix eriocephala

Phenology Flowering early May-mid Jul. Flowering early Apr-mid Jun.
Habitat Sandy or gravelly floodplains, lake margins, sedge meadows, vernal pools, alvars, open fens, marl bogs, treed bogs Gravelly or rocky stream banks, marshy fields, in mixed mesophytic woods on alluvium
Elevation 0-600 m (0-2000 ft) 0-1200 m (0-3900 ft)
Distribution
from FNA
CT; DE; IA; IL; IN; MA; MD; ME; MI; MN; ND; NH; NJ; NY; OH; PA; RI; SD; VA; VT; WI; WV; MB; NB; NL; NS; ON; PE; QC; SK; SPM
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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
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[BONAP county map]
Discussion

The Virginia plants of Salix lucida are introduced (G. W. Argus 1986).

The Salix lucida complex is a group of three weakly delimited taxa, S. lasiandra var. caudata, S. lasiandra var. lasiandra, and S. lucida. The morphological characters used to separate them (leaves amphistomatous or hypostomatous and blades glaucous abaxially or not) are usually geographically correlated, but there are exceptions. G. W. Argus (1986b) proposed, based on principal components analysis of morphological data, to treat them as a single species consisting of three subspecies. The geographic overlap of the northeastern S. lucida and the western S. lasiandra is a relatively small area in central Saskatchewan. Evidence of intergradation was based on cultivation of a plant that, in the wild, had leaves that were not glaucous abaxially but were glaucous in cultivation. It seems best to treat them as two species, S. lucida and S. lasiandra, the latter with two varieties, var. lasiandra and var. caudata.

Hybrids:

Salix lucida forms natural hybrids with S. alba and S. nigra. Hybrids with S. serissima have been reported (M. L. Fernald 1950) but no convincing specimens have been seen. Attempts to hybridize S. lucida with members of subg. Protitea (S. amygdaloides), subg. Longifoliae (S. interior), and subg. Vetrix (S. discolor, S. eriocephala, and S. petiolaris) were unsuccessful (A. Mosseler 1990).

Salix lucida × S. nigra (S. ×schneideri B. Boivin) seems to be a rare intersubgeneric hybrid between tetraploid S. lucida and diploid S. nigra. It is known only from the type specimen, an infertile plant, growing with both parents. It resembles S. lucida in bud-scale margins connate, in petiolar glands stalked or foliaceous, and in leaf shape, and S. nigra in stipules rudimentary on proximal leaves and sometimes even on early leaves, stipule apex acute, pistillate catkins relatively long and slender, and styles relatively short.

(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. 46. FNA vol. 7, p. 120.
Parent taxa Salicaceae > Salix > subg. Salix > sect. Salicaster 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. hookeriana, S. humboldtiana, S. humilis, S. interior, S. irrorata, S. jejuna, S. jepsonii, S. laevigata, S. lasiandra, S. lasiolepis, S. lemmonii, S. ligulifolia, 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 Pleiarina lucida, S. lucida var. angustifolia, S. lucida var. intonsa S. angustata, S. cordata, S. cordata var. abrasa, S. missouriensis, S. rigida, S. rigida var. angustata, S. rigida var. vestita
Name authority Muhlenberg: Ges. Naturf. Freunde Berlin Neue Schriften 4: 239, plate 6, fig. 7. (1803) Michaux: Fl. Bor.-Amer. 2: 225. (1803)
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