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weeping willow

willow family

Habit Shrubs or trees, heterophyllous or not, sometimes clonal, forming clones by root shoots, rhizomes, layering, or stem fragmentation; glabrous or glabrescent to pubescent; branching monopodial or sympodial.
Stems

branches yellow-brown to red-brown;

branchlets sparsely to moderately densely tomentose, especially at nodes.

erect to pendent; branched.

Leaves

stipules absent or rudimentary on early ones;

petiole convex to flat or shallowly to deeply grooved adaxially, 7–9 mm, tomentose abaxially;

largest medial blade lanceolate, narrowly oblong, or narrowly elliptic, 90–160 × 5–20 mm, 5.5–10.5 times as long as wide, base cuneate, margins flat, spinulose-serrulate or serrulate, apex acuminate, caudate, or acute, surfaces glabrous or sparsely short-silky, hairs straight, dull adaxially;

proximal blade margins entire;

juvenile blade reddish or yellowish green.

persistent, deciduous or marcescent, alternate (opposite or subopposite in Salix purpurea), spirally arranged, simple;

stipules present or not;

petiole present;

blade margins toothed or entire, sometimes glandular.

Inflorescences

racemose or spicate, usually catkins, unbranched, sometimes fasciculate or racemelike cymes, flowering before or as leaves emerge or year-round;

floral bract (1) subtending each flower, displaced onto pedicel or distinct, scalelike, apex entire, toothed, or laciniate;

bract subtending pistillate flower deciduous or persistent.

Peduncles

present or absent.

Pedicels

present or absent.

Flowers

usually unisexual, sometimes bisexual, usually staminate and pistillate on different plants;

sepals present or absent, or perianth modified into 1 or 2 nectaries, or a non-nectariferous disc;

stamens 1–60(–70);

filaments distinct or connate basally, slender;

anthers longitudinally dehiscent;

ovary 1, 2–7[–10]-carpellate, 1–7[–10]-locular;

placentation usually parietal, sometimes axile on intruded, fused placentae;

ovules 1–25 per ovary;

style 1 per carpel, distinct or connate;

stigmas 2–4, truncate, notched-capitate, or 2- or 3-lobed.

Staminate flowers

abaxial nectary 0.2–0.6 mm, adaxial nectary oblong or ovate, 0.4–0.7 mm, nectaries distinct or connate and cup-shaped;

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

anthers (sometimes reddish turning yellow), ellipsoid or globose.

Pistillate flowers

adaxial nectary oblong, square, ovate, or obovate, 0.4–0.8 mm;

ovary ovoid or obturbinate, beak (sometimes pilose proximally), slightly bulged below or abruptly tapering to styles;

ovules 2–4 per ovary;

styles distinct or connate 1/2 their lengths, 0.2–0.3 mm;

stigmas flat, abaxially non-papillate with rounded tip, or 2 plump lobes (almost capitate), 0.2–0.3 mm.

Fruits

capsular, baccate, or drupaceous.

Capsules

2–2.7 mm.

Seeds

sometimes surrounded by arillate coma of relatively long, silky hairs;

endosperm scant or absent.

Catkins

(flowering just before leaves emerge); staminate 13–35 mm, flowering branchlet 1–6 mm; pistillate densely flowered, stout or subglobose, 9–27 × 2.5–7 mm, flowering branchlet (0–)2–4 mm;

floral bract 1.1–1.8 mm, apex acute, rounded, or truncate, entire, abaxially sparsely hairy throughout or proximally, hairs wavy.

2n

= 76.

Salix babylonica

Salicaceae

Phenology Flowering spring.
Habitat Around settlements
Elevation ca. 50 m (ca. 200 ft)
Distribution
from FNA
AL; AR; CA; DC; DE; FL; GA; KY; LA; MD; NC; SC; TN; VA; Asia [Introduced in North America; introduced also in Mexico (Mexico City), South America]
[WildflowerSearch map]
[BONAP county map]
Nearly worldwide
[BONAP county map]
Discussion

Little is known about the origin of the strongly weeping cultivar of Salix babylonica. It was described by Linnaeus (1737[1738]) based on young garden specimens (W. J. Bean 1970–1988, vol. 4). It is thought to have originated in China, although it no longer occurs in the wild and its origin is uncertain. Selections are thought to have been transported to Europe along the trade route from China. In Tajikistan, there are three cultivated clones, one of which is staminate (A. K. Skvortsov 1999). Taxonomic treatments of S. babylonica are variable. Some botanists recognize a single species, including both pendulous and non-pendulous forms (Skvortsov), while others recognize four species: S. babylonica, with a weeping habit, S. capitata Y. L. Chou & Skvortsov, S. pseudolasiogyne H. Léveillé, and the commonly cultivated S. matsudana Koidzumi (Fang Z. F. et al. 1999), with an erect or spreading habit. Here, S. babylonica is treated in a narrow sense, including only weeping forms.

Salix babylonica is not cold tolerant and is not commonly grown in Europe (R. D. Meikle 1984) or in northern North America. In the flora area, cultivated trees with strongly pendulous branches and branchlets have been identified as S. babylonica (G. W. Argus 1985, 1986, 1993), but many are hybrids with S. alba (S. ×sepulcralis) or S. euxina (S. ×pendulina). Salix ×sepulcralis, especially nothovar. chrysocoma, with bright yellow branchlets, is the most commonly grown of these hybrids. All reported occurrences of S. babylonica need verification.

Hybrids:

Salix ×sepulcralis Simonkai: Weeping willow, S. alba × S. babylonica, is introduced from Europe and widely naturalized throughout the world. Synonyms include S. ×salamonii Carrière ex Henry and S. ×sepulcralis nothovar. chrysocoma (Dode) Meikle. It is characterized by: trees, to 12 m, stems pendulous; branches somewhat to highly brittle at base, yellowish, yellow-green, or yellow-brown; branchlets yellowish, yellow-green, or golden; stipules rudimentary or foliaceous on late leaves; petiole not glandular or with pairs or clusters of spherical glands distally or scattered throughout, short-silky adaxially; largest medial blade amphistomatous or hemiamphistomatous, narrowly elliptic to very narrowly so, margins finely serrulate or spinulose-serrulate, abaxial surface glaucous, adaxial glaucous, sparsely long-silky to glabrescent, hairs white or white and ferruginous, adaxial surface slightly glossy; catkins on distinct flowering branchlet 3–14 mm; staminate moderately densely flowered, slender, 23–53 × 3–9 mm; pistillate moderately densely to loosely flowered, slender to stout, 18–30 × 3–8 mm, flowering branchlet 3–14 mm; pistillate bracts persistent after flowering; staminate abaxial and adaxial nectaries distinct; stamens 2; anthers 0.5–0.8 mm; pistillate nectary longer than stipe; stipe 0–0.2 mm; ovaries gradually tapering to styles; ovules 4 per ovary; styles 0.15–2 mm; capsules 1–2 mm. In the flora area, it occurs in: British Columbia, New Brunswick, Nova Scotia, Ontario, Quebec; Alaska, Arizona, Arkansas, California, Connecticut, District of Columbia, Illinois, Iowa, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Missouri, Nevada, New Hampshire, New Jersey, New Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Tennessee, Utah, Virginia, and West Virginia.

The most commonly cultivated, and sometimes escaped, weeping willow with golden or yellow-green branchlets is Salix ×sepulcralis nothovar. chrysocoma (Dode) Meikle. It probably originated as S. alba var. vitellina × S. babylonica (R. D. Meikle 1984). According to F. S. Santamour Jr. and A. J. McArdle (1988), S. ×sepulcralis cv. Salamonii has a broadly pyramidal crown and is only slightly pendulous. It is not clear just how this cultivar differs from S. ×pendulina. For a discussion of the taxonomy of these and other weeping willows see J. Chmela (1983).

Salix ×pendulina Wenderoth: Weeping willow, S. babylonica × S. euxina, is introduced from Europe and grown throughout the world. It is characterized by: trees, 2.5–12 m, stems pendulous; branches highly brittle at base, yellow-brown, gray-brown, or red-brown; branchlets yellowish to brownish; stipules foliaceous on late leaves; petioles glabrous, pilose, or velvety to glabrescent adaxially; largest medial blade amphistomatous or hypostomatous, very narrowly elliptic to lanceolate, or linear, margins serrulate, irregularly so, or spinulose-serrulate, abaxial surface glaucous, adaxial slightly glossy or dull; catkins on distinct flowering branchlet, 3–14 mm; staminate loosely flowered, stout, 16–34 × 7–11 mm; pistillate densely or moderately densely flowered, slender or stout, 20–36 × 3.5–11 mm; pistillate bract persistent after flowering; staminate abaxial and adaxial nectaries connate and shallowly cup-shaped; stamens 2; anthers 0.5–0.6 mm; pistillate nectary longer than stipe; stipe 0 mm; styles 0.2–0.6 mm; ovules 4–8 per ovary; capsules 1.8–3.5 mm. In the flora area, it occurs in: Ontario; California, Connecticut, District of Columbia, Georgia, Illinois, Indiana, Maine, Maryland, Massachusetts, Michigan, Missouri, Nebraska, New Jersey, New Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Texas, Virginia, Washington, and West Virginia.

Reports of this hybrid in British Columbia and California are undocumented. Plants of Salix ×pendulina with prominent, caudate stipules are var. blanda (Andersson) Meikle; those with ovaries with patchy or streaky hairiness are var. elegantissima (K. Koch) Meikle.

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

Genera 50+, species ca. 1000 (4 genera, 123 species in the flora).

Taxonomic placement of the Salicaceae and the genera included in it have varied greatly. Some botanists (H. G. A. Engler and K. Prantl 1887–1915) treated it as a primitive member of the Dicotyledoneae and grouped it with other families having simple, apetalous, unisexual flowers arranged in catkins, the “Amentiferae.” At about the same time, others (C. E. Bessey 1915) took a different view, regarding the simple flowers as the result of reduction, and placed the taxa in Caryophyllales. As early as 1905, H. Hallier could see that there were similarities between Salicaceae and Flacourtiaceae; at the time, he was vigorously challenged by E. Gilg (1915). A. D. J. Meeuse (1975) summarized evidence for a close relationship between these families, including wood anatomy, phytochemistry, host-parasite relationships (including rust fungi), and morphology. He concluded that the Salicaceae could be combined with the Flacourtiaceae, “perhaps as a tribe.” A. Cronquist (1988) and R. F. Thorne (1992b) placed the Salicaceae, in a narrow sense, in Violales near Flacourtiaceae.

Molecular studies support a close relationship between Salicaceae and Flacourtiaceae in Malpighiales and show that Flacourtiaceae, in a broad sense, is paraphyletic. Based on a study of plastid rbcL DNA sequences, Salix and Populus were nested within a subset of 52 genera of Flacourtiaceae (M. W. Chase et al. 2002). Chase et al. proposed moving some genera from broadly circumscribed Flacourtiaceae to Salicaceae. Other studies, based on different gene sequences, came to the same conclusion (O. I. Nandi et al. 1998; V. Savolainen et al. 2000; K. W. Hilu et al. 2003; Angiosperm Phylogeny Group 2003). The discovery of the extinct fossil genus Pseudosalix (L. D. Boucher et al. 2003), from the Eocene Green River Formation of Utah, provided further support for placing some members of Flacourtiaceae in Salicaceae. The well-preserved Pseudosalix fossils, in which reproductive structures are directly associated with the leaves, occur intermixed with Populus fossils. The leaves are slender and have salicoid teeth, inflorescences are cymose, flowers are unisexual, pedicellate, tetrasepalous, and 3- or 4-carpellate, and seeds are comose, i.e., having characteristics intermediate between Salicaceae and Flacourtiaceae.

The presence, in both families, of salicoid teeth is often cited in support of their close relationship (W. S. Judd 1997b; O. Nandi et al. 1998; M. W. Chase et al. 2002; H. P. Wilkinson 2007). Salicoid teeth were first recognized and defined as having the tip of the medial vein (seta) of the tooth retained as a dark, but not opaque, non-deciduous spherical callosity fused to the tooth apex and were reported to occur in Salicaceae and Idesia of the Flacourtiaceae (L. J. Hickey and J. A. Wolfe 1975). Nandi et al. reported that a broad survey of angiosperm leaves showed that salicoid teeth occur outside of Flacourtiaceae and Salicaceae only in Tetracentraceae.

Isozyme and cytological evidence show that Populus and Salix are ancient polyploids (D. E. Soltis and P. S. Soltis 1990; Wang R. and Wang J. 1991). All Salix and Populus species contain salicin (R. T. Palo 1984).

The genera often included in Salicaceae, in the narrow sense, are Chosenia, Populus, Salix (A. K. Skvortsov 1999), and, sometimes, Toisusu. Molecular studies (E. Leskinen and C. Alström-Rapaport 1999; T. Azuma et al. 2000) show that Chosenia is nested within Salix. H. Ohashi (2001) treated Toisusu as Salix subg. Pleuradinea Kimura and Chosenia as Salix subg. Chosenia (Nakai) H. Ohashi.

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

Key
1. Flowers in catkins; sepals absent; fruits capsules
→ 2
1. Flowers not in catkins; sepals present; fruits drupes or berries
→ 3
2. Buds 3-10-scaled (usually resinous); leaf blades usually less than 2 times as long as wide, venation ± palmate (basal secondary veins strong, paired, except in Populus angustifolia); stipules caducous; catkins pendulous, sessile; floral bracts: apex deeply or shallowly cut, pistillate floral bracts deciduous after flowering; flowers without nectaries (with a non-glandular, cup- or saucer-like disc); stamens 6-60(-70); stigmas 2-4; capsules 2-4-valved, narrowly ovoid to spherical.
Populus
2. Buds 1-scaled (oily in Salix barrattiana); leaf blades often more than 2 times as long as wide, venation usually pinnate; stipules persistent or absent; catkins erect, spreading, or ± pendulous, sessile or terminating flowering branchlets; floral bracts: apex entire, erose, 2-fid, or irregularly toothed, pistillate floral bracts persistent or deciduous after flowering; flowers: perianth reduced to adaxial nectary (rarely also with abaxial nectary, then distinct or connate into shallow cup); stamens 1, 2, or 3-10; stigmas 2; capsules 2-valved, obclavate to ovoid or ellipsoid.
Salix
3. Flowers in racemelike cymes or solitary; fruits drupes, 18-25 mm
Flacourtia
3. Flowers in fascicles; fruits berries, 4-7 mm.
Xylosma
Source FNA vol. 7, p. 40. FNA vol. 7, p. 3. Authors: George W. Argus, James E. Eckenwalder, Robert W. Kiger.
Parent taxa Salicaceae > Salix > subg. Salix > sect. Subalbae
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. 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. 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
Subordinate taxa
Flacourtia, Populus, Salix, Xylosma
Name authority Linnaeus: Sp. Pl. 2: 1017. (1753) Mirbel
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